Pulsatilla vulgaris (L.) Mill.

Pasque Flower, Pulsatilla vulgaris Miller

RANUNCULACEAE
SYN.: Anemone pulsatilla L.
Status: All British populations belong to subsp. vulgaris which is classified as vulnerable
(IUCN Criterion A2ac; Cheffings & Farrell, 2005), and listed as a UK BAP Priority Species in
2007. It is currently confined to 18 sites in 19 10km squares in England. In this account
Pulsatilla vulgaris refers to subsp. vulgaris unless otherwise stated.

In partnership with:

Contents
1
1.1
1.2
1.3
1.4

Morphology, identification, taxonomy and genetics

Morphology and identification
Taxonomic considerations
Genetic implications
Medicinal properties

2
2.1
2.2
2.3
2.3.1
2.3.1.1
2.3.1.2
2.3.2

Distribution and current status

World
Europe
United Kingdom
England
Native populations
Introductions
Northern Ireland, Scotland & Wales

3
3.1
3.1.1
3.1.2
3.1.3
3.1.4
3.1.5
3.1.6
3.1.7
3.1.8
3.1.9

Ecology and life cycle

Life cycle and phenology
Flowering phenology
Flower biology
Pollination
Seed production
Seed viability and germination
Seed dispersal
Regeneration
Response to competition
Herbivory, parasites and disease

4
4.1
4.2
4.3

Habitat requirements
The landscape perspective
Communities & vegetation
Summary of habitat requirements

Management implications

Threats/factors leading to loss or decline or limiting recovery

7
7.1
7.2
7.3
7.4

Current conservation measures

In situ Measures
Ex situ Measures
Research Data
Monitoring and the Common Monitoring Standard

8
9
10
11
13
14

References
Contacts
Links
Annex 1 site descriptions
Annex 2 changes in population size, 1960-2006
Annex 3 associates

Morphology, identification, taxonomy and genetics

1.1
Morphology and identification
Hemicryptophyte; 2-15 cm, extending to ca. 45 cm in fruit. Rhizome obliquely erect or
vertical, branching, black, 2-17 mm thick, eventually decaying to form new plants. New leaf
rosettes arising from the branches of the rhizome by growth of lateral buds, producing groups
of leaf rosettes connected below ground by an anastomosing rhizome system. The remains of
old leaf bases protect the overwintering buds at the apex of the rhizome. Basal leaves up to
11 from each rhizome apex, bipinnatisect, silky-hairy and forming a rosette. Ascending,
developing during anthesis and withering in the autumn (occasionally over-wintering). Petiole
to 21 cm, blade to 11 7 cm, 2- to 4- pinnate. Cauline leaves 2(-4), sessile, 1.7-5.3 cm,
smaller and more deeply divided into linear segments (only slightly lobed), white-hairy and
reaching the perianth during most of the flowering period. All leaves silky-hairy at least
initially, the basal with pubescent petioles. On emergence the silky-villose leaves are tightly
curled around the apical growing points. Flowers solitary, terminal, erect at first but drooping
after a few days, and becoming bell-shaped (campanulate) (Fig. 1a). Buds develop during late
summer and become dormant over the winter months when they are protected by a dense
covering of silky-villose hairs and the remains of the previous years leaves. Pedicel up to 8 cm
and bent over less than 90 at anthesis, elongating to 25 cm in fruit. Perianth-segments 6, 1642 4-17 mm, deep violet-purple in British plants, more blue to violet-blue on the Continent,
darker on the inside, paler and silky on the outside (rarely white or pink), subequal, only
slightly curving outwards apically. Petals fading and becoming bleached after about a week.
Stamens numerous, 50-120 per flower, crowded, the outermost shorter, sterile, club-shaped
and secreting nectar. Staminodes and fertile stamens not more than half as long as perianth
segments, 15-25 mm; filaments stout at the base, tapering to the adnately attached anthers,
which are bright golden-yellow before fading. Styles purple, borne on a flattened, conical,
receptacle, 30-90 per flower, each with a single functional ovule. Fruit is an achene, 2-3 mm
long, with a single embryo, covered with simple, silky hairs and with a persistent feathery
style 3.5-5 cm long. The internode between the stem leaves and flower elongates considerably
after flowering, increasing the height at which the achenes are released (Fig. 1b). Flowers 13(-12) per plant, actinomorphic, hermaphrodite and protogynous in England, pollinated by a
variety of insects, especially bees (Apidae: Apis, Bombus).
1.2
Taxonomic considerations
The common name Pasque Flower has several derivations; one is reputedly from the Latin
pascha, meaning Easter, as Easter eggs were often stained by rubbing the eggs with the
flowers and leaves for celebration. Originally the flower was known by the French as the
passe-fleur, and then changed by Gerard (who considered it to be a worthy addition to the
herbaceous border) to the Pasque Flower on account of the flowers appearing around Easter
time (Smith, 1996).

Figure 1 Pulsatilla vulgaris: (a) flowers, Knocking Hoe, Bedfordshire, photo by K.J Walker;
(b) elongated scape and achenes, Devils Dyke, Cambridgeshire, photo by K.J. Walker; (c)
illustration by Stella Ross-Craig (1948).
Pulsatilla vulgaris was described by Carl Linnaeus (1753) as Anemone pulsatilla in the first
volume of his Species plantarum (p. 539)1. The Scottish botanist Philip Miller (1768) changed
the name to Pulsatilla vulgaris in the eight edition of The gardeners dictionary and this was
subsequently retained in Aichele & Schweglers (1957) monograph of the genus which
described a number of taxa within the Pulsatilla vulgaris group. As no holotype exists a
lectotype has recently been designated from Linnaeus herbarium (Herb Linn. No. 710.5;
Jarvis et al., 2005). This herbarium sheet contains a single flowering and two fruiting stems
but has no collection details and thus its origin is unknown.2
Pulsatilla is a small genus within the large, primitive family Ranunculacae. The division of the
Linnaean concept of Anemone into three genera, Anemone sensu stricto, Hepatica and
1
2

seehttp://www.biodiversitylibrary.org/page/358106#551
seehttp://www.linneanonline.org/5012/

Pulsatilla is now generally accepted based on both morphology and chloroplast DNA (e.g.
Hantula et al., 1989). Morphologically Pulsatilla taxa are distinguished by the styles
elongating greatly and becoming feathery in fruit and the presence of nectar-secreting
staminodes (both absent in Anemone sensu stricto; Akeroyd, 1993). The current worldwide
classification scheme of the Angiosperm Phylogeny Group (APG III) considers the
Ranunculaceae to be among the most basal of the derived Eudicots clade (Hill & Preston,
2002):
APG Clade:
EUDICOTS
APG Order:
Ranunculales
APG Family:
Ranunculaceae
Subfamily:
Ranunculoideae
Genus:
Pulsatilla
Species:
Pulsatilla vulgaris Mill.
The genus Pulsatilla contains around 38 species worldwide all of which occur in the Northern
Hemisphere, mainly in Europe and Asia with two species in North America. Nine species occur
in Europe (Akeroyd, 1993). Five of these are restricted to the montane regions of southwest,
central-south and southeast Europe (P. alba, P. alpina, P. halleri, P. montana, P. rubra); the
remaining four taxa are more widespread in the lowlands of northern, central and eastern
Europe extending from the Atlantic to Eurasia (P. patens, P. pratensis, P. vernalis, P.
vulgaris).
Pulsatilla vulgaris is told from all other Pulsatilla species by its sessile cauline leaves, deeply
pinnatisect basal leaves with 7-9 primary segments that wither in the autumn and erect
flowers (nodding in anthesis). It is a very variable species for which a number of poorly
defined infraspecific taxa have been described in the past. These, however, appear to
represent the more distinct of the numerous isolated populations and probably represent a
dissected continuous type of variation caused by post-glacial climatic changes and the
intolerance of the species to ploughing, shade and bad drainage (Akeroyd, 1993).
Consequently infraspecific taxa do not often appear to fall into the geographical pattern
characteristic of subspecies. The smaller and more isolated populations, particularly towards
the edge of the range of the species, are fairly homogeneous (as in Britain), but in other areas
there is considerable variation within populations. While it is likely that most populations are
separable from one another, there is a large amount of overlap between them and
intermediates occur frequently.
Today three subspecies are usually accepted. Subspecies vulgaris, to which all English
populations belong, is the most widespread taxon extending from 61 N in Sweden to c. 45 N
in the Bordeau region of France and from Gloucestershire in England to near to Poznan in
western Poland where it is replaced by subsp. grandis (Lindell, 1998). Subsp. vulgaris is
distinguished by the greater degree of lobing in the leaves (>100; <40 in subsp. grandis),
narrower perianth segments and presence of leaves at flowering (occurring after the flowers
in subsp. grandis; Akeroyd, 1993). A third taxon, subsp. gotlandica, confined to limestone
pavement on the Swedish island of Gotland (Jonsell, 2001), is dubiously distinct and placed
within subsp. grandis by some authorities (e.g. Akeroyd, 1993). On the whole British
populations exhibit little variation although plants on the Oolitic limestone are generally
larger and produce more flowers (Wells & Barling, 1971). Under heavy grazing plants also tend
to be much smaller and more compact (Bailey, 1996).

1.3
Genetic implications
The chromosome base-number of the genus is 8. Pulsatilla vulgaris is tetraploid (2n = 32) and
may have arisen following hybridization between P. patens (2n = 16) and P. pratensis (2n = 16)
(Bcher, 1934). Spontaneous hybrids between subsp. vulgaris and P. pratensis and P. patens
are known from Northern Europe (Akeroyd, 1993) and work on Swedish populations has
indicated that hybridization barriers are weak between subspecies of Pulsatilla vulgaris, and
between P. vulgaris and P. halleri (Lindell, 1998).
Bailey (1996) found a low percentage of polymorphic loci in five sites across the geographic
range of Pulsatilla vulgaris in England (Barton Hills, Knocking Hoe, Rodborough Common,
Devils Dyke, Therfield Heath), suggesting that overall genetic variation is low, although there
were significant differences for individual primers between some of the sites examined.
Further work is needed to more fully assess these differences, and in relation to overall
fitness, population size and degrees of geographical isolation.
Similar work has been undertaken in Germany on 11 populations ranging from 50 plants to
7300 plants (Hensen et al., 2005) and found a significant positive relationship between genetic
diversity and both population size and seed mass (per population). Genetic diversity was
significantly lower in smaller populations than in larger ones, suggesting that genetic drift
(random change of allele frequency within a population) has been the main cause of the loss
of diversity as populations have become smaller and fragmented following rapid land-use
change (i.e. the chance of alleles being lost from one generation to another has been greater
in smaller populations).
Hensen et al. (2005) also found a high level of within population variability which was
attributed to life-history traits likely to preserve variability, namely allogamous pollination, a
long life span and vegetative reproduction. Interestingly there was only weak, albeit
significant differentiation between populations which was correlated to the geographical
distance. This suggests that genetic drift has been the major force driving differentiation
between German populations as the former larger population became fragmented. The
weakness of this relationship suggests that the loss of genetic variability through drift has
been, to some extent, balanced by gene flow between closer populations. Since seed dispersal
in Pulsatilla vulgaris is limited it was concluded that the foraging radius of pollinators,
currently thought to be in the order of several kilometres for honeybees (Steffan-Dewenter &
Kuhn, 2003) and bumblebees (Osborne et al., 2008), is critical in offsetting genetic drift as
populations become smaller and more isolated. However, even if pollinators forage over
several kilometres it seems very unlikely that they would travel between English populations
given their small size and degree of isolation.
1.4
Medicinal properties
Pulsatilla vulgaris is unpalatable and poisonous to humans and animals due to the presence of
the glycoside ranunculin in the leaves and roots which is converted to anemonine when the
plant is dried (Plants for a Future, 2010). Small doses are taken internally in the treatment of
pre-menstrual syndrome, inflammations of the reproductive organs, tension headaches,
neuralgia, insomnia, hyperactivity, bacterial skin infections, septicaemia, spasmodic coughs in
asthma, whooping cough and bronchitis. Externally, it is also used to treat eye conditions such
as diseases of the retina, senile cataract and glaucoma. In homeopathy, extracts are used to
treat measles as well as minor complaints such as nettle rash, toothache, earache and bilious
indigestion (Plants for a Future, 2010).

Distribution and current status

2.1
World
Pulsatilla vulgaris subsp. vulgaris is endemic to Western and Central Europe and is not known
to have been introduced to other parts of the world.
2.2
Europe
Pulsatilla vulgaris belongs to the European Temperate element (Preston & Hill, 1997) with the
core of its distribution in the lowlands of western and central Europe (Fig. 2).
Its current status in Europe is given in Table 1. France appears to be the stronghold for the
plant in Europe and consequently it was not listed as a priority species for conservation in the
most recent assessment of the French flora (unlike P. vernalis; Olivier et al., 1995). In
comparison it appears to be threatened throughout the rest of its range. In Denmark,
Germany and Southern Sweden it is still relatively widespread, but appears to have declined,
especially in Sweden, where it is now classified as Vulnerable using IUCN threat criteria (A2ac;
Grdenfors, 2010), and Germany, where its populations are now small and highly fragmented
(Hensen et al., 2005). Similar trends have been reported for Austria, where only around 2000
plants now survive in 23 sites (Franz, 2005), and Switzerland where it is very rare (Pfeifer et
al., 2002) and classed as Endangered under IUCN threat criteria (Moser et al., 2002). In
Belgium it is confined to two small areas in the southern province of Wallonia, where its
habitat (limestone grassland) is under pressure from extensive tree planting (Quentin Groom,
pers. comm.). In Luxembourg it is classified as Endangered under IUCN criteria having
declined from 28 to 5 localities with population sizes now ranging from two to more than 7000
genets (Colling, 2005). It appears to be extinct in Finland, where it has not been seen since
the 1930s (Rassi et al., 2001) and Holland. Its status in Poland is unknown although recent
maps (e.g. Fig. 2) show it to be very rare. Recent records for Norway all appear to be of
casual escapes (Jonsell, 2001).

