Glossary links done & added to Big Glossary. IUCN table dSalviaone. pratensis

Meadow clary, Clari’r maes LAMIACEAE SYN.: none

Status: Schedule 8 of the Wildlife and Status in Europe: Not threatened Countryside Act 1981 Red Data Book: Lower risk - Nationally 38 (20 native or possibly native) / 233 10-km Scarce squares post 1987

Contents

Status:...... 1 1 Morphology, Identification, Taxonomy & Genetics...... 2 1.1 Morphology & Identification ...... 2 1.2 Taxonomic considerations ...... 2 1.3 Genetic studies ...... 2 2 Distribution and current status ...... 3 2.1 World ...... 4 2.2 Europe ...... 4 2.3 United Kingdom ...... 6 2.3.1. England...... 11 2.3.2. Scotland...... 12 2.3.3. Northern Ireland...... 12 2.3.4. Wales...... 12 3 Ecology & Life Cycle...... 12 4 Habitat Requirements ...... 16 4.1 Landscape perspective: Habitats where Salvia pratensis occurs ...... 16 4.2 Communities & Vegetation ...... 17 4.3 Summary of habitat requirements ...... 18 5 Management Implications ...... 19 6 Threats/ Factors leading to loss or decline or limiting recovery ...... 22 7 Current Conservation Measures ...... 23 7.1 In Situ Measures...... 23 7.2 Ex-Situ Measures ...... 26 7.3 Research Data ...... 26 7.4 Monitoring Salvia pratensis and the common monitoring standard...... 26 8 References ...... 26 9 Acknowledgements ...... 28 10 Contacts...... 28 11 Links...... 29 12 Annex 1 - Holly Court Bank Monitoring Form 2002...... 29

1 1 Morphology, Identification, Taxonomy & Genetics

1.1 MORPHOLOGY & IDENTIFICATION Salvia pratensis is a long-lived, erect perennial herb up to 80cm tall, glandular above; leaves ovate or ovate-oblong, cordate at base, doubly serrate; bracts small, green or tinged purplish; corolla 15-30mm in bisexual flowers, down to 10mm in female flowers, violet-blue. Native; calcareous grassland, scrub and wood-borders. Very local in c. 12 places in Southern England from East Kent to West Gloucs; Mons; naturalised elsewhere in central & southern Britain (Stace, 1991).

Salvia pratensis is a striking and easily recognisable member of the Lamiaceae. The only native plant with which it could be confused is Salvia verbenaca, wild clary. The two are differentiated as follows:

Bracts green, often tinged violet-blue, much shorter than Salvia pratensis flowers. Longest hairs on calyx white, eglandular; corolla with 0 or few Salvia verbenaca glandular hairs; lower leaves often distinctly lobed. Longest hairs on calyx brownish, glandular; corolla with many Salvia pratensis glandular hairs; leaves at most strongly doubly serrate.

Hybrids between the two have been reported from France but not the UK.

Figure 1 – Close up of Salvia pratensis flower (Photograph by Tim Rich, 1995).

1.2 TAXONOMIC CONSIDERATIONS None.

1.3 GENETIC STUDIES Some genetic studies have been carried out on Salvia pratensis in the UK (Kay and John, 1995). Additionally the genetics of 32 plant species of lowland grassland in Wales were studied, including Salvia pratensis. Six enzyme systems were studied using electrophoresis. Nine loci were found, of which five were polymorphic and four homozygous. The pattern of heterozygosity was variable and unexpected, suggesting that recombination might be restricted by a degree of structural heterozygosity. Kay and John found a strong correlation between geographic location and genetic identity, and

2 there was a suggestion of a cline of decreasing genetic variability from east to west, with the Monmouthshire population having least variation.

Genetic studies on 14 Salvia pratensis populations in the Netherlands found 11 polymorphic loci, and a significant positive correlation was found between population size and both the proportion of polymorphic loci and the average number of alleles observed per locus (Van Treuren et al, 1991). Another study on Dutch Salvia pratensis populations showed a significant positive correlation between phenotypic variation and population size (Ouborg et al, 1991). From this it was inferred that genetic erosion had caused allozyme loss in small populations (Bijlsma et al, 1991 & 1994). Gene flow between populations in the Netherlands was found to be zero or very low (Van Treuren et al, 1991).

In order to establish whether the genetic erosion and hence allozyme loss observed was threatening the viability of small populations of Salvia pratensis, a further study examined the relationship between population size and genetic fitness in a number of Salvia pratensis populations in the Netherlands (Ouborg & Van Treuren, 1995). Four populations from the previous studies were examined, and a number of fitness components compared in a common garden experiment. Seed size, germination rates, plant growth and reproductive success were compared between the four populations. The study showed that there was no correlation between population size and any of the fitness traits, although these did vary significantly between populations. The authors concluded that the differences were mainly maternal effects and not genetic. They did raise a note of caution suggesting that although low allozyme diversity in small populations was not currently causing any loss of fitness, this may be because these populations had only recently become small and were therefore at an early stage in the genetic erosion process. They postulated that progressive genetic erosion could lead to increased loss of alleles, increased homozygosity and hence a loss of fitness in the future.

Miles King and Plantlife International’s field volunteers known as Flora Guardians collected samples of Salvia pratensis from all available English and Welsh populations in 2001, 2002 and 2003 (Wheeler, 2001; King, 2002). Dr Johannes Vogel at the Natural History Museum in London will investigate the allozyme diversity of these samples of Salvia pratensis from the UK and compare it with that of material from a number of locations in continental Europe. Dr Vogel and his team initially analysed variation across 18 enzymes in one population of Salvia pratensis. They found five enzymes with two or three alleles, and another two enzymes that may contain more than one allele. Given that these results were from just one population, Dr Vogel is confident that it will be possible to build up a comprehensive picture of variation across the UK once material from as many sources as possible has been collected and analysed.

2 Distribution and current status CLIMATIC AND TOPOGRAPHIC LIMITATIONS TO DISTRIBUTION. Salvia pratensis occurs in Britain at the North Western edge of its European range. As a temperate species, it can be expected to cope with hard winters, but also requires warm dry summers, and may have difficulties in very wet years and particularly in wet winters. It favours south or west facing slopes and now only occurs as a probable native as far north as the north Cotswolds (Chipping Campden) and as far west as Monmouthshire. Records from further north, and into the wetter southwest, do not indicate any great length of persistence following introduction, presumably as a hay or grass-seed contaminant. It is interesting to note though that one site, at Rauceby in Lincolnshire, did support a population of two large plants between 1937 and 1951. It can occur in (sub) alpine meadows to 1920m, suggesting that it is well adapted to cold, and to long winters. Perhaps the poor survival of seedlings through wet winters determines how far north and west in Europe Salvia pratensis can occur naturally.

3 2.1 WORLD Salvia pratensis occurs in Morocco, and as far as Kazakhstan, Armenia and Azerbaijan (Wigginton, 1999).

Figure 2 – World distribution of Salvia pratensis (Hulten & Fries, 1986)

Reproduced with kind permission from the publishers Koeltz Scientific

Books.

2.2 EUROPE Salvia pratensis belongs to the temperate element of the European flora (Preston & Hill, 1997) and has a southern - continental distribution. It is rare in Northern Europe and Scandinavia, but common in many other parts of Europe (Preston et al, 2002; Wigginton, 1999). It is rare and protected in the UK and the Netherlands, where it reaches its North Western limit; it is also Rare in France and Vulnerable in south Sweden (although here it is naturalised) and occurs in Belgium and Germany. Salvia pratensis grows up to an altitude of 1920m and is common in Alpine meadows. We believe it is absent from Iceland, Ireland, Norway and Finland. Salvia pratensis also occurs in the Pyrenees in the west, to the Urals in the east (Wigginton, 1999). Table 1 shows the status of Salvia pratensis in countries where red data books are available. Its overall status in Europe is not threatened.

4 Table 1 - Country by country status of Salvia pratensis across Europe (*Country codes are taken from Flora Europaea as of 1964 with red data book listings where available [Definitions of the red list categories].)

