WATSONIA

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Watsonia, 17, 121-132 (1988) 121

Phenotypic plasticity of Elodea nuttallii (Planch.) H. St John and
Elodea canadensis Michx in the British Isles

D. A. SIMPSON

Royal Botanic Gardens, Kew, Richmond, Surrey, FW9-3AB

ABSTRACT

Elodea nuttallii (Planch.) H. St John and E. canadensis Michx show a wide range of morphological Variation, a
feature which has caused many problems of identification. In the British Isles variation within the two species is
entirely due to phenotypic plasticity. The features which most obviously demonstrate plasticity are leaf shape,
leaf size, leaf posture, internode length and plant colour. Some of these have been incorrectly used in the past
as diagnostic characters. The range of variation in these features is described. Field studies are reported which
examine the variation of material in two different localities during the growing season, and cultivation
experiments are described in which material from one genotype of each species was grown under a variety of
environmental conditions. Possible factors influencing plasticity are discussed.

INTRODUCTION

Three species of the genus E/odea Michx occur in the British Isles, namely E. canadensis Michx, E.
nuttallii (Planch.) H. St John and E. callitrichoides (Rich.) Casp. All are adventive-in the British
Isles and are native to the New World.

E. nuttallii and E. canadensis show a wide range of morphological variation, even though only a
few genotypes of each species occur in the British Isles (Cook & Urmi-K6nig 1985). This has
caused many problems of identification, and such problems were particularly evident in Great
Britain during the 1960s and 1970s, a period coinciding with the introduction and spread of E.
nuttallii. This species, first reported from Oxford in 1966 (Simpson 1984), became widespread
during the following ten years and its range of variation was great enough to suggest the possibility
of two or even three new taxa being present. In addition some material could have been a hybrid
between the two species. It was known that in America E. canadensis and E. nuttallii readily
hybridize with each other (Ernst-Schwarzenbach 1945a, 1945b; Cook & Urmi-K6nig 1985), a
feature which probably accounts for some of the variation in American specimens (Cook & Urmi-
K6nig 1985), and could also have applied to British and Irish material. It was also suggested that
material assigned to E. nuttallii was merely a phenotype of E. canadensis which had appeared as a
response to changing environmental conditions, particularly the increasing eutrophication of water
bodies (S. Haslam, pers. comm.; Harding 1979).

However, in my recent investigation of Elodea in the British Isles (Simpson 1983), it was
concluded that FE. canadensis and E. nuttallii are two distinct species, which cannot hybridize
because only female plants are present. It was also shown that both species exhibit a wide range of
phenotypic plasticity, which would explain the problems of identification. Aspects of phenotypic
plasticity in Japanese material of E. nuttallii have been examined by Kunii (1982, 1984) who
concentrated particularly on growth and phenology. In this paper I describe the range of
phenotypic plasticity shown by British material of E. nuttallii and E. canadensis, with particular
emphasis on features of taxonomic interest, such as leaf shape and size. I also report field and
laboratory studies which were carried out (i) to demonstrate the range of phenotypic plasticity in
the species in two localities over the growing season, (11) to examine the effect of differing
environmental conditions on their plasticity, and (iii) to determine whether the two species adopt a
similar morphology under certain conditions. Possible factors which influence their plasticity are
also discussed.

122 D. A. SIMPSON
MATERIALS AND METHODS

INITIAL OBSERVATIONS

Before carrying out experimental work it was necessary to identify which parts of the plant were
showing phenotypic variation. This was done by examining a range of living and herbarium
material from a variety of different habitats. A full list of the latter is kept in LANC and a list of the
herbaria consulted is given in Simpson (1986).

FIELD OBSERVATIONS

Field observations were made predominantly on E. nuttallii. The aim was to record the phenotypic
plasticity of plants at four different depths on four dates during the growing season. Two
contrasting sites were chosen, one in a shallow section of the R. Lune at Skerton, Lancaster,
Lancs., v.c. 60, and the other in deeper water at Mitchell Wyke Bay, Windermere, Westmorland,
v.c. 69. Whole plants were taken from 0-5-1 m depth in the R. Lune and 2-3 m depth in Mitchell
Wyke Bay. Observations were made on the uppermost stems of these plants, which were gathered
on 25 April, 15 June, 18 August and 23 September 1981. On 23 September, material of E.
canadensis was collected for comparison. In this case the plants were taken from the Lancaster to
Kendal canal at Burton-in-Kendal, Westmorland, v.c. 69, as well as from Mitchell Wyke Bay.
Light intensity was measured at both sites in terms of quantum flux density. Measurements were
made at the depths of the uppermost stems with a ‘Lambda’ quantum sensor adapted for
underwater use. Light intensities received by the uppermost stems of the plants varied from nearly
100% of the surface intensity for plants at 0-5 min the R. Lune and the Lancaster-Kendal Canal
(i.e. the uppermost stems were at or near the surface), to only 8% for plants at 3 m depth in
Windermere. In all cases ten plants from each depth were collected and pressed. Leaf shape, leaf
posture and plant colour were noted. Measurements were made of leaf length, leaf width at the
mid-point and internode length. Length and width of five cells in each leaf were also measured
from plants at 0-5 m in the R. Lune, and at 2 and 3 m in Mitchell Wyke Bay. All leaf data were
taken from the verticillate leaves only.

During the latter part of the fieldwork a simple transplant experiment was carried out, in which
material was transferred from shallow to deep water. Ten stem apices each 10 cm long from one
genotype of each species were planted in sand/gravel in plastic containers. One container of each
species was then placed at 2, 3 and 4 m depths in Mitchell Wyke Bay and grown for 35 days
between 18 August and 23 September 1981. The plants were obtained from c.0-5 m depth in the R.
Lune (E. nuttallii) and the Lancaster-Kendal Canal, Burton-in-Kendal (E. canadensis). Plants
growing at these localities were examined during the course of the experiment for comparison. The
characters listed above were recorded from growth which had taken place during the 35 day
period.

CULTIVATION EXPERIMENTS

Three simple laboratory experiments were carried out in which material representing one genotype
of each species was subjected to a variety of environmental conditions. The apparatus for each
experiment was similar, consisting of polypropylene bins filled with nutrient-poor sand/gravel or
nutrient-rich river sediment and tap water. The experiments were as follows.

Plants of both species were grown for 35 days under low, medium or high light intensities (2, 35
and 250 umol m~* s~') in either sand/gravel or river sediment at c.19°C. Reduced light
intensity was achieved by shading the bins with neutral density filters. Light intensities were chosen
to reproduce, as near as possible, those in the two habitats during August and September.

E. nuttallii was grown for 30 days at high light intensity (250 ymol m~s~') in sand/gravel or
river sediment at a raised water temperature of c.27°C.

Plants of both species were grown for 30 days in sand/gravel with different levels of water
nutrient enrichment. This was achieved by adding ‘Phostrogen’ 10:10:27 N:P:K fertiliser in three
different concentrations.

Ten stem apices 10 cm long were used in each treatment. These were obtained from the same
source as material used in the transplant experiment. All observations were made on growth which
had taken place during the course of the experiment. Again, material growing at the two sources
was examined during the experiment for comparison. In the first two experiments observations

PHENOTYPIC PLASTICITY OF ELODEA 123

were made on the same characters as those examined in the field work. In the third experiment leaf
shape, leaf apex shape, leaf width c.0-5 mm below the apex and plant colour were recorded. Leaf
data were again taken from the verticillate leaves. Voucher material of all field and experimental
work is in LANC.

RESULTS

RANGE OF PHENOTYPIC PLASTICITY IN E. CANADENSIS AND E. NUTTALLII

Phenotypic plasticity in both species is seen in the vegetative parts of the plant. The characters
which most obviously demonstrate plasticity are: leaf shape, leaf size, leaf posture, internode
length and plant colour.

Leaf shape has often been used as a diagnostic character for separating Elodea species, although
in many cases it has been inadequately described (cf. St John 1965). It is most variable in E.
canadensis, in which the leaves range from ovate to linear-oblong and occasionally linear-
lanceolate. In E. nuttallii the leaves are either linear or linear-lanceolate. Of particular taxonomic
importance is the shape of the leaf apex, which is broadly acute to obtuse in E. canadensis and
narrowly acute to acuminate in E. nuttallii. This discontinuity is an important character for
separating the two species.

Leaf length and mid-point width are highly variable in both species. The greatest variation of
leaf length is seen in E. nuttallii, in which the leaves may be up to 35 mm long. However, mid-point
width varies most in E. canadensis (Table 1). In many cases the leaves of E. nuttallii are longer and
narrower than those of E. canadensis, although there is a wide range of overlap between them.
Such overlap makes these characters of little use in separating the two species. However, leaf size
does yield one taxonomically important feature, namely the width of the leaf measured at a point
c.0-5 mm below the apex. This is usually greater in E. canadensis and reflects differences shown by
the shape of the leaf apex. It is the most reliable character for separating this species from E.
nuttallii.

Six types of leaf posture may be recognised in Elodea, ranging from patent to strongly recurved
(Simpson 1986). E. canadensis has only three of the types, its leaves being patent, slightly recurved
or spreading. EF. nuttallii has six types, the most frequent being those seen in E. canadensis.
However in E. nuttallii at least some leaves are strongly recurved, with their apices touching the
stem. This type of leaf posture is diagnostic for E. nuttallii.

In both species the internodes become shorter towards the apex. However internode length at
any given point along the stem is highly variable, and this appears to be closely associated with
variation in leaf size. Thus plants with the longest, narrowest leaves also have the longest
internodes and vice-versa. The greatest range of variation is seen in EF. nuttallii, which can have
internodes up to 20 mm long (Table 1).

Plant colour has often been used as a diagnostic character, with E. canadensis being described as
dark green and E. nuttallii as light green or greenish brown. However, colour variation bears little
relation to species differences and seems to be dependent on light intensity and/or nutrient status
of the water or substrate.

FIELD OBSERVATIONS

The morphology of E. nuttallii in the R. Lune was similar at both depths on 25 April, the plants
having short, broad, strongly recurved, linear-lanceolate, mid- to dark green leaves and short
internodes. As the season progressed the leaves adopted a spreading posture, together with an

TABLE 1. RANGE OF LEAF SIZE AND INTERNODE LENGTH IN ELODEA CANADENSIS AND
ELODEA NUTTALLIT

Character E. canadensis E. nuttallii
Leaf length (mm) 4.5-17.0 5.5-35.0
Leaf mid-point width (mm) 1.4-5.6 0.8-3.0
Internode length (mm) 0.5-9.1 0.5-2.0

124 D. A. SIMPSON

increase, followed later by a decrease, in leaf mid-point width (Figs 1 & 2). The internodes
remained short.

In Windermere the seasonal pattern was different, particularly at 2 m. Plants from both 2 m and
3 m started off the season with a morphology similar to those in the R. Lune, but by 15 June there
was an increase in leaf length and mid-point width. At 2 m there was a continued increase in leaf
length, coupled with a decrease in mid-point width, so that by 18 August the plants had long,
spreading, linear, light green leaves and long internodes. At the same time plants at 3 m adopted a
morphology which was intermediate between that at the start of the season and the plants at 2 m as
described above. The overall pattern was more or less the same on 23 September. None of the
Windermere plants elongated sufficiently to reach the surface of the water.

The differences in external morphology were usually reflected by differences in leaf cell size
(Table 2). The cells of plants from 0-5 and 1 m in the R. Lune were generally longer and broader
than those of the 2 m plants from Windermere. In plants at 3 m there was no definite pattern in cell
size, except that cell width was greater than in plants at 2 m.

E. canadensis showed a similar variation pattern to E. nuttallii on 23 September both in external
morphology and cell size, although the range of variation was not as wide as in E. nuttallii (Fig. 2;
Tables 2 & 3). The longest, narrowest leaved plants were again found at a depth of 2 m. Material
from the canal was dark green, whilst that in Windermere was mid-green.

In the transplant experiment, the containers at 3 m were vandalised and no result was obtained.
At 4m none of the plants survived. This was probably due to their being below the lower depth
limit for the growth of the two species. At 2 m a number of changes were noted (Fig. 3). At the
start of the transplant experiment, material of E. nuttallii had short, broad, mostly recurved, mid-
green leaves and short internodes. After 35 days the plants became similar to wild material at the
same depth, having long, narrow, spreading, light green leaves. A similar transition to longer and
somewhat narrower leaved material was also noted in E. canadensis, and again, the transplanted
material became similar to wild material growing at the same depth. In E. nuttallii internode length
showed little change, whereas in EF. canadensis it was shorter on the new growth. It is likely that the
experimental period was not long enough to allow the internodes to develop fully.

TABLE 2. CELL SIZE IN UPPER STEM LEAVES TAKEN FROM WHOLE PLANTS OF ELODEA
NUTTALLII AND ELODEA CANADENSIS ON 23 SEPTEMBER 1981 (MEAN3=S.E.)

Depth of plant (m)
0.5

Measurement (um) 2.0 3.0

E. nuttallii

Cell length 93.20+2.72 G fo ad (SFA IP) 67.60+3.20
Cell width 36.80+0.80 26.40+0.56 35.20+0.50
E. canadensis

Cell length 109.20+3.71 79 .84+3.13 76.00+2.73

Cell width 46.88+0.76 15.10+0.45 38.80+1.47

TABLE 3. COMPARISON BETWEEN THREE MORPHOLOGICAL CHARACTERS OF ELODEA
CANADENSIS ON 23 SEPTEMBER 1981 (MEAN3+S.E.)

Site and depth of plant (m)

Burton-in-Kendal Windermere
0.5 1.0 2.0 3.0
Leaf length (mm) 7.8+0.1 8.0+0.1 14.1+0.4 6.8+0.1
Leaf mid-point width (mm) 2.9+0.03 2.0+0.03 2.0+0.06 3:332-0:1
Internode length (mm) sitar Us) 8.6+0.2 10.5+0.3 5.6+0.3

PHENOTYPIC PLASTICITY OF ELODEA 125

Leaf length (mm)

Leaf mid-point width (mm)

Internode length (mm)

25 15 18 23
Apr June Aug Sep

Sampling date
FicureE 1. Variation in E. nuttallii of three morphological characters in relation to sampling date and depth.
Points are means of 50 measurements. Bars indicate + standard error. @ = 0-5mR. Lune, O = 1mR. Lune,
A = 2m Mitchell Wyke Bay, A = 3 m Mitchell Wyke Bay.

126 D. A. SIMPSON
R. Lune Mitchell Wyke Bay

Lancaster — Kendal Canal

14

Ficure 2. A: Silhouettes of upper stems from whole plants of Elodea nuttallii collected in relation to sampling
date and depth; B: Silhouettes of upper stems from whole plants of Elodea canadensis collected on 23
September in relation to depth. Scale bar = 10 mm.

| | .

m 2m 3m

PHENOTYPIC PLASTICITY OF ELODEA | ed

nutt. can.
nutt. can. nutt. can.
20 2.0

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0 0
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DAYS DAYS DAYS

Ficure 3. Changes in three morphological characters of Elodea nuttallii and Elodea canadensis 35 days after
transplanting into Mitchell Wyke Bay. Columns are means of 20 measurements. Bars = + standard error.

CULTIVATION EXPERIMENTS

At the start of all the cultivation experiments material of both species was identical in morphology
to that at the beginning of the transplant experiment. After 35 days in nutrient-poor sand/gravel at
19°C, E. nuttallii showed an overall increase in leaf length, together with a decrease in mid-point
width and internode length (Fig. 4). However, the overall variation pattern was similar to that seen
in wild material on 18 August. In particular, material grown at the medium light intensity had
similar characteristics to the plants at 2 m from Windermere, with long, narrow, spreading leaves.
E. canadensis grown under the same conditions had a similar pattern of variation, with the most
noticeable changes occurring at the middle light intensity. In this species the widest range of
variation was noted in leaf mid-point width.

When grown in nutrient-rich river sediment, plants of both species under the low and medium
light intensities showed a similar variation pattern to those grown in nutrient-poor sediment under
equivalent lighting conditions. However, at high light intensity, light transmission below water
level was reduced due to excessive phytoplankton growth. Plants grown under these conditions
were similar to those at the medium light intensity.

Under conditions of raised water temperature, the responses shown by E. nuttallii varied with
the type of substrate (Fig. 5). In both cases the plants developed linear, spreading leaves.
However, plants in nutrient-poor substrate had shorter leaves and internodes. Incidental
observations of E. canadensis under similar conditions indicated that it developed short and
markedly narrow, linear-oblong or rarely linear-lanceolate leaves.

Plants grown in nutrient-poor sand/gravel at medium and high levels of water-nutrient
enrichment showed some of the detrimental effects of such enrichment, including loss of leaves on
the lower part of the stem. After 30 days the morphology of both species was similar to
corresponding material grown under raised temperature and high light intensity conditions (Table
4). These forms developed at all levels of water-nutrient enrichment.

DISCUSSION

A wide range of phenotypic plasticity is characteristic of many aquatic taxa and this has caused
numerous taxonomic difficulties within such groups. A notable example is that of Ranunculus L.
subgen. Batrachium (DC.) A. Gray, in which Cook (1966) found a total of 302 possible names for
the 17 species that he recognised. In Elodea, St John (1965) delimited 17 species of which ten were

128 D. A. SIMPSON

E nuttallii E canadensis
24 B
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0 0 - 35
Figure 4. Changes in three morphological characters of Elodea nuttallii and Elodea canadensis after 0 and 35

days growth in differing light intensities in nutrient-poor and -rich substrates. A: high light intensity; B:
medium light intensity; C: low light intensity. Shaded columns = nutrient-poor substrate, unshaded columns =
nutrient-rich substrate. Columns are means of 20 measurements. Bars = + standard error.

PHENOTYPIC PLASTICITY OF ELODEA 129

20 2.0 20

Leaf length (mm)
=)
Leaf mid-point width (mm)
=
Intermode length (mm)
=)

0 0. 0
0 30 0) 30 0 30

Days Days Days

Ficure 5. Changes in the morphology of Elodea nuttallii after 0 and 35 days growth at high light intensity at
raised water temperature in nutrient-poor and -rich substrates. Shaded columns = nutrient-poor substrate,
unshaded columns = nutrient-rich substrate. Columns are means of 20 measurements. Bars = + standard
error.

newly described. Unfortunately he based his work purely on herbarium material, which restricted
his interpretation of the range of phenotypic plasticity. However, this is taken into account in the
revision of Cook & Urmi-Konig (1985) who reduced the number of species to five.

From the present work it is apparent that FE. nuttallii is the most variable of the species in the
British Isles, particularly in leaf size and internode length. The variation is such that the two
extremes of its morphological range have the appearance of distinct taxa, one with long, narrow,
spreading leaves, the other with shorter, broader, strongly reflexed leaves. Narrow-leaved
material, such as that seen in the Cumbrian lakes, is almost indistinguishable from E.
callitrichoides, and microscopic characters, such as the length of the teeth on the leaf margins, are
needed to separate them vegetatively. Furthermore, in the British Isles E. nuttallii has been
confused with species in closely related genera, such as Hydrilla verticillata (L.f.) Royle and
Lagarosiphon major (Ridley) Moss. Indeed for many years British and Irish material of the former
was treated as being conspecific with E. nuttallii (Tutin 1962; Scannell & Webb 1976). However
close observation shows that the two extremes of variation often occur on the same plant. These
represent responses to environmental conditions which prevail during the growth of a given part of

TABLE 4. LEAF MORPHOLOGY OF ELODEA NUTTALLII AND ELODEA CANADENSIS AFTER
30 DAYS GROWTH AT THREE LEVELS OF WATER NUTRIENT ENRICHMENT (MEANGS.E.)

Concentration of added nutrient (mg/l)

Nutrient Low Medium High

N 0.025 0.05 0.10

P 0.022 0.044 0.088
Character K 0.055 0.11 0.22
E. nuttallii
Leaf shape linear linear linear
Leaf apex shape acuminate acuminate acuminate
Leaf width 0.5 mm below apex (mm) 0.4+0.02 0.5+0.03 0.4+0.03

E. canadensis

Leaf shape

Leaf apex shape

Leaf width 0.5 mm below apex (mm)

linear-oblong
broadly acute
1.1+0.03

linear-oblong
broadly acute
1.040.04

linear-oblong
broadly acute
1.0+0.03

130 D. A. SIMPSON

the plant. Unfortunately such differences are rarely detected on herbarium specimens since only
small sections of the plant are usually collected. This has undoubtedly emphasised the taxonomic
difficulties associated with E. nuttallii, and indicates the need to have living material for
identification wherever possible.

E. canadensis has a narrower range of variation than E. nuttallii except in leaf shape, and has not
usually been confused with other species in related genera. Nevertheless, problems have
sometimes arisen in distinguishing EF. nuttallii from E. canadensis. In North America, where
hybridization occurs, this would be expected. However in the British Isles, where hybridization
cannot take place, such problems have come about because knowledge of the characters separating
the two species was inadequate.

As indicated earlier, it has also been suggested that FE. nuttallii is merely a phenotype of E.
canadensis. In relation to this, three points should be made about the comparative variation of the
two species and their distinctness under differing environments. Firstly, both species respond in a
broadly similar manner to varying environmental conditions. This is exemplified by wild material
from different depths late in the growing season. Thus in both species the leaves tend to be shorter
and broader in shallow water, longer and narrower in deeper water and intermediate between
these phenotypes at the deepest levels. The two species also show similar variation in leaf anatomy.
Secondly, the wider range of plastic response shown by E. nuttallii is demonstrated in the field by
the differences between this species and E. canadensis on 23 September, and experimentally by the
variation they show in relation to light intensity. From this the third, and taxonomically important,
point is that when the responses of the two species to each environmental factor are compared,
both E. canadensis and E. nuttallii maintain distinct morphological facies. These facies are based
on a combination of leaf shape, leaf apex shape and leaf width c.0-5 mm below the apex-tip (Table
5), and represent a clear morphological discontinuity between E. canadensis and E. nuttallii. There
is, therefore, no justification for treating E. nuttallii as conspecific with E. canadensis.

Of the environmental factors influencing phenotypic plasticity of aquatic macrophytes, light is
probably the most significant, particularly in terms of intensity. There is a sharp contrast in light
intensity between shallow and deep water, and the underwater light climate is characteristically
one of deep shade (Spence 1981, 1982). This has a marked effect on plasticity, and the adaptations
of terrestrial plants to sun and shade conditions are well known. Similar adaptations are shown by
aquatic macrophytes (Spence & Chrystal 1970a, b; Spence 1982). Temperature may also be
important and, in common with light, varies according to the time of year and depth of water. It is
also interactive with light, and responses to the two factors may be impossible to separate (Barko et
al. 1982). Other factors which might have an influence include light quality, photoperiodism,
nutrients and ontogenetic drift.

The limited amount of literature on factors influencing phenotypic plasticity in Elodea suggests
that light, temperature, substrate and water nutrients are involved (Adams et al. 1971; Spence &

TABLE 5. COMPARISON OF LEAF WIDTHS 0.5 MM BELOW THE APEX OF ELODEA NUTTALLII
AND ELODEA CANADENSIS UNDER VARYING ENVIRONMENTAL CONDITIONS (MEANGS.E.)