Figure 2 - Distribution map of Pulsatilla vulgaris subsp. vulgaris in Northwest and Central
Europe, reproduced from Hensen et al. (2005). Small filled circles are individual site records
from 1950 onwards. Small empty circles are individual site records older than 1950 or
occurrences reported as extinct. Atlas Florae Europaeae data from Jalas & Suominen (1989)
are also shown (large filled grey circles) to indicate 50 50 km squares from which no
individual occurrence data were available. The doubtful two easternmost records from Poland
are given by EUNIS (2004) but not by Wojtowicz (2001).
Table 1 The status of Pulsatilla vulgaris in Europe.
Country
IUCN Status
Austria
?
Declined to 2000 plants in 23 sites
Belgium
?
Very rare and declining
Denmark
?
Unknown
Finland
EW Formerly one site
France
?
Not threatened?
Germany
?
Widespread but declining
Holland
EW Extinct
Luxembourg
EN
Declined from 28 to 5 localities
Norway
Casual escape
Poland
?
Unknown
Sweden
VU
Ca.30% reduction in past 100 years
Switzerland
EN
Marked decline

Reference
Franz, 2005
Rassi et al., 2001
Olivier et al., 1995
Hensen et al., 2005
Colling, 2005
Jonsell, 2001
Grdenfors, 2010
Pfeifer et al., 2002; Moser et

UK

VU

Declined from ca.130 to 18

localities

al., 2002
Cheffings & Farrell, 2005

2.3
United Kingdom
2.3.1 England
2.3.1.1 Native populations
Since its discovery near to Oxford in 1551 (Druce, 1886) Pulsatilla vulgaris has been recorded
from around 130 sites in 19 English vice-counties (Table 2) but is now confined to 18 sites in
just ten vice-counties (Table 3). Former strongholds include the Cretaceous chalk downland
and ancient earthworks of Bedfordshire, Berkshire, Buckinghamshire, Cambridgeshire,
Hertfordshire and Oxfordshire and the Jurassic limestone grasslands of Gloucestershire,
Northamptonshire and Lincolnshire (Fig. 3). It was also formerly widespread in Magnesian
limestone grasslands in northern England, extending as far north as Piercebridge in County
Durham (Baker, 1906) but it is now restricted to a single site near to Wakefield (Abbott,
2005). Small clusters of populations also occurred in North Essex, Leicestershire, West Norfolk
and West Suffolk but it has not been seen in any of these counties since 1888, 1992, 1914 and
1978 respectively (Table 4).
Table 2 The number of Pulsatilla vulgaris populations recorded in English vice-counties.
Extinct.
Tota
%
Population size (2000-10 period)
l
extinc
Extinc
11201VC Vice-county name
1-10
51-200
>1000
no.
t
t
50
1000
19 North Essex
2
2
100
20 Hertfordshire
7
1
8
88
22 Berkshire
12
1
13
92
23 Oxfordshire
4
4
100
24 Buckinghamshire
2
1
3
67
26 West Suffolk
3
3
100
28 West Norfolk
1
1
100
29 Cambridgeshire
7
1
8
88
30 Bedfordshire
2
2
2
6
33
31 Huntingdonshire
1
1
100
32 Northamptonshire
7
1
8
88
East
33
17
1
2
1
1
22
77
Gloucestershire
West
34
4
1
5
80
Gloucestershire
53 South Lincolnshire
16
1
1
18
89
54 North Lincolnshire
5
5
100
55 Leicestershire
5
5
100
South-west
63
6
6
100
Yorkshire
64 Mid-west Yorkshire
7
1
8
88
66 Durham
1
1
100
Total
109
6
3
3
1
5
127
86

Of the 10 vice-counties with surviving populations, seven only have single colonies.
Bedfordshire and East Gloucestershire have the most with 4 and 5 populations respectively.
Half the English populations support less than 100 plants, including one site (Ledsham, Midwest Yorkshire) with a single individual (known since 1984). Only five sites (28%) support in
excess of 1000 plants (Fig 4a). Although total population sizes are difficult to estimate, recent
counts on these sites suggest the following totals: 20,000 at Barnack Hill and Holes, 75,000 at
Barnsley Wold, 5,000 at Barton Hills, 1,000 at Knocking Hoe and 60,000 at Therfield Heath
(Tarpey, 1999; see Annex 2 for details). These five sites account for 99.3% of the total British
population. Of the eighteen surviving populations 4 appear to be declining and are threatened
with extinction, 9 are stable and 5 appear to be increasing due to improvements in grazing
regimes (Table 3; see Annex 2 for details).

Figure 3 The hectad

distribution of Pulsatilla
vulgaris in Britain and Ireland.
Solid black circles are hectads
where P. vulgaris has been
recorded since 2000; solid grey
circles, recorded 1987-1999;
open circles, recorded before
1987; X, introduced.
Historical losses appear to have been high in all vice-counties especially East Gloucestershire,
South Lincolnshire and Berkshire where 77%, 89% and 92% of populations have gone extinct
(Table 2). The rate of loss appears to have been increased consistently since 1750 from around
1.8 populations per decade to 6.0 since 1950 (Fig. 4b) although these figures presumably
underestimate rates for earlier periods as fewer extinction events are likely to have been
detected by eighteenth century botanists (as fewer populations were then known).
Consequently the total number of populations has displayed a consistent downward trend (Fig.
4c). However, since 1960, the first period when we have detailed information on the size of

10

English populations, the overall number of plants has increased considerably from an
estimated 60,000 to over 160,000 individuals largely due to improvement in grazing regimes
on four of the best sites (Barnack Hills and Holes, Barnsley Wold, Barton Hills, Therfield
Heath; Fig. 4d). Despite these losses new populations continue to be found including a small
population of around 15 plants in Swinstead Valley (South Lincolnshire) discovered in 2009
(Jefferson & Walker, 2010).
Table 3 Details of extant populations of Pulsatilla vulgaris in England.
Size/
trend

Site

VC

10-km

Geology/management

Therfield Heath

20

TL33

Aston Upthorpe Down

22

SU58

B(r)

Steps Hill

24

Devil's Dyke

29

Chalk, some areas grazed/mown

Barton Hills
Ravensburgh Castle
Deacon Hill
Knocking Hoe

30
30
30
30

SP91
TL56/
66
TL03
TL02
TL12
TL13

Chalk, winter grazing

Chalk, ungrazed exclosure, occ.
mown
Chalk, intermittent grazing

Chalk, grazed excluding Jan-Mar

Chalk, ungrazed
Chalk, some areas grazed
Chalk, grazed
Limestone, grazed excluding MarBarnack Hills and Holes 32
TF00
Sep
Limestone, grazed excluding MarBarnsley Wold Warren
33
SP00
May
Beaumonts Hay
33
SP12 Limestone, irregular grazing
Limestone, grazed excluding AprBourton Down
33
SP13
Aug
Limestone, grazed excluding MarHornsleasow Roughs
33
SP15
May
Taylors Hill
33
SP01 Limestone, grazed
Rodborough Common
34
SO80 Limestone, light mowing/grazing
Ancaster Valley
53
SK94 Limestone, winter grazed
Swinstead Valley
53
TF02 Limestone, summer grazed
Ledsham
64
SE43
Limestone, winter grazed
Code for population sizes and trends: A, 1-10; B, 11-100; C, 101-1000; D,
10000-100000; (r), includes reintroduced plants. , population increase;
decline; population stable.

Cons.
Design
.
SSSI
SSSI

B
D
B
A
D
E
E
A
C
C

SSSI
SSSI
NNR
SSSI
SSSI
NNR
NNR
SSSI
SSSI
SSSI
SSSI

C
SSSI
B
SSSI
B(r)
SSSI
A
SSSI
A(r)
1001-10000; E,
, population

Table 4 Vice-counties in which Pulsatilla vulgaris is now extinct with reason for loss
VC Vice-county
Last recorded site
Last
Probable reason for loss
name
record
19 North Essex
Bartlow Hill
1888
Scrubbed-over
23 Oxfordshire
Upton Down
1883
Ploughed-up
26 West Suffolk
Newmarket Heath
1978
Grassland altered by horse
training
28 West Norfolk
Sporle & Tulip Hills
1914
Scrubbed-over
31 Huntingdonshire
Stibbington
1926
Possibly an error for sites in
VC32?

11

54
55
63
66

North
Lincolnshire
Leicestershire
South-west
Yorkshire
Durham

Broughton Far
Wood
Shacklewell Hollow
Smeaton Crag
Pastures
Cliffe Wood

1969
1992
1870

Scrubbed over; last plant dugup

Dumping of rubble
Quarrying/mining

1906

Landscaping of parkland

Although there is no fossil evidence of the occurrence of Pulsatilla vulgaris in Europe it was
probably much more widespread during the early post-glacial in dry, steppe grassland before
retreating to open refugia, with shallow soils and low competition, following the spread of
deciduous woodland (Hensen et al., 2005). Woodland clearance from the Neolithic onwards,
however, is speculated to have created much new habitat enabling spread from refugia,
although barriers to migration would have been present in some regions. For example,
woodland is thought to have restricted its colonisation of the southernmost chalk in England
(Pigott & Walters, 1954; Rose, 1957) although its absence from Late Glacial refugia such as
the Avon Gorge and centres of Neolithic farming activity, such as Salisbury Plain, suggests a
much more complex phytogeographical history in Britain, or at least one that is impossible to
reconstruct from modern records alone.

Figure 4 The number and size of populations of Pulsatilla vulgaris in England: (a) size of
extant populations; (b) number of populations lost per decade; (b) the number of populations

12

surviving in each year, and (c) the total number of populations and plants recorded since
1960. In (c) all populations were assumed to have been present in 1750 even if they were only
discovered much later.
2.3.1.2 Introductions
There are around a dozen deliberate or casual introductions known in England (Table 5), with
most resulting from tipping of garden waste by roadsides or in old quarries. There have also
been six deliberate introductions to either re-establish populations on sites where it become
extinct (Copper Hill Quarry, Southorpe Paddock), bolster extant populations (Aston Upthorpe,
Ancaster, Ledsham) or create entirely new populations for the purposes of research
(Hartslock) (see Table 11 for details). The most contentious site, however, is the small
population that straddles the border of Dorset and South Wiltshire on Martin Down which
interestingly appears to have increased since its discovery in 1983 (see
http://www.hantsplants.org.uk). Both Bowen (2000) and Brewis et al. (1996) suggest it might
be native given the typical nature of the habitat (ancient species-rich grassland on an
earthwork). However, in view of its isolation from other native populations and late discovery,
which is surprising given its close proximity to a path, it is probably best treated as an
introduction unless convincing evidence is produced for it being of natural occurrence
(Preston et al., 2002; Edwards & Pearman, 2004).
Table 5 Known casual records of Pulsatilla vulgaris in England. Details of translocations are
given in Table 11.
VC
VC name
Details
7
North Wiltshire
Planted on Morgans Hill in 1939 from where it possibly colonised
Cherhill Down by 1941 (Grose, 1957) but this seems unlikely.
8/9
South
Known from Martin Down since c.1983 where two small
Wiltshire/Dorset
populations grow on either side of the vice-county boundary on
ancient species-rich chalk banks (SU0518, SU0418). Likely to
have been planted given the distance to the nearest native
populations (Edwards & Pearman, 2004).
13/1 West
&
East No details; presumed planted.
4
Sussex
20
Hertfordshire
Discovered on road verges at two sites (TL1226; TL2426) where
it was presumed to have been planted (T James, pers. comm.).
21
Middlesex
No details; presumed planted.
24
Buckinghamshire Discovered in 1998 (SP8405) but assumed to have been planted.
29
Cambridgeshire
In 1994 a few plants originating from a garden were planted on
an area of downland reverting from arable (L Evans, pers.
comm.).
30
Bedfordshire
Recorded in a quarry where it had presumably been planted (C
Boon, pers. comm.).
2.3.2 Northern Ireland, Scotland & Wales
Pulsatilla vulgaris does not occur in Northern Ireland, Scotland or Wales.
3

Ecology & life cycle

3.2
Life cycle and phenology
Pulsatilla vulgaris is a monoecious, long-lived, polycarpic hemicryptophyte which appears to
reproduce mainly vegetatively by growth of adventitious buds on the branched rhizome. These

13

produce several daughter rosettes close to the parent plant each year. Although new flower
buds are produced during late summer the whole plant is dormant during the winter months
(August to March). The tightly furled leaves appear in March and unfurl at or after flowering
and wither in the autumn or after the first frosts.
Plants of Pulsatilla vulgaris are very difficult to age due to the production of new ramets from
the branched rhizome (type C perennials sensu Grubb, 1990). However, observations in
England suggest that large plants with rhizomes 15-20 mm long may be at least 20 years old
(Wells & Barling, 1971) although individual genets are potentially much longer-lived with halflife of more than 50 years (Grubb, 1990). Plants grown from seed will flower in pots in 1.5-2.5
years but in the wild this is likely to be 4-5 years.
3.1.1 Flowering phenology
Temperature appears to be a very important cue that regulates the timing and rate of flower
and fruit development (Widn & Lindell, 1996). Flowering commences when temperatures rise
above 12-15C (Kratochwil, 1988; Widn & Lindell, 1996), usually in the first week of April,
reaching a peak between 4-5 weeks later, and continues to the end of June (Fig. 4; Wells
(1994) gives the main flowering period as the 6 April to 20 May). However, the onset of
flowering and flower production does vary from year to year depending on climatic conditions.
For example, flowering and fruit maturation is known to be higher in warmer years (e.g. 1988;
Widn & Lindell, 1996) and for cultivated plants exposed to full sun light either in the
greenhouse (Widn & Lindell, 1996) or under experimental shading conditions (Fig. 5; Walker
et al., in press). In large populations, however, the behaviour of individuals is variable, some
plants flowering in successive years whilst others remain vegetative (Wells & Barling, 1971). In
addition, drought in one summer may affect flowering the following year as primordia are
initiated in late summer (Rich, 1997).