IUCN SOURCE COUNTRY* STATUS NOTES LISTING (S) Al ALBANIA Au AUSTRIA WITH LIECHTENSTEIN Az AZORES Be BELGIUM WITH LUXEMBOURG Bl ISLAS BALEARES Br BRITAIN LR Nationally scarce. Most of the Preston et INCLUDING ORKNEY, ZETLAND losses of native sites seem to al. (2002). & ISLE OF MAN have taken place before 1950, and there is little evidence for a significant decline in recent years. Introductions appear to be decreasing. Bu BULGARIA Co CORSE Cr KRITI WITH GAVDHOS, KARPATHOS, & KASOS Cz CZECHOSLOVAKIA Not Bures et al. listed (2001). Da DENMARK Not Ingelög et listed al. (1993). Fa FAEROER Fe FINLAND Not Kotiranta INCLUDING AHVENANMAA listed et al. (1998); Ingelög et al. (1993). Ga FRANCE Not Olivier et listed al. (1995). Ge GERMANY Gr GREECE Not Phitos et EXCLUDING KRITI & ISLANDS listed al. (1995). OUTSIDE EUROPE Hb IRELAND BOTH NORTHERN IRELAND &THE REPUBLIC OF IRELAND He SWITZERLAND Ho NETHERLANDS Hs SPAIN WITH GIBRALTAR & ANDORRA, EXCL BL Is ICELAND It ITALY Not Conti et al. listed (1997). Ju YUGOSLAVIA SERBIA Not Stevanović listed (1999). Lu PORTUGAL No NORWAY

5 IUCN SOURCE COUNTRY* STATUS NOTES LISTING (S) Po POLAND Not Kaźmiercza listed kowa & Zarzycki (2001); Ingelög et al (1993). Rm ROMANIA Rs (N) NORTHERN DIVISION Rs (B) BALTIC DIVISION ESTONIA Not LATVIA listed Ingelög et LITHUANIA VU al. (1993). Rs (C) CENTRAL DIVISION Rs (W) SOUTH-WESTERN DIVISION Rs (K) KRYM (CRIMEA) Rs (E) SOUTH-EASTERN DIVISION Sa SARDEGNA Sb SVALBARD (SPITSBERGEN, BYORNOYA, JAN MAYEN) Si SICILIA Su SWEDEN EN Gärdenfors (2000); Ingelög et al. (1993). Tu TURKEY Not Ekim et al. listed (2000).

2.3 UNITED KINGDOM Salvia pratensis is extant and currently thought to be native in 19 sites in England and one in Wales (Preston et al, 2002). The distribution is now concentrated in the Cotswolds, with 12 extant sites in Oxfordshire and 2 in Gloucestershire (see Table 2). Other extant sites are scattered across the southern Chalk, from the Buckinghamshire Chilterns to the North Downs, South Downs and Salisbury Plain. It is accepted as native or probably native in Kent, Surrey, Sussex, Oxfordshire, Buckinghamshire, Wiltshire, Gloucestershire and Monmouthshire (Wigginton, 1999). It is extinct in many English and Welsh counties, although almost all of these records have been determined as introductions (Rich, 1994 & 1995).

Table 2 - Known extant native sites for Salvia pratensis in Britain. (* Numbers refer to flowering plants; undifferentiated masses are called clumps – C)

VC NATURAL AREA SITE NAME GRID REF. ABUNDANCE* DATE

7 80 South Wessex Tenantry Down ST929502 1 2002 Downs 12 80 South Wessex Porton Down SU245370 50? 09/02 Downs (possibly native) 13 74 South Downs Anchor Bottom TQ205094 1? 08/03 15 69 North Downs Langdon Cliffs TR335421 4C 08/02 15 69 North Downs Boxley Warren TQ775595 0 09/02

6 VC NATURAL AREA SITE NAME GRID REF. ABUNDANCE* DATE

16 69 North Downs Queendown TQ831630/ 12C 06/02 Warren 29 16 69 North Downs Cobham wood TQ698681 4 2002 17 69 North Downs Colley Hill TQ2483521 28 2002 0 23 55 Cotswolds Abel Wood SP 403147 4C+1 06/02 23 55 Cotswolds Ardley-Middleton SP 535250 5C 05/02 Stoney Verge 23 55 Cotswolds Charlbury SP 357199 3C 06/02 (Wigwell NR) 23 55 Cotswolds Charlbury- SP 367219 0 2002 Enstone Road Verge 23 55 Cotswolds Grintley Hill Br SP 410152 65 2002 23 55 Cotswolds Cornbury Park SP 358175 1C 05/02 23 55 Cotswolds Langland Farm SP 364164 83 06/02 23 55 Cotswolds Glyme Farm SP 326264 >500 2002 23 55 Cotswolds Holly Court Bank SP 386152 >200 2002 23 55 Cotswolds Kitesbridge Farm SP 296116 1 + 10 2002 introduced. 23 55 Cotswolds Lower Farm SP 363216 36 2002 Meadow (Taston) 23 55 Cotswolds Saltway SP 358232 53 06/02 23 55 Cotswolds Stonesfield SP 390162 44 06/02 Common 23 55 Cotswolds Whitehill Bridge SP 394157 0 2002 24 65 Chilterns Dancers End SP900095 1+10? intro 05/02 24 65 Chilterns Naphill SU852977 1?+8C intro. 07/02 34 55 Cotswolds Box Farm ST865996 >30 08/03 34 55 Cotswolds Charlton Kings SO962186 3C 08/03 Common 35 - Rogiet ST45-7884 2C -

7

Figure 3 – Map of the British distribution of Salvia pratensis.

DECLINE Assessing the real decline in Salvia pratensis in the UK is very difficult given the uncertainties over its native status at so many of the extant and historic sites. It appears to have undergone a significant decline (change index value = - 0.75) between the original BSBI Atlas survey and Atlas 2000, although most of the lost sites are thought to have been introductions (Preston et al, 2002). The following table shows the vice- counties in which Salvia pratensis has appeared historically but is now extinct.

8

Table 3 - Vice Counties where Salvia pratensis has been recorded historically.

V-C NO. VICE-COUNTY NAME

1 West Cornwall 2 East Cornwall 3 South Devon 6 North Somerset 8 South Wiltshire 9 Dorset 10 Isle of Wight 11 South Hampshire 14 East Sussex 20 Hertfordshire 21 Middlesex 22 Berkshire 25 East Suffolk 26 West Suffolk 27 East Norfolk 28 West Norfolk 29 Cambridgeshire 30 Bedfordshire, 31 Huntingdonshire 32 Northamptonshire 33 East Gloucestershire 36 Herefordshire 37 Worcestershire 38 Warwickshire 40 Shropshire 41 Glamorgan 53 South Lincolnshire 54 N. Lincolnshire 55 Leicestershire 57 Derbyshire 61 SE Yorkshire 70 Cumberland

Most of these losses occurred before 1930. In the last five years native sites have been lost in Kent (Boxley Warren, last record 1997), West Sussex (Beeding Hill, last record 1998), Oxfordshire (Charlbury – Enstone road verge, last record 1998) and Gloucestershire (Daneway Banks, last known record 1994). It may also now be extinct at Whitehill Bridge (Roman Villa), where it was last seen in 2001.

Many of the extant native sites appear to have undergone population reduction in recent decades, although some of these have been reversed thanks to changes in management, and bulking-up exercises. Assessing population reductions is also confounded by difficulties in determining number of individuals in each population, given the apparent extent of vegetative reproduction.

Rich (1999) states that it has been recorded as a native or possible native in 17 10km squares since 1990, and in another 15 before 1990, with most of these disappearing

9 before 1950. Data from herbaria and county floras were collected on all historic sites (Rich, 1994 & 1995). Based on this information, the following 10km squares (shown in Table 4) held native or possibly native populations, which are now extinct. The results of ongoing research by Plantlife (2002-4) into the genetic diversity of UK populations should shed light on the native status of all extant sites.

Table 4 – Native status of extinct populations of Salvia pratensis in England.

10KM NATIVE NATIVE NATIVE SITE (S) COUNTY DATES OR (WIGGINGTON, (PRESTON POSSIBLE 1999) et al, (RICH) 2002) SO62 No No Yes Ross Heref’shire 1891 SO80 Possible Yes Yes Woodchester W. Gloucs - (34) Bulls Cross E. Gloucs - (33) Painswick E. Gloucs 1915 Selsey Hill, E. Gloucs 1930 Stroud SO81 Possible Yes Yes Sheepscombe E. Gloucs - Gloucester E. Gloucs - Docks SP00 Possible Yes Yes Meadow, 1.5 E. Gloucs 1910- miles from 1915 Cirencester SP10 Native Yes Yes Fairford E. Gloucs - Quenington E. Gloucs 1912- - SP02 Possible Yes Yes Near E. Gloucs - Salperton SP12 Native Yes & extant Yes & Old E. Gloucs 1920- post 1987 extant Hinchwick 87 1987-99 SP33 Native Yes Yes Near Sibford Oxon (23) 1910 SP80 Native Yes & extant Yes & Coombe Hill Bucks 1983 post 1987 extant 1987-99 SP81 Native Yes Yes Dancers End Bucks (24) Still NR extant ? SU58 Native Yes Yes Moulsford Berks (22) 1909- Downs 1955 Unhill Bottom SU67 Native Yes Yes Near Goring Oxon 1931- Pangbourne 1981 Hartslock NR SU94 Possible Yes Yes The Mount, Surrey (17) 1847 - Guildford 1985 TQ06 No No Yes Laleham, Surrey 1901 Lammas Land TQ15 Native Yes Yes Colley Hill Surrey Extant (should be 18471 TQ25) 937 Mickleham Leatherhead

10 10KM NATIVE NATIVE NATIVE SITE (S) COUNTY DATES OR (WIGGINGTON, (PRESTON POSSIBLE 1999) et al, (RICH) 2002) TQ67 No Yes No Cobham Kent 1994 (extant Wood 87-99) (transposition error) TQ75 No No (extant Yes Boxley Surrey 1997 (extant post 87) Warren 1990) TQ66 Native Yes No Cobham W. Kent -1980 (extant (16) 87-99) TQ85 Possible Yes Yes Thurnham E. Kent 1904- Hills, nr the (15) 1917 castle TR24 Possible Yes Yes Barham, N. E. Kent 1939 of Broome Park, E of Gravel Castle

The original native Salvia pratensis plant may still be growing at the SP 81 Dancers End site although re-introductions using seed from the original population have been carried out there by the BBOWT site manager Mick Jones. The TQ15 Colley Hill reference (Rich, 1995) is erroneous for TQ25, where Salvia pratensis still occurs on Colley Hill. TQ67 is marked as a native extant site in Wigginton, 1999 and Preston et al, 2002, although there are no records for that square in Rich, 1994 & 1995. However, this is a transposition error as the erroneous datum is TQ680701, whereas the site is recorded in Rich (1994) as TQ 701680. Similarly there are no data in Rich (1994 & 1995) for two 10km squares, SO62 and TQ06, from Preston et al 2002. These may have surfaced after Rich left the Plantlife project, and before the Atlas was completed. As both are single records, they should be treated very cautiously.