Mean width 0.5 mm below apex (mm)
Environmental variable E. nuttallii E. canadensis

a. Depth (m) 23 September
0:5

0.60.02 1.040.03
1.0 0.6+0.02 1.0+0.03
2.0 0.4+0.01 0.8+0.01
3.0 , 0.6+0.02 ie2--0:03

b. Differing light intensities, nutrient-poor substrate
High 0.4+0.05 1.8+0.16
Medium 0.340.03 0.8+0.04
Low 0.5+0.03 0.9+0.02

c. Differing light intensities, nutrient-rich substrate

High 0.3+0.04 1.0+0.03
Medium 0.2+0.02 0.8+0.01

Low 0.4+0.02 0.9+0.06

PHENOTYPIC PLASTICITY OF ELODEA 131

Dale 1978; Swann 1978; Barko et al. 1982). What might be deduced from previous and present
work about their effect on plasticity in EF. nuttallii and E. canadensis? Of the four factors, substrate
probably has the least influence, although plants grown experimentally in nutrient-poor and
nutrient-rich substrates do show slight variation. In the case of water nutrients, studies by Adams et
al. (1971) of E. canadensis growing in a variety of habitats indicated that with increasing water
nutrient enrichment, the species showed increases in leaf length and internode length, together
with decreases in leaf mid-point width. Unfortunately no account was taken of variation in the light
and temperature conditions of these habitats. In addition, the present work suggests that, when in
cultivation under the same light and temperature regime, there is little response to water nutrient
enrichment in-either species. Although slight organic pollution of the water occurred at both
sampling sites, comparison with equivalent pollution-free sites showed that a similar pattern of
variation was taking place in both species during the growing season.

From both the literature and the present work it is clear that both species react markedly to
varying light intensity and temperature. For example Spence & Dale (1978) noted a general
increase in internode length of E. canadensis with increasing temperatures at low light intensities,
whereas there is a decrease with increasing light intensities at all temperatures. Barko et al. (1982)
indicated a similar relationship by showing that stem length increases with increasing temperature
up to 28°C. Lawrence (1976) noted a decrease in leaf width in E. canadensis maintained at a mean
temperature of 25-6°C, while Swann (1978) observed that E. canadensis plants in a static water
tank growing in temperatures fluctuating between 22 and 30°C showed a 25% decrease in leaf
length. Barko et al. (1982) concluded that E. canadensis is most responsive to differences in light
when at optimum temperature levels and vice versa. In the present work, E. nuttallii and E.
canadensis grown in the same substrate under differing light intensities at 19°C showed variation
somewhat similar to that of wild plants in August and September, particularly in leaf shape and
size. The light and temperature regimes in these experiments were designed, within the limits of
the equipment available, to reproduce conditions equivalent to that of the sampling sites in late
August or September. Therefore it is likely that the phenotypic plasticity shown by the two species
is in response to variation in light and temperature.

The effects of these two factors might be as follows: except at the lowest light intensity, an
interaction between the two factors produces longer, narrower leaves and longer internodes. At
the highest light intensity, a high temperature (c.27°C and above) is required to produce the
longer, narrower-leaved form, but as the light intensity decreases the optimum temperature
similarly decreases. Such high temperatures are rarely experienced by plants in the British Isles.
Therefore material receiving the highest light intensity will usually have short, broad, mid- or dark
green leaves and short internodes. Where light intensity is reduced, ideal conditions will occur for
the production of long, narrow, light green leaves and longer internodes. Thus the shorter,
broader-leaved phenotype typically occurs when the plants are close to the water surface, whereas
the long, narrow-leaved phenotype occurs in deeper water. The latter may also be present near the
surface when light intensity is reduced due to shading or turbidity of the water.

The phenotypes described above demonstrate obvious adaptations to sun and shade conditions.
However, a problem arises in relation to material growing at the lowest light intensities in deep
water, such as that found at 3 m in Windermere. It appears that at the lowest light intensity, plants
show characters which are intermediate between the short, broad-leaved and long, narrow-leaved
phenotypes. This suggests that another environmental factor, not previously considered, might be
involved. One such factor is pressure. It has been argued that pressure may limit the depth
distribution of aquatic macrophytes (Gessner 1952), although this has been disputed (Bodkin et al.
1980; Spence 1982). However it is possible that morphological and anatomical adaptations might
occur which combine extra rigidity with the need to maintain leaves capable of utilising low light
intensity in an efficient manner. This may be worthy of further investigation.

ACKNOWLEDGMENTS
I wish to thank Dr G. Halliday for his guidance during the course of this work, and the staff of the

Freshwater Biological Association for providing facilities enabling me to carry out work in
Windermere. I also thank Dr J. Parnell for reading an earlier version of the manuscript.

132 D. A. SIMPSON
REFERENCES

ApaMs, F. S., MACKENZIE, D. R., Cote, H., & Price, M. W. (1971). The influence of nutrient pollution levels
upon element constitution and morphology of Elodea canadensis Rich. in Michx. Environ. Pollut., 1:
285-298.

Barko, J. W., Harbin, D. G. & MatTTHEws, M. S. (1982). Growth and morphology of submerged macrophytes
in relation to light and temperature. Can. J. Bot., 60: 877-887.

BopkIN, P. C., PoLuszny, V., & DALE, H. M. (1980). Light and pressure in two freshwater lakes and their
influence on the growth, morphology and depth limits of Hippuris vulgaris. Freshwater Biol., 10: 545-552.

Cook, C. D. K. (1966). A monographic study of Ranunculus (L.) subgen. Batrachium (DC.) A. Gray. Mitt.
Bot. StSamml. Miinch., 6: 47-237.

Cook, C. D. K., & Urmi-Konie, K. (1985). A revision of the genus Elodea (Hydrocharitaceae). Aquat. Bot.,
21: 111-156.

ERNST-SCHWARZENBACH, M. (1945a). Zur Bliitenbiologie einiger Hydrocharitaceen. Ber. schweiz. bot. Ges.,
55: 33-69.

ERNST-SCHWARZENBACH, M. (1945b). Kreuzungsversuche an Hydrocharitaceen. Arch. Julius Klaus-Stift.
Vererbforsch., 20: 22-41.

GESSNER, F. (1952). Der Druck in seiner Bedeutung fiir das Wachstum submerser Wasserpflanzen. Planta, 40:
391-397.

HarpInaG, J. P. C. (1979). River macrophytes of the Mersey and Ribble basins. Tech. Rept. North-West Water
Authority, no. TS—BS—79-1.

Kunu, H. (1982). Biology of Elodea nuttallii (Planch.) St John with special reference to its growth. Ph.D thesis,
Tokyo Metropolitan University.

Kuni, H. (1984). Seasonal growth and profile structure development of Elodea nuttallii (Planch.) St John in
Pond Ojaga-Ike, Japan. Aquat. Bot., 18: 239-247.

LawreNceE, D. K. (1976). Morphological variation of Elodea in western Massachusetts: field and laboratory
studies. Rhodora, 78: 739-749.

St Joun, H. (1965). Monograph of the genus Elodea: summary. Rhodora, 67: 155-181.

SCANNELL, M. J. P. & Wess, D. A. (1976). The identity of the Renvyle Hydrilla. Ir. Nat. J., 18: 327-331.

Simpson, D. A. (1983). Experimental taxonomic studies of Elodea Michx in the British Isles. Ph.D. thesis,
University of Lancaster.

Simpson, D. A. (1984). A short history of the introduction and spread of Elodea in the British Isles. Watsonia,
15: 1-9.

Simpson, D. A. (1986). Taxonomy of Elodea Michx in the British Isles. Watsonia, 16: 1-14.

SPENCE, D. H. N. (1981). Light quality and plant responses underwater, in SmiTH, H., ed. Plants and the
daylight spectrum, pp. 245-275. London.

SPENCE, D. H. N. (1982). The zonation of plants in freshwater lakes. Adv. Ecol. Res., 12: 37-124.

SPENCE, D. H. N., & Curysta_, J. (1970a). Photosynthesis and zonation of freshwater macrophytes, I: depth
distribution and shade tolerance. New Phytol., 69: 205-215.

SPENCE, D. H. N. & Curystat, J. (1970b). Photosynthesis and zonation of freshwater macrophytes, II:
adaptability of species to deep and shallow water. New Phytol. , 69: 217-227.

SPENCE, D. H. N. & DALE, H. M. (1978). Variations in the shallow water form of Potamogeton richardsonii
induced by environmental factors. Freshwater Biol., 8: 251-268.

SwANn, E. L. (1978). Elodea canadensis and Elodea nuttallii. Trans. Norfolk and Norwich Nat. Soc. , 24: 183-
192.

TutTin, T. G. (1962). Elodea Michx, in CLAPHAM, A. R., Tutin, T. G. & WarRBuRG, E. F. Flora of the British
Isles, 2nd ed. pp. 937-938. Cambridge.

(Accepted October 1987)

Watsonia, 17, 133-138 (1988) 133

Alchemilla gracilis Opiz, a species new to the British flora

G. A. SWAN
81 Wansdyke, Morpeth, Northumberland, NE61 3O0Y
and
S. M. WALTERS

Inland Close, 46 Mill Way, Grantchester, Cambs., CB3 5ND

ABSTRACT

Alchemilla gracilis Opiz, a widespread species in northern and Central Europe not previously known with
certainty in the British Isles, was found in 1976 in Northumberland. The diagnostic characters of the species are
described, and its status in Britain is discussed in the light of the distribution of related Alchemilla species.

INTRODUCTION

In May 1976 G.A.S. was informed by Mr M. Braithwaite, a local botanist, of an interesting site
near Cockplay, north-west of Hexham, Northumberland, v.c. 67, where the sheep-grazed turf of
an old lime quarry contained some local plant species. He visited the locality on 13th June 1976 and
made a detailed species list, which included Alchemilla filicaulis subsp. vestita (earlier identified by
Mr Braithwaite) and a second, quite distinct Alchemilla which G.A.S. was unable to identify. He
therefore collected a small quantity of flowering material and sent the pressed specimens at the end
of the season to S.M.W. for determination. These were, unfortunately, not examined by S.M.W.
until December 1977, but when they were finally dealt with, they turned out to be the European
Alchemilla species, A. gracilis Opiz, never previously certainly reported in Britain. On 16th June
1978, S.M.W., together with G.A.S. and some botanical colleagues, visited the site and studied the
population in detail; they also saw two other nearby sites where the species had been discovered by
Mr Braithwaite since the confirmation of the plant at the original locality.

(S.M.W. apologises for the inordinate delay in publishing details of this remarkable discovery,
and wishes to make clear that he has been solely responsible for that delay.)

ALCHEMILLA GRACILIS AND ITS DISTINCTION FROM OTHER SPECIES OF ALCHEMILLA

Alchemilla gracilis (A. micans Buser) belongs to a group of microspecies of the Alchemilla vulgaris
aggregate which are reasonably well characterised on morphological, distributional and ecological
grounds. They are in the subseries Hirsutae H. Lindb., species 41-53 in the treatment in Flora
Europaea (Walters 1968; see also Lippert & Merxmiiller 1975). Three of them constitute a
remarkable group of continental Alchemilla species centred on Upper Teesdale in Britain: these
are A. monticola Opiz, A. subcrenata Buser and A. acutiloba Opiz, all medium to large plants with
spreading hairs present on stems and petioles and (at least to some extent) on both surfaces of the
leaves. In many parts of Central and northern Europe these three species are frequently found
growing together, in hay-meadows, on roadsides, etc., and are often accompanied by the fourth
member of the group, A. gracilis. In sub-alpine regions of Central Europe, and in parts of
Scandinavia, the common ‘vulgaris’ Alchemillas belong to this group, A. monticola being in
general the commonest of all.

134 G. A. SWAN AND S. M. WALTERS

i

PLATE 1. Herbarium specimen of Alchemilla gracilis collected in Teesdale in 1924. (Photograph by P. F. Yeo).

ALCHEMILLA GRACILIS NEW TO BRITAIN 135

In view of this general pattern of correlated distribution in Europe, the possibility that A. gracilis
was also a member of the Upper Teesdale flora has long been in the minds of British botanists
studying Alchemilla; thus Walters (1949, 1952) wrote ‘‘A. gracilis Opiz . . . should be looked for in
Teesdale, particularly in view of the fact that there is a single inadequate specimen (Druce in Herb.
Mus. Brit.) which is doubtfully referable to this species.’ This specimen, annotated by A. J.
Wilmott as doubtfully attributable to A. gracilis by S.M.W. in 1947, and seen again and annotated
by S.M.W. in 1953, lay neglected in the British Herbarium at BM until G.A.S. re-found it after his
discovery of the living plant in Northumberland. The reason for this neglect was two-fold. S.M.W.
was in any case cautious in identifying the plant from a single gathering, and the very detailed study
conducted during the 1950s by Dr Margaret Bradshaw failed to discover any A. gracilis in the
Upper Teesdale area. (See Bradshaw (1962, 1985) for comments on this.)

The sheet in question is, however, undoubtedly referable to A. gracilis. It was collected (as ‘A.
pastoralis’, i.e. A. monticola) by Francis Druce at Langdon Beck in Upper Teesdale on 12th
August 1924. The material is quite adequate for critical determination, although much of the ripe
fruit has already been shed from the inflorescences. It is probably ‘second-growth’ material from a
rather disturbed, perhaps road-side, habitat. Such plants growing with A. monticola and A.
acutiloba are not easy to detect. This sheet remains the sole evidence that A. gracilis grows (or
once grew) in Teesdale (Plate 1).

The unequivocal identification of Alchemilla microspecies depends on the availability of well-
grown summer flowering specimens complete with some basal leaves. Grazed plants, even if
flowering, and second growth late in the season may be unidentifiable. On the other hand, the
characters of importance in Alchemilla taxonomy, especially the hairiness and leaf-shape, are
retained very satisfactorily in well-prepared pressed specimens, so that detailed study is entirely
practicable in the herbarium. It is further true that some Alchemilla species have very characteristic
appearances in the living state so that, with experience, field identification of technically
inadequate material may be entirely reliable. This is the case with A. gracilis, and it is interesting
that G.A.S., who had never seen the plant before, saw that even in its grazed state it was quite
distinct from A. filicaulis subsp. vestita with which it was growing.

The characters shared by the group of four widespread European Alchemilla species related to
A. monticola can be listed as follows:

1. Petioles and (at least) lower part of stems with erecto-patent or patent hairs (i.e. spreading at
approx. 45° or 90°). This separates A. glabra, a common species in Northern England and
Scotland, which has sparse, appressed hairs on petioles and the lower part of stems.

2. Upper surface of mature summer leaves hairy at least in the folds. This separates the other
common northern British Alchemilla, A. xanthochlora, which has glabrous upper leaf surfaces.

3. At least the pedicels of the flowers glabrous. This separates A. filicaulis subsp. vestita, the most
widely-distributed British species, which has some hairs even on the pedicels, and also the (very
local) A. glaucescens, a relatively small plant with dense indumentum throughout all parts of the
plant.

Plants possessing leaves with hairs on the upper surface are therefore worth careful inspection: if

the pedicels (and especially if the inflorescence in general) are glabrous, then the species is likely to

be a rare or local one in Britain.

Within this group of species related to A. monticola, discrimination of A. gracilis is easy, because
the hair covering, especially on the petioles, is erecto-patent (45° angle), not more or less patent
(90°). This quality in the indumentum is shown also on the leaf-surfaces and particularly on the
veins beneath the leaf, and gives the live plant a characteristic silky sheen (referred to in the name
A. micans given to the species by Buser). Indeed, the indumentum sometimes approaches that of
the arctic-alpine species A. glomerulans, though leaf-shape and particularly hypanthium-shape
easily distinguish the two species. Growing in grazed turf mixed with A. filicaulis, A. gracilis can
readily be picked out by the silky erecto-patent hair covering: as G.A.S. demonstrated in the field
in 1978, individual leaves could also be distinguished in that they were usually 9-lobed, whereas the
A. filicaulis leaves were 7-lobed.

Two other diagnostic features are shown by A. gracilis. The first, which seems to be very reliable
even in grazed material which may have only poorly-developed inflorescences, is the shape of the
hypanthium in flower. Uniquely in A. gracilis this is narrow with a base cuneate in outline (see Fig.
1). If the inflorescence is reasonably well-grown, the other diagnostic feature becomes evident: the

136 G. A. SWAN AND S. M. WALTERS

|

a b

FicureE 1. Diagram of hypanthium shape in (a) Alchemilla gracilis, (b) other British species of the subseries
Hirsutae. (Adapted from Lippert & Merxmiiller 1975.) Scale bar = 1 cm.

branching of the inflorescence is rather sparse, and the whole inflorescence is narrow and relatively
few-flowered. Combined with the erecto-patent or even sub-appressed indumentum, this gives the
inflorescence a very characteristic appearance (Plate 1). Leaf-shape, as in most Alchemilla species,
is too variable to be a very reliable diagnostic character, but a well-grown summer leaf of A.
gracilis is reniform to reniform-orbicular in outline, with a narrow but open sinus (that is, the lobes
do not overlap to cover the junction of the blade and the petiole). In this respect the leaf is rather
intermediate in shape between the reniform leaf of A. filicaulis, which usually has quite a wide
sinus, and the more or less orbicular leaf of A. monticola in which the basal lobes usually overlap.
(The coloured illustrations of leaves of Alchemilla in Garrard & Streeter (1983) include A. gracilis,
and are recommended for their accuracy.)

The early flowering of A. gracilis seems to differentiate it from most other Alchemilla species
with which it might be confused. Thus, G.A.S. recorded it in full flower at the Cockplay locality on
16th May 1984, at least one week in advance of A. filicaulis subsp. vestita growing with it. Plants
which S.M.W. has cultivated (from one of the Coldwell localities) since 1978 also come into flower
early every year. If this phenological difference is consistent, it may explain why the species
remained undetected for so long in mixed populations, for the inflorescences in cultivation are
shedding ripe seed and becoming relatively inconspicuous by late June or early July, and the whole
plant could become covered by competing vegetation in the wild.

DESCRIPTION OF THE LOCALITIES

In the locality near Cockplay, where G.A.S. first detected the plant, the population in June 1978
was estimated to contain in the main site some 100 separate plants (or small clumps) mostly
concentrated in an area of some 30 X 10 m of heavily-grazed, more or less closed, grassland turf.’
Nearby, in the same field, a smaller population in more broken turf on a small limestone outcrop
contained an estimated 30 or 40 plants. In the main site, there was some admixture with A.
filicaulis subsp. vestita, and in the subsidiary site the two species of Alchemilla seemed about
equally common. The history of the site is not known, but is presumed to have been worked for
lime; it is on the junction of limestone and the Whin Sill rock, and the surface limestone explains
the exceptionally species-rich grassland turf. Associated species listed on 13th June 1978 were:
Agrostis capillaris, Anthoxanthum odoratum, *Avenula pratensis, Bellis perennis, Briza media,
Cerastium fontanum, Cynosurus cristatus, Deschampsia cespitosa, Festuca ovina, F. rubra,
*Galium verum, Holcus lanatus, *Koeleria macrantha, Lotus corniculatus, Plantago lanceolata,
*Plantago media, Potentilla erecta, Ranunculus bulbosus, Sanguisorba minor, Taraxacum cf.
brachyglossum, Thymus drucei, Trifolium pratense, T. repens, Viola riviniana. (*Species indicating
relatively calcareous soil.)

The two other localities discovered in 1978 are near Coldwell, about 1 km from the original one.
Both sites were visited on 13th June 1978 and in one of these there was a small population of

' This estimate was, apparently, too small: Dr A. J. Richards surveyed the population on 8th May 1984 and
estimated the flowering plants of A. gracilis to number 800 + 100.

ALCHEMILLA GRACILIS NEW TO BRITAIN 137

A. gracilis in a damp corner of a roughly-grazed pasture. At the other locality nearby, there were a
few clumps of separate individual plants with lax, somewhat etiolated inflorescences up to 50 cm
tall growing in lush ungrazed meadow with competition from tall coarse grasses, especially Dactylis
glomerata. Associated spp. here were: Alopecurus pratensis, Bellis perennis, Cerastium fontanum,
Cirsium arvense, Cynosurus cristatus, Festuca rubra, Lolium perenne, Plantago lanceolata,
Plantago major, Poa annua, Poa trivialis, Ranunculus acris, Rumex acetosa, Rumex obtusifolius,
Taraxacum sp., Veronica chamaedrys, Veronica serpyllifolia.

Neither locality showed any unusually rare or local species, and indeed seemed quite
unremarkable. In the ungrazed meadow, the lack of grazing was possibly a recent factor: some
species (including the Alchemilla itself) seemed rather etiolated and not well adapted to the
vigorous competition by the grasses.

After 1978, G.A.S. kept a sharp look-out during extensive botanical study of Northumberland
for new localities for A. gracilis, but it was not until 1985 that he was successful. The new locality
near Sharpley is in the general area of the 1976 discoveries, and consists of grassland on a roadside
verge, where A. gracilis was found growing with A. filicaulis subsp. vestita. The adjacent field had
relatively species-rich calcareous grassland with much Poterium sanguisorba and other calcicoles
such as Plantago media, but G.A.S. could not find A. gracilis in the field.

STATUS AND DISTRIBUTIONAL SIGNIFICANCE

A. gracilis, as we have explained, has a wide distribution in Central and northern Europe
essentially similar to that of the other three related species A. monticola, A. subcrenata and A.
acutiloba; such distributions are characterised as Northern-Montane (Matthews 1955) and are
shown by several other British species, including e.g. Primula farinosa. Its discovery in the north of
England is therefore not unexpected. On the other hand, the three other species are centred in the
Upper Teesdale area (if we neglect two apparently casual and non-persistent records for A.
monticola elsewhere), where the flora already has a remarkable assemblage of rare and local
species, including two other Alchemilla species, A. glomerulans and A. wichurae, with montane
distributions in Britain.

Bradshaw (1962) discusses the status of the Northern-Montane group of Alchemilla species in
the Teesdale area, and concludes that a hypothesis of survival and secondary expansion from a
wider Late-Glacial distribution, which might reasonably be applied to the Arctic or Arctic-alpine
species A. wichurae and A. glomerulans, has little to support it for the Northern-Montane group,
though clearly it cannot be ruled out. She points to the almost complete absence for these three
species of any natural habitats. They are strongly synanthropic, both in Teesdale and also in
Scandinavia where much information is available (see especially Sjérs (1954) and Samuelsson
(1943)). She considers the possibility of accidental introduction as hay-meadow ‘weeds’, or even
intentional introduction as medicinal ‘Alchemilla vulgaris’ in historic time, and concludes that the
question of their origin remains an entirely open one.

Samuelsson (1943) gives a detailed discussion of the Scandinavian distribution of A. gracilis, and
shows that it is not significantly different from that of A. monticola and A. acutiloba — except that
there is a curious, unexplained absence from Western Finland, a feature not shown by the other
two species. Even more so than A. monticola, A. gracilis is a ruderal plant in Scandinavia, being
particularly common in artificial park grassland and on roadsides, but readily colonizing from such
sites into hay meadows and pastures. The spread of the species in recent historic time is reasonably
well documented in particular localities both in Norway and Sweden where the Alchemilla flora
has been critically surveyed over a sufficiently long period to record such a spread. Thus
Samuelsson himself was certain that the species spread in his own life-time in the vicinity of his
birthplace (the town of Strangnas in S6dermanland) from an initial record in 1910. This assessment
accords well with the fact that Sjérs (1954) does not record A. gracilis from the (relatively old)
synanthropic hay meadow communities of Dalarna (Sweden) where he made a special study.
Indeed, it seems that throughout Fennoscandia A. gracilis occurs rarely in even semi-natural
habitats until one reaches the Finnish-Russian frontier region, where it is recorded in open
deciduous woodland and assessed as a truly native species (Fagerstrom 1939-40; cited in
Samuelsson 1943, p. 35). In European Russia, the species is common in a variety of habitats; for

138 G. A. SWAN AND S. M. WALTERS

example, Tikhomirov (1969) states that in the Moscow region: “‘it often grows in abundance” and
“possesses perhaps the broadest ecological range among our [species of Alchemilla], being often
found in dry meadows, on hillsides, in shady forests, etc.”” The species extends into W. Siberia,
outside the boundary of Europe as adopted for Flora Europaea (see Juzepczuk, 1941, p. 348).