Number of site records

60
50
40
30
20
10
0
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21

Weeks from 1st March

Figure 4 Phenology of Pulsatilla vulgaris in England based on the number of historical

records for sites held in the Vascular Plants Database, regardless of location, year & recorder.
3.1.2 Flower biology
Each plant produces 1-3(-12) bell-shaped flowers with numerous carpels with long purple
styles (30-150) and bright yellow stamens, the outer smaller, sterile and secreting nectar (Fig.
1a). These are produced on very short stalks and face the sky for about 48 hours after first
opening, only closing at night or during rain (Wells & Barling, 1971). Flowers are
hermaphrodite and markedly protogynous with the female reproduction organs reaching

14

maturation before the male (andro- and gyno-monoecious and dioecous forms have been
reported from the continent but are likely to be rare). Both stamens and styles continue to
grow during anthesis increasing the distance between the anthers and stigmatic papillae
(Jonsson et al., 1991). On cultivated plants the obligate female phase (the time from the
beginning of anthesis to the opening of the first pollen sac) lasts just over a day whereas
pollen is produced for about 4 days and remains viable for 7-8 days (Jonsson et al., 1991). The
obligate female phase appears to be much shorter late in the flowering season and is directly
correlated with an increase in temperature (Kratochwil, 1988; Jonsson et al., 1991). In most
populations the length of the obligate female phase is likely to be sufficient for some crosspollination to take place since the great proportion of flowers receive pollen before the onset
of the male phase (Jonsson et al., 1991). However, geitonogamous pollination is probably also
common as most populations include plants with 2 or more flowers, including flowers on
identical (daughter) genets growing close-by (Warden, 2001).

Figure 5 Effects of experimental shading on (a) the % survival and (b) flowering
performance of Pulsatilla vulgaris grown at Monks Wood Experimental Station,
Cambridgeshire, 1970-80 (Walker et al., in press). Plants were planted in a chalk subsoil to a
depth of 1 m and exposed to three levels of shading (dense, medium, light) simulated using
black gauze screens erected over each plot. Each plot consisted of 12 regularly spaced plants
25 cm apart. Three replicates of each treatment and a control (no shade) were arranged in a
randomised latin square and the survival of transplants and flowering performance recorded in
April and May of each year. The plots were fenced to exclude rabbits and weeded to remove
competition.
3.1.3 Pollination
The flowers of Pulsatilla vulgaris are insect-pollinated mainly by aculeate Hymenoptera and
bees (Apidae: Apis, Bombus). Many other insects have been recorded visiting flowers in
England and on the continent but appear not to contribute significantly to pollination (e.g.
flies, beetles, Lepidoptera, non-Apoidea Hymenoptera; Wells & Barling, 1971; Kratochwil,
1988; Warden, 2001). Experiments with exclusion of pollinators showed that Pulsatilla vulgaris
is mainly an outbreeding plant due to the absence of spontaneous selfing (index of selfincompatability (ISI) <0.1, n = 18; ISI cited in Henssen et al., 2005). However, work on Swedish

15

populations has shown that dichogamy is incomplete, since artificial selfing produces a small
amount of fertile seed although selfed flowers produced fewer seeds than those that cross
(Jonsson et al., 1991; Lindell, 1998). Seed set from self-pollination is variable with reported
values of 18% in England (Warden, 2001) and 31-50% in Sweden (Lindell, 1998). Insects seem to
be necessary even for self-fertilisation as enclosed flowers do not set seed automatically
(Wells & Barling, 1971; Kratochwil, 1988; Widn & Lindell, 1996) and early flowering plants
and those in small populations have lower seed-set due to pollinator-limitation (Widn &
Lindell, 1996).
3.1.4 Seed production
Between 30-90 carpels are borne on a flattened, conical receptacle, each containing a single
embryo which when fertilised develops into an achene 2-3 mm long with a persistent feathery
style (3-5 cm) borne on stems that elongate after anthesis (Fig. 1b). Achenes ripen 4-5 weeks
after the flowers open and fruits may be ripe from mid May to mid June. There is considerable
variation in the number of achenes produced per flower and total achene production per
plant. For example, Wells & Barling (1971) found 23-74 achenes per flower in five English
sites, with an average of 46, though not all may have been fertile (Rich, 1997) whereas
Warden (2001) found an average of 62 at Aston Upthorpe in Berkshire of which ca.30-35 were
viable (filled). In Sweden Widn & Lindell (1996) found a similar average of 18 and 28 viable
(undamaged) seeds per flower at two sites excluding achenes damaged by seed-boring insects.
In Germany Kratochwil (1988) found fruit set to exceed 70% over four years, even though only
a few hours of favourable conditions occurred in most years. However, intermediate flowering
plants appear to be more fecund than earlier or late flowers, because early flowers encounter
a paucity of pollen vectors and suffer from early frosts whereas late flowers suffer from
greater predation and may have to compete with developing leaves for resources (Widn &
Lindell, 1996; Warden, 2001). Later-flowering plants also appear to develop seeds more
rapidly due to higher temperatures (Widn & Lindell, 1996). Flowers in most populations
produce infertile achenes containing no embryo, usually towards the centre of the flower;
these weigh about 0.6 mg whereas fertile achenes weigh 3 mg (Wells & Barling, 1971)
although seeds containing no embryo can sometimes weigh more than those that are viable
(Bailey, 1996). The cause of this infertility is unknown but is presumably caused by cold,
inclement weather at the time the flower is produced and the abundance of flying insects.
(e.g. 1996 in England; Bailey, 1996).
3.1.5 Seed viability and germination
The viability of fresh seeds is generally high with reported germination values at room
temperature of 47% for Swedish material (Widn & Lindell, 1996) and 59% (Bailey, 1996), 75%
(Warden, 2001) and 90% (Wells & Barling, 1971) for English material. Under more controlled
laboratory conditions 80% germination was been achieved on 1% Agar solution at 20C, 8/16
hours light and dark and 95% at 21C, 12/12 hours light and dark (RBG Kew, Wakehurst Place).
In comparison, only 16% of seeds germinated in a field germination trial increasing to 29%
when seeds were covered with 3 mm of soil (Wells & Barling, 1971).
Tests on the viability of seed stored at room temperature have shown variable declines with
age. Wells & Barling (1971) found between 15-65% germination after 8-9 months with no
germination occurring after 2.5 years. However, only a small decreases in viability has been
reported for seed stored at 4C for one year (Warden, 2001) and dried to low moisture
contents and stored for 13 years (RBG Kew, Wakehurst Place). Germination appears to be
unaffected by light levels or storage at temperatures as low as -7C (Wells & Barling, 1971).

16

3.1.6 Seed dispersal

Achenes of Pulsatilla vulgaris are adapted for wind-dispersal by modification of the style into
a feathery pappus and the elongation of the fruiting stems (by a process of cell elongation),
increasing the height at which the ripe seeds are released to above the general level of the
surrounding vegetation (Fig. 1b). For example Wells & Barling (1971) give a mean scape height
at fruiting of 18.4 cm for 50 plants at Barnsley Wold in 1966 as opposed to 4 cm for the
surrounding grassland. Despite these adaptations, long-distance dispersal is most probably
absent or very rare (Tackenberg et al., 2003). Observations in England suggest that seeds
disperse less than 20 cm from parent plants often because the entire head remains intact or
the feathery tail becomes entangled in the surrounding vegetation (Wells & Barling, 1971;
Warden, 2001). In addition simulations of aemochorous dispersal showed that 90% of the seeds
land within the mother population with only 0.05% of seeds being dispersed over a distance of
more than 100m (Tackenberg, 2001). According to Rder & Kiehl (2006) Pulsatilla seeds also
have a high potential for epizoochory, including dispersal on machinery, which might be a
more important strategy for long distance dispersal in modern landscapes. Establishment of
new populations in this way is likely to be rare, but may have occurred at one site in
Cambridgeshire where a new population was discovered 300 m from the nearest population
(Leslie, 2004; see Devils Dyke, Annex 1). Dispersal by machinery is the most likely vector at
this site as the new colony occurs on a mown road verge inaccessible to livestock.
3.1.7 Regeneration
Despite the production of viable, highly germinable seeds, establishment of new individuals
from seed appears to be rare (seedlings observed at only one site by the author) and most
regeneration, is probably achieved by the disintegration of the branched rootstock (Wells &
Barling, 1971). Plants reproduce vegetatively by growth of adventitious buds on the rhizome 2
cm or more below ground level, producing a small rosette of leaves close to the parent plant.
Two or three new rosettes may be produced each year and excavation of groups of plants has
revealed as many as seven distinct rosettes from a single rootstock, the younger plants being
clustered around the parent plant. Consequently, the density of flowering stems can reach up
to 67 plants m-2, although an average of ca.4 plants is probably more typical in short, well
managed grassland (Wells & Barling, 1971).
Pulsatilla vulgaris appears to have a short-lived, transient seed bank (Thompson et al., 1997)
and so most recruitment is likely to occur shortly after the seed is dispersed. Once the seed
reaches the soil surface hygroscopic movement of the style may help bury the seed, but this
has not been observed. Germination is epigeal, the green cotyledons expanding 3-6 mm above
the soil surface. The slender primary root of the seedling grows vertically downwards reaching
a depth of 15 cm within 6 months of germination by which time it has formed an extensive
root system. At this stage the underground stem is ca.2 mm long with 2 or 3 leaves arising
from axillary buds at the apex of the rootstock.
The establishment of new plants appears to be a rare event as seedlings have only been
observed a few times in recent decades. For example seedlings were found on only one site
during a recent survey of all extant British populations (Fig. 6). At this site (Bourton Down)
seedlings were confined to bare soil in very open grassland with abundant limestone rubble at
or on near the soil surface (see Fig. 9k).
Soil moisture appears to be critical for establishment as seedlings only appear to survive in
wet summers (Wells, 1994; Rich, 1997). For example, Wells & Barling (1971) showed that

17

higher germination occurred where seeds were covered with a thin layer of soil but that all
seedlings suffered high mortality due to physical disturbance and herbivory. In addition,
seedling development was extremely slow and even after two years plants were still very
small.
These factors, combined with low seed production and frequent dessication in drought-prone
soils, probably explain the apparent inability of this species to spread to new sites and the
rarity of seedlings in the wild.

Figure 6 Seedlings of Pulsatilla vulgaris recorded at Bourton Down in Gloucesterhire on 30th

April, 2005.
3.1.8 Response to competition
Pulsatilla vulgaris occurs at highest density where soils are shallowest and above ground
competition (biomass) is low (Fig. 7a). Flower production also decreases as competition from
tall grasses increases, with a sharp fall occurring when the surrounding vegetation reaches 1015 cm (Fig. 7b). This was first observed in grazing exclosures (Wells, 1968) and has been
attributed to the smothering effect of leaf litter of Bromopsis erectus and increased
competition from tall grasses. For example, Mitchley (1988) showed a positive correlation
between interference ability and mean height of the rosette leaves in Bromopsis erectus
grassland. This was attributed to the ability of dominant species such as B. erectus to project
their leaves into the upper canopy thereby reducing the amount of photosynthetically active
radiation able to reach lower levels (Mitchley & Willems, 1995). Bromopsis erectus is also
known to generate high shoot thrust, enabling it to attain early dominance in the vegetation
by physically restricting the growth of surrounding species, especially low-growing
hemicryptophytes such as Pulsatilla vulgaris (Campbell et al., 1992). Attempts to simulate
these effects through shading have shown significant decline in longevity, survivorship and
flowering performance at increasing levels (Fig. 5). Likewise transplants of Pulsatilla vulgaris
have significantly greater survivorship and performance (number of leaves) when competition
from surrounding vegetation is removed (Warden, 2001). These findings highlight the critical
role of above-ground competition plays in restricting the amount of light reaching the base of

18

the sward and thereby determining the relative abundance of hemicryptophytes such as
Pulsatilla vulgaris.
(a)

(b)