It is clear that many records that have been classed as native extinct sites are in fact doubtful. Some records are based on a single sighting, such as at Barham, Kent or the two separate sightings many years apart near Box Hill, Surrey. For others there is clear evidence of a long history of recorded presence such as Hinchwick, Gloucs and the Oxfordshire/Berkshire borders near Goring. Others occurred immediately adjacent to currently extant sites, e.g. Coombe Hill, Bucks and Cobham and Thurnham Hills, Kent.

2.3.1. ENGLAND Salvia pratensis is thought to be native in eight vice counties across southern and central England, (and South Wales). Oxfordshire holds most of the current UK population, though many populations are small.

Most extinct Salvia pratensis sites that are thought to have been native were in the Cotswolds Natural Area. As the Cotswolds are still the main stronghold for Salvia pratensis in Great Britain this is not unexpected. Most of these Cotswolds populations became extinct before 1930, with two surviving into the 1980s. These were at Hinchwick, Gloucs and at Finstock, Oxon, near the Manor House. The latter was known for many years (mentioned in Druce, 1886) and only became extinct around 1993 (J Dunn, pers. comm.). The former was once a large population, with several hundred plants found in 1966, although even then they occurred in scrub. Before that, there were sites in East Gloucestershire near Stroud, Painswick, Cirencester and Fairford. A single most likely native plant was seen in 1964 on the road verge between Charlbury and Chipping Norton, at SP336252; it may well have been an outlier from the Saltway colony, but is now extinct (J Dunn, pers. comm).

11

Three native historic sites are known from the Chilterns Natural Area; in the north, the site described as Coombe Hill may have been the one next to Dancer’s End BBOWT nature reserve, at SP898099, and is probably best dealt with as part of the Dancer’s End population. However, there is a possible complication in that there are two sites called Coombe Hill in SP80, the other being the Coombe Hill at SP840060. There were two populations close together on the border between the southern Chilterns and the North Wessex Downs Natural Area, at Hartslock BBOWT nature reserve, where Salvia pratensis was known from 1931-1981, and Moulsford Down / Unhill Bottom, west of Goring, known from 1909-1955.

In the North Downs Natural Area, there are records at the Mount, Guildford, and from Mickleham (1847) and Leatherhead (1937). All are single records and should be treated cautiously. The Boxley Warren site is now extinct, with the last plant seen in 1997; this site has a long recorded history. It also appears that the population from the long- recorded site at Cobham may have extended into adjacent squares in the past, but has contracted more recently. Local colonisation from Cobham, Queendown or Boxley could also explain the relatively short-lived population at Thurnham Hills. A single record from Barham in 1939 could also have been caused by local colonisation from the nearby Langdon Cliffs colonies, near Dover.

In the South Downs Natural Area, there have only been two recorded native sites very close to each other at Beeding Hill and Anchor Bottom. The latter is still extant; the former, a single plant just a few hundred metres from the main population, has not been seen since 1998 and is presumed extinct. Again these ought to be treated as part of the same population.

2.3.2. SCOTLAND Not recorded.

2.3.3. NORTHERN IRELAND Not recorded.

2.3.4. WALES There is only one extant site in Wales, at Rogiet in Monmouthshire. There are no former native sites. This extant site could be considered as an outlier of the Cotswolds population, and it does appear to be genetically more similar to these populations than others (Kay & John, 1995). It may have been introduced from a Cotswold site (Rich, 1994).

It may never be possible to determine whether these sites were actually native or not, or whether other sites which have been discounted as introductions were in fact native, short-lived sites.

3 Ecology & Life Cycle A detailed study of the ecology of Salvia pratensis was carried out (Scott, 1988) and much of this section draws on this study.

Flowering - Salvia pratensis is a large rosette-forming perennial and can live for many years; Dutch studies suggest that individuals can live for one to a few decades (Ouborg, 1993). Salvia pratensis flowers from May to September. Flowers appear in whorls around flowering shoots that may be branched and each plant may have one or more flowering shoots. The proportion of flowering plants in a given population varies, depending on the age of the plants, shade and density of plants. Some evidence suggests that plants are vegetative at the beginning and end of their lifecycle (Berko, 1974). Scott found no

12 correlation between flower shoot height and soil depth, but did find a positive correlation with sward height, suggesting that plants tend to produce more flowering shoots in unmanaged tall grassland. The largest plants observed by Scott had over 80 flowering shoots. Scott found that slightly over half of all flowering shoots were un-branched, but that branched shoots were taller and supported more flowers. Flowering shoots that appeared after cutting were shorter; less branched and had a lower density of flowers than on uncut plants.

Figure 4 – Salvia pratensis at

the Plantlife reserve in Kent,

Queendown Warren.

(Photograph by Joe Sutton).

Floral biology – Salvia pratensis is self-compatible, although insect pollinators are required for within-flower selfing. Flowers are typically dark blue, but occasionally flowers can be white, light blue or pale violet. Flowers are arranged in two opposite and much contracted cymes composed of three flowers each, forming an apparent whorl or verticil. Adjacent flowers are arranged perpendicularly, on the upper stem, some of which are branched. Within each whorl the central flower of each cyme opens first and whorls open from the base up the spike. The terminal spike begins the flowering process, but as flowering ends spikes synchronise their flowering. Individual flowers last two or three days, before the lower lobe withers and stigmas brown. Flowering period varies from site to site and between years, from May through to September. Peak flowering occurs in late June.

Salvia pratensis is gynodioecious, with most of the plants observed by Scott to be hermaphrodite, and less than 10% female (male sterile) only, with a similar proportion falling into the intermediate category. In the Netherlands male sterile proportions vary from 0-4% in small populations to 21-24% in large ones (Van Treuren et al, 1993).

Stamens of hermaphrodite flowers have a swollen anther full of pollen borne on a long filament contained within the upper corolla lobe. Corollas in hermaphrodite plants are typically 22mm long, with calyx length around 7mm. In male sterile plants both stamens are aborted, while the shrunken anthers appear translucent or brown. These are held on incompletely elongated filaments at the corolla mouth. In male sterile plants the corolla is shorter, about 17mm long, and the calyx is also smaller.

13 Flower shoots can also consist of a mixture of hermaphrodite and male sterile flowers and individual flowers can have one aborted and one normal stamen, with a 20mm intermediate length corolla. Scott observed that male sterile or intermediate flower shoots were more likely to occur in shaded locations or when plants were damaged. This apparent sex change in response to changing conditions has been observed elsewhere and this phenomenon clearly needs further investigation.

Gene flow, inbreeding depression and genetic drift - Gynodioecy promotes outcrossing as male sterile plants must outcross; male sterile plants apparently achieve high levels of seed set, suggesting hermaphrodite plants must outcross to an extent. Overall, Salvia pratensis has relatively high outbreeding rates – 40-80% have been observed (Van Treuren et al, 1993). Gynodioecy is favoured when male sterile plants are fitter mothers than hermaphrodites. This can occur when a type of inbreeding depression called heterosis occurs, when selfed offspring of hermaphrodites are less variable than outcrossed male sterile plants, as male sterile plants expend less energy on stamen and corolla production (removal of male load, Ross 1978). Some work suggests that male sterile plants produce slightly heavier and therefore possibly more viable seeds, than hermaphrodite plants (Holden, unpublished). In a relict population under unfavourable management, it is possible that the proportion of male sterile plants could increase, due to demographic genetic drift, until no seed at all is set (G. Oostermeijer, pers comm).

Pollination, pollen robbing and flower predation - The insects that pollinate Salvia pratensis are mainly long-tongued bumblebees, including Bombus hortorum, B. lucorum, B. pascuorum, B. pratorum, B. ruderarius and Bombus terrestris. Pollen is also robbed by the pollen beetle Meligethes aeneus, and this can in places affect pollination success. Flower robbing was also observed by large skipper and marbled white butterflies, and by some species of bumblebee. Various insects lay eggs on Salvia pratensis flowers on which their larvae feed. These include sawflies of the genus Aglaostigma, and geometrid moths. Aglaostigma sp. larvae caused up to 20% loss of flowers on some plants.