The nearest Continental localities to Britain are in the Low Countries; here there seems to be
some evidence that A. gracilis is still extending its range, though the belated recognition of the
species in the flora of a particular country cannot, obviously, by itself be taken as evidence of its
recent introduction. For a discussion of the status of the species in Belgium see Sougnez &
Lawalrée (1956, 1959), and in Luxembourg see Reichling (1969).

What light does this throw on the newly-discovered British occurrences of A. gracilis? Firstly, we
can say that, since they are all within a small area, a relatively recent introduction and spread from
a Continental source seems quite probable. On the other hand the Cockplay locality — grazed
limestone turf — is emphatically not a crudely artificial new habitat, and the population there
certainly gives the impression of relative age. Secondly, we have to admit that there is a great deal
of northern England for which systematic and careful Alchemilla recording, such as Bradshaw
(1962) undertook for Teesdale and Weardale, has not been done, so that we have no idea whether
the newly-discovered Alchemilla is really so rare and local. If we have to conclude with Bradshaw
that the case for the native status of the Northern-Montane Teesdale Alchemillas is wholly
uncertain, the same doubt must remain over the Northumberland records of A. gracilis. It is a
challenge to sharp-eyed northern botanists to extend the known range of this interesting addition
to the British flora.

REFERENCES

BRADSHAW, M. E. (1962). The distribution and status of five species of the Alchemilla vulgaris L. aggregate in
Upper Teesdale. J. Ecol., 50: 681-706.

BrapsHaw, M. E. (1985). Studies on the flora of Teesdale. Naturalist, 110: 3-21.

FaGerstrOm, L. (1939-40). Ett bidrag till kannedomen om vegetation och flora i Terijoki socken pa Karelska
naset. Mem. Soc. Fauna Flora Fennica, 15 (1938-39).

GARRARD, I. & STREETER, D. (1983). The wild flowers of the British Isles. London.

JUZEPCZUK, S. (1941). Alchemilla, in Komarov, V. L., ed. Flora U.R.S.S., 10: 348. Leningrad.

LipPERT, W. & MERXMULLER, H. (1975). Untersuchung zur Morphologie und Verbreitung der Bayerischen
Alchemillen (II). Ber. Bayer. Bot. Ges., 46: 5—46.

MATTHEWS, J. R. (1955). Origin and distribution of the British flora. London.

REICHLING, L. (1969). Notes floristiques. Bull. Soc. Nat. Luxembourg, 64: 3-52.

SAMUELSSON, G. (1943). Die Verbreitung der Alchemilla-Arten aus der Vulgaris-Gruppe in Nordeuropa. Acta
Phytogeog. Suecica, 16: 1-159.

SyOrs, H. (1954). Slatterangar i. Grangarde, Finnmark. Acta Phytogeog. Suecica, 34: 5—126.

SOUGNEZ, N. & LAWALREE, A. (1956). Alchemilla gracilis Opiz en Belgique. Bull. Jard. Bot. Etat Bruxelles, 26:
247-252.

SOUGNEZ, N. & LAWALREE, A. (1959). Les Alchemilla de Belgique. Bull. Jard. Bot. Etat Bruxelles, 29: 389-423.

TIKHOMIROV, V. N. (1969). Alchemilla-species of the Moscow region (a systematic review). Proc. Study Fauna
Flora USSR, Sect. Botany, 13 (XXI): 98-151. (In Russian.)

Watters, S. M. (1949). Alchemilla vulgaris L. agg. in Britain. Watsonia, 1: 6-18.

Watters, S. M. (1952). Alchemilla subcrenata Buser in Britain. Watsonia, 2: 277-8.

Wa ters, S. M. (1968). Alchemilla L., in Tutin, T. G. et al., eds. Flora Europaea, 2: 48-64. Cambridge.

(Accepted August 1987)

Watsonia, 17, 139-143 (1988) 139

The coastal ecodeme of Parnassia palustris L.

R. J. GORNALL

Botany Department, The University, Leicester, LE] 7RH

ABSTRACT

Cultivation and breeding experiments have shown that Parnassia palustris L. var. condensata Travis &
Wheldon (Parnassiaceae), a coastal ecodeme, retains its characteristic, dwarf, clumped habit and larger ratio of
flower diameter to stem height in cultivation, and breeds true from seed. Its taxonomic rank of variety is
therefore upheld, and its name is lectotypified by a specimen collected by J. A. Wheldon from W. Lancs., v.c.
60. It has two cytodemes, with 2n = 18 and 36.

MORPHOLOGICAL VARIATION

The existence of a distinctive, coastal ecodeme of Parnassia palustris L. in the British Isles had
been known for some time (Hall 1839; Wheldon & Wilson 1907; Druce 1911) before it was given
taxonomic rank as var. condensata (Travis & Wheldon 1912). It was described from ‘“‘the sand-
dunes on the coasts of Lancashire and Cheshire’’, and was distinguished from var. palustris by the
following characters: shorter stature; branched rhizome producing more (4-20) and thicker (1:5—
1-6 mm in diameter) stems; more tufted basal leaves, the laminae of which are usually as long as or
longer than the petioles (rather than shorter); a cauline leaf immersed amongst, or only slightly
protruding above the basal leaves (rather than elevated); larger flowers (25-37 mm or more in
diameter); and a larger capsule (12-16 mm long). In order to examine these differences,
measurements were made of British and Irish specimens of P. palustris at BM, CGE, DBN, E, LIV,
LTR, NMW, RNG and TCD for the characters listed by Travis & Wheldon (1912). It became clear
that these authors underestimated or, in some cases, misjudged the amount of variation to be
found within each variety, such that the actual differences between the characters are less clear-cut
than they claimed (Table 1), and a distinction between the varieties must be made on the
characters considered in combination. For example, variation in stem height and especially in
flower diameter is such that any distinction based on these characters individually is either
impossible or at best very difficult. However, when the two characters are considered together, the
pattern of variation in each variety is different. In var. palustris short specimens usually have
proportionally smaller flowers, whereas in the dwarf var. condensata, the reduction in flower size is
much less marked. Thus, when flower diameter is considered as a ratio of stem height, the two
varieties can be distinguished much more reliably, and indeed this ratio is probably the best
discriminant for use in the field (Table 1).

CULTIVATION AND BREEDING EXPERIMENTS

Cultivation and breeding experiments were conducted to discover whether the coastal ecodeme is
merely an environmentally-induced variant (Melvill 1912; Marshall 1913), or whether its features
are due directly to genetic factors (Wheldon & Travis 1913). Previous attempts at cultivation have
produced different results, partly perhaps because, by general agreement, the species is difficult to
grow and maintain in cultivation. Wheldon & Travis (1913) reported that samples of var.
condensata from S. Lancs., grown by Prof. K. Graebner in Berlin, retained their characteristic
features. In contrast, a later but similarly undocumented report stated that var. condensata
reverted to “‘average size and habit” after two years in cultivation, becoming indistinguishable
from var. palustris (Salisbury 1952: 271, 278).

140 R. J. GORNALL

TABLE 1. CHARACTERS USED TO DISTINGUISH VAR. CONDENSATA FROM VAR. PALUSTRIS
Measurements of wild plants are from herbarium specimens. Measurements of cultivated plants are quoted as
means and standard deviations; the statistical tests were applied to these data only.

Wild Cultivated*

Character palustris condensata palustris condensata
Stem height, mm 100—330(-512) 30-—150(-230) 186+60 102+66*
Height of cauline leaf

above ground, mm (10-)20-170 5—40 37.5£21.4 30.2+18.0 -
Basal leaf petiole

length: lamina length ratio (1-)2-6 0.4-3(4) 1.7+0.4 1.9+0.6
Rhizome branched usually usually usually usually

sparingly extensively sparingly extensively

Number of basal leaves” — — 14+6 28+10*
Number of stems 1-6(-32) (1-)4-15(-34) 2..0221-0 3:92:3.2
Stem diameter, mm 0.5-1.5 0.5-2.0 1.3+0.4 d5e:05
Flower diameter, mm 15-35 15-35 22.8+4.2 25.0+6.0
Flower diameter: stem

height ratio 0.04-0.23 0.15—0.76 0.14+0.04 0.40+0.32**
Capsule length, mm (6-)9-13 8—15(-20) 924-23 1021 1.7,

*Mann-Whitney U-test on cultivated plants:— **significant difference, P<0.01; *significant difference, P<0.05;
all other characters not significantly different. Origins of ramets, sample sizes in parentheses:— Var.
condensata: Ainsdale, S. Lancs., v.c. 59 (5); Tentsmuir, Fife, v.c. 85 (3); Loch Spiggie, Shetland, v.c. 112 (2).
Var. palustris: Monks Dale, Derbys., v.c. 57 (3); Malham Tarn, Mid-W. Yorks., v.c. 64 (4); Lochranza, Clyde
Is., v.c. 100 (3).

> Character not used by Travis & Wheldon (1912).

In view of these conflicting reports, I undertook a small cultivation experiment in which
transplanted material of both var. condensata and var. palustris was grown side by side in a
glasshouse (Table 1). Ramets taken from plants in the wild were planted in a mixture of 3:1
compost and sand in 7-5 cm plastic pots, and grown in a mist unit; I found this to be the most
satisfactory method of maintaining plants in a healthy condition. The duration of cultivation
varied, but was usually about 12 months (up to three years in some cases), and the period always
included the development of the flowering phenotype from the dormant rhizome.

Three features of var. condensata remained distinctive in cultivation: its shorter stems; its larger
flower size relative to stem height; and its greater degree of rhizome branching, a character
reflected also in the larger number of basal leaves (Table 1). The other differences seen in the field,
particularly the position of the cauline leaf and the relative lengths of the petiole and lamina of the
basal leaves, tend to disappear in cultivation, with var. condensata becoming like var. palustris.

Most plants were cultivated for about a year, after which they were pressed, but it is worth
noting that the individuals of var. palustris from Malham Tarn and of var. condensata from
Tentsmuir, which were grown for three years, retained their different morphologies as outlined
above.

Confirmation that the flower, stem and rhizome differences are under genetic and not
environmental control came from a breeding experiment. Self-pollination of cultivated plants of
var. condensata, collected from Ainsdale, produced full capsules of seeds, showing the plants to be
self-compatible. Twelve S,; seedlings were randomly selected to be raised to maturity. Five of them
flowered in their first year, five succumbed to attack by vine weevil and two have remained in a
vegetative state. The five flowering plants were all very similar and showed the characteristically
large flowers and short stature. The only important difference between the S, generation and the
parental plants was in the number of stems produced. All S; plants had but a single stem, although
examination of the rhizomes revealed them to have branched into three or four shoots; these may
eventually each produce a flowering stem of their own, thereby generating the clumped
appearance typical of var. condensata. Seed collected from a wild population of var. condensata in
Orkney produced similar plants. In contrast, plants of var. palustris grown from seed collected in

COASTAL ECODEME OF PARNASSIA PALUSTRIS 141

5 cm

b 4

Figure 1. Silhouettes of representative specimens of Parnassia palustris grown from selfed seed; (a) var.
palustris, from an Argyll parent; (b) var. condensata, from an Ainsdale parent.

Argyll were taller, had smaller flowers relative to stem height, and rhizomes that were less
branched (1 (—4) flowering shoots). Representative specimens of both varieties grown from seed
are illustrated in Fig. 1.

CONCLUSIONS

The conclusions to be drawn from the herbarium studies and the cultivation and breeding
experiments are: i) that P. palustris var. condensata is a coastal genoecodeme, which constitutes
the characteristic variant of the species in machair-type habitats, usually dune-slacks, or in the
short turf on the tops of sea-cliffs; and ii) that it is appropriate to award it taxonomic recognition at
the rank of variety, because it is a local rather than a consistently regional variant (Du Rietz 1930).

Some comment should be made here on the contrary results obtained by Salisbury (1952)
mentioned earlier. It is possible that more than one coastal variant exists, and that Salisbury’s
material represented an environmentally induced, phenotypic copy of var. condensata. Such
plastoecodemes sometimes occur in coastal populations with an otherwise genetically fixed
phenotype (Akeroyd, in press), but how frequent such plants may be in populations of var.
condensata is unknown.

142 R. J. GORNALL

CHANNEL ISLANDS
PLOTTED ON
UTM GRID

Ficure 2. Distribution of Parnassia palustris var. condensata in the British Isles (all records).

TAXONOMIC ACCOUNT

The following description is based entirely on specimens from Great Britain and Ireland.

PARNASSIA PALUSTRIS L. var. CONDENSATA Travis & Wheldon in J. Bot., Lond., 50:256
(1912). TYPE: N. of St Annes, [W. Lancs., v.c. 60], Sep. 1900, J. A. Wheldon (Lectotype:
NMW,, designated here).

Rhizome usually much-branched. Stems (1—)4—-15(—34), usually less than 15(—23) cm tall, diameter

0-S—2 mm. Basal leaf lamina 8-25 x 7-20 mm, petiole 7—30(-40) mm, usually less than 1-5 times as

long as the lamina. Cauline leaf cordate or amplexicaul, rarely deltoid, 5—25(-30) x 3-25 mm,

COASTAL ECODEME OF PARNASSIA PALUSTRIS 143

positioned 5-40 mm from base of stem, the apex of the leaf often level with or below the point to
which the apices of the basal leaves reach when positioned alongside the stem. Flowers 15-35 mm
in diameter, usually 1/5 or more of stem height; sepals 4-10 x (1-)2-4 mm; petals 6-16 x 5-12
mm, with 7-11 veins; staminode lamina 2-3 mm, longer than broad, narrowed abruptly or tapering
to a claw, bearing 7-14 filiform divisions, 2-4 mm long; filaments 3-6 mm; anthers 2-3 mm.
Capsule ovoid to globose, obtuse or rounded at the apex, 8-15 (—20) x 9-16 (-21) mm.
Chromosome number, 2n = 18, 36 (Gornall unpublished, based on native material).

There are no Travis or Wheldon specimens predating the protologue in W. G. Travis’s
herbarium at LIV. Therefore, I have designated the Wheldon specimen from the latter’s
herbarium at NMW as the lectotype because it is the only one which obviously pre-dates the
protologue and is fairly characteristic of the variety. Far better material, collected from Freshfield,
S. Lancs., v.c. 59, was widely distributed by Wheldon (1913) as part of a Botanical Exchange Club
distribution, but I have been unable to discover whether it was collected before the protologue was
published (both are dated August 1912).

DISTRIBUTION

The distribution of var. condensata in the British Isles, based on herbarium specimens, is shown in
Fig. 2. Records are from the following vice-counties: 4, 27, 52, 58-60, 62, 67, 68, 82, 85, 90, 100,
107-09, 111, 112, H21, H27, H28, H34, H35, H39 and H40. Some populations of P. palustris from
cliff-tops on the coast of Co. Durham, v.c. 66, are referable to var. palustris and grow in a wet
meadow habitat, rather than in short turf. Some plants of var. palustris from exposed, inland
populations (often at high elevations) may resemble var. condensata in their dwarf, compact habit,
although they usually do not have the characteristically large ratio of flower diameter to stem
height. Whether this dwarfness is genetically fixed or not is unknown.

In Europe, var. condensata has been collected from dune-slacks in north-western France
(Brittany), the Netherlands and Denmark, and, like many coastal ecotypic variants (Akeroyd in
press), it appears to be restricted in its occurrence to north-western Europe; southwards and
eastwards it grades into var. palustris.

ACKNOWLEDGMENTS

I would like to thank the curators of the following herbaria for loaning or allowing me access to
specimens: BM, CGE, DBN, E, LIV, NMW, RNG, TCD. I should also like to thank E. R. Bullard,
Dr T. C. G. Rich, W. Scott and A. A. Slack for help in acquiring living plant material.

REFERENCES

AKEROYD, J. R. (in press). Genetic variation in European coastal plant species, in VAN DER MAAREL, E., ed.
Dry coastal ecosystems. Amsterdam.

Druce, G. C. (1911). The international phytogeographical excursion in the British Isles, III. The floristic
results. New Phytol., 10: 306-328.

Du Retz, G. E. (1930). The fundamental units of biological taxonomy. Svensk Bot. Tidskr., 24: 333-428.

Hatt, T. B. (1839). A Flora of Liverpool, p. 15. London.

MarsHaLl, E. S. (1913). Parnassia palustris, L., var. condensata, Travis and Wheldon. Rep. botl Soc. Exch.
Club Br. Isl., 3: 252.

MELVILL, J. C. (1912). Parnassia palustris L. var. (or forma?) condensata. J. Bot., Lond. , 50: 376-377.

SALISBURY, E. J. (1952). Downs and dunes, their plant life and its environment. London.

Travis, W. G. & WHELDON, J. A. (1912). A new variety of Parnassia palustris. J. Bot., Lond. , 50: 254-257.

WHELDON, J. A. (1913). Parnassia palustris, L., var. condensata, Travis and Wheldon. Rep. botl Soc. Exch.
Club Br. Isl., 3: 252. ;

WHELDON, J. A. & Witson, A. (1907). The flora of west Lancashire. Liverpool.

WHELDON, J. A. & Travis, W. G. (1913). Parnassia palustris var. condensata. J. Bot., Lond., 51: 85-89.

(Accepted July 1987)

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Watsonia, 17, 145-148 (1988) 145

Conyza sumatrensis (Retz.) E. Walker established in England

B. WURZELL

47 Rostrevor Avenue, London, N15 6LA

ABSTRACT

Conyza sumatrensis (Retz.) E. Walker has achieved a coherent distribution pattern through the Thames
estuary to eastern London. The implications are examined, together with a résumé of its earlier colonization of
France and the Channel Islands. The habitat requirements of C. sumatrensis, C. bonariensis (L.) Cronq. and C.
canadensis (L.) Cronq. are compared, and a key is given for their identification.

DISCOVERY IN LONDON

On 26th September 1984, I first observed Conyza sumatrensis (Retz.) E. Walker on the English
mainland, well established by the River Lea towpath from Old Ford to Bromley-by-Bow, in the
London Borough of Newham. Previously, the British literature had reported this robust fleabane
only from Guernsey (Jee 1962) as C. bonariensis (L.) Cronq., and from Jersey (McClintock 1975)
and Sark (Ryan 1977) under the present name. In 1980, I noted its widespread occurrence in the
first two of these islands, and, as it self-sowed annually in my garden since then, it was still possible
to compare fresh Sarnian material with the new Lea-side plant. D. McClintock (pers. comm.) also
confirmed that they belonged to the same species.

PRESENT DISTRIBUTION IN EASTERN AND CENTRAL LONDON

Between October 1984 and October 1986, I found C. swmatrensis occupying over 40 urban sites,
sometimes abundantly, and often close to the River Thames and/or its associated waterways. My
current records for each London borough are summarized as follows.

Newham, v.c. 21: from Old Ford (GR 51/378.832) to Bromley-by-Bow (GR 51/382.823), plentiful
along c. 2 km of River Lea towpath, in ancient concrete fissures and wall crevices, with much
Chenopodium ambrosioides and Hirschfeldia incana.

Tower Hamlets, v.c. 21: a) Carpenter’s Road (GR 51/374.845). One large plant in factory
forecourt. It had more deeply sinuately-cut lower leaves, rather Coronopus-like, and smaller,
less pubescent capitula, appearing somewhat intermediate to C. canadensis (L.) Crong., but too
sturdy, stout and fertile for an F, hybrid. I have seen similar plants of Spanish origin in BM. b)
Regent’s Canal from Victoria Park (GR 51/348.835) south-eastwards to Limehouse (GR 51/
364.812), scattered individuals on and below the towpath wall. c) Limehouse Basin (GR 51/
364.811), abundant and very vigorous on quayside, mostly at foot of south-facing brick wall. d)
from Ratcliff (GR 51/360.810) to Shadwell (GR 51/354.808), frequent by pavements and on
concrete dump. e) Shadwell Basin (GR 51/352.807), frequent on stony waste ground. f)
Wapping (GR 51/345.801), two plants on roadside. g) Wapping (GR 51/343.804), several plants
on gravelly waste ground. h) Stepney (GR 51/353.817), pavement weed with Sorghum
halepense.

Hackney, v.c. 21: a) Upper Clapton (GR 51/355.867), one casual plant on concrete path by
Middlesex Wharf. b) Stamford Hill (GR 51/337.878), one casual plant on pavement by school
wall. c) Stoke Newington (GR 51/332.870), stony waste ground behind Abney Park Cemetery,
five large plants amid a mixed colony of Atriplex prostrata and A. prostrata x longipes. d)
Shacklewell (GR 51/341.858), two plants on sheltered footpath. e) Graham Road (GR 51/

146 B. WURZELL

344.847), three plants as garden weeds in warm, sheltered hollow. f) London Fields (GR 51/
345.845), pavement and shrubbery weed, several plants surprisingly mixed with Rumex palustris
and Atriplex littoralis. g) Regent’s Canal, from Victoria Park (GR 51/348.835), north-westwards
to De Beauvoir Road (GR 51/328.838), small scattered colonies frequent along brick walls. h)
Homerton (GR 51/355.845), widespread street weed.

Islington, v.c. 21: Regent’s Canal, from De Beauvoir Road (GR 51/328.838), south-westwards to
Danbury Street (GR 51/318.835), several small colonies on brick walls.

Haringey, v.c. 21: by Finsbury Park (GR 51/313.874), one plant atypically and casually on rough
grassy verge of Parkland Walk, the most north-westerly of all European records to date.

Lambeth, v.c. 17: by Westminster Bridge, opposite County Hall (GR 51/306.796), two small plants
at the foot of a concrete wall.

Southwark, v.c. 16: Peckham Rye Park (GR 51/345.754), one plant on a roadside.

Logically, one would now expect frequent occurrences close to the Thames in the Bermondsey,
Rotherhithe, Deptford, Greenwich, Isle of Dogs, Poplar and Canning Town areas; further
investigations are needed. It should also be more frequent in central London than the above
records indicate, but it has not yet been found naturalized anywhere west of London.

OTHER THAMES-SIDE RECORDS

Palmer (1983) not only reported C. bonariensis (now redetermined as C. sumatrensis) from stony
waste ground near Blackfriars, in the City of London, but also referred to a strong colony
established at least since 1977 at Chalkwell, S. Essex, v.c. 18. The latter was first found in 1974 by
R. M. Payne (Rand 1975), and E. J. Clement (pers. comm.) further states that R. M. Payne had
identified two colonies at Westcliff-on-Sea. Additionally, R. B. Hastings (Burton 1985) detected a
large stand of the species on Rainham Marsh in 1984, and G. D. Kitchener (pers. comm.)
discovered a plant at Swanscombe, N. Kent, v.c. 16, in 1985.

True C. bonariensis, which is closely related, has, to the best of my knowledge, only occurred in
Great Britain as an ephemeral shoddy alien.

FRENCH RECORDS

C. bonariensis, presumably sensu stricto, has long been known as an established alien in the south
of France, either by the name Erigeron crispus Pourret (Bonnier & Layens 1909) or as Conyza
ambigua DC. (Le Maout & Decaisne 1855). Rouy (1927) also noted its occurrence in Corsica.
More significantly, De Langhe et al. (1973) mention ‘‘Erigeron crispus Pourret’”’ as an adventive in
Belgium, Luxembourg and northern France, but there is no description given and thus no way of
ascertaining which Conyza species is intended; at such temperate latitudes, C. bonariensis s.s.
would indeed be adventive, but one would, by that date, expect C. sumatrensis to have naturalized
itself locally in the region. Fournier (1961) had already made a clear distinction between the two;
the latter, as Erigeron naudinii (Bonnet) G. Bonnier, is described as very rarely naturalized in Var,
Aude, Pyrénées Orientales, etc., and of unknown origin. An interesting footnote claims that “‘les
vrais Conyza ont les styles plus longs que les corolles; ce qui n’est pas le cas ici.”