Figure 7 The effects of competition on Pulsatilla vulgaris. (a) The relationship between
depth of soil and above-ground biomass on the density of Pulsatilla vulgaris at Barnsley Wold,
Gloucestershire. Means based on 50 samples at each soil depth. (b) The relationship between
sward height and the % of Pulsatilla vulgaris plants in flower in a grazing exclosure at Barton
Hills, Bedfordshire. Each dot represents a year with overall numbers increasing from 138
following the withdrawal of grazing 1964 to 654 in 1969. Both figures adapted from Wells &
Barling (1971).
3.1.9 Herbivory, parasites and disease
Pulsatilla vulgaris is a plant of sheep and rabbit grazed downland and is well adapted to
grazing by herbivores. The plant is dormant during the winter, and at other times the
vegetative and floral buds and leaves are generally at ground level and therefore not readily
available to grazing animals. Occasionally whole flowers are grazed by rabbits, or the flower
or part of it are eaten by caterpillars, pheasants (Bowen, 1997) or small mammals (wood
mouse Apodemus sylvaticus, field vole Microtus agrestis) (Carter, 1967). Although the leaves
contain anemonin they are eaten by sheep, rabbits, slugs and snails. Caterpillars of two leafmining moths of the genus Cnephasia have also been recorded on leaves in the UK (Database
of Insects and their Food Plants, accessed August 2010). At least two other leaf-mining insects
have been reported on the continent and nematode worms of the genus Meloidogyne have
been reported to cause root galls in the Netherlands caused by (Wells & Barling, 1971).
In Sweden achenes are attacked by the larvae and pupuria of a range seed-boring flies
belonging to two Diptera families, the Agromyzidae and Cecidomyiidae, and are a potentially
serious factor reducing seed production by up to 36% (Widn & Lindell, 1996). Many small flies
have been reported visiting flowers of Pulsatilla vulgaris but no evidence of attack of seed has
so far been observed (Warden, 2001). Parasites on Pulsatilla vulgaris include Cuscuta
epithymum, which has been recorded on plants at Barnsley Wold. The fungal rust
Coleosporium pulsatilla has also been recorded on Pulsatilla species on the Continent (Aichele
& Schwegler, 1957), but it is not known if hosts include Pulsatilla vulgaris.
4

Habitat requirements

4.1

The landscape perspective

19

In England Pulsatilla vulgaris is confined to moderately steep south to southwest facing slopes
where insolation is high, soils are shallow and competition from other species is restricted
(Fig. 8). Most populations are confined to escarpment grasslands of Chalk and Jurassic
limestone (Fig. 9c, 9e-g, 9i-l) but it also occurs in similar grasslands in old quarries (e.g.
Hornsleasow Roughs, Fig. 9b) and on ancient earthworks such as hill-forts (e.g. Ravensburgh
Castle, Fig. 9d) and Anglo-Saxon dykes (e.g. Devils Dyke, Fig. 9a). In old quarries and on
earthworks plants tend to occur on relatively shallow slopes on a greater range of aspects,
although even on these plants are generally found on more southerly aspects than those facing
northwards (Wells & Barling, 1971). Plants usually occur at greatest density on the steepest
(mid) part of the slope where soils are shallowest, rather than on deeper soils at the top or
bottom. For example, Wells & Barling (1971) recorded densities of 7.5 plants m-1 where the
soil was 5 cm deep but only 1.2 plants m-1 at soil depths greater than 100 cm (Fig. 7a).
(a)

(b)

Figure 8 The aspect (a) and slope (b) of Pulsatilla vulgaris populations in England. The
figures include values for the 29 populations included in Wells & Barling (1971) and five
populations discovered since then.
With the exception of Hildersham Furze Hills (see below), Pulsatilla vulgaris occurs on shallow
(5-15(25) cm) drought-prone calcareous rendzinas over chalk and limestone with relatively
high surface soil pH (average 7.5; range 7.1-7.8; Table 6). These soils are very friable and
porous with high contents of calcium carbonate (mean 67%; range 34-91%), organic matter
(mean 15%; range 8-22%) and total nitrogen (mean 0.5%; range 0.3-0.7%) (Wells & Barling,
1971). All are base-saturated with calcium being the most abundant cation present (320-830
mg 100g-1 dry soil) with other mineral nutrients, especially phosphorous, occurring at low
levels (Table 6). The clay/silt/sand fractions of these soils is also low (<17%).
Pulsatilla vulgaris is confined to similar calcareous strata over most of its European range and
in Sweden, Jutland and Zealand it also occurs on sandy, acid soils. At Hildersham Furze Hills,
Cambridgeshire, where it was last seen in 1990, a small population also grew on soil derived
from glacial sands and gravels mixed with chalk flints and chalky boulder clay. The soil is
calcareous sandy loam with no free calcium carbonate, a pH of 5.2 and high sand content
(Wells & Barling, 1971; Trist, 1988; Table 6). This site therefore differs considerably from all
others in Britain (see Fig. 10b) and provides an important link with continental populations in
Sweden and Denmark where Pulsatilla vulgaris grows in acidophilous communities on soils
with pH values of 4.8-5.7 with low calcium content and virtually no free calcium carbonate.

20

Table 6 Chemical and mechanical analysis of soil from Pulsatilla vulgaris sites in England and
five sites on sandy soils in Sweden for comparison. These summarise data presented for 10
sites in Table 1 of Wells & Barling (1971).
Limestone
Calc.
Chalka
Mean
Range
Sweden
b
sandc
pH
7.6
7.4
7.5
7.1-7.8
5.2
5.0
Organic
matter
13
8-22
8
(%)
15
11
8.5
Nitrogen (%)
0.5
0.4
0.5
0.3-0.7
0.2
0.3
K (mg 100g-1)
18
21
19
11-31
0.9
18
Ca (mg 100g-1)
654
492
582
320-830
170
44
Mg (mg 100g-1)
13
11
12
7-17
12
6
P (mg 100g-1)
1.5
1.0
1.3
0.6-2.3
0.8
1.1
CaCO3 (%)
71
62
67
34-91
0.8
<0.5
Clay (%)
14
13
13
3-21
17
Low
Silt (%)
4
13
8
1-42
3
Low
Sand (%)
10
17
12
5-34
79
High
The ten sites included by Wells & Barling (1971) are as follows (with indication whether
Pulsatilla vulgaris is extinct): a chalk, Aldbury Nowers (extinct), Aston Upthorpe, Devils Dyke,
Knocking Hoe, Pitstone Hill (extinct); b limestone, Barnack Hills & Holes, Barnsley Wold,
Holywell Mound (extinct), Shacklewell Hollow (extinct); c calcareous sand, Hildersham Furze
Hills (extinct).
All English populations are lowland with an altitudinal range from ca.15 m in Cambridgeshire
(also formerly in North Lincolnshire and Northamptonshire) to 245 m at Hornsleasow Roughs in
Gloucestershire, with ca.70% of recently recorded populations occurring between 41-160 m.
Like many other central and southern European species present in Britain the 16 C mean
monthly isotherm for July marks it northwestern limit (mean of 16.5 C for all extant
populations). It grows in regions where the annual total precipitation ranges from 520 mm to
at least 1088 mm; the mean for extant populations in England is 700 mm. High summer
temperatures combined with low summer rainfall therefore restrict its distribution southwards
whereas its eastern and northeastern limits are probably determined by its sensitivity to low
winter temperature. Perring (1956) speculated that higher humidity might be responsible for
its absence from the southern most chalk in England based on an analysis of
precipitation/saturation deficit ratios but this has been rejected by Wells & Barling (1971)
who cite the past history of chalklands as a much more plausible explanation (but see section
2.3.1.1).

21

Figure 9 Habitats of extant populations of Pulsatilla vulgaris in England (arrows mark the
position of small populations): (a) Anglo-Saxon earthwork, Devils Dyke, Cambridgeshire; (b)
shallow limestone quarry workings, Hornsleasow Roughs, Gloucestershire; (c) Knocking Hoe
and Spiranthes Bank, Bedfordshire; (d) Ravensburgh Castle Iron Age hill fort, Bedfordshire;
(e) Ledsham Banks, Mid-west Yorkshire; (f) Steps Hill (Incombe Hole), Buckinghamshire.

22

Figure 9 continued. Habitats of Pulsatilla vulgaris in England: (g) Beaumonts Hay,

Gloucesterhire; (h) Barnack Hill and Holes, Northamptonshire; (i) Barnsley Wold,
Gloucestershire; (j) Barton Hills, Bedfordshire; (k) Bourton Down, Gloucestershire; (l) Church
Hill, Therfield Heath, Hertfordshire.
4.2
Communities & vegetation
In England Pulsatilla vulgaris is almost entirely confined to short, herb-rich Bromion grasslands
on calcareous soils derived from the Chalk or Oolitic Limestone (Rodwell, 1992). The main

23

National Vegetation Classification (NVC) grassland type on both chalk and limestone is Bromus
erectus grassland (CG3), including grazed species-rich and ungrazed species-poor subcommunities (CGa-b and CG3d respectively) (Fig. 10a). It occurs less frequently in Bromus
erectus-Brachypodium pinnatum (CG5) and Brachypodium pinnatum (CG4) grasslands, mainly
on limestone (Rodwell, 1992). These three communities form the core alliance of the
Brometalia in western Europe and are broadly characterised by the grasses Brachypodium
pinnatum, Bromopsis erectus, Helictotrichon pratense, Koeleria macrantha and a wide range
of herbs (Rodwell et al., 2007). It occasionally occurs in other calcareous grasslands,
especially Festuca ovina (CG2) grasslands on chalk, usually where heavy grazing by sheep and
rabbits has reduced the cover of Bromopsis erectus to very low levels. In all these
communities the vegetation is typically very diverse (23 species m-2) with a high cover of
broad-leaved herbs (44 % m-2) and constant associates include Bromopsis erectus, Carex
flacca, Cirsium acaule, Festuca ovina, Helianthemum nummularium and Sanguisorba minor
and associated (see Annex 3).

Figure 10 Grassland communities in which Pulsatilla vulgaris occurs in England: (a) National
Vegetation Classification (NVC) grassland types; and (b) ordination plot (Detrended
Correspondence Analysis) showing overall variation in species composition. Both plots include
quadrat data for 30 sites including all 29 visited by Wells & Barling (1971) plus Swinstead
Valley discovered in 2009. Twenty-nine of these quadrats were recorded in 1964-1968 and 27
recorded since 2000. * includes sub-communities 2a & 2d; ** includes sub-communities 3a, 3b
and 3d.
Overall there is remarkably little variation between the composition of these grasslands on
chalk or limestone (Fig. 10b), the main difference being the almost complete replacement of
Brachypodium pinnatum by Bromopsis erectus on nearly all the sites on the chalk (Wells &
Barling, 1971). Two notable exceptions, positioned to the extreme left of the first axis of the
ordination plot, are Hildersham Furze and Unhill Bottom. Both were rank, species-poor swards
dominated by Festuca rubra by the time of the 1964-68 survey. At one of these sites,
Hildersham Furze Hills, Pulsatilla vulgaris formerly grew on an acidic soil derived from glacial
sands and gravels, although the vegetation was distinctly calcicolous due to the presence of
free calcium. Pulsatilla vulgaris occurs on soils with a similar pH in Sweden and Denmark, but
these are much more base-poor and consequently the vegetation is dominated by acidophiles
such as Calluna vulgaris and Deschampsia flexuosa (Wells & Barling, 1971). Pulsatilla vulgaris
probably once grew in similar acid communities in Breckland as a few former sites (e.g.

24

Cavenham Heath) support extensive areas of Festuca ovinaAgrostis capillarisRumex

acetosella acid grassland (Erodium cicutarium-Teesdalia nudicaulis U1c sub-community), a
species-rich types with close affinities to calcareous grassland (Rodwell, 1992).
Pulsatilla vulgaris tends to grow in short swards (7.5 0.7 cm) though it can persist for long
periods in taller grassland (>20 cm) in the absence of grazing, cutting or burning. However,
recent population trends suggest that a sward height of between ca.5-10 cm is ideal and that
in taller swards competitive exclusion by coarse grasses eventually leads to an overall decline
in plant numbers (Fig. 11a). Consequently taller swards lead to a reduction in overall species
diversity (Fig. 11b) and an increase of in the dominance of Bromopsis erectus (Fig. 11c).
Bailey (1996) provides a useful list of species positively and negatively associated with the
open conditions favoured by Pulsatilla vulgaris.