Seed production - Each flower contains four ovules which once fertilised grow at the base of a horizontally held calyx. Scott found that seed set for hermaphrodite plants varied from 84% to 95% on the sites he observed, with nearly 1500 seeds on one plant. Seed set declines later in the season as the number of pollinating insects decreases. Few male sterile plants were studied, though these had lower seed set rates of around 70%. Seed takes around 30 days to ripen, at which point it drops from the open calices, with the help of wind or passing animals.

Most seed falls within a metre of the parent plant, though this can increase down steep slopes. Seeds are covered in sticky mucilage that can help them adhere to animal coats, aiding dispersal. Seed dispersal may also be aided by glandular hairs on the calyx, which promote adherence to grazing animals (Weeda et al, 1988). Seed dispersal beyond the immediate environs of adult plants presumably occurred in the past through a combination of movement via animal coats and movement of hay containing ripe seed. Neither dispersal mechanism is likely to have taken the seed more than a few kilometres, except occasionally, when stock was driven to market, for example.

Hay making before seed is completely ripe tends to trap the seed in the calyx, unless the hay is thoroughly dry. Small mammals predate unripe seed; in one site Scott found over 50% set unripe seed had been predated; ripe seed does not appear to be predated, possibly because the calyx tends to eject the seed under compression when opened. Scott found seed viability to be around 72% for fully ripened seed, but only 34% for unripe seed, which was after-ripened. Germination occurred immediately after wetting, with 95% of seed germinating after 12 days. Immediate germination suggests that a persistent seed bank is unlikely to be established, especially as the mucilage formed on germination anchors seed to soil particles preventing further movement. Salvia pratensis does not apparently have a persistent seed bank (Thompson, Bakker & Bekker, 1997),

14 although seed produced late in the summer can germinate the following spring. Seedlings, identifiable by their persistent cotyledons, appear from July onwards.

Seedling recruitment - Seedling recruitment appears to be most successful on site where bare ground is present, either caused by shading by large rosettes of Cirsium vulgare or Salvia pratensis, or rabbit scrapes, anthills, worm casts, small mammal runs or human paths. Most seedlings found by Scott were on active anthills, though these tended to die early, as they were buried by more earth. Seedling establishment was much reduced where significant litter had developed, and dense patches of Brachypodium pinnatum produced too much litter and hence shade, for seedlings to develop. Seedling establishment was also very poor in Festuca rubra - Bromopsis erecta tall-ungrazed grassland. Festuca rubra also forms a dense thick litter layer. Short, rabbit-grazed grassland with little leaf litter accumulated and more exposed mineral soil produced far more seedlings, though occasionally moss such as Pseudoscleropodium purum built up and prevented seedling establishment. In a recent study in the Netherlands, seedlings were mainly found in populations in late-mown grasslands, and six out of seven sites with seedlings were mown, with only one site grazed; this study also found bare ground was a strong indicator of seedling presence (Hegland et al, 2001).

Distinguishing individual plants - Salvia pratensis plants can be very large and can grow very close together, making it difficult to determine the boundary of individual plants. Scott found it possible to distinguish individuals by morphological characters, particularly flowers, sex and growth form. He also dug around roots of individual plants to investigate these distinctions. Some plants excavated showed that shoots up to 7cm apart above ground were from the same plant, while other shoots separated by only 1cm were from different plants. Scott did not excavate into the C horizon and so could not be sure that Salvia pratensis did not form deep-rooted clones covering large areas. He was confident however that this did not occur to any great extent. Grazed or non-flowering plants are particularly difficult to tell apart, compared with flowering plants.

Rootstock and shoot production - Scott found that small single-shoot individuals have a taproot growing vertically downwards, while the roots of plants with two or three shoots grow horizontally for 4-5cm. The upper rootstock had very dense fibrous lateral roots. Older thicker tap roots of flowering shoots degenerated in the centre forming a cavity bounded by root tissue persisting as a series of fibrous, sometimes spirally coiled, strands that rejoin near the inflorescence. Vegetative shoots can continue to grow from one season to another, while flowering shoots terminate on flowering, forcing adjacent lateral buds to develop. These dormant buds only develop when the main flowering shoot is damaged or has finished flowering. From July to early September, flowering shoots grow one or more vegetative shoots at their base. These can die back completely in hard winters, but can also over-winter as a slightly wintergreen rosette (Salisbury 1952).

Vegetative growth - It is not known to what extent, if at all, that Salvia pratensis spreads vegetatively. In the Netherlands no vegetative propagation has been observed (Hegland, et al) and Scott saw no vegetative propagation either, though rooting at the nodes of decumbent stems has been reported, and it is possible to propagate stem cuttings, as well as leaf cuttings. Salvia pratensis has been observed growing massed together in a manner that suggests vegetative spread similar to other labiates (Peter Marren, pers. comm.). Some populations have been observed to spread under heavy grazing conditions, which prevent flowering. This indicates some degree of vegetative propagation occurs, but further research is needed to confirm this and its significance relative to sexual propagation. For this reason, the current approach to censuses of Salvia pratensis populations in the UK is to differentiate between flowering plants and clumps, which may be a single large vegetatively spreading plant, or a number of separate plants growing very close together.

15 Response to grazing and hay cutting - Summer hay-cutting significantly reduces the number of flowering shoots and flowering plants, but new leaves are grown following cutting, and in some cases flowering can also occur after hay cut. Following cutting, the site of new growth depends on the height at which the stem is cut – growth can either originate from the basal leaf axils of cut stems, or from rootstock buds.

Both cattle and sheep graze Salvia pratensis flower shoots and leaves. Sheep selectively graze Salvia pratensis, first removing flower shoots and then unripe calices and leaves. Rootstock buds at ground level escape grazing, and develop following the loss of apical meristems. Shoots which had re-grown following grazing were adpressed and much smaller than those produced in un-grazed conditions. Grazing caused flowering shoots and approximately half the vegetative shoots to produce new vegetative shoots, but no new flowering shoots. Cattle unselectively grazed leaves and inflorescences, intense cattle grazing also causing small adpressed shoots, as with sheep. Rabbits may bite through inflorescences, but do not apparently feed on leaves. Deer apparently have more impact on flowering spikes than rabbits. Molluscs, e.g. slugs or snails, also account for the loss of some inflorescences and may be responsible for up to 40% loss of flowering shoots.

4 Habitat Requirements

4.1 LANDSCAPE PERSPECTIVE: HABITATS WHERE SALVIA PRATENSIS OCCURS Salvia pratensis currently occurs in a very small number of localities in the UK, and in its native localities it is now confined to south, southwest or west-facing slopes on unimproved calcareous grassland in the Cotswolds, Salisbury Plain, the Chilterns, North Downs and South Downs. Many sites are isolated, while others form a small number of clusters (Gloucestershire, Chilterns, North Downs), with the largest cluster around the Wychwood Forest area of Oxfordshire.

Salvia pratensis occupies a relatively discrete group of habitats, all associated with traditional farming techniques. It occurs in hay meadows; sheep, cattle or horse-grazed pastures; green lanes and road verges; scrub, and woodland clearings. All of these habitats share similar features: the management, or lack of management enables the Salvia pratensis plants to flower and set seed, to a greater or lesser extent, each year. Recruitment from seedlings into populations is dependent on appropriate levels of soil disturbance, and adequate light levels. Salvia pratensis at unmanaged sites can be threatened by decreasing light levels, as a result of scrub development and consequent shading, as well as a lack of appropriate soil disturbance.

Salvia pratensis occurs in Britain solely on calcicolous soils, derived either from chalk or oolitic limestone. Sites in the Cotswolds and Monmouthshire are usually on brown rendzina soils over Oolite, while the Chiltern, North and South Downs populations occur on rendzina soils over chalk. Some populations also occur on slightly deeper calcareous brown earth soils. High levels of calcium, as would be expected in soils derived from calcareous parent material, but also magnesium, were found (Scott, 1989 & Cauwood, 1981) in some sites supporting Salvia pratensis, although the contention that high magnesium levels are required by Salvia pratensis (Cauwood, 1981) is not supported by observations by Scott (1989). In the Netherlands Salvia pratensis is only found in dry floodplain grasslands, where its ability to colonise newly formed river dunes suggests it may be water-dispersed in this habitat (Hegland et al, 2001). In continental Europe Salvia pratensis is not restricted to calcicolous soils, and it has been suggested that the British distribution is more linked to a need for well-drained warm conditions, rather than for a particular chemical basis (Salisbury, 1952). Scott found Salvia pratensis preferred locations on shallow soils and steeper slopes within occupied sites.