Neither Mérat (1812) nor Cosson & Germain (1845) nor even Jeanpert (1977) mention any
Conyza species other than C. canadensis occurring in the Parisian region. However, in October
1984, personal observations of C. sumatrensis growing plentifully on gravelly roadsides in the
Aulnay-sous-Bois, Bondy and Blanc-Mesnil suburbs (about 15km north of Paris) revealed that its
recent northward progress had extended on a broader west-to-east front than is indicated in Jovet
& Vilmorin (1975). This suggests that De Langhe et al. (1973) may either have overlooked or
misnamed it in the countries listed above. It also supports the hypothesis that our present Thames
estuarine and metropolitan populations arose from wind-blown seed originating in north-eastern
France rather than the Channel Islands. Clearly, further records from northern Europe generally
would now be very helpful in placing our London colonies into a wider context.

CONYZA SUMATRENSIS IN ENGLAND 147

It is also relevant to recall that the decade 1974-1984 enjoyed exceptionally frequent hot
summers, with prolonged spells of dry, south-easterly winds. These conditions favoured not only
the establishment of C. sumatrensis here, but also encouraged other highly drought-resistant
Continental species such as Picris hieracioides and Hirschfeldia incana to extend their British range
and abundance quite dramatically. Nowadays, they often grow together in London, an association
unheard of ten years ago.

AUTECOLOGIES OF THE THREE CONYZA SPECIES

From my own research carried out near Los Angeles, California, in June 1984, it became apparent
that naturalized C. bonariensis would seek the extra warmth of narrow pavement cracks and tight
wall crevices, even given a climate more tropical than that prevailing in the Mediterranean. C.
sumatrensis, also a marked thermophile, again favours arid, sun-baked niches, but to a lesser
degree, and this has enabled it to spread as far northwards as London. C. canadensis, the least
demanding of the trio, profusely colonizes cold, bare, loose soil and gravel in rural and urban areas
alike, but still remains adaptable enough to produce stunted, rather woody (sometimes biennial?)
plants in the same stone and concrete heat-traps which are actually sought by the other two. As a
result, I have found it almost always accompanying C. sumatrensis, both in London and in Paris,
the relative vigour of each species doubtlessly influenced by the relative heat-retentiveness of each
substrate.

A marked degree of moisture intolerance in C. canadensis is implied by the Atlas (Perring &
Walters 1962) whose numerous distribution dots cluster most closely in the driest south-eastern
and East Anglian regions; the species is not recorded from Scotland or Ireland at all. For a similar
reason, London may now represent the north-western limit of the Eurasian distribution of C.
sumatrensis, its recent extension of range aided by a number of particularly rainless, warm,
continental summers. Having arrived, it may slowly retreat again, faced with the return of a normal
Atlantic climate. Or it may evolve morphologically indistinguishable variants more tolerant of
dampness, so as to hold its own or even move a little further northward and westward. After all,
Conyza species do vary considerably in response to environmental conditions, and C. sumatrensis
is Sometimes seen to produce casual examples in unusual habitats such as shady gardens, exposed
spoil heaps, and windswept grassy banks. This tendency is more noticeable towards the outer edge
of its present urban distribution, and therein could lie evolutionary potential.

Be that as it may, C. sumatrensis has, so far, shown no capacity for using British Rail’s rolling stock
as aseed distribution vector. Unlike C. canadensis, Senecio squalidus and Epilobium ciliatum, which
have exploited this facility to a spectacular degree in southern England, it still finds loose trackside
ballast uncolonizable; the limited ability of that medium to hold heat at night could well explain why.
Bearing in mind also the small likelihood of favourable long-term habitats existing in cooler,
damper, country areas, I would, at the present time, anticipate only a remote chance that this tall
subtropical species might permanently invade a major part of the United Kingdom.

KEY TO CONYZA SPECIES IN ENGLAND

1. Inflorescence + columnar. Mature capitula 3-5 mm broad, + bell-shaped. Involucral bracts
yellowish-green, glabrous or subglabrous. Tubular florets short, yellow, four-lobed.
Leaf-blades relatively thin, their borders sparsely but usually conspicuously hispid-ciliate
CLM UO OMG epee eee eae Se ene a ae Mutant Nocscesecut eStats cy etneedeistens C. canadensis

1. Inflorescence + pyramidal. Mature capitula 5-10 mm broad, + straight-sided. Involucral
bracts pale grey-green, shortly and evenly pubescent. Tubular florets longer, yellow
below, whitish above, five-lobed. Leaf-blades relatively thick, their borders densely
appressed-pubescent, sometimes with a few soft longer cilia towards the base
OTN Me ene Seek ones ey ese e tea Sets wo tnisccuoesesswacGenicesnmoectecetceeeneteese reece es 2

De Plant often robust (10—) 30-200 cm. Inflorescence finely grey-pubescent throughout, not
glandular, widely and profusely branched, the lateral branches not overtopping the main
axis. Mature capitula 5-7 mm broad, pappus pale grey. Shortly ligulate florets present.
Widely naturalized in London, preferring hot, stony substrates ................ C. sumatrensis

148 B. WURZELL

2. Plant seldom exceeding 60 cm. Inflorescence glandular, sticky, often becoming purplish, the
branches less numerous, with laterals often overtopping the main axis. Mature capitula 6—
10 mm broad, pappus dull grey-brown. True ligulate florets absent. Naturalized in the
Mediterranean region, occurring in England only as a temporary shoddy alien
eS TRS ATS tte AO, NE OUT ES PU On USS cc ee kd Sen Gate. SLi t te C. bonariensis

ACKNOWLEDGMENTS

I wish to thank D. McClintock and J. B. Marshall for their encouragement and advice, and R.
Burton and D. Bevan for providing photocopies of several important references.

REFERENCES

Bonnier, G. E. M. & LAYENS, G. DE (1909). Flore complete portative de la France et de la Suisse. Paris.

Burton, R. M. (1985). Botanical records for 1984. Lond. Nat., 64: 113-124.

Cosson, E. S.-C. & GERMAIN, J. N. E. (1845). Flore descriptive et analytique des environs de Paris. Paris.

De LANGHE, J.-E. et al. (1973). Nouvelle flore de la Belgique, du Grand-Duché de Luxembourg, du nord de la
France, et des régions voisines, 3rd ed. Bruxelles.

Fournier, P. (1961). Les quatre flores de France. Paris.

JEANPERT, H. E. (1977). Vade-mecum du botaniste dans la région parisienne. Paris.

JEE, N. (1962). Botanical section report for 1961. Rep. Trans. Soc. Guernésiaise, 17: 96.

Jovet, P. & Vi_morin, R. DE (1975). Flore descriptive et illustrée de la France, 3¢me supplément. Paris.

Le Maout, J. E. M. & Decaisng, J. (1855). Flore élémentaire des jardins et des champs. Paris.

McCuntock, D. (1975). Wild flowers of Guernsey. London.

Merat, F. V. (1812). Nouvelle flore des environs de Paris. Paris.

PaLMER, J. R. (1983). Conyza bonariensis (L.) Cronq. in central London. B.S.B.I. News, 37: 16.

PERRING, F. H. & WALTERS, S. M., eds. (1962). Atlas of the British flora. London.

RAND, M. W. (1975). Botanical report, 1974. Rep. S. Essex nat. Hist. Soc., 1974: 35, 37.

Rovuy, G. (1927). Conspectus de la flore de France. Paris.

Ryan, P. (1977). Botanical section report for 1976. Rep. Trans. Soc. Guernésiaise, 20: 26-28.

(Accepied October 1987)

Watsonia, 17, 149-152 (1988) 149

The reintroduction of Cirsium tuberosum (L.) All. in
Cambridgeshire

C. D. PIGOTT

University Botanic Garden, Cambridge, CB2 1J]F

ABSTRACT

The Tuberous Thistle (Cirsium tuberosum (L.) All.) became extinct at its only locality in Cambridgeshire in
1974. A plant from the population was transferred to the University Botanic Garden at Cambridge in 1954 and
has been used to provide plants which, in 1987, were planted in grassland dominated by Arrhenatherum elatius
and Festuca rubra close to the original locality.

INTRODUCTION

The Tuberous Thistle (Cirsium tuberosum (L.) All.) is a sub-Atlantic species with its main area of
distribution at altitudes below 1450 m in southern and south-western Germany, northern
Switzerland and eastern and central France (Hegi 1928; map in Schonfelder 1970). There are a few
localities in adjacent parts of Austria and northern Italy, and scattered localities in northern France
and northern Belgium to latitude 50°20’N (van Rompaey & Delvosalle 1972). The localities in
Glamorgan and Wiltshire are now the northern limit of the species, but the most northerly locality
was that in Cambridgeshire at latitude 52°09'N.

THE LOCALITY IN CAMBRIDGESHIRE

Cirsium tuberosum was discovered in Cambridgeshire by Dr W. H. Miils in July 1919. In their
report of the discovery Mills & Evans (1922) refer to the possibility that the species had been found
much earlier, but confused with C. pratense Hudson (= C. dissectum (L.) Hill). At that time these
two species were not distinguished. The record in Professor C. C. Babington’s manuscript, to
which they refer, though from the ““Wimpole District” was, in fact, from Hayley Wood and it
seems unlikely that this could be the same locality.

In 1919 there were many plants of the thistle growing on an ancient grassy trackway and on the
grassy headland of an adjacent field, but already by 1922 some of the plants in the field had been
destroyed by ploughing (Mills & Evans 1922). The precise position of the original locality was for
many years kept a closely-guarded secret; it was, in fact, on the Mare Way above Little Eversden,
and in the adjacent field on the northern side at GR 52/363.519. The Mare Way is an ancient
ridge-way, forming the boundaries between parishes (mere, a boundary), and this with the upper
edge of the field were regarded by Mills & Evans (1922) as original downland that had escaped
being ploughed after enclosure at the beginning of the nineteenth century, when so much pasture
was lost to “the rapacity of modern agriculturists” (Babington 1860, pp. xiv—xv). In 1925 two other
populations of the thistle were found on the southern slope of the hill about 750 m from the original
locality, probably at GR 52/356.520 on the edge of a small plantation (Sharp Hill Plantation) and
on the boundary between the fields north of this plantation (unpublished records of Dr W. H.
Mills).

During the second World War and in the years immediately after, the Mare Way became choked
by scrub and brambles spreading in from the two hedges, and ploughing encroached further on the
headland of the adjacent field. By the 1950s the thistle had disappeared from the trackway and was
greatly reduced in the field, and, by the 1960s, only two patches of the plant remained. These were

150 Cx7D; PIGOEE

almost in the base of the broad hedge of Crataegus monogyna and Prunus spinosa in a grassland
composed of the following species: Arrhenatherum elatius, Dactylis glomerata, Deschampsia
cespitosa, Festuca rubra, Lolium perenne and Trisetum flavescens. It also contained scattered
plants of: Agrimonia eupatorium, Cirsium arvense, C. vulgare, Galium verum, Knautia arvensis,
Picris echioides, Primula veris, Senecio erucifolius and Sison amomum. This represents a well-
defined regional variant, containing such species as Picris and Sison, of the Festuca rubra
subcommunity of the Arrhenatherum elatius community recognised by the National Vegetation
Classification. The soil at the site was a heavy clay containing small pebbles of erratic rocks over
broken chalk.

The two remaining plants were seen year after year until 1973 when, in spite of strenuous efforts
to conserve the site, the whole area was ploughed and even the hedge on the north side of the Mare
Way was destroyed. The leaves reappeared in 1974, but thereafter no plants were seen and now so
little suitable land remains, even though the trackway has been reinstated, that it seems certain the
original population is lost.

No information about the thistle at the localities disco. -red in 1925 has been recorded, but a
search in 1987 revealed neither the plant, nor any suitable grassland in or beside the plantation.
The field boundary to the north has disappeared and crops entirely cover it. At the western end of
Sharp Hill Plantation a narrow bank of suitable grassland remains between two arable fields. This
is dominated by Arrhenatherum elatius and Festuca rubra and contains such species as Primula
veris, Viola hirta, Blackstonia perfoliata and Clinopodium vulgare, but a search failed to discover
any sign of C. tuberosum.

In 1954 a small lateral shoot of one of the plants was taken to the University Botanic Garden in
Cambridge by Dr S. M. Walters and has been propagated vegetatively. This plant is
hermaphrodite and, in 1986, capitula were bagged and selfed by V. Morgan and produced a few
fertile achenes from which two plants were successfully raised.

The gift of the Wimpole Estate to the National Trust in 1976 made it feasible to consider re-
establishing the Tuberous Thistle close to the original sites on land in ownership favourable to
conservation. Most of the Park is grazed by sheep or cattle and, although the thistle might survive
in a stunted vegetative state, it would be unlikely to flower in these conditions. The Arrhenatherum
elatius community in which it originally grew is characteristically a lightly grazed or mown
grassland. A search of the estate disclosed a few small areas of grassland which are not grazed by
stock. One such area carried a community similar to that at the original site. In 1986, after detailed
discussions, the National Trust agreed to the thistle being planted on this site.

The site selected is a low bank facing west on the gentle south-facing slope. The grassland is
composed of the species shown in Table 1 and belongs to the same subcommunity as did the
original site; there are scattered bushes of Acer campestre and suckers of elm.

By midsummer the sward has an average height of 20-35 cm with an ill-defined mosaic of slightly
shorter patches. Although there is a risk of the site being affected by drift of herbicides or
fertilisers, the present composition of the grassland suggests that the area has until now escaped
significant damage.

The soil is Brown Calcareous Soil with a variable depth (20-35 cm ) of ochre-coloured, heavy
clay, overlying a horizon containing fragments of chalk, flints and small erratic rocks derived from
the Great Chalky Boulder Clay. In summer the upper horizon cracks vertically into columns; part
of the area is kept moist by seepage of water. Although in Britain C. tuberosum normally grows on
free-draining soils, in Germany it is also characteristic of seasonally wet soils which support
vegetation containing Molinia caerulea (Oberdorfer 1981).

METHOD OF REINTRODUCTION

Six plants of C. tuberosum, four of the original stock and two grown from achenes produced by
selfing, were washed clean and repotted in 1986 in soil obtained from the site at Wimpole. On 7
May 1987, these plants and the soil were planted into holes cut in the turf and their exact positions
recorded on a plan. The plants were heavily watered at planting and ten days later; they showed no
sign of wilting in a short drought in July and thereafter the summer of 1987 was exceptionally wet.
By June two of the plants had been damaged by slugs and some of the developing inflorescences

REINTRODUCTION OF CIRSIUM TUBEROSUM 151

TABLE 1. SPECIES COMPOSITION AND PERCENTAGE COVER-ABUNDANCE OF THE SITE OF
REINTRODUCTION OF CIRSIUM TUBEROSUM

Arrhenatherum elatius 50 Galium verum 3
Brachypodium sylvaticum 5 Glechoma hederacea 5
Bromus erectus 1 Heracleum sphondylium 6)
B. sterilis 5 Knautia arvensis 1
Carex flacca 3 Mercurialis perennis 5
Dactylis glomerata 10 Picris echioides 1
Festuca rubra 30 Rumex conglomeratus 1
Holcus lanatus 2 Taraxacum officinale 3
Phleum nodosum 1 Veronica chamaedrys 3
Poa pratensis 5 Viola hirta 3
Trisetum flavescens 2

Brachythecium cf. rutabulum 10
Cirsium arvense 3 Eurhynchium swartzii 10
C. vulgare 1 Fissidens cristatus 1
C. eriophorum 2 Pseudoscleropodium purum 1

Weissia cf. microstoma 1

were bitten off, probably by hares (Lepus capensis L.). By August four plants were vigorous and
well established, one was weak and one had been lost after being undermined by a run of field vole
(Microtus agrestis L.). This plant will be replaced.

DISCUSSION

In general, plants should not be introduced into sites where they appear to be native because it may
confuse knowledge of their natural distribution, of their natural genetic variability and of their
ecology. There may also be unforeseen and undesirable consequences. The decision to attempt to
return C. tuberosum to the wild was not taken lightly and was felt to be justified on the following
grounds:

1) The species is very rare in Britain and the original localities in Cambridgeshire were isolated
and far removed from the nearest localities in Wiltshire.

2) The population was probably long-established and apparently not hybridised with other
species, particularly C. acaule, as is the case in some populations in Wiltshire and the
population at Nash Point, Glamorganshire.

3) The destruction of the population and its habitat was witnessed and now so little suitable
ground remains that it is almost certain that the plant is extinct.

4) As the population was reduced and increasingly threatened, an off-shoot was transferred to the
University Botanic Garden where it has been propagated vegetatively, so that there has been
no possibility of hybridisation. An individual from the original population was thus available.

5) Even with a plant grown as easily as C. tuberosum there is a risk in a large collection that it
could be lost, or its origin confused. Maintenance of pure stocks in gardens requires constant
care, is labour-intensive and therefore expensive.

6) Ina garden a plant is isolated from its natural environment and not subject to selection imposed
by that environment. C. tuberosum is gynodioecious, but the particular plant is hermaphrodite
and self-fertile. Allowing it to reproduce by seed in a garden may lead to selection for garden
conditions and, for this reason, fruiting heads have normally been cut off before maturity.

7) It has been possible to find a site which closely resembles the original as judged by the plant
community and soil, and which is under sympathetic ownership and treatment.

8) Establishment of a population from one individual is no different from natural establishment
from a single fruit.

9) If successfully established the species becomes a member of a community of plants and animals
including its natural pollinators and predators. Not least, the plant of local origin is preserved
and may be seen and enjoyed in as near as possible its natural habitat.

152 C. D. PIGOTT

Introductions must be adequately recorded. The exact locality is not given in this report to allow
the plants to become established without disturbance. Those who chance upon the plants are asked
to avoid disturbing them. The progress in establishment will be recorded and, when appropriate,
the precise locality published.

ACKNOWLEDGMENTS

I am grateful to the National Trust for granting permission for the experiment. I also thank Dr S.
M. Walters, who originally collected the plant, and Mrs G. Crompton for their permission to use
unpublished information, and Mr Ian Young, the Park Warden, for his assistance.

REFERENCES

BaBINGTON, C. C. (1860). Flora of Cambridgeshire. London.

Heal, G. (1928). Illustrierte Flora von Mittel-Europa, 6: 2. Minchen.

MiLis, W. H. & Evans, A. H. (1922). Short notes: Cirsium tuberosum in Cambridgeshire. J. Bot., Lond., 60:
21

OBERDORFER, E. (1981). Pflanzensoziologische Exkursions-Flora. Stuttgart.

SCHONFELDER, P. (1970). Siidwestliche Einstrahlungen in der Flora und Vegetation Nordbayerns. Ber. der
Bayerischen Bot. Ges., 42: 17-100.

VAN Rompaey, E. & DELVOSALLE, L. (1972). Atlas de la flore Belge et Luxembourgeoise. Bruxelles.

(Accepted December 1987)

[Editors’ note: the B.S.B.I. Panel for Introductions should be consulted before similar projects
are undertaken; contact the Conservation Committee in the first instance. ]

Watsonia, 17, 153-157 (1988) 153

Two subspecies of Molinia caerulea (L.) Moench
in the British Isles

P. J. O. TRIST
Glovers, High Street, Balsham, Cambs., CB1 6DJ
and
P:.D. SELL

Botany School, Downing Street, Cambridge, CB2 3EA

ABSTRACT

Two subspecies of Molinia caerulea (L.) Moench are recognised in the British Isles, the history of their
discovery set out, their characters given and their nomenclature explained. Known localities for subsp.
altissima (Link) Domin are listed in vice-county order.

INTRODUCTION

Many infraspecific taxa of Molinia caerulea (L.) Moench have been described, but in Flora
Europaea (Tutin 1980) only two subspecies are recognized. Subsp. caerulea is found on heaths,
moorland and mountain grassland, as well as in depressions on river banks, in peat fens, on stony
shores of lochs and on mountain cliff ledges up to 1220 m. The plants are often found in isolation in
open situations and not in competition with other vegetation; but in bog conditions, plants of
subsp. caerulea are often crowded.

Subsp. altissima (Link) Domin (subsp. arundinacea (Schrank) H. Paul) is a much larger plant in
all its parts, and is more specific in its habitat. It is a plant of open fens and fen river valleys and
occasionally in fen scrub reversion. The texture of the soil on which it grows is a light peat or coarse
sand over peat with a fluctuating water level which is high in winter. Its associated species are tall,
strong plants which create dense ground cover and competition and include Angelica sylvestris,
Cladium mariscus, Eupatorium cannabinum, Filipendula ulmaria, Galeopsis tetrahit, Juncus
effusus, J. subnodulosus and Phragmites australis. It is sometimes shaded by Betula and Salix. The
tall stem of subsp. altissima has the ability to push its way into the above mentioned vegetation and
the long, lax panicle is often hidden from the casual observer.

HISTORY

Subsp. altissima was first described as Melica caerulea var. major by Roth in April 1789 and as a
species, M. arundinacea Schrank, later in the same year. Var. major was taken up by Babington
(1856) and Syme (1872) and has been used in some County Floras. The earliest reference to it in
British Floras which we can trace, however, is as Monilia caerulea var. sylvatica by S. F. Gray (1
November 1821) which is presumably based on Enodium sylvaticum Link (ante 30 June 1821).
Conert (1961) has discussed the morphology and taxonomy of the plant at some length.

The current distribution of this subspecies is imperfectly known. A search in herbaria under M.
caerulea var. major, var. robusta, var. arundinacea and M. litoralis has produced 58 sheets of
subsp. altissima from 47 sites: 42 from England & Wales, three from Scotland and two from
Ireland. Of these records, only six refer to finds prior to 1900. From 1900 to 1955, only 31 sites

154 P. J.O. TRIST AND P. D. SELL

were found and only ten new sites have been reported between 1955 and 1985. For this record
there are two possible reasons. First the general lack of interest in infraspecific taxa may mean that
it has been ignored, but secondly, and more likely, its specific habitat (needing a fluctuating water
level) has had interference or has been destroyed by drainage and more ruthless maintenance of
ditches since the Second World War.

NOMENCLATURE

The first problem concerns the generic name. Aira caerulea L. (i.e. Molinia caerulea) (1753) was
selected as the lectotype of Aira L. (1753, 1754) by Britton & Brown (1913) and is accepted by
Index Nominum Genericorum (1979). This typification is undesirable in that it does not allow Aira
to be used in the sense in which it is most commonly accepted. It can be rejected on the grounds
that it was made on a largely mechanical basis (Art. 8). Hitchcock (1920) selects Aira caespitosa L.
(=Deschampsia caespitosa). This also is contrary to common usage. The author says he selected it
arbitrarily, but considering Linnaeus’ description of the genus it does appear to be logical.
Hitchcock & Green (1929), Philips (1951) and Clayton & Renvoize (1986) all choose A. praecox L.
as the lectotype of Aira. This is clearly the most sensible attitude, but A. praecox surely needs
conserving as the lectotype to make sure neither A. caerulea or A. caespitosa can be used. The
lectotype of Molinia Schrank is M. varia Schrank which is a superfluous nomen illegitimum of M.
caerulea (L.) Moench.

Aira caerulea was described by Linnaeus on page 63 of Species Plantarum in 1753. The diagnosis
‘Aira foliis planis, panicula coarctata, floribus pedunculatis muticis convoluto subulatis’ is taken
verbatim from his Flora Suecica. It is thus a species he knew in his native Sweden and which he
would himself have collected. In the Linnaean herbarium is a sheet (Savage Cat. 85/1) labelled
“caerulea 3”’. It fits the original diagnosis and is an excellent specimen of what is regarded as the
most common form of Molinia caerulea. We designate it as the lectotype of Aira caerulea L. This
establishes the application of the nominate subspecies.