Figure 11 Differences in (a) sward height (cm), (b) species diversity and (c) % cover of
Bromopsis erectus ( 1 SE) on extant Pulsatilla vulgaris sites in England (n = 14) in relation to
population trends, 1968-2006. Means with the same latter are not significantly different from
one another (Walker et al., in press).
4.3
Summary of habitat requirements
A number of habitat features are important to Pulsatilla vulgaris in England and these are
summarized in Table 7.
Table 7 - A Summary of habitat features important to Pulsatilla vulgaris in England.
Type
Description
Physical and
Lowland (<240 m)
topographical
Escarpments, old quarries and earthworks
Moderately steep slopes with a southwesterly aspect
Insolation high (full sun)
Shallow soils (<15 cm) derived from calcareous substrates (chalk,
limestone, calcareous sand)
Vegetational and Open CG2-5 calcareous grassland on chalk or limestone
structural
Very open vegetation with 30% bare ground
Low vegetation ca.5(-10) cm with low cover of either Bromopsis
erectus or Brachypodium pinnatum
Processes
Ideally grazing by sheep and rabbits to reduce the dominance of
coarse grasses (especially Bromopsis erectus) and to create bare

25


Chemical

microsites for seed germination and seedling establishment

Transfer of seed between sites by herbivores and/or mowing
machinery
Transfer of pollen between populations by insects
Soils with pH ca. 7.5 (formerly 5)
Infertile soils with high calcium carbonate content

Management implications

Pulsatilla vulgaris is a long-lived polycarpic perennial herb which mainly reproduces

vegetatively, and rarely from seed. It is a poor competitor and therefore unable to persist in
ungrazed, closed swards or amongst scrub where the general height of the vegetation exceeds
15 cm and there is a build-up of litter from coarse grasses, especially Bromopsis erectus and
Brachypodium pinnatum. Its survival is therefore dependent on grazing, although short-term
fluctuations in grazing intensity or short-term neglect are unlikely to be critical in the longerterm. For example, one of the largest populations in England survived 100 years of sheep
grazing, 13 years of very heavy sheep grazing, 22 years of no grazing including a severe fire
during the flowering season (Wells, 1968).
Management should aim to produce a sward of ca.5(-10) cm during the flowering period with
up to 30% bare ground although there is much flexibility in how this can be achieved (Rich,
1997). The preferred management is sheep grazing from August to April (< 5 sheep ha-1),
although low intensity grazing, including by cattle or horses, during the flowering season
appears to have no ill-effect. All its current sites are also subject to moderate to high levels
of rabbit grazing throughout the year and, although this can cause localized extinction where
severe, it has probably been vital in maintaining open conditions on sites with intermittent or
low levels of stock grazing. It has also survived a variety of cutting treatments ranging from
cutting once a year in spring, summer or autumn, to three times a year as well as annual
burning (swaling) in February or March to reduce the dominance of coarse grasses such as
Bromopsis erectus and Brachypodium pinnatum. On some sites it has also benefited from
occasionally burning and trampling by humans which appears to stimulate the development of
the deep-seated adventitious root buds.
On some sites where over-grazing, mainly by rabbits, is perceived to be a problem fencing
exclosures have been erected although this appears to have been detrimental due to the
build-up of coarse grasses.
Given the poor dispersal ability of Pulsatilla vulgaris and the high degree of isolation of many
populations the movement of livestock and cutting machinery between extant and potentially
suitable sites is likely to be critical in promoting gene-flow and in enabling colonization to
take place.
7

Threats/factors leading to loss or decline or limiting recovery

In England Pulsatilla vulgaris has undergone a rapid and dramatic range contraction over
recent millennia and its current distribution now represents just 27% and 14% of its former
range at the 10-km and site-population scales respectively. A similar magnitude of decline has
been reported for Pulsatilla vulgaris in central Germany (Hensen et al., 2005), Austria (Franz,
2005), Switzerland (Pfeifer et al., 2002) and Luxembourg (Colling, 2005) and for Pulsatilla
patens in Finland where 60% of populations have been lost since the 1930s (Uotila, 1996;

26

Kalliovirta et al., 2006). There are also striking similarities with the decline of Orchis ustulata
in England (Foley, 1992), which might be expected given that the two species formerly grew
together on a number of sites (and still do at one site in Bedfordshire).
In England historical records show that Pulsatilla vulgaris has been declining since at least
1750 (Fig. 4; Table 8). The majority of the losses before 1900 (64%) were due to ploughing-up
of common lands following Parliamentary Enclosure awards, mainly between 1750 and 1850
(Wells, 1968; Jones, 1969; Wells, 1969). These earliest losses were described most vividly by
the Cambridgeshire botanist Charles Babington (1860), who noted that Until recently (within
60 years) most of the chalk district was open and covered with a beautiful coating of turf,
profusely decorated with Anemone Pulsatilla [Pulsatilla vulgaris], Astragalus Hypoglottis
[Astragalus danicus], and other interesting plants. It is now converted into arable land, and its
peculiar plants mostly confined to small waste spots by road-sides, pits, and the very few
banks which are too steep for the plough. Although this probably over-states its former
abundance, it was certainly occurring in other parts of southern England (e.g. Druce, 1886,
p.xxxii) due to increased demand for home-grown corn fuelled by a rapidly increasing urban
population. During the same period many populations were also lost to quarrying and mining
activities on coal fields in the Midlands and northern England (Table 8).
Table 8 Reasons for the loss of Pulsatilla vulgaris populations in England between 1750 and
2000. % loss figures are based only on localities where the cause of loss is known and therefore
exclude Unknown and Possible error/duplicate record.
17501801185119011951Total
Reason for loss
1800
1850
1900
1950
2000
(%)
Ploughing
2
8
10
7
1
28 (42)
Lack of grazing/scrub
encroachment
1
1
1
14
17 (25)
Quarrying/mining/mineral
extraction
3
1
3
7 (10)
Improvement/overgrazing
7
7 (10)
Building/urban development
2
1
1
1
5 (7)
Afforestation
1
1
2 (3)
Landscaping of parkland
1
1 (1.5)
Unknown
1
1
14
13
7
36
Possible error/duplicate record
2
3
5
Total
9
12
31
26
30
108
Rate of loss per decade
1.8
2.4
6.2
5.2
6.0
4.3

27

Figure 12 Sites where Pulsatilla vulgaris is now extinct (all photos taken 2003-2006; year of
last record indicated): (a) Fleam Dyke, Cambridgeshire (1973); (b) Shacklewell Hollow,
Leicestershire (1992); (c) Hildersham Furze Hills, eastern (Sand) hill, Cambridgeshire (1990);
(d) Honnington Camp, Lincolnshire (1992); (e) Pitstone Hill, Buckinghamshire (1996); (f)
Southorpe Roughs, Northamptonshire (1990).

28

Losses to ploughing and quarrying continued into the twentieth century and by the 1960s only
29 populations remained (Wells & Barling, 1971). Losses since then have mainly been caused
by increased above-ground competition, particularly with Bromopsis erectus, as a result of
under-grazing (Walker et al., in press). Ironically this was first observed in grazing exclosures
that were erected to protect populations from over-grazing (e.g. Carter, 1967; Wells, 1971).
Of the 20 sites where Pulsatilla vulgaris has either declined or gone extinct since the 1960s,
16 have been ungrazed for over 40 years or have only received intermittent grazing, mowing
or burning (Table 9). These include small areas of grassland on earthworks or steep banks that
are now completely isolated within arable landscapes and are now partly or completely
scrubbed-over (e.g. Fleam Dyke, Fig. 12a; Honnington Camp, Fig. 12d), small exclosures
erected to exclude livestock, deer and rabbits on over-grazed sites (e.g. Aston Upthorpe
Down, Fig. 13a) and privately managed nature reserves where resources for management have
been limited (e.g. Ancaster Valley, Fig. 13).
This decline of grazing frequency and intensity on many sites has been due to the reduced
profitability of livestock farming in arable areas since the 1950s which has made it difficult to
sustain appropriate grazing management on unproductive sites (Nisbet & Shere, 2006). On
mixed farms the conversion of grassland to arable has made farmers less committed to grazing
small areas, whereas their inaccessibility has deterred potential graziers because of increased
transport and infrastructure costs. Even on livestock farms these problems have prevented the
grazing of small fragments of semi-natural grasslands perceived to be unproductive and/or
difficult to manage. In addition, there has been a decline in rabbit grazing since the spread of
myxomatosis in 1953 which is known to have caused dramatic successional changes, especially
on sites with no history of livestock grazing (Sumpton & Flowerdew, 1985).
Table 9 - Management of Pulsatilla vulgaris populations in relation to trends in abundance,
1968-2006. 1-5 represent a decline in the intensity of grazing management.
Extant
Extinc
Management since 1968
Total
Increas
t
Stable
Decline
e
1. Winter + some spring/summer/autumn
grazing
3
2
5
2. Winter grazing (since 1980)
2
1
3
3. Irregular grazing, mowing, burning
4
2
2
8
4. Over-grazed then under-grazed
2
3
5
5. Ungrazed for over 40 years
7
7
6. Improved/destroyed
4
4
7. Unknown
1
1
Over the last 40 years many grassland sites have also been agriculturally improved and
reseeded to increase productivity. Other threats have included the digging-up of plants for
horticulture including over 1000 from Knocking Hoe in 1948 (Hope-Simpson, 1948) and more
recently at Barton Hills (M. Gurney, pers. comm.). In addition, the last known plants were
dug-up at two former sites (Broughton Far Wood, Fleam Dyke; Marren, 1999) although other
factors were more important in causing declines. Other direct threats, including forestry,
building developments, etc. have only caused a few losses, including the dumping of rubble on
the last Leicestershire colony. More recently there has also been concern, but no direct
evidence, that atmospheric nitrogen deposition may be adversely affecting some populations
through eutrophication of infertile grassland (Crawley, 2005; Rich et al., 1993).

29

The current range of Pulsatilla vulgaris in England is characterised by a high level of

fragmentation, since numbers and size of populations have declined considerably over the
past 40 years. During this time the average distance between neighbouring populations has
increasing from 8 km to 22 km. Most significantly eight populations are now separated by more
than 10 km, five by more than 35 km and the most isolated site, Ledsham in Yorkshire, by 102
km. Consequently, gene-flow between populations is likely to be very low or none existent
making populations vulnerable to genetic drift and inbreeding depression, and therefore local
extinction due to genetic, demographic or environmental problems (Schaffer, 1987; Lande,
1988). In these circumstances a decrease in genetic diversity is to be expected (Soul, 1986)
accompanied by a reduction in reproductive performance as shown for Pulsatilla vulgaris in
central Germany, as well as a number of other European grassland species that have suffered
similar declines (e.g. Oostermiejer et al., 1995; Ouburg & van Treuren, 1995; Vergeer et al.,
2003). The results from the German study suggests that due to life-history characters
Pulsatilla vulgaris may have maintained higher levels of genetic variation than would be
expected for a declining species, although small populations are likely to display lower levels
of genetic variation than larger ones, even where gene exchange is occurring between sites in
close proximity. As the distance between half of English populations is well beyond the
foraging radius of most pollinators (>10 km) we would expect to see a significant
differentiation between isolated or outlying populations, both in terms of genetic variation
and reproductive performance. Further work is needed to test these hypotheses.
7

Current Conservation Measures

7.1
In Situ Measures
Pulsatilla vulgaris is currently classed by as Vulnerable using IUCN criteria but is not currently
included in Schedule 8 of the Wildlife and Countryside Act, 1981 (Cheffings & Farrell, 2005). It
was also included as a priority Biodiversity Action Plan species in 2007 (NERC Act 2006) and is
listed as one of 47 priority Crop Wild Relatives in the UK for which action plans have been
produced (Maxted et al., 2007).
With the exception of Taylors Hill, Gloucestershire, all extant populations of Pulsatilla
vulgaris occur on SSSIs or NNRs (Table 10). Seven sites appear to be in favourable condition
from the perspective of Pulsatilla vulgaris (but not necessarily for other designated site
features), with some form of grazing management to maintain calcareous grassland features
of interest including Pulsatilla vulgaris. Populations have therefore increased in size or
remained stable over the past 40 years. Seven sites appear to be in poor condition or are
recovering from poor condition due to the introduction of management. These include sites
that were under-grazed for a long period and have recently been cleared of scrub and grazing
re-introduced. The most dramatic example is Ancaster Valley, South Lincolnshire (Fig. 13). By
1981 dense Ulex europaeus scrub had eradicated most of the Pulsatilla vulgaris population (as
well as Carex ericetorum) and only a few plants remained. Since then the scrub has been
completely removed and the condition of the grassland improved through winter sheepgrazing. A similar project to stock-proof and reintroduce grazing to the Devils Dyke in
Cambridgeshire is also benefiting Pulsatilla vulgaris largely through the reduced dominance of
Bromopsis erectus (Fig. 14d). The recent re-introduction of grazing is also having positive
results at Rodborough Common (cattle) and Deacon Hill, where a flying flock of Hebridean
sheep are used to graze a number of Pulsatilla vulgaris sites managed by the Wildlife Trust.
Table 10 - Extant populations of Pulsatilla vulgaris in England.
Site
Cons Condition

30

.
(a) Sites in good condition, grazing regime ideal
Barnack Hills and NNR Good condition, population temporary fenced to reduce
Holes
disturbance
Barnsley
Wold SSSI Good condition
Warren
Barton Hills
NNR Good condition, flowering poor in some years due to overgrazing
Bourton Down
SSSI Good condition, seedlings present
Hornsleasow
SSSI Good condition, some parts under-grazed
Roughs
Knocking Hoe
NNR Good condition, severe rabbit disturbance in some years
Therfield Heath
SSSI Good condition, some parts under-grazed
(b) Sites in poor condition or recovering from poor condition
Ancaster Valley
SSSI Scrub clearance and grazing reintroduced
Deacon Hill
SSSI Under-/over-grazed in the past; severe rabbit disturbance
Devil's Dyke
SSSI Under-grazed but grazing introduced to some areas
Beaumonts Hay
SSSI Grazing irregular
Taylors Hill
Grazing irregular
Rodborough
SSSI Under-grazed
Common
Swinstead Valley
SSSI Under-grazed, encroachment by Brachypodium pinnatum
(c) Sites in poor condition, plant likely to go extinct
Aston
Upthorpe SSSI Most plants in ungrazed exclosure; rest of site overgrazed
Down
Ledsham
SSSI Only a single plant; likely to go extinct
Ravensburgh Castle SSSI Ungrazed; likely to go extinct
Steps Hill
SSSI Undergrazed, threatened by scrub encroachment
As noted above, exclosures have been erected at a few sites to protect populations from overgrazing (e.g. Aston Upthorpe, Knocking Hoe, Pitstone Hill) but in general these had lead to
declines due to increased growth of Bromopsis erectus. At Aston Upthorpe annual mowing has
been introduced to counter this and maintain sward heights at the appropriate height (Fig.
14a). At Therfield Heath temporary fencing is used to rotationally graze common land and to
ensure that areas containing Pulsatilla vulgaris are grazed appropriately. At Barnack Hills and
Holes single strand fencing is used to reduce disturbance by visitors during the flowering
period (Fig. 14b). At the most recently discovered site, Swinstead Valley, a temporary grazing
exclosure erected during the flowering period led to an increase from 4 flowers in 2009 to 18
in 2010.