16 IS SALVIA PRATENSIS NATIVE? The current and recent historic distribution of Salvia pratensis is somewhat disjunct, and the lack of any historic records from large areas of Southern and Central England, where suitable habitat survived until relatively recently, is intriguing. No late 19th or 20th century records from the East Anglia chalk or from Bedfordshire, Northamptonshire, Leicestershire, Dorset, Somerset, and particularly, Hampshire have been accepted as native. Did Salvia pratensis really become extinct in Hampshire and most of Wiltshire before the mid-19th Century? If it was a recent introduction, and the first record as a wild plant is quite late at 1699 (Preston et al, 2002), then it must have been introduced very locally, or undergone a rapid expansion, followed by a rapid reduction in range. It was regarded as rare in the 19th century (French, 1875; Druce, 1886) and these authors attributed its decline to the ploughing up of downland (Scott, 1989).

All the native extant and recently extinct sites for Salvia pratensis lie very close to or on pre Roman or Roman roads, or mediaeval drove-ways. One view is that the plant could have been introduced by the Romans, intentionally or otherwise, and spread along the Roman roads of southern England, and that widespread ploughing of downland in the 19th century would not leave a scattered distribution with such a strong relationship with the ancient road system. There is an almost linear distribution of sites from the Roman port of Dover, along Watling St, across the North Downs Ridgeway, to Akeman St, through Buckinghamshire, Oxfordshire and into Gloucestershire. Several populations occur along saltways in Oxfordshire and Gloucestershire, others fall on the Icknield Way in Buckinghamshire and south Oxfordshire. Other Gloucestershire sites lie near Ryknild St, near Cirencester, the Fosse Way, Ermin Way and the Roman port of Gloucester. Even isolated sites fit the trend; Anchor Bottom is only 4km from a Chichester-bound Roman road, while Rogiet is within 2.5km of the Roman road from Caerwent to Caerleon. Most of the Oxfordshire sites occur within 2km of a Roman road (mainly Akeman Street) or even closer to Roman villa sites. Some sites still occur on the edges of Roman roads.

It is also possible that Salvia pratensis was spread by people along the ancient road network, but not by the Romans. The ancient ridgeways and Roman road network was used for trade and travel well into modern times. It was not until the 18th century when the turnpike roads were built that this network was expanded; and it was only superseded when the 20th century trunk road and motorway network was built. It is possible that Salvia pratensis seed contained within hay from local meadows or trapped in the coats of animals on the move could have been scattered along this road network and colonised road verges and nearby fields. An alternative theory is that the distribution can be explained by the association of suitable unploughed banks, where it can and does survive, with these ancient tracks. Perhaps the current genetic studies, or more detailed DNA fingerprinting in the future, will shed further light on the origins of Salvia pratensis and its unusual distribution.

4.2 COMMUNITIES & VEGETATION Salvia pratensis occurs in Britain in grazed and un-grazed grasslands, hay meadows, road verges, green lanes, scrub and woodland clearings. It has been recorded in a wide range of communities identified in the National Vegetation Classification (Rodwell et al, 1991), considering how few sites there are. These are:

CG2 Festuca ovina – Helictotrichon pratensis grassland. CG3 Bromopsis erecta grasslands. CG4 Brachypodium pinnatum grassland. CG5 Bromopsis erecta – Brachypodium pinnatum grassland. MG1 Arrhenatherum elatius coarse grassland. MG7 Lolium perenne leys. W21 Crataegus monogyna - Hedera helix scrub.

17 Insufficient data have been collected to date to place Salvia pratensis in any sub- communities of these communities.

Scott carried out ordination on vegetation quadrats from 10 sites (Scott, 1989). These clustered into 3 groups equivalent to unmanaged CG4/CG5, which is prevalent at Holly Court Bank in Oxfordshire; CG3 grazed at moderate to high density; and cut or lightly grazed MG1 type grassland, which is typical of road verge sites. He found several sites to have communities intermediate between CG3 and MG1, and recent data collected from Holly Court Bank also suggests intermediate communities between CG3, CG4, CG5 and MG1. Although Salvia pratensis plants are known to occur within the scrub edge community of W21, they cannot reproduce effectively in dense shade, and adult plants eventually also succumb under dense scrub.

Recent research in the Netherlands has investigated the relationship between population structure, community composition and site management (Hegland et al, 2001). Salvia pratensis in the Netherlands is rare and protected, and only found in a small number of sites within a dry floodplain grassland community that is not present in the UK. This study showed that there were three different types of Salvia pratensis population; dynamic populations, which were characterised by a large proportion of young individuals (seedlings, juveniles and immature plants); normal populations, with a relatively higher proportion of adults but still plenty of immature individuals; and regressive populations, in which adult stages, especially large flowering individuals, dominated and rejuvenation hardly occurred. Population size and plant density were highest in dynamic populations, and lowest in regressive populations. The vegetation communities in which Salvia pratensis occurred were associated with the type of population, with dynamic populations occurring in species-rich grassland on nutrient- poor soils, while regressive populations were found in species-poor nutrient-rich conditions. Population structure was also associated with site management, with the optimal management regimes being a late hay cut, which supported the largest and most dynamic populations.

4.3 SUMMARY OF HABITAT REQUIREMENTS

Figure 5 – Salvia pratensis habitat at Kings Glyme Farm (Photograph by Miles King).

18 Table 5 - Habitat features important to Salvia pratensis in Britain.

TYPE DESCRIPTION Physical & Usually occupies south, south-west or west-facing slopes. Topographical Prefers moderate to steep slopes. Low altitude in UK, highest site at circa 300m in Gloucestershire. Can grow up to 1920m in the Alps. Restricted to calcareous soils derived from chalk or oolitic limestone. Soils are mainly rendzinas, rarely calcareous Brown Earths. Susceptible to rot in moister conditions/locations. Vegetation/Structural Adult plants are strong competitors in managed grassland. Tall flower shoots attain canopy height in managed or unmanaged grassland. Can survive in transitional scrub. Shading affects flowering performance. Able to withstand intense grazing, although this prevents flowering. Can thrive in hay meadows if hay cut is sufficiently late. Seedling germination requires bare soil patches. Litter accumulation in unmanaged grassland prevents seedling establishment. A component of several different calcareous grassland NVC communities, coarse grassland and scrub. Chemical Neutral-calcareous to calcareous soils; putative association with high magnesium content.

5 Management Implications As a dry grassland species, the typical management activities of grassland, grazing, mowing, scrub management, soil disturbance and altering soil fertility, are all important for Salvia pratensis in the UK. Table 7 shows the current management at native extant Salvia pratensis sites in the UK.

Table 6 - Current management at native extant Salvia pratensis sites in the UK (after Davis, 1999).

VICE SITE MANAGEMENT IN 2002 COUNTY Tenantry Down South Winter cattle grazing. Wilts Porton Down North None. Extensive rabbit and deer grazing. Hampshire Exclosures. Anchor Bottom West Cattle grazing. Sussex Boxley (extinct) East Kent Scrub clearance. Queendown Warren East Kent Sheep and cattle grazing. Langdon Cliffs East Kent Some amenity mowing. Cobham Wood West Kent Scrub clearance. Colley Hill, Surrey Surrey Scrub clearance and cattle grazing, with exclosures. Asthall Leigh Oxon Light cattle grazing. Glyme Farm Oxon Rotational sheep grazing, with exclosures.

19 VICE SITE MANAGEMENT IN 2002 COUNTY Wigwell, Charlbury Oxon Late hay cut, winter grazing with sheep and cattle. Saltway Oxon Scrub clearance, mowing and scarification by ANHSO. Cornbury Park Oxon Late summer + autumn sheep grazing. Lower Farm Meadow Oxon Late hay cut, autumn and winter sheep grazing. Langland Farm Oxon Horse grazing. Holly Court Bank Oxon Mostly deer and rabbit grazing. Stonesfield Common Oxon Summer/autumn cattle grazing; exclosures. Whitehill Bridge (Roman Oxon Occasional safety scrub management. Villa) Abel Wood Oxon Scrub clearance. Grintley Hill Bridge (Coombe) Oxon Scrub clearance on Network Rail part, no management in field. Ardley – Middleton Stoney Oxon Verge cutting, some scarification and weed road verge control. Dancers End Bucks Sheep and cattle grazing with exclosures. Naphill Bucks Scrub clearance and cattle, with deer/rabbit grazing. Charlton King's Common East Amenity mowing. Gloucs Box Farm Reserve East Cattle grazing. Gloucs Rogiet Monmouth Hay cut with cattle grazing.

Grazing of Salvia pratensis sites is the normal mode of management and consists of either formal grazing, with domesticated stock, such as cattle, sheep or horses, or informal grazing, by wild herbivores, principally rabbits and deer, or a combination of the two. Scott suggested that rabbit grazing was a good management approach for Salvia pratensis as it created the bare ground needed for seedling establishment, although it did also lead to the development of a bryophyte layer where seedlings could not establish (Scott, 1989). Where cattle are used, it has often been considered necessary to build permanent exclosures around plants to prevent the cattle from removing flower shoots. There are problems with this approach though, as although Salvia pratensis can survive well as an adult plant for many years without producing flowers, the overall population will decline without regular recruitment of seedlings, entering the regressive state as defined in Hegland et al (2001). Permanent exclosures may protect adult plants from grazing damage, but they will also allow a build-up of litter and coarse vegetation, preventing seedling establishment. Exclosures should therefore only be used during the flowering period, then opened once seed has ripened, to allow stock in to graze around the adult plants, thus creating the bare ground needed for seedling establishment. The timing of opening exclosures will be critical, and dependent on the season and the flowering behaviour of individual plants.