The second subspecies was first recognized as Melica caerulea var. major Roth in 1789. Between
then and 1829 it received no fewer than six specific names. H. Paul (1937) gave it the rank of
subspecies, basing it on M. litoralis Host (1827). In the following year he changed his mind and
made a new combination based on M. arundinacea Schrank (1789). Although Schrank’s name was
earlier it also was at the rank of species so that M. caerulea subsp. litoralis (Host) H. Paul was in
fact earlier.

Tutin (1980) used the name M. caerulea subsp. arundinacea. However, there is an even earlier
subspecific name, M. caerulea subsp. altissima (Link) Domin (1935), which was in fact given as a
synonym in Flora Europaea. The types of these names have not been examined, but from their
original descriptions and general usage there seems to be little doubt that they refer to the large
variant of Molinia caerulea.

MOLINIA Schrank, Baier. Fl. 334 (1789) (Lectotype: M. varia Schrank nom. illegit. = Aira
caerulea L., i.e. Molinia caerulea (L.) Moench, vide Hitchcock, U.S. Dept. Agric. Bull., 772: 50
(1920)).

Enodium Pers. ex Gaudin, Agrost. Helv., 1: 145 (1811) (Holotype: E. caeruleum (L.) Gaudin);
superfl. nom. illegit. pro Molinia Schrank ab lectotyp.

Monilia S. F. Gray, Nat. Arrang. Brit. Pl., 2: 110 (1821) (Holotype: M. caerulea (L.) S. F. Gray);
superfl. nom. illegit. pro Molinia Schrank ab lectotyp.

M. CAERULEA (L.) Moench, Meth. 183 (1794).

a) subsp. CAERULEA

Aira caerulea L., Sp. Pl. 63 (1753) (Lectotype: Savage Cat. 85/1 (LINN), designated here).
Melica caerulea (L.) L., Mantissa Alt. 325 (1771).

Molinia varia Schrank, Baier. Fl. 334 (1789), superfl. nom. illegit. pro Aira caerulea L.
Molinia variabilis Wibel, Prim. Fl. Werth. 115 (1799).

Festuca caerulea (L.) DC. in Lam. & DC., Fl. Fr., 3rd ed., 3: 46 (1805).

Enodium caeruleum (L.) Gaudin, Agrost. Helv., 1: 145 (1811).

TWO SUBSPECIES OF MOLINIA CAERULEA 155

Arundo agrostis sensu Lapey., Hist. Abr. Pl. Pyr. 52 (1813).

Hydrochloa caerulea (L.) Hartm., Genera Gram. 8 (1819).

Monilia caerulea (L.) S. F. Gray, Nat. Arrang. Brit. Pl., 2: 110 (1821).

Molinia depauperata Lindley, Syn. Brit. Fl. 307 (1829) (Holotype: Clova Mountains, Forfar, v.c.
90, D. Munro (CGE)).

Molinia minor Holandre, Fl. Moselle 813 (1829).

Molinia obtusa Peterm. in Flora (Regensb.), 27: 235 (1844).

A densely tufted, strong-rooted perennial forming clumps, which consist of a number of single
culmed plants. Culms 8-65 cm, erect, strong. Sheaths smooth and ribbed, the basal internode
clothed with short sheaths 4-10 cm and scales of 1-2 cm. Leaves 10-50 (—63) cm x (1.5)3-6 (—8)
mm, long-pointed, flat, with very short hairs only, or some hairs of 2-3 mm scattered on the upper
surface, rough on the margins, deciduous in winter. Ligule 0, but mouth of sheath ciliate, the cilia
0.5—2.0 mm. Panicles 1-30 cm, dense to widely interrupted. Spikelets 3.0-5.5 mm, dark mauve to
light green, lanceolate, up to 4-flowered but mainly 1-2. Glumes persistent, unequal to equal,
lanceolate to ovate; lower 1.5—3.0 mm, upper 2.2—3.5 mm. Lemma 3+4 mn, lanceolate.

(Hubbard’s (1968) description of this taxon is not comparable as his measurements make it clear
that those of subsp. altissima are included.) The height of plants and the length of panicles are
influenced by sheep grazing.

b) subsp. ALTISSIMA (Link) Domin in Preslia, 13-15: 39 (1935).

Melica caerulea var. major Roth, Tent. Fl. Germ., 2(1): 103 (April 1789).

Molinia arundinacea Schrank, Baier. Fl., 1: 336 (June-Dec. 1789).

Aira atrovirens Thuill., Fl. Par., 2nd ed., 1: 37 (1800).

Enodium sylvaticum Link, Enum. PI. Hort. Berol. Alt., 1: 80 (ante 30 June 1821).
Monilia caerulea var. sylvatica (Link) S. F. Gray, Nat. Arr. Brit. Pl., 2: 110 (1 Nov. 1821).
Enodium atrovirens Dumort., Obs. Gramin. Belg. 108 (1824).

Molinia litoralis Host, Fl. Austriac. 118 (1827).

Molinia altissima Link, Hort. Berol., 1: 197 (1827).

Molinia sylvatica (Link) Link, Hort. Berol., 1: 197 (1827).

Enodium litorale (Host) Kunth, Enum. Pl., 1: 380 (1833) nom. in syn.

Molinia caerulea var. arundinacea (Schrank) Ascherson, Fl. Brand. 837 (1864).

Molinia caerulea var. robusta Prahl, Krit. Fl. Schl. Holst., 2: 257 (1890).

Molinia caerulea b. arundinacea (Schrank) Richter, Pl. Eur., 1: 72 (1890).

Molinia caerulea subsp. litoralis (Host) H. Paul in Ber. Bayer. Bot. Ges., 22: 18 (1937).
Molinia caerulea subsp. arundinacea (Schrank) H. Paul in Ber. Bayer. Bot. Ges., 23: 154 (1938).

A densely tufted and extensive, strong-rooted perennial, building a tussock of 20 cm high. Culms
very variable and variation presumably related to moisture supply. Culms (63—) 75—124 (—162) cm,
erect. Sheath smooth and ribbed, the basal internode clothed with short sheaths and scales slightly
longer than in subsp. caerulea. Leaves (28—) 40-62 (—78) cm X 4.0-8.5 mm, long-pointed, flat,
tapering from near the base, with short hairs scattered on the upper surface or glabrous, margins
smooth, deciduous in winter. Panicle (23—) 35-53 (-65) cm, generally widely interrupted with many
unequally long branches 4-15 cm, patent to erecto-patent, dark green to light mauve. Spikelets
variable in length, (3—) 4-7.5 mm; glumes unequal to equal, lanceolate, pointed; the lower 1.6—3.2
mm, upper 2.3-4 mm. Lemma (3.2-) 3.5—5.4 (-5.7) mm, lanceolate.

DISTRIBUTION OF SUBSP. ALTISSIMA

v.c. 1, W. Cornwall, Pengerswick Castle, 1873, J. Cunnack (K); Trelawney Mill, 1919, E.
Thurston (K).

v.c. 2, E. Cornwall, Boscastle, 1915, E. Thurston (K); Pencarrow Wood, Bodmin, 1957, C. C.
Townsend (CGE, K).

v.c. 4, N. Devon, Banks of R. Lyn, Watersmeet near Lynmouth, 1902, S. H. Bickham (CGE,
NMW).

Ps J7 OF TRISTAND P-D:? SELL

. 5, S. Somerset, Shapwick, 1936, F. K. Makins (K).
. 6, N. Somerset, Walton in Gordano, 1908, C. Bucknall (RNG); Wood at Penselwood, 1916,

W. Herridge (EXR); Beacon Hills, Shepton Mallet, 1942, J. P. M. Brenan (K); S. of Ashcott
Railway station, 1944, J. P. M. Brenan (K).

. 9, Dorset, Arne Heath, Wareham, 1899, L. V. Lester Garland (K).

. 10, Wight, St Helen’s, 1857, A. G. More (CGE).

. 11, S. Hants., Near Milton, 1913, J. Comber (BM, K, NMW).

. 17, Surrey, Near Woking, 1900, H. J. Riddelsdell (BM); Wimbledon Common, 1894, J. Fraser

(K); Esher Common, 1928, C. E. Hubbard & V. S. Summerhayes (K); Richmond Park, 1929,
V. S. Summerhayes & H. M. Montford (K); Brookwood bank of Basingstoke Canal, 41/
952.572, 1985, P. J. O. Trist (CGE, Herb. P.J.O.T.).

. 19, N. Essex, Middlewick, S. of Colchester, 1924, G. C. Brown (CGE, K, NMW).
. 22, Berks., Parson’s Moor, near Cothill, 1943, C. E. Hubbard (K); Cothill Fen, near Oxford,

1955, F. White (EXR).

. 23, Oxon, North Leigh Heath, 1944, C. E. Hubbard (K).
. 24, Bucks., Wilstone, near Thame, 1913, G. C. Druce (NMW); East Burnham Common,

1928, V. S. Summerhayes & E. Nelmes (K).

. 26, W. Suffolk, Palmers Heath, near Brandon, 52/742.847, 1972, P. J. O. Trist (Herb.

P.J.O.T.); Lt. Eriswell Hall, Lakenheath, 52/721.708, 1976, P. J. O. Trist (Herb. P.J.O.T.);
Pashford Poors Fen, Lakenheath 52/736.836, 1974, 1976, 1986, P. J. O. Trist (CGE, Herb.
P:J.O-2.).

. 28, W. Norfolk, Dersingham Common, 1928, C. E. Hubbard (K); Rockland marshes, 1935, F.

Ballard & C. E. Hubbard (K); Cranberry Fen, Wolferton, 1921, A. R. Horwood (NMW);
Breckles Heath, Stow Bedon, 1971, C. E. Hubbard (K); Little and Middle Fens, South
Lopham, 52/042.795 and 52/052.798, 1985, P. J. O. Trist (Herb. P.J.O.T., CGE); Gooseberry
Lane, Castle Rising, 1975, E. L. Swann (K).

. 29, Cambs., Great Heath Wood, Gamlingay, 1910, R. S. Adamson (BM).

. 33, E. Gloucs., Driffield, 1942, H. K. Airy Shaw (K).

. 35, Mons., South-west of Bassaleg, 1934, A. E. Wade (NMW).

. 36, Herefs., Bucknall’s Wood, Madley, 1905, H. J. Riddelsdell (NMW).

. 43, Rads., Llandrindod Wells, 1916, W. C. Barton (K).

. 46, Cards., Esgair Elan, east of Devil’s Bridge, 1984, A.G. de R. Channer (RNG).

. 54, N. Lincs., Epworth, 1945, J. M. Taylor (K).

. 55, Leics., Bradgate Park, 1944, R. M. Payne (K).

. 59, S. Lancs., Simonswood Moss, 1897, 1898, J. A. Wheldon (NMW); Risley Moss, 1907, J. A.

Wheldon (NMW).

. 63, S. W. Yorks., Thorne Waste, near Doncaster, 1942, S$. P. Rowlands (K).

. 80, Roxburghs., Newcastleton State Forest, S. of Tweedenhead, 1934, D. H. S. Davis (K).
. 100, Clyde Is., West of Brennan Head, Arran, 1934, R. Mackechnie (RNG).

. 104, N. Ebudes, Loch Ainort, Skye, 1934, H. M. Montford (K).

. H1, S. Kerry, Between O’Sullivan’s Cascade and Shehy Mountains, 1935, S. Ross-Craig, B.

L. Burtt & J. R. Sealy (K).

. H11, Co. Kilkenny, Kilcoron House, Kilkenny, 1937, N. L. Bor (K).

ACKNOWLEDGMENTS

We are grateful to the curators of the herbaria at BM, EXR, K, NMW and RNG for the loan of
specimens, to A. O. Chater and S. M. Walters for reading and commenting on the text, and to D.
H. Kent for information on distribution and nomenclature.

REFERENCES

BaBINGTON, C. C. (1856). Manual of British botany, 4th ed. London.
Britton, N. L. & Brown, A. (1913). An illustrated Flora of the northern United States, 2nd ed. New York.
CLayTton, W. D. & RENvoizE, S. A. (1986). Genera graminum. London.

TWO SUBSPECIES OF MOLINIA CAERULEA 157

ConerT, H. J. (1961). Die systematik und Anatomie der Arundinea. Weinheim.

Domin, K. (1935). Plantarum Cechoslovakiae enumeratio. Preslia, 13-15: 1-305.

Gray, S. F. (1821). A natural arrangement of British plants. London.

Hitcucock, A. S. (1920). The genera of grasses of the United States. U.S. Dept. Agric. Bull., 772: 47-50, 114—
116.

Hircucock, A. S. & GREEN, M. L. (1929). Proposed standard species of Linnean generic names. Proposals of
British Botanists.

Host, N. T. (1827). Flora austriaca, 1: 117-118. Vienna.

Hussarp, C. E. (1968). Grasses, 2nd ed. Harmondsworth.

LINNAEUS, C. (1753). Species plantarum. Holmiae.

Linnaeus, C. (1754). Genera plantarum, 5th ed. Holmiae.

Pau, H. (1937). Beitrage zur Kenntnis des Formenkreises des Besenriedes Molinia caerulea Moench in
Bayern. Ber. Bayer. Bot. Ges., 22: 15-22.

Pau, H. (1938). Nachtag zu ‘Beitrage zur Kenntnis des Formenkreises des Besenriedes Molinia caerulea
Moench in Bayern’. Ber. Bayer. Bot. Ges., 23: 154.

Puiips, E. P. (1951). The genera of South African flowering plants, 2nd ed. Pretoria.

Rotu, A. W. (1789). Tentamen florae germanicae, 2(1): 103. Lipsiae.

SCHRANK, F. P. von (1789). Baiersche Flora. Minchen.

SyME, J. T. B. (1872). In Sowersy, J. E. English botany, 3rd ed., 11: 89-91. London.

TutTin, T. G. (1980). Molinia Schrank, in Tutin, T. G. et al., eds. Flora Europaea, 5: 254. Cambridge.

(Accepted December 1987)

na

adie
,

Watsonia, 17, 159-162 (1988) 159

Notes on Avenula pratensis (L.) Dumort. in Britain

J. M. DIXON

School of Environmental Science, The University, Bradford, BD7 1DP

ABSTRACT

It is suggested in the literature that Avenula pratensis (L.) Dumort. from the mountains of Scotland and
northern England might represent a taxon, described as Avena alpina Sm., distinct from that of lowland
Britain. Morphological observations made on A. pratensis from sites throughout Britain do not support this
suggestion, nor do specimens from Scotland and northern England agree with the published description of
alpine Avena alpina.

INTRODUCTION

A rider to the entry for Avenula pratensis (L.) Dumort. in Flora Europaea (Holub 1980) states that
“Plants from the mountains of Scotland and N. England, with larger more strongly coloured
spikelets, described as Avena alpina Sm., may represent a distinct taxon’’.

In Britain, A. pratensis is found to 366 m in Derbyshire, to 457 m in Yorkshire and to 838 m in
Scotland (Wilson 1956). Smith (1811) described his specimens, collected from the summits (307 to
945 m) of the Clova mountains, Angus, as forming compact tufts with the roots ‘‘not inclined to
creep” (cf. Avenula pubescens). The leaves resembled those of A. pratensis, being non-hairy and
having serrated edges, but differed from the latter in their rough and greatly elongated sheaths.
The flowers differed from A. pratensis ‘‘not only in their much greater size”’ (no figures given),
“but in their partial stalk, or rachis (rachilla?), the hairiness of which I observe to be crowded up
into a very dense tuft, towards the base of each floret, not dispersed over the whole rachis’. Smith
made no mention of the colour of the spikelets.

Hegi (1967) described Avena alpina as having sheaths not, or only slightly, rough and with the
culm rough only in the panicle, whereas the culms of A. pratensis are upwardly rough. A. alpina
usually has two basal branches to its panicle, as opposed to one, seldom two, in A. pratensis. Like
Smith, he specified that the spikelets of A. alpina are larger than those of A. pratensis (but again no
figures are given), with 6 to 8 florets. He states that the glumes are overlaid with violet, and that
the lemmas bear an awn which is “up to a little over 1.5 cm long’, compared with A. pratensis,
whose awns are “up to almost 2.0 cm long”’.

MORPHOLOGICAL OBSERVATIONS

Numerous specimens of A. pratensis from the British Isles have been examined (site details are
given in Table 1) and the following observations made:—

1. Culms of specimens examined were mostly smooth in the lower part — occasionally small
protuberances were noticed. When examined microscopically these were found to occur at right
angles to the culm until the panicle was almost reached, when they resolved into minute upward
pointing teeth. The rachis of all specimens had upwardly pointing teeth, variable in size and
quantity; all pedicels were minutely toothed.

2. None of the specimens examined had noticeably rough or elongated sheaths.

3. Spikelet measurements from 55 different sites indicated that there is indeed a significant
difference (at p <0.05) in mean spikelet length between Scotland plus the north of England
(defined as Yorkshire and Lancashire northwards), and the Midlands, Wales and the south of
England. Mean spikelet lengths are given in Fig. 1. However, all the British material examined

TABLE 1. SITES FROM WHICH AVENULA PRATENSIS WAS EXAMINED

Location

Scotland

Aberfeldy, Perthshire

Beinn Laoigh, Argyllshire
Ben Lui, Perthshire

Braid, Edinburgh
Cockburnspath, Berwickshire
Craig Dorney, Aberdeenshire
Creag an Lochan, Perthshire
Forres, Morayshire

Glen Clova, Forfarshire
Glen Doll, Forfarshire

Glen Lyon, Perthshire
Golspie, Sutherland

Loch Lyon, Perthshire

Loch Pattack, Invernesshire
Relugas, Morayshire
Tomintoul, Banffshire

North of England

Arnside Knott, Cumbria
Aysgarth Falls, N. Yorkshire
Giggleswick, N. Yorkshire
Gilsland, Northumberland
Gordale Scar, N. Yorkshire
Helmsley, N. Yorkshire
Malham Tarn, N. Yorkshire
Overdale, N. Yorkshire
Scout Scar, Cumbria

Shap, Cumbria

Skirethorn, N. Yorkshire
Sproxton, N. Yorkshire
Sutton Bank, N. Yorkshire
Teesdale, Durham

Wilson Scar, Cumbria

Midlands, Wales and the South of England

Bakewell, Derbyshire
Ballard Down, Dorset
Boxhill, Surrey

Collyweston, Northamptonshire

Coombe Hill, Buckinghamshire
Corfe Castle, Dorset
Cosgrove, Northamptonshire
Devil’s Ditch, Cambridgeshire
Dover, Kent

Epsom Downs, Surrey

Firle Beacon, Sussex

Kithurst Hill, Sussex

Little Ormes Head, Gwynedd
Litton, Derbyshire
Littonfields, Derbyshire
Llysfaen, Clwyd

Monksdale, Derbyshire
Monyash, Derbyshire
Newport, Gwent

Portland, Dorset

Risby Heath, Suffolk

Saffron Walden, Essex
Salisbury Plain, Wiltshire
Winter Hill, Berkshire

Grid
Reference

27/855 .473
27/262.274
27/266.266
36/255.698
36/790.712
38/403 .352
26/596.392
38/020.592
37/320.745
37/243.775
27/654.505
28/823 .957
27/450.424
27/545 .789
38/995 .487
38/170.190

34/453.777
44/016.890
34/810.648
35/634.668
34/915.638
44/585 .840
34/885 .694
44/840.715
34/488 .921
35/560.173
34/963.648
44/601.819
44/515 .826
35/902.280
35/548.182

43/184.670
40/030.815
51/178.516
53/010.002
42/848 .066
30/964 .823
42/792.426
52/581.645
61/300.398
51/217.584
51/485 .060
51/082.127
23/812.823
43/156.748
43/175.759
23/888.774
43/135.738
43/158.663
31/354.906
30/690 .697
52/790.683
52/570.405
41/055.451
41/873 .863

Vice-County

Approximate
altitude (m)

NOTES ON AVENULA PRATENSIS 161

14.3
18.3
21.0
20.1 16.5
19.0 15.5
i 2
15.5 16.3 as 14.5
19.7
~—F
18.4 19.5N. peiaizole
20.4 20.7
18.8
12.
15.4 12.3
14.3 15.
16.0 5.0 11.0
17.0
14.3
16.8; 21.7
16.6
15.3
20.2
19.0
18.6 18.5
14.0

Figure 1. Variation in spikelet length of Avenula pratensis in the British Isles. Figures are mean values (mm) of
spikelets taken from the middle of the panicle. Sample size varied between 5 and 30 inflorescences per
population. Maximum standard error for any sample was + 1.1 mm.

162 J. M. DIXON

falls within the range given for A. pratensis in Flora Europaea, namely 12—23(—28) mm. In only
nine out of 90 spikelets measured from Scottish sites were the lengths greater than 23 mm, and 23
out of 420 spikelets measured from England and Wales had lengths greater than 23 mm. Eleven of
these, interestingly, came from one site in Derbyshire. On the other hand, spikelets of 14.0 mm
were recorded from Scotland, and one depauperate specimen from East Anglia had spikelets of
only 10.0 mm. Thus, spikelets encompassing more or less the entire range enumerated in Flora
Europaea for A. pratensis have been found throughout Britain.

4. None of the specimens examined had an unusually hairy rachis, and no noticeable differences
were apparent in the quality or quantity of rachilla hairs.

5. The maximum number of florets per spikelet found was six, including rudimentary florets.

6. No distinct north/south variation in spikelet colour has been observed, and deeply coloured
specimens were found at several sites throughout Britain.

7. Awns varied from 12 to 27 mm in length with the longer awns generally matching the larger
spikelets of Scotland and the north of England, but again with almost the full range throughout
Britain.

8. Many specimens had two basal branches to their panicles. This character seems to depend on
soil depth/fertility. Specimens collected from cliffs, screes and poor shallow soils tend to have few
paired branches, whilst specimens from deeper ‘richer’ soils had more. From the 25 sites examined,
the populations having the maximum number of paired basal branches came from Salisbury Plain
and a population cultivated in the author’s garden, where all 20 panicles collected from each site
had paired basal branches. The minimum number came from a scree population at Gordale Scar,
North Yorkshire, and a population growing on a thin rendzina above Malham Tarn, North
Yorkshire where the 20 panicles collected from each site had no paired basal branches.

Comparing the above observations with the descriptions given for Avena alpina by Smith (1811)
and Hegi (1967) shows there to be no good correspondence, nor do any of the characters examined
fall outside the definitive limits given in Flora Europaea for Avenula pratensis. It is therefore
suggested that although A. pratensis occurring in Scotland and the north of England has on average
larger spikelets and longer awns, these features are not sufficient to refer them to a different taxon.

ACKNOWLEDGMENT

My grateful thanks go to Dr M. R. D. Seaward for his comments on a draft of the paper.

REFERENCES

Hea, G. (1967). Illustrierte Flora von Mittel-Europa, 3rd ed. Munich.

Ho us, J. (1980). Avenula (Dumort.) Dumort., in Tutin, T. G. et al., eds. Flora Europaea, 5: 210-216.
Cambridge.

SmiTH, J. E. (1811). An account of several plants recently discovered in Scotland by Mr George Don A.L.S.
not mentioned in the Flora Britannica nor English Botany. Trans. Linn. Soc. London, 10: 333-346.

Witson, A. (1956). The altitudinal range of British plants. Supplement to N. West Nat., 2nd ed.