31

1981

1992

1999

2006

Figure 13 The restoration of Pulsatilla vulgaris habitat at Ancaster Valley, South

Lincolnshire. The extant population is located on the left hand slope covered in Ulex
europaeus in 1981. The scrub was cleared in the 1980s and winter-grazing reintroduced.
Table 11 Details of translocations of Pulsatilla vulgaris in England.
Site
No.
%
Year(s)
of
Source
surviv
Details of translocation attempt
of intro plan
of seed
al
ts
Augmentation of existing colony.
Aston Upthorpe,
1999
87
56
Aston
Berkshire
(2009) Upthorp Plants protected within exclosure
but few flower due to predation by
e
rabbits or voles.
Hartslock,
1998
78
80
Barnack Experimental introduction to
Oxfordshire
(2008)
investigate translocation
techniques (Warden, 2001).
1999
12
0
Barnack Re-introduction to a former site.
Southorpe
Plants failed to survive more than 2
Paddock,
years.
Northamptonshir
e
Ancaster, South
199268
32
Ancaster Augmentation of existing colony.
Lincolnshire
2002
(2005)
Plants protected by wire cages
during the flowering period.
Copper Hill
199468
0
Ancaster Re-introduction to a former site.

32

Quarry, Ancaster

1996

Most scratched out by rabbits and

pheasants within a few days of
planting. Four plants survived for
two years.
Ledsham, Mid2000
1
0
Barnack A single plant was introduced to
west Yorkshire
encourage cross-pollination of the
sole remaining plant in northern
England. This died within a couple
of years.
In England attempts to re-introduce Pulsatilla vulgaris to former sites or augment existing
populations have been carried out at six sites with variable success (Table 11). Transplants
have survived at Aston Upthorpe, Ancaster and Hartslock plants within protective exclosures
where the surrounding vegetation has been removed to reduce competition (Fig. 14c) and the
surrounding vegetation has been mown or grazed. However, at both Aston Upthorpe and
Hartslock very few plants manage to set seed due to predation by rabbits or voles (Warden,
2009). All other attempts have been failures with plants only surviving for one or two years.
This included an attempt to translocate a single plant to within 4 metres of the sole surviving
plant in northern England (at Ledsham) to encourage cross-pollination (A. Headley, pers.
comm.). Similar high rates of translocation failures have been reported for Switzerland
(Pfeifer et al., 2002).

Figure 14 Examples of in situ conservation work for Pulsatilla vulgaris in England: (a)
fencing to exclude livestock, Aston Upthorpe Down, Berkshire; (b) temporary fencing to

33

reduce trampling by visitors, Barnack Hills & Holes, Northamptonshire; (c) cages to protect
transplants from rabbits (note clearance of vegetation from around the plant), Ancaster
Valley, South Lincolnshire; and (d) effects of the reintroduction of grazing to the Devils Dyke,
Cambridgeshire, following the erection of stock-proof fencing.
7.2
Ex situ Measures
Plants of wild origin are known to be cultivated in the Oxford Botanic Garden and Kews
garden at Wakehurst Place. Plants originating from Barnack Hills and Holes were also grown by
Terry Wells at Monks Wood Experimental station between 1970 and 1980 for the purposes of
research, some of which were transferred to his garden in Upwood. Many plants originating
from wild British populations are likely to be cultivated in other private gardens. A small
amount of viable seed is currently stored in the Kew Millennium Seed Bank at Wakehurst Place
(Royal Botanic Gardens Kew, 2008).
Pollen from the single Ledsham plant has been used to cross-pollinate five plants from
Barnack Hills and Holes, with the intention of growing on plants for reintroduction to the wild
site, but without success (A. Headley, pers. comm.).
7.3
Research Data
Research is urgently required to assess genetic variation within English populations and the
extent to which small population size, limited gene-flow (due to extreme population
fragmentation), genetic drift and inbreeding depression are affecting the performance of
English populations. Likewise studies that assess dispersal and pollination mechanisms, in
particular the potential for dispersal via livestock and machinery, may provide valuable
insights into gene-flow and how best to conserve Pulsatilla vulgaris in the future. Studies on
the genetic relationships between British and continental populations would also be useful
help to inform European conservation priorities.
Research on the regeneration niche of Pulsatilla vulgaris is needed in order to establish the
optimum conditions under which field germination and seedling establishment takes place.
Further work is also needed to assess factors controlling the abundance of Pulsatilla vulgaris,
in particular the affects of the removal of competitive dominance in grassland swards e.g.
responses to environmental perturbations and the removal of dominant species and leaf litter
(Bailey, 1996).

Figure 15 Fluctuations in the number of Pulsatilla vulgaris in fixed plots at Barnack Hills
and Holes, Northamptonshire. Data supplied by Chris Gardner (Natural England).

34

7.4
Monitoring and the Common Monitoring Standard
Traditionally Pulsatilla vulgaris has been monitored by simple counts of flowering crowns or
estimates based on densities in small areas (e.g. Tarpey, 1999). These provide an overall
assessment of year to year variation but are limited for a number of reasons. First, crowns
do not represent discrete genets due to the production of offshoots close to parents. The total
count is therefore likely to overestimate the total size of the population possibly by orders of
magnitude. Second, grazing can have a marked influence on estimates especially on sites
where livestock are present during the flowering period. Third, such counts do not take into
account the life-stage structure of the population or whether recruitment is taking place
through the production of seed, presence of juveniles, etc.
Possible alternative approaches include counts within fixed plots as has been carried out at
Barnack Hills and Holes in Northamptonshire since 1975 (Fig. 15). This should preferably
include a number of demographic parameters, including the individuals in different lifestages, flowering performance, seed production etc., and relate these environmental
variables such as sward height, % cover of bare ground, hits on dominant grasses, etc. Simple
counts of rosettes rather than flower stems is probably a more preferable performance
parameter given the dramatic fluctuations in the numbers of flowers observed on some sites
(e.g. Fig. 15). These are less affected by seasonal factors and grazing and therefore provide a
more accurate assessment of the size of populations (Bailey, 1996). However, all these
approaches are likely to be time consuming to carry-out and therefore dependent on available
expertise, time and resources.
8

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Preston, C.D., Pearman, D.A. & Dines, T. 2002. New Atlas of the British and Irish flora.
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Rich, T.C.G. 1997. The Management of Semi-natural Lowland Grassland for Selected Rare and
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Warden, K. 2001. An investigation into techniques for the propagation and augmentation of a
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pp.138-139.

39

Contacts

Botanical Society of the British Isles

97 Dragon Parade
Harrogate
North Yorkshire
HG1 5DG.
Plantlife International
The Wild Plant Conservation Charity
14 Rollestone Street
Salisbury
Wiltshire
SP1 1DX.
Tel: 01722 342730

10

Kevin Walker - kevinwalker@bsbi.org.uk

Tim Wilkins - tim.wilkins@plantlife.org.uk

Links

BSBI Maps Scheme: http://www.bsbimaps.org.uk/atlas/main.php

Records via NBN Gateway: http://www.nbn.org.uk

Global Biodiversity Information Facility: http://www.gbif.org

Pasqueflower is a Species of Principal Importance in England (S41 of NERC Act 2006):

http://www.ukbap.org.uk/NewPriorityList.aspx

The UK list of threatened (red listed) vascular plants is maintained by JNCC:

http://jncc.defra.gov.uk/page-3354

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11

Annex 1 site descriptions

Descriptions of the 34 English population of Pulsatilla vulgaris thought to be extant in the

1960s including 29 included in Wells & Barling (1971) and five discovered subsequently. With
the exception of Swinstead Valley (discovered in 2009) all have been resurveyed by Walker et
al. (in press) since 2000. Sites are listed in vice-county and then alphabetical order.
Conservation designations are given in parentheses. Denotes where Pulsatilla vulgaris is
thought to be extinct.
HERTFORDSHIRE (v.c.20)
Aldbury Nowers (SSSI)
During the 1960s a few plants were recorded in rank Bromopsis erectus grassland (CG3b) and
scrub overlying chalk adjacent to an Iron Age camp. The site was ungrazed for many years and
consequently scrubbed over (recently cleared) except in a few places where small fragments
of chalk grassland remain. Pulsatilla has not been seen at this site since 1969 despite
repeated searches (Trevor James, pers comm.).
Therfield Heath (SSSI)
The second largest British colony largely confined to very short, rabbit grazed turf on the
steep south to southwest-facing slopes of a narrow chalk spur known locally as Church Hill
(TL332395). A few plants also occur in rank Bromopsis erectus grassland (CG3a) on the
adjacent Pen Hill (TL332397) and to the east of the tumuli marked on the OS map (TL332399).
All occur on common land in the roughs of a golf course which were ungrazed for many
years. Church Hill has a long history of heavy rabbit grazing and in the 1980s small areas were
also mown in late February or early March using a tractor and Flymo. Since 1986 rotational
winter grazing has been carried out across much of the site using temporary grazing
compartments and, on Church Hill at least, this has led to an increase in overall population
numbers, which now exceed 60,000 and up to 12 rosettes m-2 (Tarpey, 1999). Recreation
pressures include trampling on the spine of Church Hill and the occasional removal of flowers
or whole plants. The population near the tumuli may have originated from seed spread by a
local farmer (W.H. Darling) in 1953/54.
BERKSHIRE (v.c.22)
Aston Upthorpe Down (SSSI)
The sole relic of a once extensive population that stretched along the Berkshire Downs (Druce,
1886; Crawley, 2005). By the 1960s around 300 plants were confined to a northwest-facing
chalk slope although flowering was limited due to heavy grazing by rabbits. As a consequence,
a small exclosure was erected in 1961 with an inner fence to exclude rabbits (repaired 1976
and replaced in 1998). This resulted in a gradual decline in the numbers of Pulsatilla vulgaris
due to increased competition from coarse grasses and the grazing off of inflorescences by
wood mice (Carter, 1967). The population continued to decline into the 1970s and 1980s
despite occasional mowing/burning to control the growth of Bromopsis erectus. Mowing was
re-introduced in 1996 and this led to an increase in flowering performance although by this
time only around 13 plants remained (Warden, 2001). Only a few plants have been recorded
outside this exclosure since 1994 due to very heavy rabbit grazing. Eighty-seven plants grown
from Aston Upthorpe seed were planted inside the new exclosure in October 1999 (Warden,
2001); 55 (63%) were still present in 2004 (Kathy Warden, pers. comm.).

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Lockinge
In 1962 two plants were discovered in rough grassland above Lockinge. It has apparently been
seen again since 1987 (Crawley, 2005) but no plants or suitable habitat could be located by
the author in 2005 (SU427846).
Unhill Bottom
This site was the remnant of an extensive population, possibly totalling over 55,000 plants,
which spread over much of the adjacent chalk downland of Deans Bottom (Crawley, 2005).
Much of this area was used for military training during the 1939-45 war and was subsequently
ploughed-up. The population had declined to around 5000 plants by the late 1950s (Bowen,
1968) and then to around 750 plants by 1967 when Pulsatilla was confined to a small area of
ungrazed Festuca rubra grassland (CG6) on a NNW-facing slope. It apparently survived in this
locality till at least 1978 but eventually disappeared when the grassland was improved and
intensive sheep grazing was introduced (Warden, 2001).
BUCKINGHAMSHIRE (v.c.24)
Clipper Down (SSSI)
In the 1960s a few plants were recorded on a steep SSW-facing chalk slope which was being
heavily grazed by sheep from November until April. The intensity of grazing was increased
during the 1970s and 1980s (excluding November to March) leading to deterioration in the
condition of the site (Everett, 1988). The population was fenced (c.1986) for a number of
years until grazing levels were reduced on the surrounding slope. The last reliable record was
in 1976 although there are unconfirmed records of flowering in 1986 and more recently as
vegetative plants (Greves, 1997). No plants have been found in recent years despite repeated
searches during the flowering period (L. Trowbridge, pers., comm.). This site currently falls
within the Ivinghoe Hills SSSI complex and is managed favourably by a tenant farmer for the
National Trust.
Pitstone Hill (SSSI)
In the 1960s a few plants occurred on the steep S-facing slope of a mediaeval ditch in
relatively short Bromopsis erectus grassland. It persisted in tightly-grazed grassland although
plants were very difficult to locate in the mid-1980s due to heavy grazing pressure. An
exclosure was erected around the site in 1986 and this has led to the build up of coarse
grasses and scrub. The last record was at the top of the S-facing bank in 1996; it has not been
recorded since despite repeated searches.
Steps Hill (SSSI)
In the 1960s over 100 plants were recorded in short, Festuca ovina turf on a steep SW-facing
spur of chalk downland above Incombe Hole. The site was formerly very heavily grazed by
sheep and cattle leading to localised poaching and the removal of most of the flowerheads
during the spring (R. Maycock, pers. comm.). Grazing levels were subsequently reduced and
this led to a gradual increase in Bromopsis erectus (CG3). Sheep now lightly graze the site
from autumn to early spring (Greves, 1997) and as a result vegetation remains fairly rank (>15
cm). In 2005 there approximately 50 plants were located in a limited area (20 15-m)
surrounded by hawthorn scrub and many freely regenerating whitebeams.
CAMBRIDGESHIRE (v.c.29)
Devils Dyke (SSSI)