Exclosure of stock using flexi-netting is a more sophisticated management approach, more akin to hay meadow management. Temporary fences are placed around the Salvia pratensis colonies, while grazing continues outside these areas. Temporary fences are then removed once seed has ripened, enabling the vegetation within the temporary exclosures to be grazed and bare ground created. This approach appears to be successful at Glyme Farm in Oxfordshire. Sheep graze at this site, and this also has the benefit of seed being dispersed into new areas of the farm in the sheep’s fleeces.

20 Salvia pratensis plants have also been protected, with some success, on summer-grazed pastures e.g. at Stonesfield Common, Oxfordshire, using cut, spiny scrub branches laid over the plants. This prevents stock and rabbits from eating the flower shoots, while allowing the positive benefits of grazing to continue through the flowering and seeding season. This approach also appears to be successful in preventing removal of flower shoots from plants growing in a woodland glade at Abel Wood, Oxfordshire, where rabbits and deer have prevented seed set in the past.

Cattle, sheep and horse grazing are all effective management techniques for Salvia pratensis, although timing is critical. A site which is only winter-grazed by sheep, for example, can suffer from a build up of litter, and the gradual replacement of a fine species-rich sward, with coarse grasses. This can be a particular problem if Brachypodium pinnatum is present, as this is relatively unpalatable to sheep in the winter, and can form dense mounds with much litter. Spring grazing by sheep is very effective at controlling Brachypodium pinnatum and other coarse grasses, and if stock is removed before Salvia pratensis flower buds appear, no damage is done to the flowering performance. Returning stock after seed is set, in late summer, can then ensure bare ground is created for seedling establishment, as well as continuing to control coarse vegetation. Scott suggests summer grazing every four of five years is acceptable; this is generally true although a site with a regressive population that has been unmanaged or under-managed for years may benefit from an initial restoration phase where summer grazing was repeated until the vegetation composition and structure had been restored to conditions optimal for seedling establishment.

Mowing for hay, either on its own, or in combination with aftermath grazing, is also a very effective technique for managing Salvia pratensis. Hay meadows should be cut in late July or early August, depending on the season (Scott, 1989). This ensures that there is at least some ripe seed available. A less rigid mowing schedule, which takes the temporal and spatial variation among populations into account, seems the best way to maintain or promote the remaining population of Salvia pratensis, according to researchers in the Netherlands (Hegland et al, 2001). Late mowing followed by aftermath grazing provides the combination of flowering and seed-set, suppression of competitive vegetation, and creation of disturbance patches for seedling establishment, which will enable Salvia pratensis populations to be maintained.

Mowing of a different kind is used on road verge sites and where Salvia pratensis occurs in an amenity context. Road verge mowing is constrained by other factors such as road safety, but good working relationships between local conservation groups, such as the Ashmolean Natural History Society of Oxfordshire and Cotswold Rare Plants Group, and Oxfordshire County Council, have ensured that cutting times at the last surviving road verge site between Ardley and Middleton Stoney allow Salvia pratensis to set seed. Unfortunately this approach does not create the bare patches needed for seedling establishment, while the absence of grazing has allowed coarse grasses and tall herbs to become dominant. Post-flowering scarification to create bare patches around these plants is planned for summer 2003, along with herbicide control of more competitive ruderals. Similar approaches have been adopted in other sites where grazing has not been possible, such as Langdon Cliffs in Kent and Charlton Kings Common Gloucestershire. In both these sites large mature adult plants are present but no seedlings have been found for many years. The future of these sites under this management regime is very uncertain.

Scrub control and management is necessary on grassland Salvia pratensis sites where grazing has ceased or has been insufficient, or on the handful of extant sites in woodland glades, rides and clearings. Salvia pratensis can survive as an adult plant in the interface between tall grassland and scrub, also known as “saum”. Although it can flower successfully in these conditions, seed does not appear to be able to germinate and seedlings are not able to establish, as the closed sward has no bare ground and the shade reduces seed germination (Scott, 1989). Scott observed that shade cast by scrub

21 and woodland also reduced the likelihood of successful pollination, thus reducing the amount of viable seed produced (Scott, 1989). He also observed that shade increased the proportion of male sterile flowers within the total flower population, though the reasons for, and mechanism underlying this change, are unclear. Scrub control, followed by the placing of cut scrub species branches over plants to protect them from grazing animals, has been used with some success at Abel Wood, Oxfordshire, where no formal grazing is possible.

Salvia pratensis can survive as an adult plant in the interface between tall grassland and scrub, also known as “saum”, and it can flower successfully in these conditions, because the grassland / scrub matrix prevents flower shoots from being grazed (Scott, 1989). However, seed does not appear to be able to germinate and seedlings are not able to establish, as the closed sward has no bare ground, and the shade discourages seed germination.

The indiscriminate use of herbicides or artificial fertiliser has a damaging effect on individual plants and whole populations. Although mature adult plants can survive under enriched conditions, seedling establishment does not occur, so effectively these populations have no future. Herbicide use has led to the loss of plants at Cobham wood, and although unintentional, this has significantly reduced what is already a small and vulnerable population. A closer working relationship between Plantlife and the land manager is being fostered there and this will hopefully ensure that there is no further unintentional damage.

6 Threats/ Factors leading to loss or decline or limiting recovery

Table 7 – Summary of threats to the survival of Salvia pratensis in Britain.

TYPE THREAT Habitat loss Ploughing of unimproved grassland. Fertiliser & herbicide application. Neglect Scrub encroachment. Lack of grazing. Litter accumulation. Small population effects Increased susceptibility to extinction. Genetic complications due to inbreeding effects. Dispersal Seed cleaning. Abandonment of hay production and movement. Abandonment of drove roads to move stock.

HABITAT LOSS In the past, habitat loss through ploughing of unimproved grassland, or fertiliser and herbicide application, led to the loss of a number of sites for Salvia pratensis. Ploughing of downland was considered to be the principal reason for Salvia pratensis being such a rarity by the end of the 19th century (French, 1875; Druce, 1886). Only one site is thought to have been lost to ploughing in recent decades – at Hinchwick in Gloucestershire.

CHANGES IN HABITAT MANAGEMENT Neglect - As mentioned in the previous section, a lack of grassland management leading to scrub encroachment can adversely affect the chances of Salvia pratensis surviving on a given site. This is also true for woodland glade sites, where some form of management is necessary to prevent the glade from disappearing. Salvia pratensis sites where formal grazing ceases, usually because these sites are agriculturally irrelevant, are under severe threat. The process of scrub encroachment can take several decades,

22 as the un-grazed grassland with its accumulation of litter is a very inhospitable place for seedling establishment for scrub species, as well as for Salvia pratensis. During this time adult plants can survive for many years, allowing the opportunity for restoration management. It seems that the large population at Hinchwick in Gloucestershire had declined due to scrub encroachment, before the last plants were ploughed out.

Small population effects - When a population declines in numbers to a single plant, there is the danger of random events leading to the loss of these singletons, and the extinction of the site. These stochastic events can combine with genetic effects, which become more significant with small populations. For example, if small populations have a higher proportion of male sterile plants, indeed if the last surviving member of a population is a male sterile plant, then self pollination is impossible, and no more seed will be produced without human intervention. It is possible, although as yet unclear, that male sterile plants outlive their hermaphrodite companions in regressive populations, due to removal of male load. If this were the case it would have serious consequences for all regressive populations. A further possibility is that demographic genetic drift could lead to a higher proportion of male sterile plants in small populations. Even where hermaphrodite plants were present in small populations, inbreeding depression caused by the allee effect, could also be a possible concern, as this would reduce overall seed viability and hence seedling recruitment (Hegland et al, 2001). It is hoped that the current studies of allozyme diversity in Salvia pratensis populations will help answer some of these questions.

Dispersal - Salvia pratensis does not appear to be dispersed in the same way as it was in the past. The dramatic decline in hay meadows and unimproved pasture and the loss of hay as an economically important crop, as well as changes in the way hay is moved around the country, have all probably played a part in the decline of Salvia pratensis. Old drove roads which would have allowed Salvia pratensis seeds to be spread in the coats of domestic stock to new sites, have fallen into disuse, as stock are moved by lorry rather than by foot. Advances in seed cleaning technology in the 19th century have also led to a decline in casual introductions with grass seed from continental Europe (Davis, 1999). Nevertheless there are occasional new records for Salvia pratensis, and it is assumed that these originate from garden throw-outs, and the increasing use of wildflower seed mixes, most of which originate in Europe.