(Accepted July 1987)

Watsonia, 17, 163-181 (1988) 163

Short Notes

PUTATIVE REYNOUTRIA JAPONICA HOUTT. xX FALLOPIA BALDSCHUANICA
(REGEL) HOLUB HYBRIDS DISCOVERED IN BRITAIN

Reports that seed taken from naturalized plants of Reynoutria japonica Houtt. in Britain is
frequently the result of hybridization with the commonly grown garden plant Fallopia
baldschuanica (Regel) Holub (Russian Vine) have been the subject of two B.S.B.I. exhibition
meeting displays (Bailey & Conolly 1984; Bailey 1987). These exhibits were accompanied by pleas
to those interested to try to find such plants growing in the wild. Experience at Leicester had shown
that although hybrid seed was capable of surviving the winter and germinating in the spring, such
seedlings would be unlikely to survive a British winter, and so the best chance would be to look for
seedlings under R. japonica plants in spring. These requests have, I am happy to report, borne
fruit, and have resulted in B.S.B.I. member D. Bevan, who had seen our exhibits, finding the first
R. japonica X F. baldschuanica growing in the wild; furthermore, the discovery was not just of a
seedling, but of a clump of some considerable size, at Railway Fields, Haringey, Middlesex, v.c. 21
(GR 51/317.882) in 1987. This has prompted me to give a fuller account of the discovery, incidence
and morphology of this new addition to the British flora.

Bailey & Conolly (1985) reported that the most commonly found variant of R. japonica in
Britain was octoploid (2n=88) and female. Persistent reports that such plants were capable of
setting some seed in the absence of male-fertile Reynoutria plants led to comparative examination
of the chromosome complements of the octoploids with their seedling offspring from eleven
localities in Britain (Table 1).

TABLE 1. LOCALITIES OF OCTOPLOID FEMALE R. JAPONICA PLANTS FROM WHICH SEED
WAS COLLECTED AND GROWN, AND THE NO. OF SUCH SEEDLINGS THAT HAD 54

CHROMOSOMES
No seedlings No seedlings
Location with 2n = 54 grown
V.c. 11, S. Hants. Petersfield GR 41/744.234 5 5
V.c. 12, N. Hants. Itchen Abbas GR 41/541.329 2) 5
V.c. 40, Salop Ironbridge GR 33/671.033 3 4
V.c. 48, Merioneth Boston Lodge GR 23/589.382 4 5
V.c. 48, Merioneth Tyn Coed GR 23/67.18 Z, 2
V.c. 48, Merioneth Ynys GR 23/597.353 2 2
V.c. 49, Caerns. Criccieth GR 23/492.381 4 4
V.c. 49, Caerns. Pentre’r-felin GR 23/526.396 3 5
V.c. 49, Caerns. Pwllheli GR 23/374.350 5 6
WiC255, Leics, Sileby GR 43/602.153 + 4
NViC295, Leics: Stoughton GR 43/644.026 5 3%

* artificial hybridization with F. baldschuanica

Since the female parent, R. japonica, had in all the above localities 88 chromosomes, it was
something of a surprise that all seedlings counted had 54 chromosomes. Furthermore, whatever
was going on was a very regular and widespread phenomenon, occurring as it did in widely
separated parts of the country. Cytologically the seedling karyotype was rather distinctive in that
ten large chromosomes could be readily distinguished from 44 smaller and more uniform
chromosomes typical of Reynoutria. Working on the assumption that these seeds were the result of
a fertilization event (rather than some bizarre meiotic aberration) one would be looking for a
pollen parent with 20 chromosomes. The cytological data in combination with the leaf shape and
the semi-twining habit of the seedlings pointed to the involvement of the diploid climber F.
baldschuanica (2n=20) as a putative pollen donor.

164 SHORT NOTES

Artificial hybridization between a female R. japonica from Stoughton and F. baldschuanica gave
rise to several seeds; three seedlings were subsequently grown on and found to have the same
chromosome number and karyotype as well as being virtually identical morphologically with the
plants grown from wild-collected seed. A further indication of the ubiquity of this phenomenon
came later, when I contacted Richard Scott of I.T.E., Merlewood, who had been conducting
research into the suitability of Reynoutria taxa as biomass producers. During the course of this
work he had collected seeds from female R. japonica growing next to a plant of R. sachalinensis,
thinking not unnaturally that they would grow into the interspecific hybrid. However, when I
examined these plants it quickly became apparent that they were hybrids between R. japonica and
F. baldschuanica.

Plants grown outdoors at Leicester for three years are becoming more vigorous but are
extremely reluctant to flower; some flower-buds were initiated on one plant this year (1987) but
were aborted before they reached maturity. In contrast, the plant at Haringey is extremely
vigorous, covering over 10 m’, and is strongly rhizomatous and with several densely flowered
inflorescences. Judging from its size, it must have been established for some time, since it is many
times larger than the three-year-old Leicester plants.

DESCRIPTION OF R. JAPONICA X F. BALDSCHUANICA

Superficially similar to R. japonica, but with stems much thinner and with smaller leaves. Stems
herbaceous, up to 2 m long, slender with red blotches, bending over almost to touch the ground,
reducing the height of the plant to c. 1.5 m. Leaves acuminate, ovate to narrowly ovate-oblong, to
13 X 6.5 cm; petioles slender, 2-2.5 cm long. Inflorescence of axillary and terminal panicles.
Flowers hermaphrodite (?), resembling R. japonica more than F. baldschuanica; the three outer
perianth segments more broadly keeled than in Reynoutria; anthers 0.60.7 mm long; style trifid,
with fimbriate, club-shaped stigmas (intermediate between the fimbriate and capitate stigmas of its
respective parents). Younger plants without rhizomes, or only weakly rhizomatous; established
clumps may have tough, woody rhizomes up to 2 cm in diameter. Flowering occurs very late in the
season (late September to early October); there are no reports of any seed set in this hybrid.

ACKNOWLEDGMENTS

I am indebted to Ann Conolly for her invaluable assistance and encouragement, to Professor Clive
Stace for his good advice, and to Professor Smith for permitting me the use of the Department’s
facilities. I would also like to thank the warden of Railway Fields Nature Conservation Park, Dave
Perry, for his kind assistance. This work was assisted in part by a Leicester University Research
Board Grant.

REFERENCES

BaILey, J. P. & CoNno.ty, A. P. (1984). A putative Reynoutria x Fallopia hybrid from Wales. Watsonia, 15:
162-163.

BalILey, J. P. & Cono.ty, A. P. (1985). Chromosome numbers of some alien Reynoutria species in the British
Isles. Watsonia, 15: 270-271.

BaILey, J. P. (1987). Reynoutria hybrids in the British Isles. B.S.B.I. News, 45: 36.

Jo Ps BAILEY
Department of Botany, The University, Leicester, LE] 7RH

SHORT NOTES 165
THE VASCULAR PLANT HERBARIUM OF E. C. WALLACE

The herbarium of the late Edward Charles (Ted) Wallace (1909-1986) was massive and
exceptionally well curated. His bryophyte collections were bequeathed to NMW and his lichens to
the British Lichen Society. The vascular plants were presented to RNG by Mr Alan Crundwell to
whom they had been bequeathed (Jury 1987).

The vascular plant herbarium was kept in an upstairs bedroom of his semi-detached house in
Surrey in four purpose-built cupboards with a total of 137 shelves! Even so this was not enough and
the specimens of Carex were housed separately in 15 boxes. The collection was arranged according
to Dandy (1958) with the sheets of each species in strict vice-county order. No trace of any insect
damage was found, except in some unmounted duplicate material which was destroyed.

Although still avidly collecting specimens to near his departing, no backlog of material awaiting
processing or incorporating was found: Ted was too organized. A recent obituary (Richards 1987)
gives details of his life, travels and great dedication to botany. Examination of his specimens also
shows just how great his botanical passion was.

The job of incorporating the herbarium has now been completed and a substantial number of
duplicates removed and exchanged. This was a large task, for RNG is fortunate in also having the
herbarium of the late J. E. Lousley, who had exchanged many specimens with Wallace, and a good
deal of material was duplicated. Both men had been members of the Botanical Exchange Club and
the Watson Botanical Exchange Club and had acquired specimens from these sources.
(Incidentally, it is very interesting to note that a very large number of specimens collected by
Lousley which had not been kept by Lousley himself were found in Wallace’s herbarium. )

All sheets were stamped on incorporation into RNG: ‘Herbarium E.C. Wallace (1909-1986).
Presented to Herbarium RNG 1986” so they will be identifiable in the future. (Specimens from
Lousley’s own herbarium have also been similarly annotated.)

While examining and checking each specimen some simple statistics were recorded (Table 1).
Kent & Allen (1984) record that his herbarium contained 25,000 gatherings and had significant
holdings from Surrey, v.c. 17, and Scotland. Wallace’s vascular plant herbarium in fact contained
26,190 gatherings, of which 1,458 were foreign.

The Lousley herbarium contained 3,570 (14.6%) specimens from Surrey (Jury 1977), and with
the addition of another herbarium also known to be rich in Surrey specimens a note was kept of
these. In fact of the total 22,855 British gatherings incorporated, no less than 4,692 (20.5%) were
from Surrey. Botanists working on the flora of v.c. 17 are therefore advised that they are likely to
find it worthwhile consulting RNG. A total of 5,951 gatherings (24.5%) from Scotland were
incorporated.

The foreign material is almost all from his recent travels (listed by Richards 1987), e.g. Carex
illegitima Cesati from Rodhos collected in April 1987 (see photograph in Briggs 1986).

Table 1 shows what genera were especially well represented in his collection. The comparable
figures for Lousley’s herbarium are also given. It is interesting to note how both shunned the
apomictic genera Rubus and Taraxacum, though Lousley had a fine collection of Hieracium
specimens. In many cases the two herbaria have proved to be complementary.

Only two single specimens of Agrostis have been found, both only recently collected and
unnamed by Wallace. Also missing were specimens of the genera X Agropogon, Aira, Alopecurus,
x Ammocalamagrostis, Ammophila, Anthoxanthum, Apera, Corynephorus, Cynodon, Digitaria,
Echinochloa, Eragrostis, Gastridium, Hierochloé, x Hordelymus, Lagurus, Mibora, Milium,
Nardus, Parapholis, Phalaris, Phleum, Polypogon, Setaria and Spartina. Were these loaned by
Wallace to someone? Information on their whereabouts would be gratefully received.

ACKNOWLEDGMENTS

Special thanks are due to J. R. Akeroyd, H. J. M. Bowen, S. Thornton-Wood and M. F. Watson
who have all spent many hours voluntarily working on this collection, removing duplicates, etc.
Without their help Ted Wallace’s specimens would not now be available for consultation and
research.

166 SHORT NOTES
TABLE 1. PARTIAL BREAKDOWN OF THE WALLACE VASCULAR PLANT HERBARIUM

Number of gatherings Number of

gatherings in
Se a ESTAS
Surrey Rest of the Rest of Rest of Total herbarium

v.c. 17 Scotland British Isles Europe the World number where known
Pteridophyta 144 429 681 72 58 1384
Gymnospermae 19 63 19 6 9 116
Monocotyledons
Cyperaceae
Carex 204 863 1162 205 67 2501 949
Scirpus 34 33 95 13 7 182 118
Juncaceae
Juncus 70 194 186 42 11 503 301
Luzula 31 69 65 9 5) 177 71
Liliaceae
Allium i 7 43 - 61 68
Orchidaceae
Dactylorhiza 2 11 24 37 116
Orchis 9 2 28 39 41
Poaceae
Agrostis Z 2 156
Bromus 55 22 147 19 2 245 271
Festuca ois) 69 104 16 Japp 99
Poa 34 165 88 40 + 331 219
Vulpia IS 3 48 5) 72 104
Potamogetonaceae
Potamogeton 69 121 316 3 + 513 242
Others 346 558 i127, 120 40 2190
Total monocotyledons 909 DAA, 3435 476 238 7075
Dicotyledons
Asteraceae
Centaurea 14 2 54 6 76 108
Hieracium 69 7, 76 8 1 7A 644
Senecio 32 18 100 6 156 105
Taraxacum 3 16 21 1 4] 35
Boraginaceae
Myosotis 35 32 80 3 150 96
Brassicaceae
Lepidium 18 9 47 3 ai, 120
Caryophyllaceae
Arenaria 18 15) 137 8 8 248 56
Sagina A 123 116 1 261 ey
Silene 21 65 243 10 2 317 114
Chenopodiaceae
Atriplex i 24 tae 1S 2 110 HZ
Chenopodium 42 5 123 8 178 308
Euphorbiaceae
Euphorbia 38 6 78 12 1 135 114
Fabaceae
Medicago 18 8 81 9 1 117 218
Trifolium 47 25 156 17 245 387

Vicia 34 39 122 Z 2 199 179

SHORT NOTES 167
Table 1. continued

Number of gatherings Number of
gatherings in
eae eee kl a a ORES oa los ke LLOUSICY S
Surrey Rest of the Rest of Restof Total herbarium
v.c. 17 Scotland British Isles Europe the World number where known

Fumariaceae

Fumaria 23 12 85 2 122 115
Gentianaceae

Gentianella 16 36 45 6 2 105 83
Geraniaceae

Erodium 1 D) 27 33 152

Geranium 30 24 116 1 171 132
Hypericaceae

Hypericum 49 31 77 3) 160 89
Lamiaceae

Mentha 210 73 335 5 623 176
Onagraceae

Epilobium 94 80 140 3 2 319 145
Polygonaceae

Polygonum 153 65 155 q 2 382 326

Rumex 41 50 129 8 228 1126
Ranunculaceae

Ranunculus 49 35 182 7 2 Dp 246
Rosaceae

Alchemilla 3 27 47 i 78 64

Potentilla 37 54 65 13 3 172 92

Rosa 24 1 10 35 66

Rubus 8 23 12 43 3)
Rubiaceae

Galium 52 58 104 11 4 229 138
Salicaceae

Salix 90 309 188 12 i 600 181
Scrophulariaceae

Euphrasia 30 107 119 2) 261 298

Veronica 83 101 135 11 330 208

Verbascum 10 il 26 Si) 95
Solanaceae

Solanum 3 3 24 30 105
Violaceae

Viola 156 155 422 10 2 745 204
Others 3429 2726 7365 500 101 14121
Total dicotyledons 4007 3786 9125 581 118 17617

Total in herbarium 5079 6395 13260 1135 323 26192 24465
Duplicates extracted 387 444 1046 10 1 1888

Total specimens added
to RNG 4692 5951 12212 1125 322 24304

168 SHORT NOTES
REFERENCES

Briccs, M. (1986). Ted Wallace - a memory. B.S.B.I. News, 44: 5-6.

Danpy, J. E. (1958). List of British vascular plants. London.

Jury, S. L. (1977). The herbarium of J. E. Lousley. Watsonia, 11: 312.

Jury, S. L. (1987). The herbarium of the late Ted Wallace. B.S.B.I. News, 45: 13-14.
Kent, D. H. & ALLEN, D. E. (1984). British and Irish Herbaria. London.

RIcHARDS, P. (1987). Obituary: E. C. (Ted) Wallace 1909-1986. J. Bryol., 14: 603-608.

S; JURY
Botany Department, The University, Whiteknights, Reading, RG6 2AS

TRIFOLIUM OCCIDENTALE D. E. COOMBE, NEW TO WALES

On 15 May 1987 a single, profusely-flowering patch of Trifolium occidentale was found in cliff-top
turf near Tears Point in the Gower Peninsula, Glamorgan (v.c. 41, GR 21/407.872) by Jo Dunn.
Subsequent searches by Rosemary John, E. M. Kay and Q. O. N. Kay of suitable habitats on the
coast of south-western Gower between Port-eynon Point and Rhosili showed that T. occidentale
grew both on Tears Point and on similar south-facing limestone cliff-tops between Fall Bay and
Mewslade Bay (GR 21/413.872 to 21/421.873); none was found elsewhere.

On Tears Point (the headland on the south-western side of Fall Bay near Rhosili) it is apparently
confined to one minor headland, where several (c. 12) small scattered plants of varied size grow
within an area of about 20 m? in dense, closely sheep-grazed and species-rich Festuca rubra turf on
the level cliff-top, about 3-7 m from the edge of the cliff. The sloping limestone cliffs are about
45 m high here. Two more small patches were found about 25 m west of the first site on Tears Point
(Table 1).

On the other side of Fall Bay, 7. occidentale was found to be locally more abundant between
Lewis Castle (the headland to the east of Fall Bay) and Mewslade. Here it grows in floristically
similar but less intensively grazed grassland near limestone outcrops on exposed cliff-tops, and also
on the upper cliff-slopes facing the sea. The total population in this area was estimated to be 350—
500 plants. On Lewis Castle, for example, at least 50 patches of T. occidentale were seen in about
60 m? of Festuca rubra turf (Table 1) at the seaward end of the headland, about 50 m above sea-
level. Most plants were more vigorous than those on Tears Point and many were flowering
strongly; on 7 June 1987 some flower-heads were still open but others had formed mature seed, so
had probably started flowering in late April. Here and at Tears Point scattered patches of T.
repens, not yet in flower on 7 June, grew locally intermingled with T. occidentale, mainly on the
less exposed landward side of the site.

The Gower plants closely resembled plants of T. occidentale from Cornwall and the Isles of Scilly
both morphologically and in their overall isoenzyme profiles, although the latter showed some
distinctive local features (N. A. Ab-Shukor, unpublished). Chromosome numbers of 2n=16 were
determined for three plants from Mewslade and Tears Point, in agreement with those reported
from other populations of this diploid species.

When it was first described, T. occidentale, which closely resembles the tetraploid species T.
repens and had previously been confused with it, was known in Britain only from western Cornwall
and the Isles of Scilly (Coombe 1961). Its occurrence in the Channel Islands and north-eastern
Spain suggested that it should be widespread on the western coast of France, and it was
subsequently found both there and on the northern coast of Portugal (Coombe & Morisset 1967;
Géhu 1972). Its discovery in 1979-81 on the eastern coast of Ireland as far north as Skerries in
northern Co. Dublin (Akeroyd 1983) and in 1983 in a small population at Welcombe Mouth in
Devon (Milton 1984a), about 70 km north-east of its nearest Cornish site, though only just in
Devon, suggested that it should occur further north on the coast of England and Wales. Milton
(1984b) drew attention to this in a widely-circulated paper in which he described and illustrated T.
occidentale, but the present report appears to be the first discovery since then of T. occidentale in a

SHORT NOTES 169

TABLE 1. SPECIES ASSOCIATED WITH TRIFOLIUM OCCIDENTALE AT SITES IN THE GOWER
PENINSULA (DOMIN COVER-ABUNDANCE SCORES)
Species that were recorded with T. occidentale in Cornwall and the Channel Islands by Coombe (1961) are
marked + + + if they occurred in at least three of Coombe’s four noda, + + if they occurred in two and + if they
occurred in one with a constancy of 2 or more.

Site

Tears Point E. Tears Point W. Lewis Castle
Species (21/407.872) (21/407.872) (21/413.873)

,
WW

Trifolium occidentale 3-4 4-5
Armeria maritima+ ++

Bellis perennist+ + +

Bromus hordeaceus subsp. ferronii+ ++
Cerastium diffusum+ ++

Cerastium fontanum+ +

Cynosurus cristatus+ + +

Dactylis glomeratat+ ++

Festuca ovina+ ++

Festuca rubra+ ++

Helianthemum nummularium
Hieracium pilosella

Koeleria macrantha+ + +

Leontodon taraxacoides+ ++

Lotus corniculatus+ ++

Luzula campestris++

Plantago coronopus+ + +

Plantago lanceolata+ + +

Poa pratensis+ +

Ranunculus bulbosus+ + +

Sanguisorba minor

Scilla verna+ +

Taraxacum erythrospermum

Thymus praecox+++ —
Trifolium dubium+ + 3-4
Trifolium repens+ 2
Trifolium striatum+ 2-3 —

Nea Pywernnnnt | Ponanwfne
WS Bp! On
HTae| J Rewafrvaeeanes |] ove |

ig
w

NNeENnnT Ponwn | Lefrvanoefrurnnnlt

Additional species found at only one site: Tears Point E., Centaurium erythraea+ + 1, Cirsium vulgare++ 1,
Poa annua 3—4; Tears Point W., Polygala vulgaris++ 1, Stachys officinalist++ 1; Lewis Castle, Aira
caryophyllea+++ 3, Carex caryophylleat++ 1, Galium verum++ 1, Lolium perenne+++ 3, Sherardia
arvensis 1, Trifolium scabrum++ 1-2.

locality to the north of its known range in Great Britain. The Gower sites are the first to be found
in Wales and are about 70 km north-north-east of Welcombe Mouth, separated from it by the
Bristol Channel.

The maritime Festuca rubra turf of the Gower sites differs from that of the sites in Cornwall and
Guernsey described by Coombe (1961) chiefly in the presence of a few calcicoles (e.g. Sanguisorba
minor) and the absence of some calcifuges (e.g. Jasione montana and Sedum anglicum); otherwise
they are very similar in floristic composition, with Armeria maritima as the chief halophytic species
in both cases.

REFERENCES

AKEROYD, J. R. (1983). Further notes on Trifolium occidentale D. E. Coombe in Ireland. /r. Nat. J., 21: 32-34.

CoomsE, D. E. (1961). Trifolium occidentale, a new species related to T. repens. Watsonia, 5: 68-87.

CoomsE, D. E. & Morisset, P. (1967). Further observations on Trifolium occidentale. Watsonia, 6: 271-275.

GEuHu, J. M. (1972). Trifolium occidentale D. E. Coombe: éspéce nouvelle pour le littoral du nord du Portugal.
Agron. Lusit., 34: 197-204.

170 SHORT NOTES
Mitton, J. N. B. (1984a). Trifolium occidentale D. E. Coombe (Atlantic Clover) new to North Devon.

Watsonia, 15: 120-121.
Mitton, J. N. B. (1984b). Trifolium occidentale D. E. Coombe, an overlooked clover. B.S.B.I. News, 37: 8-9.

Q. O. N. Kay & N. A. AB-SHUKOR
School of Biological Sciences, University College, Swansea, SA2 8PP

REDISCOVERY OF HALIMIONE PEDUNCULATA (L.) AELLEN IN BRITAIN

In northern Europe Halimione pedunculata (L.) Aellen is a rare annual of saltmarshes and tidally
inundated dune-slacks. It appears to have undergone considerable decline during the present
century, and is now listed as an ‘endangered’ species in Europe (Council of Europe 1983). H.
pedunculata was believed to have become extinct in Britain in the 1930s. There are old records of
the plant from at least 16 localities on the eastern coast between Kent and Lincolnshire (Perring &
Walters 1962; Perring & Farrell 1983), but this century it has been observed at only three of these:
at Pegwell Bay, Kent (many records, last seen about 1924); on the Wash at Freiston, Lincolnshire
(1932, specimen at NOT); and at Walberswick, Suffolk (last seen 1938 (J. E. Lousley, note at
RNG)).

On 29th September 1987 a colony of H. pedunculata was discovered in S. Essex, v.c. 18, GR 51/
9.8, by S. J. L. and S. A. Wolfe-Murphy in the course of a Nature Conservancy Council survey
of grazing marshes in Suffolk and Essex. The species occurs here on a narrow strip of relict
saltmarsh about 100m back from the sea-wall, where it is associated with Puccinellia maritima,
Aster tripolium, Spergularia media, Suaeda maritima and Triglochin maritima (Table 1). In
phytosociological terms this vegetation clearly belongs to the Puccinellietum maritimae, and is
similar to stands containing H. pedunculata described from elsewhere in Europe (Géhu & Meslin
1968; Géhu 1969; Westhoff & Den Held 1969). The presence of Parapholis strigosa is of particular
interest, since on the Continent this species helps to typify the subassociation of the Puccinellietum
in which H. pedunculata characteristically occurs (Beeftink 1977).