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Historically Pulsatilla has been recorded sporadically along 7 km of an Anglo-Saxon chalk ditch
and bank between Burwell and where the Cambridge-Newmarket railway crosses the dyke to
the SE. The majority of plants are confined to the SW-facing banks particularly where the
grassland has been kept open by rabbits, infrequent mowing or burning. A few plants also
occur on the top of the dyke, where grass growth is reduced by trampling, and on the
adjacent Newmarket Heath which is cut fortnightly, although no plants have been recorded
from the heath since 1984. The dyke itself was not grazed for many decades leading to the
development of a thick sward of Bromopsis erectus. Accidental burning of one section in the
1970s led to a dramatic increase in flowering numbers although numbers subsequently
declined to pre-1970 levels. Since then localised mowing has led to localised increases on
some sections (e.g. adjacent to Newmarket Heath) and in 2003 the introduction of wintersheep grazing (and stock-fencing) led to increased flowering on the southern section. In the
same year four plants appeared (with Himantoglossum hircinum) on the steep SE-facing bank
of the A14 c. 300m to the N of the nearest plants on the Dyke itself (TL603632; Leslie, 2004).
These presumably established from seed dispersed to the site on mowing machinery.
Fleam Dyke (SSSI)
In the 1960s a small population was recorded on an ungrazed SW-facing bank of a mediaeval
chalk ditch and bank immediately to the NW of a small tumulus. It was last seen in April 1973
when the plants were dug-up by the farmer in order to reduce the number of botanists visiting
the site (Marren, 1999).
Hildersham Furze Hills (SSSI)
A small population formerly grew (with Dianthus deltoides) on the spine of the easterly (Sand)
hill on a sandy soil (pH 5.2) derived from glacial sands and gravels (Wells & Barling, 1971).
During the 1950s there were never more than 3-4 clumps with 20-30 flowers (Trist, 1988).
Annual rabbit damage was intense, especially during droughts, although this declined
following the advent of myxomatosis in 1954. Three plants survived on the spine of the hill
into the mid-1980s despite the spread of rank grasses and blackthorn scrub (Trist, 1988). By
1990 only one, non-flowering plant remained and it has not been recorded since despite
improvements in the condition of the grassland and the removal of scrub (Compton, 2010
onwards).
BEDFORDSHIRE (v.c.30)
Barton Hills (NNR)
Several large sub-populations occur towards the S end of this reserve in exceptionally rich
chalk downland turf on steep, predominantly S- to SW-facing slopes and spurs where soils are
very shallow and competition from coarse grasses is restricted (Wells, 1968; Welsh, 1983). A
few plants also occur on ancient field systems at the N end of the reserve where grazing is
more variable. Historically the site was heavily grazed by sheep but management was
neglected from 1934 to 1954 resulting in the development of a dense sward of Bromopsis
erectus (Wells, 1968). This was burnt-off in 1954 and sheep grazing re-introduced for nine
months of the year (2-3 ac-1), although rabbit grazing declined dramatically due to
myxomatosis. Since the mid-1980s the site has been grazed by rabbits all year (although
populations have increased in recent years) and sheep (0.4 ha-1) from May to December
(inclusive) although this has been relaxed in recent years due to a rise in the rabbit population
(Bailey, 1996). This has resulted in an increase in the flowering population to over 5,000
crowns although flowering is suppressed in some years by heavy grazing (G. Bellamy, pers.
comm.). The Barton population of Pulsatilla is well known and attracts many visitors during

43

the flowering period. In the past many flowers were traditionally gathered at Easter, although
this activity it is much less common today. However, in 2006 around 70 plants were illegally
removed (M. Gurney, pers. comm.).
Deacon Hill (SSSI)
Since 1960 small numbers of Pulsatilla have been recorded sporadically from several localities
in the vicinity of Deacon Hill on steep, SW to NW-facing chalk downland slopes and spurs: (1)
the NE slope of Barn Hole (TL119294); (2) narrow spur adjacent to Pegsdon Firs (TL121297);
(3) combe NE of Pegsdon Firs (TL123296); and (4) the celtic field system below Deacon Hill
itself (TL12SP98). Historically the northern half of the site (Deacon Hill) has been heavily
grazed by sheep and rabbits. In contrast, grazing has been much lower in the southern half of
the reserve leading to a decline as a result of scrub encroachment and build-up of coarse
grasses from the 1950s onwards. In recent years, scrub clearance and the reintroduction of
grazing have led to an improvement in the quality of the grassland although no plants have
been found in Barn Hole despite repeated searches.
Knocking Hoe (NNR)
Between 500-1000 plants grow on the SW-facing slope of a small chalk hill (the Hoe;
TL130139) in exceptionally rich calcareous turf with a number of other national rarities
(Hypochaeris maculata, Orchis ustulata, Seseli libanotis and Tephroseris integrifolia). A few
plants also occur on a Celtic field system (within a grazing exclosure) just to the east
(Spiranthes bank; TL131308), and more sporadically within an abandoned exclosure on a
steep S-facing slope to the W of Tingley Wood (TL132306). It was lost from other parts of the
reserve that were ploughed for a short period in 1956 but it has not re-colonised this area.
The site was sheep grazed throughout the nineteenth century although this declined after
1931 and since then it has had periods of heavy and light grazing by sheep, rabbits and cattle
with occasional mowing and raking of cut material to maintain a tight sward. Few plants were
recorded during the 1970s due to the decline in the rabbit population and the growth of
coarse grasses (Bromopsis erectus). However, the population increased dramatically following
the introduction of mowing and the recovery of the rabbit population in the 1980s although
these caused severe localised erosion in places (e.g. Spiranthes Bank) (Bailey, 1996). The Hoe
itself is currently heavily-grazed by rabbits with additional winter sheep grazing and
management is now considered to be ideal with over 1000 flowers counted in 2005 (G.
Stevens, pers. comm.). In contrast, numbers within the two less heavily grazed exclosures
remain low due to greater competition with coarse grasses. The site receives few visitors but
there are reports of over 1000 plants being dug-up in the past (Hope-Simpson, 1948).
Ravensburgh Castle (SSSI)
In the 1960s two sub-populations were recorded from adjacent chalk slopes of an Iron Age hill
fort dating back to 400BC. The smaller of the two populations occurred immediately below
the fort on a steep W-facing slope (TL098295); the larger colony extended eastwards from a Sfacing spur overlooking the first site (TL097296). Over 300, mainly vegetative plants were
recorded on this slope in 1981 and 30 below the hill fort (Welsh, 1983). In 1964 the grassland
was very rank but by 1980 the vegetation was very short with much bare chalk due to heavy
grazing by sheep and rabbits. It has not been grazed since and is now dense Bromopsis erectus
grassland with scattered hawthorn scrub. As a consequence, no plants have been seen on the
W-facing slope for many years and only a few plants survive on the S-facing spur where the
grassland is more exposed and the soils are very shallow.
Smithcombe Hill (SSSI)

44

In the 1960s a small population occurred in rank Bromopsis erectus grassland (CG3a)
surrounded by scattered Viburnum lantana scrub on a steep SSE-facing chalk slope. By the
1980s the scrub had become very dense and only a single vegetative rosette could be found
(Welsh, 1983). It has not been recorded since despite scrub clearance and the introduction of
goat grazing.
NORTHAMPTONSHIRE (v.c.32)
Barnack Hills and Holes (NNR)
Many thousands of plants (>20,000) occur over a large area of mediaeval limestone quarry
workings where they are confined to shallow soils on S and SW-facing slopes. Grazing ceased
in 1914 and by the 1960s the grassland was very rank Bromopsis erectus-Brachypodium
pinnatum grassland with a dense layer of leaf litter. Sheep grazing (September to lateDecember) was re-introduced in 1978 resulting in a dramatic increase in flowering. Monitoring
in fixed plots showed an increase from 260 in 1978 to 4,727 crowns by 2005. The entire
population is now estimated to exceed 20,000 crowns (C. Gardiner, pers. comm.). The
management is now considered ideal (Rich, 1997) and as a result P. vulgaris has spread to new
areas of the reserve where it may have established from seed (C. Gardiner, pers comm.).
Southorpe Paddock (SSSI)
In the 1960s a small colony was recorded in rough Brachypodium pinnatum grassland. The site
has received variable amounts of grazing in the past and the last confirmed record was in
1995. In 1999 12 plants originating from Barnack seed were planted out but none survived for
more than a year (C. Gardner, pers. comm.).
Southorpe Roughs (SSSI)
A few plants formerly occurred in rank Bromopsis erectus-Brachypodium pinnatum grassland
in an area of old quarry workings. The site was not grazed for many years and by the 1980s
plants were very difficult to locate amongst the rank grass and invading scrub. It was
apparently last seen by Franklyn Perring in 1990 (L. Farrell in litt. to T.C.E. Wells). It has not
been seen since despite the recent reintroduction of sheep grazing (2004) and tree felling to
reduce shade.
Sutton Heath
A single flowering plant was discovered on a small ridge of limestone turf close adjacent to
Sutton Bog in 1972 (Wells, 2003) where it was seen again in 2003. The grassland shows signs of
agricultural improvement and is heavily grazed. No plants have been found since despite
repeated searches, although Astragalus danicus still occurs on the same ridge.
EAST GLOUCESTERSHIRE (v.c.33)
Barnsley Wold Warren (SSSI)
The largest population in the UK with the majority of plants occurring at high density in
exceptionally rich limestone turf on a steep (c.20) SW-facing slope where soils are very
shallow and there is much bare soil and rubble. Small numbers of plants have also occurred
sporadically on a W-facing slope nearby. Historically the site has been heavily grazed by
sheep, cattle and horses and the vegetation burnt (swaled) in February or March. This
practice was abandoned in 1971; since then the site has been grazed by sheep, at high
density, between February to March and then from June onwards. This management has

45

resulted in an increase from around 50,000 to an estimated 75,000 crowns (D. Barling, pers.
comm.).
Beaumonts Hay (SSSI)
This site forms part of the Brassey Reserve and Windrush Valley SSSI and supports a small
population of Pulsatilla on a relatively steep SW-facing limestone slope. Grazing has been
increased in recent years producing a much shorter, species-rich sward.
Bourton Down (SSSI)
Around 500 plants are restricted to a small area of sheep-grazed, W-facing downland with
extremely shallow soils. Historically the site has been heavily grazed by sheep, although since
1982 stock have been removed from March to late summer in order to allow Pulsatilla to
flower and currently the population appears stable. This management is similar to Barnsley
Wold but with more variable winter grazing (Rich, 1997). In the 1966 two patches of Pulsatilla
were recorded in ungrazed Brachypodium pinnatum grassland about 1 km to the NW of the
reserve (SP132321) but this area is now dense hazel scrub.
Hornsleasow Roughs (SSSI)
At this site Pulsatilla is scattered across a large area of former limestone quarries (hills and
holes) that have had periods of heavy (1960s) and light grazing (1970-80), mostly by sheep
but with a few cattle. It is currently lightly grazed by sheep (<3 ha-1) for most of the year
(excluding March-May) and although this is not considered ideal management there are still
around 100 plants scattered over a hectare of grassland but at low density. Since the
relaxation in grazing the grassland changed from tightly grazed (c.2 cm) Festuca ovina turf
(CG2d) to moderately rank (c.6 cm) but species-rich Bromopsis erectus grassland (CG3).
Sweetslade Farm
The majority of this site was ploughed-up in 1966. However, a small area of short limestone
grassland (CG5a) was preserved to protect a few plants of Pulsatilla on WNW-facing slope.
There are no subsequent records and the site is now completely covered by gorse scrub
(M.A.R. and C. Kitchen, pers comm.).
Taylors Hill
Although this population was originally discovered in 1847 it was not seen again until C.S.
Downer found a few plants growing on a SW-facing limestone slope in 1971 (Holland, 1986).
These had increased to 110 plants by 1982 whereas only 15 were located over a 200 m stretch
of downland in 1996 (SP003157 to SP005156) (M.A.R. and C. Kitchen, pers comm.). This site
currently has no conservation designation.
WEST GLOUCESTERSHIRE (v.c.34)
Rodborough Common (SSSI)
In the 1960s Pulsatilla was recorded from two adjacent limestone slopes on Bear Hill
(SO849027 and SO847029) and in rough grassland below the Iron Age hill fort on Rodborough
Hill (SO848038). One of the colonies on Bear Hill still survives amongst rough Brachypodium
pinnatum grassland (CG4) on a gentle S-facing slope. The other sites were lost due to scrub
encroachment and lack of grazing (M.A.R. and C. Kitchen, pers comm.). Historically there has
been light grazing by sheep and cattle and occasionally burnt (swaling) in February or March
by commoners but both practices have declined in recent decades. As a consequence the Bear
Hill slope was mown in the 1990s and since 1999 winter grazing by cattle has been re-