7 Current Conservation Measures

7.1 IN SITU MEASURES Species protection In 1992 Salvia pratensis was added to Schedule 8 of the Wildlife and Countryside Act 1981 (as amended) and is therefore afforded special protection under Section 13(2). This makes it an offence to pick, uproot or damage plants unless as an incidental result of a lawful operation that could not reasonably have been avoided. Any activities involving the collection of any part of the plant, including growing wild plants from seed for re-introduction, must be carried out under licence from either English Nature or the Countryside Council for Wales.

Site protection Just over 50% of native extant sites are SSSI and these support the bulk of the UK population of Salvia pratensis in terms of numbers of individuals. Some of these SSSI are also possible Special Areas of Conservation (pSAC) under the EU “Habitats Directive”.

23

Table 8 - Salvia pratensis sites within SSSI / possible Special Area of Conservation (pSAC) / County Wildlife Site (CWS).

SITE PROTECTION Tenantry Down SSSI & pSAC Porton Down (possibly native) SSSI pSAC Anchor Bottom SSSI Langdon Cliffs SSSI & pSAC Queendown Warren SSSI & pSAC Cobham wood SSSI Colley Hill SSSI Abel Wood CWS Ardley-Middleton Stoney Verge CWS Charlbury (Wigwell NR) LNR Grintley Hill Br CWS Cornbury Park CWS Langland Farm CWS Glyme Farm SSSI Holly Court Bank SSSI Kitesbridge Farm CWS Lower Farm Meadow (Taston) CWS Saltway CWS Stonesfield Common SSSI Whitehill Bridge CWS Dancers End SSSI & pSAC Naphill CWS Box Farm Glos Wildlife Trust reserve Charlton Kings Common SSSI Rogiet SSSI

Agri-environment schemes Agri-environment schemes such as Countryside Stewardship, Environmentally Sensitive Areas, and the proposed new Entry Level and Higher Tier scheme, have played or will play a useful role in providing support for land managers of Salvia pratensis sites. At least one Oxfordshire site is managed under the Countryside Stewardship Scheme and there is no reason why others should not benefit from this approach. This would apply equally well to designated SSSI and unprotected sites. Only the Sussex site appears to fall within an Environmentally Sensitive Area, the South Downs. This site is already an SSSI and managed by the landowner working with English Nature.

Other initiatives Members of the Cotswold Rare Plants Group, and the Rare Plants Group of Ashmolean Natural History Society of Oxfordshire carry out management work and monitoring at all the extant Oxfordshire sites. Scrub control is carried out regularly at Abel Wood and the Saltway sites while soil scarification is undertaken by these groups at the Ardley - Middleton Stoney road verge and Saltway. A new approach to monitoring Salvia pratensis populations, based on the methodology developed by Hegland et al, was piloted at Holly Court Bank by the Project Officer and the Rare Plants Group of the Ashmolen Natural History Society of Oxfordshire in June 2002 (see Annex 1).

24

Figure 6 – Holly Court Bank (Photograph by Miles King).

Plantlife volunteers, known as Flora Guardians, carry out management and monitoring at Naphill, Bucks; Colley Hill, Surrey; and Langdon Cliffs, Kent, the latter also being owned by the National Trust. Kent Wildlife Trust manages the Plantlife reserve Queendown Warren in Kent, while the Berkshire Buckinghamshire and Oxfordshire Wildlife Trust (BBOWT) owns and manages Dancers End in Buckinghamshire. Gloucestershire Wildlife Trust own and manage the Box Farm reserve. Plantlife is also working with the land manager at Cobham Wood, Kent, to clear scrub at this woodland site. Defence Estates manage the Wiltshire site, which is part of the Salisbury Plain Training Area, and the Defence Science Technical and Logistics Agency (DSTL) manage Porton Down, which may be a site at which Salvia pratensis is native. Wigwell in Charlbury is a Local Nature Reserve managed by Charlbury Town Council working with a local farmer who also manages Lower Farm, Taston. Stonesfield Common is managed by the Parish Council, who have recently re-introduced cattle grazing at this SSSI. Other sites are managed by private landowners and managers, who in Oxfordshire work in partnership with the Ashmolean Natural History Society of Oxfordshire Rare Plants Group, the Cotswold Rare Plants Group, the Plantlife Project Officer, Oxfordshire County Council and English Nature. CCW work with Plantlife, the National Museum of Wales and the landowner at the only Welsh site, at Rogiet in Monmouthshire.

Bulking up small populations The populations of several sites, that previously had only one or a handful of plants remaining, have been increased through systematic seed collection and propagation programmes. A single male-sterile plant was left at Naphill in Bucks. Pollen was collected from the Saltway and taken to Naphill where it was used to fertilise the male sterile flowers. A small amount of seed was produced, which was collected and germinated ex- situ. The resulting seedlings were raised by volunteers, and planted out on site in subsequent years. This population is still small but now has at least a chance of survival, with half the genetic material of the original plant having been conserved. A similar exercise took place at Colley Hill, Surrey, although there were hermaphrodite plants

25 extant at this site, so no cross-pollination was needed. The re-introduced plants are currently doing very well. Further bulking-up exercises are planned for Whitehill Bridge, Oxon, Tenantry Down, Wiltshire and Cornbury Park, Oxon. Rogiet is also a good candidate for bulking-up. Further detailed information on this programme can be found in King, 2002.

7.2 EX-SITU MEASURES Seed-banking Seed from the following sites is held at the Millennium Seed Bank, Kew: Box Farm Nature Reserve, Gloucs; Cobham Wood, Kent; Queendown Warren, Kent; Glyme Farm, Oxfordshire; Langland Farm, Oxfordshire; Lower Farm meadow, Taston, Oxfordshire; Holly Court Bank, Oxfordshire; The Saltway, Oxfordshire; Colley Hill, Surrey; Anchor Bottom, Sussex, and Tenantry Down, Wiltshire.

Material held in cultivation Plants of known provenance are in cultivation at Oxford University Botanic Garden. (Davis, 1999).

7.3 RESEARCH DATA The allozyme analysis project run by the Natural History Museum has already been described in section 1.3.

7.4 MONITORING SALVIA PRATENSIS AND THE COMMON MONITORING STANDARD Plant counts have been undertaken in the past (Rich, 1994, 1995). More recently estimates or counts have been made using the Plantlife monitoring form. Results from such counts are difficult to interpret though, given the difficulty in determining exactly what constitutes an individual, particularly when plants are not flowering. The Dutch methodology (in Hegland et al, 2001) took an altogether different approach, identifying 6 life stages based on numbers and size of leaves, and presence of flowering shoots. This methodology does assume that no vegetative reproduction occurs. The methodology was used to establish a baseline against which future monitoring of the Holly Court Bank population could take place. The trial took place in June 2002 with the ANHS/CRPG. 18 1m square quadrats were randomly located within the site and the number of Salvia pratensis plants in each age category counted, as well as data on vegetation composition and structure. If considered successful this methodology could be used elsewhere on Salvia pratensis sites in the UK.

8 References Bijlsma, R., Ouborg, N.J. & Van Treuren, R. (1991). Genetic and phenotypic variation in relation to population size in two plant species: Salvia pratensis and Scabiosa columbaria. Species Conservation: A Population-Biological Approach (eds. A. Seitz & V Loeschke): 89-101. Birkhauser Verlag. Basel. Bijlsma, R., Ouborg, N.J. & Van Treuren, R. (1994). On genetic erosion and population extinction in plants: A case study in Scabiosa columbaria and Salvia pratensis. Conservation genetics (eds V. Loeschke, J. Tomiuk & S. Jain): 255-272. Birkhauser Verlag. Basel. Bures, et al. (2001). Black and Red List of Vascular plants of the Czech Republic – 2000. Príroda, Praha, 18: 1-166, 2001. Cauwood, D. (1981). Salvia pratensis: an ecological survey. BSc thesis, University College Wales, Aberystwyth. Conti, F., Manzi, A. & Pedrotti, F. (1997). Liste Rosse Regionali delle Piante d’Italia. Associazione Italiana per il World Wildlife Fund & Società Botanica Italiana, Camerino, Italy. Druce, G.C. (1886). Flora of Oxfordshire. Parker & Co. London. Ekim et al. (2000), Red Data Book of Turkish Plants. Ankara.