TABLE 1. PLANT SPECIES GROWING IN ASSOCIATION WITH HALIMIONE PEDUNCULATA
Cover-abundance is listed on the Domin scale.

Quadrat No. (1x1 m)

Species 1 2 3 4 ) 6
Puccinellia maritima 8 8 8 7 7 7]
Aster tripolium 7. 6 7 q 7 7
Spergularia media 3 4 - 3 + 3
Halimione pedunculata 5 5 2 3 3 4
Suaeda maritima 3 1 3 2 1 3
Salicornia ?ramosissima 1

Triglochin maritima 1 5 6 3
Parapholis strigosa 5 2 3
Atriplex prostrata 1

Juncus gerardii 4
Bare ground (%) 5 it 5 2 5
Vegetation height (cm) 12 8 15 13 9 12

On 2nd October J. P. Doody, L. Farrell and S. J. L. revisited the site to carry out a population
count of H. pedunculata and to map its distribution. A total of 1,714 plants were found within an
area of 95m*. They were distributed very unevenly, with large numbers in areas of more open
vegetation and around the edges of slight depressions where seed had collected, possibly as a result
of flooding. Most of the plants were in full fruit, with pedicel lengths frequently exceeding the ‘up
to 12mm’ given in the Floras (e.g. Tutin 1987): a sample of 33 pedicels had an average length of

SHORT NOTES 171

13mm, and a maximum of 25mm. Plant height was also very variable, with a sample of 41 fruiting
plants having an average height of 14.2 cm and a range of 1.7-32.5 cm.

It is tempting to speculate on the possible origins of H. pedunculata at this site. Has it always
been here or is it a recent arrival? The plant’s centre of distribution in Europe is on the Danish
coast (Jalas & Suominen 1980) and this also happens to be the major ‘refuelling’ area for Brent
Geese on their autumn migration to south-eastern England (Owen et al. 1986). Huge flocks of
these birds pass through Denmark in late September — just as the seeds of H. pedunculata are
ripening — and arrive on the Essex coast in early October. Is it possible that H. pedunculata could
have made the journey across the North Sea in the belly of a Brent Goose?

On the other hand, if it is indeed a site of long standing, how is it that the plant has remained
undetected for so long? H. pedunculata is a notoriously unreliable species in that its numbers can
vary dramatically from year to year (Bennett 1905; Géhu 1969), and this is bound to make finding
the plant more a matter of luck than judgement. In addition, it only becomes visually obvious in
late September or early October after the pedicels have lengthened. Most botanists are in the field
earlier in the year when H. pedunculata could easily be overlooked.

This is the first record of H. pedunculata in Britain for 50 years, and the first ever record for
Essex. Yet the vegetation in which it occurs is otherwise quite unremarkable, and there are
certainly large areas of suitable looking habitat in other parts of Essex, and in Kent, Norfolk and
Suffolk. It is likely that — if searched for late enough in the season — H. pedunculata could be found
elsewhere in south-eastern England.

ACKNOWLEDGMENTS

I thank the owners for permission to visit the site, and S. A. Wolfe-Murphy, J. P. Doody and L.
Farrell for assistance with the field survey. I am grateful to J. D. Pratt, C. D. Preston, A. Smith
and N. F. Stewart for help with the Continental literature. I also wish to thank C. D. Preston for
supplying details of old records, and for useful discussion during the preparation of this note.

REFERENCES

BEEFTINK, W. G. (1977). The coastal salt marshes of western and northern Europe, in CHAPMAN, V. J., ed.
Ecosystems of the World, Volume 1: Wet coastal ecosystems. Amsterdam.

BENNETT, A. (1905). Distribution of Sonchus palustris L. and Atriplex pedunculata L. in England. Trans.
Norfolk Norwich Nat. Soc., 8: 35-43.

COUNCIL OF Europe (1983). List of rare, threatened and endemic plants in Europe. Nature and Environment
series, No. 27. Strasbourg.

GéEuHu, J. M. (1969). La persistance de /’Obione pedunculata en baie du Mont-Saint-Michel et en quelques
points du littoral du nord de la France. Sa signification biologique. Le Monde des Plantes, No. 359: 1+.

Géuu, J. M. & MEsLINn, R. (1968). Sur la répartition et l’écologie d’Halimione pedunculata (L.) Aell.
(Dicotyledoneae, Chenopodiaceae) en France. Bull. Lab. Marit. Dinard, 1 (1): 116-136.

JaLas, J. & SUOMINEN, J. (1980). Atlas florae Europaeae. Distribution of vascular plants in Europe. Volume 5:
Chenopodiaceae to Basellaceae. Helsinki.

Owen, M., ATKINSON-WILLES, G. L. & SALMON, D. G. (1986). Wildfowl in Great Britain, 2nd ed. Cambridge.

PERRING, F. H. & FarreELL, L. (1983). British Red Data books: 1. Vascular plants, 2nd ed. Lincoln.

PERRING, F. H. & WALTERS, S. M., eds (1962). Atlas of the British flora. London.

TutTIn, T. G. (1987). Halimione Aellen, in CLAPHAM, A. R., TuTIn, T. G. & Moore, D. M. Flora of the British
Isles, 3rd ed., p. 160. Cambridge.

WESTHOFF, V. & DEN HELD, A. J. (1969). Planten Gemeenschappen in Nederland. Zutphen.

S. J. LEACH
Nature Conservancy Council, Northminster House, Peterborough, Cambs., PEI 1 UA

172 SHORT NOTES

ON CONYZA SUMATRENSIS (RETZ.) E. WALKER AND CERTAIN HYBRIDS IN THE
GENUS

The first object of this note is to establish that the correct name for a Conyza now well naturalized
in the Channel Islands (Marshall & McClintock 1972; McClintock 1975) and now in and near
London (Badmin 1986; Wurzell 1988) is C. sumatrensis (Retz.) E. Walker (Marshall 1974).
This epithet was first published by Retzius in 1789 for such a plant in Sumatra — which may seem
unexpected for one said to be native of Peru. It is, however, now at least, a common weed in
Malaysia (Kalkmann & Van Steenis, in litt., 12 Sept. 1976) and may well have started spreading
across the globe before the late 18th century. There are objections, however, to the use of this
epithet:
a) Burtt (1948) considered Retzius’ description did not give sufficient detail to enable certain
identification to be made. But his alternative, C. bonariensis (L.) Cronq., is a separate species
which has never been known in that part of the world (Kalkman & Van Steenis, in litt., 12 Sept.
1976).
b) Retzius described the stems as red, whereas in northern Europe they are usually green. Guédés
(1984) dismissed the epithet sumatrensis solely because of this, even though in other places,
including Sumatra, plants with red stems also occur — the latter presumably due to the Sumatran
sun.
c) There is no type or other specimen of Retzius’ to check the application of the name (Guédés, in
litt., 17 Jan. 1985). This is of course true of many published names and we here designate a
neotype:

Conyza sumatrensis (Retz.) E. Walker in J. Jap. Bot., 46: 72 (1971).
TYPE: Sumatra, Berastagi, February 1921, H. N. Ridley (Neotype: K, designated here).

Later epithets which have been used for this species include the following:

a) C. floribunda Kunth. This is rejected by Guédés & Jovet (1975) on the ground that it should be
considered a distinct species, having smaller, glabrescent, finally chestnut-brown capitula and
glabrescent or glabrous leaf-surfaces with setiferous margins. Marshall (1973) called this taxon C.
floribunda var. subleiotheca (Cuatr.) J. B. Marshall, using much the same characters, but he now
considers it within the range of C. sumatrensis which, with its wide distribution, not surprisingly, is
variable.

b) C. albida Willd. ex Sprengel. This was favoured by Guédés & Jovet (1975) and by Jovet &
Vilmorin (1975), but in neither of these publications is sumatrensis considered.

c) C. erigeroides DC. (see Guédés & Jovet 1975).

Other synonyms are listed by Marshall (1973, 1974).

HYBRIDS

The boundaries of Conyza species are not always clear, the plants becoming modified according to
the conditions in which they grow. McClintock (1985) mentioned the opinion of Cronquist (1976)
to whom C. bonariensis and C. floribunda (sumatrensis) are conspecific. Furthermore, hybrids
have been claimed to occur in south-western France and north-eastern Spain, notably under names
published (but not all validly so) by Sennen (1904, 1908, 1912, 1916, 1929). We have studied
numerous specimens from BM, BOG, BRI, E, GH, K, LP, RB and US with such names and most of
them seem to us to fit under C. sumatrensis (see synonymy in Marshall (1974)). Two taxa do,
however, appear to be distinguishable:

a) C. X daveauiana Sennen (said to be C. naudinii Bonnet (sumatrensis) X ambigua DC.
(bonariensis)); all the samples come from the neighbourhood of Barcelona, but it is not clear how
his later C. xX barcinonense (with the same postulated parentage) differs from it. All eight
specimens we have seen look distinct; indeed Sennen (1912) himself wrote “‘peut-étre la plante
n’est pas hybride”’. We do not know how fertile it is, nor indeed if any of the claimed hybrids are. _
None have been made experimentally.

b) Another alleged hybrid of note is C. X mixta Foucaud & Neyraut (= C. ambigua (bonariensis) X
canadensis (L.) Cronq.); we know of no specimen so named but the drawing the authors gave of it
from near Bordeaux (Foucaud & Neyraut 1902) suggests the postulated parentage may be correct.

SHORT NOTES 173

CONCLUSION

We have found that we can place any plants we have seen from the British Isles, and most of those
from the Continent, under one of three species: C. canadensis, C. bonariensis or C. sumatrensis
(cf. Hansen 1972). Nevertheless, members may still like to look out for possible hybrids.

REFERENCES

BapmMIN, J. (1986). Annual exhibition. Kent Fld Club Bull., 31: 18.

Burtt, B. L. (1948). On Erigeron bonariensis L. Kew Bull., 3: 369-372.

CronauisT, A. (1976). Conyza Less., in TuTin, T. G. et al., eds. Flora Europaea, 4: 120. Cambridge.

Foucaup, J. & NeyrauT, E. J. (1902). Une hybride nouveau. Ann. Soc. Sci. Char. Inf., 7: 109-111.

Guépés, M. (1973). Conyza altissima (C. naudinii) et C. X rouyana a Tours. Le Monde des Plantes, 378:
4

Guépés, M. (1984). Conyza bonariensis (L.) Cronq. and Conyza albida Willd. ex Sprengel. B.S.B.I. News, 38:
19.

GuépEs, M. & Jovet, P. (1975). Conyza albida Willd. ex Sprengel, the correct name for “‘C. altissima’’ Naudin
ex Debeaux (C. naudinii Bonnet). Taxon, 24: 393-394.

Hansen, A. (1972). Contributions to the flora of the Canary Islands. Cuad. Bot. Canar., 14/15: 59-60.

JoveT, P. & VILMORIN, R. DE (1975). Conyza, in Coste, H. Flore descriptive et illustrée de la France, 3-eme
supplement, pp. 187-192, 308-311. Paris.

MarsHAaLt, J. B. (1973). Conyza — taxa found in Britain. Watsonia, 9: 372-373.

MarsHALL, J. B. (1974). A note on Conyza sumatrensis (Retz.) E. Walker (C. floribunda Kunth). Watsonia,
10: 166-167.

MarsHALL, J. B. & McCuintock, D. (1972). Conyza in Britain. Watsonia, 9: 201-202.

McCuintock, D. (1975). The wild flowers of Guernsey, pp. 203-204. London.

McCuintock, D. (1985). Conyza. B.S.B.I. News, 40: 11.

SENNEN, E. C. (1904). Note sur le Conyza naudinii Bonnet et sur quelques hybrides. Bull. Soc. bot. Fr., 51:
426.

SENNEN, E. C. (1908). Plantes d’Espagne, années 1906 & 1907. Bull. Acad. Int. Geog. Bot., 18: 469-470.

SENNEN, E. C. (1912). Quelques formes nouvelles ou peu connues. Bol. Soc. Aragon., 11: 194.

SENNEN, E. C. (1916). Liste des plantes . . . adventices . . . de Barcelone. Bol. Soc. Aragon., 15: 97-98.

SENNEN, E. C. (1929). Sobre dos plantes nuevas en la flora Espafiola. Bol. Soc. [bér., 28: 66-67.

WURZELL, B. (1988). Conyza sumatrensis (Retz.) E. Walker established in England. Watsonia, 17: 145-148.

D. McC.Lintock & J. B. MARSHALL
Bracken Hill, Platt, Kent, TN15 8JH

A NEW BRAMBLE FROM SKYE AND THE OUTER HEBRIDES
Rubus ebudensis A. Newton, sp. nov.

R. grato similis, sed proprie differt:— Turiones acutiores, profundius sulcati; aculei validiores,
curvati e basi lata (vice recti declinati e base angustata); foliola terminalia elliptica vel elliptico-
rotundata, apice paulatim acuminata, petioli aculeolis validis curvatis armati; inflorescentia saepe
uno vel nonnullis foliis simplicibus + rotundatis supra ornata; rachis aculeis validis curvatis
declinatis armata.

Flores erubescentes 2.5 cm diam. vice rosei 3.5 cm; petala corrugata antherae epilosae. Habitat
praecipue in insulis Skyense et Ebudibus extremis.

Similar to R. gratus particularly in panicle structure, but differing as follows:— Stems more sharply
angled and more deeply grooved; stem prickles stouter, curved rather than straight, declining from
broad rather than narrow bases; terminal leaflets elliptic or elliptic-rotund, gradually acuminate
rather than obovate; petioles with several strongly curved prickles.

Panicle often with one or more roundish simple leaves above the 3-nate leaves, the rachis armed
with several stout-based curved declining prickles. Flowers pinkish and 2.5 cm diam. rather than
pink and 3.5 cm, petals crumpled, anthers glabrous. Found in Skye and the Outer Hebrides; not
noted as yet on the mainland.

174 SHORT NOTES

HOLOTYPE: Side of B884 near Colbost above Loch Dunvegan, Skye, v.c. 104, GR 18/212.492,
18/8/1966, B. A. Miles 66/250 (Herb. A. Newton).

Specimens of this species have previously been named as R. gratus or given the manuscript name of

‘false gratus’. I have also seen the following material:

S. Uist, N. Glendale, GR 08/791.177, 8/1980, R. Pankhurst & A. Chater 377 (BM).

Benbecula, 8/1940, J. W. Heslop-Harrison (BM).

S. Harris, Leverburgh, GR 18/020.855, 8/1980, R. Pankhurst & A. Chater 351 (BM); Ardvey, GR
18/125.928, 8/1980, R. Pankhurst & A. Chater 347 (BM); Obbe, 7/1894, W. A. Schoolbred (as
R. gratus, det. W. M. Rogers) (BM).

Lewis, Eishken, GR 19/327.120, 8/1980, R. Pankhurst & A. Chater 354 (BM).

Skye, near Loch Treaslane, GR 18/3.5, 8/1966, B. A. Miles (CGE).

A. NEWTON.
10 The Fairways, Leamington Spa, Warwickshire, CV32 6PR

RORIPPA X HUNGARICA BORBAS (R. AMPHIBIA X R. AUSTRIACA) (CRUCIFERAE)
NEW TO THE BRITISH ISLES

In June 1987, while investigating Rorippa populations along the River Roding, Chigwell, Essex
(v.c. 18), we found a single plant intermediate between R. amphibia (L.) Besser and R. austriaca
(Crantz) Besser. Further morphological analysis indicated that it was probably a hybrid between
the two species and material was therefore sent to Professor B. Jonsell, who confirmed the
identification (pers. comm. 1987). This hybrid has been reported rarely from Europe
(Javirkova-Kratochvilova & Tomiovic 1972; Jonsell 1975) and this is the first record from the
British Isles. Voucher specimens have been placed in BM, LTR, herb. T.C.G.R. and herb. B. W.

We have accepted the name R. X hungarica as the binomial for R. amphibia xX R.
austriaca (Borbas 1879) but have not seen the material from which it was described.

TABLE 1. CHARACTERS OF RORIPPA AMPHIBIA, R. AUSTRIACA AND R. X HUNGARICA
BASED ON MATERIAL FROM CHIGWELL
Measurements are ranges made on fresh material

Character R. austriaca

habit

stem T.S. below
inflorescence

middle stem leaves
auricles

teeth

sepal length (mm)
petal length (mm)
shape at base

pedicels
immature ovaries
stipe (mm)
shape
length (mm)
width (mm)
fruit set

R. amphibia
clumps with ascending
stems

hollow with thin,
supple, dark green cells

small, but clasping stem

deep, outwards-
pointing
(3.3) 3.44.3

(4.3) 4.8-6.2
clawed

spreading to deflexed

0.40.5
oblong
2.0—3.0
0-8-1.2

setting good fruit

R. X hungarica

+ clump-forming

loosely packed with
thick, firm whitish-
green cells

obvious and nearly
projecting
medium, generally
forward-pointing
3.2-3.6

4.8-5.7

+ unclawed

spreading to deflexed

¢.10:3

elliptic
1.3-1.5
0.9-1.2

not maturing, racemes
elongating markedly

patch-forming with
erect stems

solid with dense, hard
foamy white cells

prominent, projecting
well behind stem

shallow, forward-
pointing
2.7-3.0

3.7-5.0
unclawed

ascending, angle c. 40°

sessile

+ round
1.2-1.5
1.1-1.4

setting good fruit

SHORT NOTES 175

a b

C nye
Figure 1. Leaves and immature ovaries of (a) R. amphibia, (b) R. X hungarica and (c) R. austriaca.

The plant was found growing amongst tall vegetation on the steep bank of the river within 20 m
of both parents. Rorippa amphibia was frequent along the river bank and one patch of R. austriaca
was found in dry grassland; this appears to be a new site for the latter although there are three
other colonies known nearby in the Roding Valley. Other Rorippa taxa present along the river
here are R. palustris, R. sylvestris and R. amphibia X sylvestris (the latter also confirmed by
Jonsell).

Characters of R. X hungarica and its parents from Chigwell are given in Table 1.
Drawings of leaves and immature ovaries are given in Fig. 1, and a plot of fresh sepal length against
fresh petal length for individual flowers of all the taxa is given in Fig. 2. These illustrate the general
intermediate nature of the hybrid, though any one character may vary towards either parent. In
addition, Professor Jonsell examined pollen under a microscope and states (pers. comm.) “the R.

65
e
e
6:0 @
@
x e @
— x
= 55 xx x e e
E x @x
x, x @
= x © @
= se
oO
SG 5-0
il}
x e
zal) o
. 45 ;
oO o
oe 40 o
Oo
40 o0
7 oO
oO

255 3:0 35 40 45

SEPAL LENGTH(mm)

FiGure 2. Fresh sepal length plotted against fresh petal length for individual flowers of (@) R. amphibia, (Q) R.
austriaca and (X) R. X hungarica, all from Chigwell.

176 SHORT NOTES

amphibia X austriaca hybrid seems to produce only inviable pollen . . . in the parent species pollen
quality is, as is very often the case, far from excellent, but there are at least a number of well
developed presumably functioning grains’.

ACKNOWLEDGMENTS

Our thanks to Ken Adams who suggested we should investigate this section of river in the first
place, to Professor B. Jonsell for examining the material, and to Mike Mullin for help in checking
nomenclature.

REFERENCES

Borsis, V. (1879). Floristicai adatok kiilénds tekintellel a Roripakra. Ertek. természettudom. Kérébél, 9(15):

JAVURKOVA-KRATOCHViLovA, A. & TomSovic, P. (1972). Chromosome study of the genus Rorippa Scop. em.
Reichenb. in Czechoslovakia. Preslia, 44: 140-156.

JONSELL, B. (1975). Hybridization in yellow-flowered European Rorippa species, in WALTERS, S. M.& KING,
C. J., eds. European floristic and taxonomic studies. Faringdon.

T. C. G. Rich & B. WuRZELL
Biological Records Centre, Monks Wood Experimental Station, Abbots Ripton, Huntingdon,
Cambs., PE17 2LS

CEPHALANTHERA RUBRA (L.) RICH. IN HAMPSHIRE

On 20th July 1986 the flower-spike of a Cephalanthera was found in bud, in a beech-yew wood on
the chalk in N. Hants., v.c. 12. When the first flower opened a few days later, it was clear that this
was a plant of Cephalanthera rubra (L.) Rich. of which no satisfactory records exist for Hampshire.
Druce (1927) reported that it was “recorded from near Selborne, Hants., but in error, the
specimen being a Helleborine (i.e. Epipactis)’’. It is not mentioned in the Flora of Hampshire
(Townsend 1904), but in a supplement (Rayner 1929, p. 99) C. rubra is recorded as having been
found in 1926 in District VI(1) (Upper Test Catchment) on the authority of Miss E. Williams.
Nothing further is known of this report. We have received oral information that Anne Pratt, the
Victorian botanist, reported C. rubra from the Hampshire Chalk Hangers in the 1880s, but we
cannot trace this record in print.

The present-day locality for C. rubra in Hampshire is under rather deep shade, on a north-west-
facing slope. The single plant that flowered in 1986 was a fine one, with a total of 13 flower-buds,
but some of these aborted. Several, however, opened widely enough to produce flowers in which
the narrow labellum and pubescent ovary were well displayed. Careful search of the site revealed
that at least nine leafy, but non-flowering, shoots of this species were present over some 50m of the
bank. Botanists have visited this area over some years previously, in general botanical surveys, but
C. rubra has not been reported before. It is however known that a few trees were felled higher up
the bank a few years ago; this may well have allowed enough (almost horizontal) morning sunlight
to penetrate the site and to stimulate one plant to produce a flower-spike.

In early August this flower-spike unfortunately was found to have been broken off. Efforts at
pollinating the flowers having met with no success, the spike was preserved, using the sulphur
dioxide method, by F. R.

The site has little ground flora apart from sparse ivy; conservation work has been undertaken,
with the cooperation of the owners, to thin out the tree canopy, particularly the yews and smaller
scrubby beeches, to let in more light. In its localities in France and Germany, C. rubra only flowers
well in situations where it gets some hours of sunlight on each sunny day, e.g. in glades or on the

SHORT NOTES 1a

northern sides of minor roads or trackways. Similar management has been carried out in other
British sites, with considerable success in stimulating several plants to flower. It is known that in
some continental sites there are large numbers of rhizomes present that do not even produce leafy
shoots, but in later years have inflorescences if light is let in.

REFERENCES

Druce, G. C. (1927). New county and other records. Rep. botl Soc. Exch. Club Br. Isl., 8: 134.
Rayner, J. F. (1929). A supplement to Townsend’s Flora of Hampshire, p. 99. Privately printed.
TOWNSEND, F. (1904). Flora of Hampshire, 2nd ed. London.

F. Ros—E & A. BREwIS
Benhams House, Benhams Lane, Blackmoore, Liss, Hants., GU33 6BE

THE IRISH VICE-COUNTY MAP IN WATSONIA: A CORRECTION

I wish to draw attention to the map of vice-counties of Ireland which has been published in most
issues Of Watsonia from 1976 onwards. In this map the boundary line limiting West Cork, v.c. H3,
to the east and Mid Cork, v.c. H4, to the west is incorrect and not in accord with the reference
vice-county map for Ireland published by Praeger (1901). This map is reproduced in both editions
of Census Catalogue of the flora of Ireland (Scannell & Synnott 1972, 1987).

Webb (1980) set out very clearly the historical background to the vice-counties. In this paper a
number of ambiguities and errors are discussed and attention is drawn to the changes in the
administrative county boundaries in 1898-1900, and the effects of those on the biological scheme.
Webb reaffirms the principle adopted by the Praeger Committee of the Royal Irish Academy — that
the biological vice-counties should be defined in accordance with the frontispiece map of Irish
Topographical Botany (Praeger 1901). Webb delineates in words and by line-drawing the
boundaries of the vice-counties and in particular those lines drawn to subdivide the larger counties.
In the case of West Cork, v.c. H3, and Mid Cork, v.c. H4, the dividing line runs from the median
part of the Bandon estuary, along the Bandon River to the bridge at Bandon, from thence by a
straight line to the centre of Macroom, from there by a straight line to Millstreet and then along the
Mallow-Killarney railway line to the Kerry border. This is not the line presented in the map
published in Watsonia.