46

introduced by the National Trust leading to a reduction in the cover of Brachypodium

pinnatum and a gradual increase in the numbers of Pulsatilla.
SOUTH LINCOLNSHIRE (v.c.53)
Ancaster Valley (SSSI)
A small colony occurs in lightly grazed limestone grassland (CG3) on an E-facing slope of a
narrow limestone valley. The site was not grazed for many years leading to the development
of rank Brachypodium pinnatum grassland and dense gorse scrub. The latter was removed
during the late 1980s and the site has subsequently been annually winter-grazed by c.30 sheep
(J. Welhan, pers. comm.). Due to improvement in the condition of the grassland 68 plants
(originating from Ancaster seed) were re-introduced to the same slope (in 1992, 1993, 1998
and 2002) where they are protected by wire cages during the flowering period. All plants have
been monitored annually since the early 1990s; in 2005 6 native plants and 22 (32%)
introduced plants remained.
Holywell Mound (SSSI)
In the 1960s around 200 plants were present in species-rich limestone turf on a SW-facing
slope although few plants flowered due to heavy grazing and the occasional picking of
flowers by local school children (J. Gibbons in litt. to T.C.E. Wells). In the 1960s the grassland
was very short, herb-rich Bromopsis erectus-Brachypodium pinnatum grassland (CG5) with
abundant Thesium humifusum. A few plants survived into the 1980s although the site was
subsequently sprayed and partially improved. Pulsatilla has not been recorded for many years
and the grassland is now a species-poor Bromopsis erectus grassland (CG3b) with few herbs.
Honington Camp (SSSI)
In the 1960s a small colony occurred in rank Bromopsis erectus grassland (CG3d) on the Wfacing outer embankment of an Iron Age hill fort surrounded by arable land. The site has been
infrequently grazed leading to the development of a thick sward of Brachypodium pinnatum
amongst scattered Crataegus scrub. No plants have been seen since 1992 despite repeated
searches.
Swinstead Valley (SSSI)
On 25th April 2009, a single flowering plant of Pulsatilla vulgaris was discovered at a new site
by Richard Jefferson and Fiona Hart in Swinstead Valley, South Lincolnshire (Jefferson &
Walker, 2010). Subsequent searches revealed a total of four plants in 2009 and 18 in 2010,
following the erection of a temporary grazing exclosure. This population is located on a
moderately steep (c.30) west-facing slope (280-310N) of limestone grassland on the east side
of a narrow valley where the vegetation had been kept very short by rabbit and sheep grazing.
The vegetation is species-rich Brachypodium pinnatum-Bromus [Bromopsis] erectus grassland
(74% fit to CG5a, typical sub-community) with abundant Bromopsis erectus, Festuca ovina,
Helianthemum nummularium, Sanguisorba minor and Thymus polytrichus and Astragalus
danicus growing close-by.
NORTH LINCOLNSHIRE (v.c.54)
Broughton Far Wood (Clap-gate Pit) (SSSI)
In the 1960s 3-4 plants were recorded amongst ungrazed grassland and scrub in an abandoned
limestone quarry. The plants were last recorded in 1969 when they were dug-up (I. Weston,
pers. comm.).

47

LEICESTERSHIRE (v.c.55)
Shacklewell Hollow (SSSI)
In the 1960s a small colony was recorded in species-rich limestone grassland (CG5a) on a
shallow SW-facing slope. Three plants were re-discovered at this site in 1992 when the
grassland was very rank due to the lack of grazing. However, the population was subsequently
destroyed when the farmer dumped rubble over the area (D. Isaac, pers. comm.).
MID-WEST YORKSHIRE (v.c.64)
Ledsham (SSSI)
In 1984 a single plant was discovered in heavily grazed limestone grassland (CG4a) on a SEfacing limestone bank. For many years flower heads were grazed-off and so cages were placed
around the plant to protect it from rabbits. In 1997 the plant was hand-pollinated with pollen
collected (the same day) from Barnack Hills and Holes but the plant failed to set seed due to
predation by slugs. As a result pollen from the Ledsham plant (male parent) was used to crosspollinate five plants from Barnack (site 4) but without success (A. Headley, pers. comm.). A
single Barnack plant was then translocated to within 4 metres of the Ledsham plant in order
to encourage pollination but disappeared within a couple of years.

48

13

Annex 2 changes in population size, 1960-2006

Changes in the numbers of Pulsatilla vulgaris on 33 sites included in Walker et al. (in press).
Totals are the maximum number of crowns recorded in any one year for each decade and
were derived from a variety of sources.
Site-population
(a) Increasing
Barnack Hills and Holes
Barnsley Wold
Barton Hills
Knocking Hoe
Therfield Heath

1960-69

1970-79

1980-89

1990-99

2000-06

2000
50000
1100
300
1000

1300
50000
685
50
2897

14200
50000
4000
460
?

15000
50000
5000
300
60000

20000
75000
5000
1000
60000

(b) Stable
Ancaster Valley1
Beaumonts Hay
Bourton Down
Deacon Hill
Devils Dyke
Hornsleasow Roughs
Taylors Hill
Ledsham 3

6
<100
500
1
250
200
?
?

5
?
?
?
1500
?
42
?

16
25
500
5
1000
?
110
2

11
?
?
7
500
100
15
2

6
10
471
1
250
100
?
1

(c) Declining
Aston Upthorpe2
Ravensburgh Castle
Rodborough Common
Steps Hill

300
39
300
500

101
0
3
100

35
300
?
30

10
5
40
10

26
4
37
52

(d) Extinct
Aldbury Nowers
17
0
0
0
Broughton Far Wood
9
0
0
0
Clipper Down
15
1
?
?
Fleam Dyke
25
2
0
0
Hildersham Furze Hills
4
14
5
1
Holywell Mound
200
?
1
?
Honington Camp
53
?
10
1
Lockinge
2
?
?
?
Pitstone Hill
1
1
?
2
Shacklewell Hollow
56
?
?
3
Smithcombe Hill
30
?
1
0
Southorpe Paddock
8
2
3
5
Southorpe Roughs
5
6
10
Present
Sweetslade Farm
7
?
?
0
Sutton Heath
?
1
?
?
Unhill Bottom
750
3
0
0
1
Excludes 68 plants introduced since 1992. 2 Excludes 87 plants introduced since 1999.

0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0

49

14

Annex 3 associates

Vegetation composition and structure of Pulsatilla vulgaris populations in England (in 16 of

the 18 extant populations - Taylors Hill & Swinstead Valley not included). For associates
figures are frequency in quadrats (n = 62) with the mean % cover given in parentheses. Species
recorded in <10% of quadrats are listed after the table. Constant species (> 80 % of quadrats)
are highlighted in bold. For composition and structure variables figures are means with 1SE
in parentheses. Figures are presented for (a) all 16 sites and for (b) limestone and (c) chalk
sites only (n = 7 and 9 respectively).
All sites
(a) Associates
Pulsatilla vulgaris
Bromopsis erectus
Festuca ovina
Sanguisorba minor
Helianthemum nummularium
Carex flacca
Cirsium acaule
Leontodon hispidus
Pilosella officinarum
Briza media
Helictotrichon pratense
Koeleria macrantha
Carex caryophyllea
Brachypodium pinnatum
Ctenidium molluscum
Lotus corniculatus
Campanula rotundifolia
Scabiosa columbaria
Plantago lanceolata
Scleropodium purum
Filipendula vulgaris
Thymus polytrichus
Thymus pulegioides
Centaurea nigra
Plantago media
Polygala calcarea
Hippocrepis comosa
Campyliadelphus
chrysophyllus
Campanula glomerata
Viola hirta
Homalothecium lutescens
Asperula cynanchica
Ranunculus bulbosus
Carlina vulgaris

Limestone only

Chalk only

V
V
V
V
V
V
V
IV
IV
IV
IV
IV
IV
III
III
III
III
III
III
II
II
II
II
II
II
II
II

(4.2)
(24.2)
(12.9)
(8.8)
(6.0)
(4.2)
(4.2)
(2.8)
(2.5)
(1.5)
(1.3)
(1)
(0.9)
(5.1)
(3.6)
(1.2)
(0.9)
(0.8)
(0.7)
(2.0)
(1.6)
(1.0)
(1.0)
(0.8)
(0.6)
(0.6)
(0.6)

V
V
IV
V
V
V
V
IV
IV
IV
III
III
IV
V
III
III
II
III
III
III
I
III
II
II
II
II
I

(3.6)
(24.7)
(8.5)
(8.7)
(7.2)
(3.7)
(5.3)
(2.8)
(3.2)
(1.9)
(0.6)
(0.6)
(0.9)
(7.4)
(3.4)
(1.6)
(0.9)
(1.1)
(0.7)
(3.5)
(0.5)
(1.8)
(1.0)
(0.4)
(0.9)
(0.9)
(0.5)

V
V
V
V
IV
V
V
IV
III
III
V
V
IV
I
III
III
V
II
III
II
IV
I
III
III
III
I
III

(4.9)
(23.7)
(17.6)
(8.9)
(4.7)
(4.7)
(3.1)
(2.9)
(1.8)
(1.0)
(2.0)
(1.4)
(0.9)
(2.7)
(3.7)
(0.8)
(1.0)
(0.5)
(0.6)
(0.3)
(2.9)
(0.1)
(1.0)
(1.3)
(0.4)
(0.3)
(0.8)

II
II
II
II
II
II
II

(0.5)
(0.5)
(0.3)
(0.3)
(0.2)
(0.2)
(0.2)

II
I
III
II
II
II
II

(0.2)
(0.2)
(0.3)
(0.4)
(0.2)
(0.3)
(0.3)

I
II
II
I
II
II
I

(0.9)
(0.8)
(0.3)
(0.1)
(0.2)
(0.2)
(0.1)

50

Annex 3 continued.
All sites
Pimpinella saxifraga
Linum catharticum
Crataegus monogyna
Galium verum
Anthyllis vulneraria
Primula veris
Fissidens sp.
Thesium humifusum
Fissidens cristatus
Succisa pratensis
Tephroseris integrifolia
Taraxacum officinale agg.
Prunella vulgaris

II
II
II
II
I
I
I
I
I
I
I
I
I

(b) Vegetation composition/structure

Vegetation height (cm)
7.5
Bare ground (%)
6.2
Number of species
23.0
Number grasses & sedges
6.2
% cover of grasses & sedges
49.1
Number herbs
14.6
% cover of herbs
44.1
Diversity
1.0
Evenness
0.7
% cover of Bromopsis erectus
24.2
Number of positive indicators
10.2
% cover of positive indicators
63.3
% fit to CG3 or CG5
60.5

Limestone only

(0.2)
(0.1)
(0.1)
(0.1)
(0.6)
(0.4)
(0.3)
(0.2)
(0.2)
(0.2)
(0.2)
(0.1)
(0.1)

II
III
II
II
II

(0.2)
(<0.1)
(0.1)
(<0.1)
(1.1)

II
II
I
I

(0.3)
(0.4)
(0.1)
(<0.1)

II
I

(0.7)
(1.2)
(0.7)
(0.2)
(1.9)
(0.6)
(2.1)
(0.0)
(0.0)
(2.4)
(0.4)
(2.5)
(2.0)

7.0
7.0
23.1
6.4
46.0
14.0
46.0
1.1
0.8
24.7
10.4
69.1
54.7

Chalk only
I
II
II
I

(0.2)
(0.1)
(0.2)
(0.2)

(0.1)
(<0.1)

II
I
I
I
I
II
I
II

(0.9)
(0.2)
(<0.1)
(0.3)
(0.4)
(0.5)
(<0.1)
(0.2)

(0.8)
(1.7)
(1.1)
(0.3)
(3.0)
(0.9)
(3.0)
(0.1)
(0.0)
(3.8)
(0.5)
(3.8)
(2.7)

7.9
5.3
22.8
6.0
52.0
15.2
42.0
1.0
0.7
23.7
10.0
57.1
66.8

(1.1)
(1.6)
(1.0)
(0.2)
(3.0)
(0.9)
(3.0)
(0.0)
(0.0)
(2.8)
(0.5)
(3.0)
(2.5)

Sites and number of quadrats recorded (1968, 2006): Ancaster Valley (2,1), Aston Upthorpe (1,1),
Barnack Hills and Holes (4,3), Barnsley Wold (3,2), Barton Hills (2,4), Beaumonts Hay (2,1), Bourton
Down (3,2), Devil's Dyke (2,2), Hornsleasow Roughs (1,2), Knocking Hoe (3,1), Knocking Hoe - Spiranthes
Bank (1,1), Ravensburgh Castle (2,1), Rodborough Common (4,1), Steps Hill (1,1), Therfield Heath Pen
Hill (2,1), Therfield Heath - Church hill (3,1). Species recorded in less than 10% of quadrats: Aceras
anthropophorum, Arrhenatherum elatius, Blackstonia perfoliata, Carex ericetorum, Centaurea
scabiosa, Cerastium fontanum, C. pumilum, Cirsium arvense, C. vulgare, Cladonia sp., Cynosurus
cristatus, Dicranum scoparium, Euphrasia nemorosa, Festuca rubra, Galium aparine, G. mollugo,
Gentianella amarella, Gymnadenia conopsea, Helictotrichon pubescens, Heracleum sphondylium,
Hieracium exotericum, Himantoglossum hircinum, Hylocomium splendens, Hypnum cupressiforme,
Hypochaeris maculata, H. radicata, Leucanthemum vulgare, Luzula campestris, Medicago lupulina,
Neckera complanata, Onobrychis viciifolia, Ononis repens, Ophrys apifera, Orchis mascula, O. morio,
Pastinaca sativa, Phleum bertolonii, Picris hieracioides, Poa compressa, Polygala vulgaris, Polytrichum
sp., Prunus spinosa, Quercus robur, Rhamnus cathartica, Rhinanthus minor, Rhytidiadelphus
squarrosus, R. triquetrus, Rosa canina, Seligeria sp., Senecio jacobaea, Serratula tinctoria, Seseli
libanotis, Sonchus asper, Thuidium assimile, Trifolium pratense, T. repens, Trisetum flavescens,
Veronica chamaedrys, Weissia sp.

51

Written by Kevin Walker, BSBI

Edited by Plantlife International
First draft dated Sept 2010
Last revised 19 April 2011

52