26 French, A. (1875). The distribution of Salvia pratensis Linn. in Oxfordshire. J. Bot. 13: 292-294. Gärdenfors, U. (2000). Rödlistade arter i Sverige 2000 – The 2000 Red List of Swedish Species. ArtDatabanken, SLU, Sweden. Gledhill, D. (1985). The names of plants. Cambridge University Press. Hegland, S.J., Van Leeuwen, M.V. & Oostermeijer, J.G.B. (2001). Population structure of Salvia pratensis in relation to vegetation and management of Dutch dry floodplain grasslands. J. Appl. Ecol. 38: 1277-1289. HM Government. (1981). Wildlife and Countryside Act (1981) and amendments. London, HMSO. Hulten, E. & Fries, M. (1986). Atlas of the North European Vascular Plants north of the Tropic of Cancer. Vol II. Koeltz Scientific Books, Konigstein. Ingelög, T., Andersson, R. & Tjernberg, M. (1993). Red Data Book of the Baltic Region: Part 1 – Lists of threatened vascular plants and vertebrates. Swedish Threatened Plants Unit, Uppsala, Sweden. Kay,Q.O.N. & John, Rosemary F. (1995). The conservation of scarce and declining plant species in lowland Wales: Population genetics, demographic ecology and recommendations for future conservation in 32 species of lowland grassland and related habitats. CCW Science report no. I 10. Bangor, Countryside Council for Wales. Kaźmierczakowa, R. & Zarzycki, K. (2001). Polish Red Data Book of Plants. Polish Academy of Sciences, Cracow, Poland. Kotiranta, H., Uotila, P., Sulkava, S. & Peltonen, S.L. (eds). (1998). Red data book of East Fennoscandia. Ministry of the Environment, Finnish Environment King, M.P. (2002). Meadow clary Salvia pratensis in 2001. Plantlife Report No. 195, London. Lambrick, C.R. (1994). Meadow clary (Salvia pratensis) in Oxfordshire 1994. Ashmolean Natural History Society of Oxfordshire, Rare Plants Group report for Plantlife. Plantlife, London. Olivier, L., Galland, J-P. & Maurin, H. (1995). Livre Rouge de la Flore Menacée de France. Tome I: Espèces Prioritaires. Paris. Ouborg, N.J., and Van Treuren, R. (1995). Variation in fitness-related characters among small and large populations of Salvia pratensis. Evolution, 83: 369-380. Ouborg, N.J, Van Treuren, R. & Van Damme, J.M.M. (1991). The significance of genetic erosion in the process of extinction: 2 Morphological variation and fitness components in populations of varying size of Salvia pratensis L. and Scabiosa columbaria L. Oecologia, 86: 359-367. Perring, F.P, & Walters, S.M. eds. (1962). Atlas of the British Flora, 1st Edition. BSBI, London. Phitos, D. et al. (1995). The Red Data Book of Rare and Threatened Plants of Greece. WWF. Athens. Plantlife. (1998). Report on work undertaken on Meadow clary (Salvia pratensis) in 1997/98. Plantlife Report no. 103. Plantlife, London. Preston, C.D. & Hill, M.O. (1997). The geographical relationships of British and Irish vascular plants. Botanical Journal of the Linnean Society, 124: 1-120. Preston, C.D., Pearman, D.A. & Dines, T.D. (2002). New Atlas of the British & Irish Flora. University Press, Oxford. Rich, T.C.G. (1995). The status of Meadow clary (Salvia pratensis L.) in Britain in 1994. vols. 1 & 2. Plantlife Report no. 44. Plantlife, London. Rich, T.C.G., Hill, D. McNab, C., McVeigh, A. & Davis, R. (1997). Meadow clary in 1996. Plantlife Report no. 76. Plantlife, London. Rich, T.C.G., Lambrick, C. & McNab, C. (1999). Conservation of Britain's Biodiversity: Salvia pratensis L. (Lamiaceae), Meadow Clary. Watsonia, 22: 405-411. Rich, T.C.G. & McNab C., (1996). Meadow clary (Salvia pratensis L.) in 1995. Plantlife Report no. 72. Plantlife, London. Rodwell, J.S. et al, eds. (1991). British plant communities vols. 1-5. Cambridge University Press. Ross, M. D. (1978). The evolution of gynodioecy and subdioecy. Evolution, 32: 174-188.

27 Scott, A. (1989). The ecology and conservation of meadow clary (Salvia pratensis L.) MSc Thesis, University College, London. Stace, C.A. (1991). New Flora of the British Isles. Cambridge, Cambridge University Press. Stevanović, V. (1999). The Red Data Book of Flora of Serbia, 1: Extinct and critically endangered taxa. Belgrade. Stewart, A., Pearman, D.A. & Preston, C.D. (comps & eds). (1994). Scarce Plants in Britain. Peterborough: Joint Nature Conservation Committee. Sturt, N. (1995). Wild clary (Salvia verbenaca) in churchyards. BSBI News, 68: 28-29. Thompson, K., Bakker, J.P. & Bekker, R.M. (1997). Soil seed banks of North West Europe: methodology, longevity and density. CUP. Cambridge. Van Treuren, R., Bijlsma, R., Van Delden, W. & Ouberg, N.J. (1991). The significance of genetic erosion in the process of extinction. 1. Genetic differentiation in Salvia pratensis and Scabiosa columbaria in relation to population size. Heredity, 66: 181-189. Van Treuren, R., Bijlsma, R., Ouberg, N.J. & Van Delden, W. (1993). The effects of population size and plant density on out-crossing rates in locally endangered Salvia pratensis. Evolution, 47:1094-1104. . Wigginton, M. ed. (1999). British Red Data Book 1: Vascular plants. 3rd Edition. JNCC, Peterborough. Wheeler, B. (1998). Salvia pratensis (meadow clary), work on potential restoration sites undertaken during 1998. Plantlife Report no. 110, London. Wheeler, B. (1999). Meadow clary (Salvia pratensis): Progress report on the 1999 project, Volumes 1 & 2. Plantlife Report nos. 137 & 137a, London. Wheeler, B. (2001). Meadow Clary Salvia pratensis in 2000. Plantlife Report no. 176, London.

9 Acknowledgements

Thanks to Miss Jo Dunn of the Cotswolds Rare Plants Group, Dr. Alan Showler, Flora Guardian at Naphill, Buckinghamshire and Dr Derek Hill, Flora Guardian at Colley Hill, Surrey for their valuable comments on the draft of this dossier.

10 Contacts

Plantlife International Andrew Byfield - [email protected] 14 Rollestone Street Or Salisbury Amanda Miller - [email protected] Wiltshire SP1 1DX. Tel: 01722 342730 Miles King [email protected]

Plantlife International’s Back from the Brink (species recovery) programme is supported by: English Nature, Scottish Natural Heritage & the Countryside Council for Wales.

Work on Salvia pratensis is supported by:

28

11 Links ARKive species web page for Salvia pratensis http://www.arkive.org/.

Original draft by Miles King, Edited by Plantlife International First draft dated June 2003 Last revised 29 April 2004

12 Annex 1 - Holly Court Bank Monitoring Form 2002.

Date Quadrat National Grid Reference (10 fig. + error, from GPS) Recorders Slope Aspect Vegetation height (cm) 1. 2. 3. 4. 5. Avge.

SPECIES IN THE QUADRAT Species English DOMIN Species English DOMIN Achillea Yarrow Linum Fairy flax millefolium catharticum Agrimonia Agrimony Lotus Bird’s-foot eupatoria corniculatus trefoil Agrostis canina Common Malva moschata Musk bent grass mallow Anacamptis Pyramidal Medicago lupulina Black pyramidalis orchid medick Arrhenatherum False-oat Mercurialis Dog’s elatius grass perennis mercury Briza media Totter grass Ononis repens Restharrow Brachypodium Tor grass Pastinaca sativa Wild pinnatum parsnip Bromopsis erecta Hairy brome Plantago Ribwort grass lanceolata plantain Carex flacca Glaucous Plantago media Hoary sedge plantain Centaurea nigra Common Poa trivialis Rough knapweed meadow grass Centaurium Common Poterium Salad erythraea centuary sanguisorba burnet Chaerophyllum Rough Potentilla reptans Cinquefoil temulum chervil Cirsium acaule Stemless Primula veris Cowslip thistle Cirsium Woolly Prunella vulgaris Self heal eriophorum thistle Cirsium palustre Marsh Prunus spinosa Blackthorn thistle

29 Corylus avellana Hazel Senecio jacobaea Ragwort Clematis vitalba Traveller’s Sonchus arvensis Field joy sowthistle Conium Hemlock Stachys sylvatica Hedge maculatum woundwort Dactylis Cock’s foot Thymus Wild thyme glomerata polytrichum Fraxinus Ash Veronica Germander excelsior chamaedrys speedwell Galium aparine Goosegrass Viburnum opulus Wayfaring tree Geranium molle Dove’s foot Viola hirta Hairy violet crane’s bill Galium verum Ladies Bare ground bedstraw Glechoma Ground ivy Moss hederacea Helianthemum Rockrose Litter nummularium Heracleum Hogweed sphondylium Hypericum Hairy St hirsutum John’s wort

Hypericum Perforate St perforatum John’s wort Lapsana Nipplewort communis Leontodon Rough hispidus hawkweed Leucanthemum Oxeye daisy vulgare

DOMIN Scores: 1=<4% with few individuals; 2=<4% with several individuals; 3=<4% with many individuals; 4=4-10%; 5=11-25%; 6=26-33%; 7=34-50%; 8=51-75%; 9=76-90%; 10=91-100%. Vertical projection of all live above ground parts of the plants.

Number of Salvia pratensis (meadow clary) plants in quadrat Seedlings Juveniles Immature Vegetative Small Large plants Adults Generative Generative Adults Adults

30