Where then did the H3/H4 line in the Watsonia map come from? In 1949 the Ordnance Survey of
Ireland published a large-scale map (10 miles to one inch) setting out the biological divisions.
Colonel Niall MacNeill, an authority on dragonflies, then in charge of Ordnance Survey,
supervised the work. In this map the H3/H4 line is based on barony boundaries and appears as an
undulating line from the sea to the Kerry border. This is the line presented in the Watsonia map.
Webb (1980), however, has discussed this map and has given reasons for its rejection; he also
points out that MacNeill’s boundary between West Cork and Mid Cork deviates from Praeger’s by
as much as 8 km in two places. The boundary line is also incorrect in the overlay issued with Atlas
of ferns of the British Isles (Jermy et al. 1978).

The map in Druce (1932) is correct as regards the boundary line and the letter-press on the back
of Druce’s map describes the geographic points on the line. Dandy’s work on the vice-counties
(Dandy 1969) refers to Great Britain only.

It is recommended that the correct map be published and that attention be drawn to Webb
(1980) on the Irish vice-counties.

ACKNOWLEDGMENT

I wish to thank Professor David Webb for his comments on an earlier draft of this note.

178 SHORT NOTES
REFERENCES

Danpy, J. E. (1969). Watsonian vice-counties of Great Britain. London.

Druce, G. C. (1932). The comital flora of the British Isles. Arbroath.

JermMy, A. C., ARNOLD, H. R., FARRELL, L. & PERRING, F. H. (1978). Adlas of ferns of the British Isles. London.
PRAEGER, R. L. (1901). Irish topographical botany. Proc. R. Ir. Acad., 3rd series, 23: 61-94.

SCANNELL, M. J. P. & Synnotr, D. M. (1972). Census catalogue of the flora of Ireland. Dublin. (2nd ed., 1987).
Wess, D. A. (1980). The biological vice-counties of Ireland. Proc. R. Ir. Acad., 80 B(12): 179-196.

M. J. P. SCANNELL
Herbarium, National Botanic Gardens, Glasnevin, Dublin 9

POLYGONUM HYDROPIPER L. VAR. DENSIFLORUM A. BRAUN

While walking along the edge of the Ouse Washes near Welches Dam, Manea, Cambridgeshire,
v.c. 29, on 13th November 1986, P.D.S. saw in some quantity two completely different looking
Polygonums growing with Urtica dioica subsp. glabrescens, Bidens tripartita, Atriplex prostrata,
Erysimum cheiranthoides , Senecio aquaticus and Rumex spp. In some places they formed separate
patches, in others they were close together, but not intermingled. There were no intermediates.

One of them (Sell 86/229, colour slide 423/16 in CGE) had narrow, wavy leaves and very lax
inflorescences, and was recognizable at a glance as Polygonum hydropiper L.; the peppery taste of
the leaves confirmed the identification. The other plant (Sell 86/228, colour slide 423/15 in CGE)
had broader, non-wavy leaves and much denser inflorescences, many of which came from the axils
of the leaves. The plant was on the whole much larger, taller and more branched and straggly. On
first tasting this plant P.D.S. got no reaction, but later did obtain some reaction, as did others who
tried it. An examination of the plant in detail indicated that it must be included in P. hydropiper,
but it is a distinct variant of that species.

Consultation of the literature showed that C. E. Britton (1926, 1933) had recorded such a plant
from Brox in Surrey, v.c. 17, under the name P. hydropiper var. densiflorum A. Braun, Flora
(Regensb.), 7: 352 (1824), and that this determination was upheld by Salmon (1931). Grose (1957)
recorded it from a cultivated field near Dewey’s Water and a damp field near Garsdon, both in
Wiltshire. Timson (1966), in his Biological Flora account, included only these records and gave no
indication that he had seen the plant himself. This variant is also mentioned in Lousley & Kent
(1981).

During the autumn of 1986, J.R.A. independently received material of Polygonum for
determination, from Mrs M. E. R. Martin (v.c. 72) and L. J. Margetts (v.c. 3), that appeared to be
P. hydropiper var. densiflorum. Mrs Martin had annotated the specimen: “‘More elegant plant.
Brown glands not so numerous. Biting taste less obvious. Not drooping tips.’’ Mr Margetts noted
(in litt. to J.R.A.) that “the inflorescences were not drooping.’’ P.D.S. later confirmed this
identification.

In the Cambridge University herbarium (CGE) there is a specimen collected by the Rev. E. S.
Marshall, no. 2487 from the head of Loch Long, Dunbartcn in 1900, which was associated with
normal P. hydropiper and P. persicaria L. It is annotated: “I do not think that this is a hybrid, but I
believe it may be P. hydropiper L. var. densiflorum Braun in Bot. Zeit. 1824; but I have not access
to his paper, but take it from Bosch’s Flora Batavica. Ar. Bennett in litt. 31/12/1900.” E. F. Linton
has annotated the specimen: “‘Too strongly fertile in my opinion for a hybrid Polygonum. Reminds
me of forms of Hydropiper which simulate P. mite. I do not know the varietal name. I have one
nearly as dense-flowered (from Nayland, Suffolk), but never a specimen with such broad-round-
basal leaves! I consider I have P. mite X hydropiper from Sway, not entirely barren, but fruits
rather intermediate. The fruit and perianth of your plant are out and out Hydropiper, not a trace of
P. persicaria.”’ A duplicate of Marshall’s gathering (BM) bears the annotation “Taste insipid’,
which fits the observations of P.D.S. and Mrs Martin noted above. We consider both Marshall’s
Dunbarton plant and Linton’s Suffolk plant to be var. densiflorum.

SHORT NOTES 179

Although more difficult to recognize when pressed, a number of specimens in CGE and
elsewhere seem to be referable to P. hydropiper var. densiflorum. These are listed below. The
Nayland plant is presumably that to which Linton referred. P.D.S. did not comment on the
Flitwick plant, which is typical of the variety, when he collected it, other than that it had an acrid
taste: it may be this variety that Saunders (1883) called P. minus Hudson and which Dony (1953)
later corrected to P. hydropiper. Both collections were from a ditch (stream) at Flitwick Moor.
v.c.3. §S. Devon. Starcross, side of small stream, 22.9.1900, S. H. Bickham, CGE; near Exeter,

Columbjohn, by River Culm (GR 20/95.99), 30.9.1986, L. J. Margetts, RNG.

v.c.13. W. Sussex. Near Lyminster, farmyard, 23.9.1928, E. C. Wallace, RNG.

v.c.17. Surrey. Brox, 2.10.1926, C. E. Britton 2988, BM, RNG; between Kew and Richmond, by
River Thames, 10.9.1932, J. E. Lousley, RNG; Frensham Great Pond, dried bed of lake,
12.9.1941, E. C. Wallace, RNG.

v.c.23. Oxon. Oxford, Port Meadow, 1891, J. Baker, OXF; Lechlade, 9.1894, G. C. Druce, OXF.

v.c.25. E. Suffolk. Stoke-by-Nayland, lower park gate, roadside, 9.1889, J. D. Gray, CGE;
Nayland, ibid., 12.9.1889, E. F. Linton, BM.

v.c.27. E. Norfolk. Near Cromer, Felbrigg Woods, 8.1896, A. Wallis, CGE.

v.c.29. Cambs. Manea, near Welches Dam, edge of Ouse Washes, 13.11.1986, (GR 52/472.860)
P. D. Sell 86/228, CGE.

v.c.30. Beds. Flitwick Moor, side of stream, 12.9.1962, P. D. Sell 62/805, CGE.

v.c.37. Worcs. Bransford, ditch, 1.10.1910, R. F. Towndrow, CGE.

v.c.41. Glam. Near Cardiff, Llanedyrne Road, ditches, 5.10.1935, A. E. Wade, CGE.

v.c.71. Man. Rushen, marsh by Croak Mooar, 16.9.1952, D. E. Allen, CGE.

v.c.7/2. Dumfries. Near Dumfries, riverside sandy gravel (GR 25/97.76),13.9.1986, M. E. R.
Martin, RNG.

v.c./9. Dunbarton. Head of Loch Long, Arrochar, 28.8.1900, E. S. Marshall 2487, BM, CGE.

v.c.83. Midlothian, Crookster, 1849, J. B. Syme, BM.

REFERENCES

Britron, C. E. (1926). Polygonum hydropiper L. J. Bot., Lond., 64: 328.

Britton, C. E. (1933). British Polygona, section Persicaria. J. Bot., Lond., 71: 90-98.

Dony, J. G. (1953). Flora of Bedfordshire. Luton.

Grose, D. (1957). The flora of Wiltshire. Devizes.

LousLey, J. E. & Kent, D. H. (1981). Docks and knotweeds of the British Isles. London.

SALMON, C. E. (1931). Flora of Surrey. London.

SAUNDERS, J. (1883). On the flora of South Bedfordshire. J. Bot., Lond., 21: 310-312.

Timson, J. (1966). Biological Flora of the British Isles, 103. Polygonum hydropiper L. J. Ecol. , 54: 815-821.

P. D. SELL & J. R. AKEROYD
Botany School, Downing Street, Cambridge, CB2 3EA

LECTOTYPIFICATION OF FIVE NAMES IN RANUNCULUS L. SUBGENUS
BATRACHIUM (DC.) A. GRAY

This group is bedevilled with names difficult to apply, many of which need to be lectotypified, and
progress is necessarily piecemeal. Five of these names are dealt with below.

R. hederaceus L., Sp. pl. 556 (1753)

The diagnosis given by Linnaeus is, in fact, taken verbatim from the Hortus Cliffortianus of 1738,
and the type should, if possible, be selected from material available to him at that time. There is
unfortunately no specimen in the Hortus Cliffortianus in BM. It is therefore necessary to consider
the synonyms and any specimens in his herbarium (LINN) up to 1753. Benson (1954) designated

180 SHORT NOTES

the sheet of the Savage Catalogue 715/74 (LINN) as the lectotype. Cook (1966) says he has seen
this specimen and has no reason to doubt Benson’s designation. P.D.S. has labelled the specimen
as the lectotype.

R. hederaceus forma natans Moss, Camb. Br. Fl., 3: 139, pl. 144 (1920)

No type was designated, but judging from the distribution given, there were obviously a great
many syntypes. The original drawing made by E. W. Hunnybun for pl. 144 is in the Botany School,
University of Cambridge. It is labelled ‘‘Hemingford Common [Huntingdonshire], May, 1900”,
but there is no specimen to go with it in CGE. P.D.S. has therefore selected as lectotype a
specimen in CGE which is cited with the original description. It is labelled ‘‘Rheinflache bei Kruft,
15 Aug. 1861, Dr Wirtgen Herb. Pl. Crit. Fasc. 10, no. 546”, on which Moss has written, “‘R.
hederaceus L., floating form. Not R. homiophyllus Ten.’’ Cook (1966) considers that forma natans
has no genotypic basis.

R. hederaceus var. grandiflorus Babington, Man. Br. Bot. 5 (1843)

There are two syntype gatherings in Babington’s herbarium in CGE made before 1843 and labelled

by him as var. grandiflorus.

1) Floating on the water in a small pit near the Monastery, Charnwood Forest, Leicestershire, 20
June 1837, C. C. Babington.

2) Needwood Forest, Staffordshire, 15 June 1837, C. C. Babington.

Both are R. omiophyllus Ten. P.D.S. has selected the first as the lectotype. The second becomes a

paralectotype.

R. floribundus Babington in Ann. Mag. nat. Hist., ser. 2, 16: 397 (1855)

All the material on which Babington based his description is in his herbarium in CGE and is so
labelled in his own handwriting. There are six syntypes:

1) In a pond by the road on the common at Denver, Norfolk, 2 June 1853, C. C. Babington.
2) In a pit on the common at Denver, Norfolk, 2 June 1853, C. C. Babington.

3) On the mud by the pond on the common at Denver, Norfolk, 2 June 1853, C. C. Babington.
4) In a pond near Legge’s Farm near Hatfield, Herts., 12 June 1855, C. C. Babington.

5) Hedon, near Hull, 7 Sept. 1853, W. W. Newbould.

6) In stagnis Siciliae, Prof. Gasparinii.

In selecting a lectotype, uncertain specimens and those not fulfilling the required characters should
be rejected first. Number 6 can be rejected as Babington himself says it only “‘appears to be
floribundus’’. Number 3 has no floating leaves and therefore shows only some of the characters.
The remaining four sheets are excellent specimens. A careful comparison of these specimens with
the original description enables 4 and 5 to be rejected on one or more grounds and for the Denver
plants to be regarded as those that best fit the description. Both are good specimens, but number 2
shows the rooting at the lower nodes and good sepals. It thus fits the description exactly and is
designated as the lectotype. It is R. peltatus Schrank and R. floribundus must be reduced to a
synonym of that species. Sheets 1 and 4 are also R. peltatus. Sheet 5 and possibly 3 are R. baudotii
Godron. Sheet 6 is R. aquatilis L.

R. peltatus var. (vel forma) lacerus Druce in Rep. botl Soc. Exch. Club Br. Isl. , 5: 272 (1919)
Druce stated that this plant “‘differs from the type in the floating leaves being irregularly cut into
acute, wedge-shaped segments, sometimes with the apices prolonged into long, comb-like parts.”’
There are three syntypes in Druce’s herbarium in OXF collected from the River Don, Alford, N.
Aberdeen by Druce in 1918 and labelled by him as ‘‘R. peltatus var. (vel forma) lacerus’’, all of
which are R. peltatus Schrank. One of these can be rejected because the laminar leaves are not
prolonged into capillary appendages. Both of the remaining sheets are good specimens and
S.D.W. has designated one of them as the lectotype. This specimen has the longer capillary
appendages of the two and thus best fits the description. This sheet is marked ““Don” in Druce’s
handwriting.

SHORT NOTES 181
REFERENCES

BENSON, L. (1954). Supplement to a treatise on the North American Ranunculi. Amer. Midl. Nat. , 52: 328-
369.

Cook, C. D. K. (1966). A monographic study of Ranunculus subgenus Batrachium (DC.) A. Gray. Mitt. bot.
StSammil., Mtiinch., 6: 47-237.

S. D. WEBSTER & P. D. SELL
Nature Conservancy Council, All Saints House, High Street, Colchester, COl 1UG

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Watsonia, 17, 183-198 (1988) 183

Plant Records

Records for publication must be submitted to the appropriate vice-county Recorder (see Vice-county Recorders
(1985)), and not the Editors. The records must normally be of species, hybrids or subspecies of native or
naturalized alien plants belonging to one or more of the following categories: 1st or 2nd v.c. record, 1st post-
1930 v.c. record; only extant v.c. locality, or 2nd such locality; a record of an extension of range by more than
100 km. Such records will also be accepted for the major islands in v.cc. 102-104 and 110. Only 1st records can
be accepted for Rubus, Hieracium and hybrids. Records for subdivisions of vice-counties will not be treated
separately; they must therefore be records for the vice-county as a whole. Records of Taraxacum are now being
dealt with separately, by Dr A. J. Richards, and will be published at a later date. New records for Irish vice-
counties are published elsewhere, usually in the Jrish Naturalists’ Journal.

Records are arranged in the order given in the List of British vascular plants by J. E. Dandy (1958) and his
subsequent revision (Watsonia, 7: 157-178 (1969)). All records are field records unless otherwise stated. With
the exception of collectors’ initials, herbarium abbreviations are those used in British and Irish herbaria by
D. H. Kent & D. E. Allen (1984).

The following signs are used:

* before the record: to indicate a new vice-county record.

+ before the species number: to indicate that the plant is not a native species of the British Isles.

+ before the record: to indicate a species which, though native in some parts of the British Isles, is not so in the
locality recorded.

[] enclosing a previously published record: to indicate that the record should be deleted.

1/4. LYCOPODIUM CLAVATUM L. 12, N. Hants.: Warren Heath, GR 41/76.59. Bank of pond.
C. R. Hall, 1985, herb. A. Brewis. Only extant record.

4/1. EQUISETUM HYEMALE L. *12, N. Hants.: Westlands Copse, GR 41/80.45. Stream-bank.
R. B. Gibbons, 1985, herb. A. Brewis, det. F. Rose.

4/7. EQUISETUM SYLVATICUM L. *§2, Anglesey: Bryn Mwcog, GR 23/46.83. Damp grassland.
L. J. Colley, M. R. Davies & R. H. Roberts, 1987, NMW.

4/7 X 10. EQUISETUM SYLVATICUM L. X E. TELMATEIA Ehrh. *11, S. Hants.: Minstead, GR
41/29.10. Roadbanks. R. P. Bowman, 1986, E, det. C. N. Page. 1st ever record of this hybrid.

4/8. EQUISETUM PRATENSE Ehrh. 99, Dunbarton: Lang Craigs, GR 26/43.76. Grassy rock
ledge on basalt crags. A.McG. Stirling, 1987, E. 2nd record.

4/10. EQUISETUM TELMATEIA Ehrh. *79, Selkirks.: Newhouse Bank, Ettrick, GR 36/37.22.
Flushed woodland and roadside verge. R. W. M. Corner, 1987, herb. R.W.M.C.

7/1. HYMENOPHYLLUM TUNBRIGENSE (L.) Sm. *67, S. Northumb.: Bull Crag, GR 35/66.86.
M.O. Hill, 1987, herb. G.A. Swan.

+17/1. MATTEUCIA STRUTHIOPTERIS (L.) Tod. *79, Selkirks.: R. Ettrick at Thirlestane, GR
36/28.15. Shaded river-bank. *80, Roxburghs.: Linthill, GR 36/54.26. Woodland by burn.
Both C.O. Badenoch, 1987, herb. R.W.M. Corner.

21/6 X 7. DRYOPTERIS CARTHUSIANA (Vill.) H. P. Fuchs x D. pitatata (Hoffm.) A. Gray
*70, Cumberland: Culgaith, GR 35/60.31. Cleared woodland. R. W. M. Corner, 1983, det. C. R.
Fraser-Jenkins.

21/8. DRYOPTERIS AEMULA (Aiton) O. Kuntze *8, S. Wilts.: Langley Wood, GR 41/22.20.
Streamside. F. Rose & R. J. Hornby, 1985. *11, S. Hants.: Wilverley Inclosure, GR 41/24.01.
Drainage ditch in Pseudotsuga plantation. Mrs A. Bolton, 1987, herb. R. P. Bowman, det. R.P.B.

25/1 X int. POLYPODIUM VULGARE L. X P. INTERJECTUM Shivas *6, N. Somerset: Nye Drove,
Winscombe, GR 31/40.60. E. M. McDonnell & Lady R. FitzGerald, 1985, det. R. H.

184 PLANT RECORDS

Roberts. *75, Ayrs.: Ailsa Craig, GR 25/02.99. Rock crevice. A.McG. Stirling, 1987, herb.
A.McG.S.

+32/kae. LARIX KAEMPFERI (Lamb.) Carriére *99, Dunbarton: Near Rosneath Point, GR 26/
26.80. Forest trackside. A.McG. Stirling & A. Rutherford, 1984. Regenerating.

40/1. ACONITUM NAPELLUS L. +*73, Kirkcudbrights.: Below Carsluith Castle, GR 25/49.54.
Wood. O. M. Stewart, 1987.

46/12 < 13. RANUNCULUS FLAMMULA L. X R. REPTANS L. *98, Main Argyll: Loch Awe
opposite Kilchurn Castle, GR 27/13.27. Damp loch shore. A.McG. Stirling, 1977, LTR, det. R. J.
Gornall.

46/22c. RANUNCULUS PENICILLATUS (Dumort.) Bab. subsp. PSEUDOFLUITANS (Syme) S. Webster
var. PSEUDOFLUITANS (Syme) S. Webster *46, Cards.: Afon Teifi above Henllan, GR 22/37.40.
River. A. O. Chater, 1986, NMW, det. S. D. Webster.

50/2. THALICTRUM ALPINUM L. *64, Mid-W. Yorks.: Giggleswick Scar, Richardsons Scar &
Gordale, GR 34/8.6. N. J. Winch, annotated Flora in LINN. Rocks above Settle, GR 34/8.6.
J. Tatham, 1841, BM, det. J. M. Mullin. lst and 2nd records. See BSBI News, 47: 20-21 (1987).

62/1. CHELIDONIUM MAJUS L. +*103, Mid Ebudes: Dervaig, Mull, GR 17/43.52. B. Rae,
1987, det. J. W. Clark & A. Slack.

+63/1. ESCHSCHOLZIA CALIFORNICA Cham. *12, N. Hants.: Stoke, GR 41/40.50. G. D. Field,
1984.

+65/1. CORYDALIS SOLIDA (L.) Sw. *39, Staffs.: Colwich, GR 43/02.20. Roadside spinney.
M. B. Jennings, 1968, still there in 1986.

66/7. FUMARIA DENSIFLORA DC. 73, Kirkcudbrights.: Castle Douglas, GR 25/75.62. Waste
ground. P. Stanley, 1987. 1st post-1930 record.

+68/1. ERUCASTRUM GALLICUM (Willd.) O. E. Schultz *69, Westmorland: M6 road west of
Bendrigg, GR 34/58.90. Waste ground. C. E. Wild, 1987, LANC, det. T. C. G. Rich.

+71/1. HIRSCHFELDIA INCANA (L.) Lagr.-Foss. 39, Staffs.: Pensnett, GR 32/91.88. Spoil heap.
C. B. Westall, 1987. 1st post-1930 record.

75/1. CRAMBE MARITIMA L. 59, S. Lancs.: Between Hall Road and Hightown, GR 34/29.02.
Rubble above high-tide mark. V. Gordon, 1987. Ist post-1930 record.

+76/2. RAPISTRUM RUGOSUM (L.) All. 69, Westmorland: Heversham, GR 34/49.82. Copse.
M. Baecker, 1986, LANC, det. T. C. G. Rich. 2nd record.

+79/den. LEPIDIUM DENSIFLORUM Schrader 17, Surrey: Sutton, GR 51/25.64. Churchyard.
M. J. Dyke, 1987. 1st record since 1920.

81/2. CARDARIA CHALEPENSIS (L.) Hand.-Mazz. *25, E. Suffolk: Felixstowe, GR 62/3.3.
Roadside. Herb. A. H. Maude, 1919, BM, det. T.C.G. Rich.

88/5. COCHLEARIA DANICA L. +*12, N. Hants.: Aldershot, GR 41/86.52. Central reservation
of A325 road. T. Dove, 1985.

94/4. DRABA MURALIS L. +98, Main Argyll: Ormaig House, Loch Craignish, GR 17/82.03.
Garden wall. B. H. Thompson, 1987. 2nd record.

+97/7. CARDAMINE RAPHANIFOLIA Pourret *99_ Dunbarton: Loch Lomond, near Balloch, GR
26/38.82. Marshy ground. J. H. Penson, 1966. Upper Helensburgh, GR 26/30.83. Wet stream-
bank. A. McG. Stirling & A. Rutherford, 1972. 1st and 2nd records.

+98/3. BARBAREA INTERMEDIA Bor. *98, Main Argyll: Bishop’s Glen, Dunoon, GR 26/16.76.
Disturbed ground. B. H. Thompson, 1987, herb. B.H.T., det. T.C.G. Rich.

102/1 xX 2. NASTURTIUM OFFICINALE R.Br. X N. MICROPHYLLUM (Boenn.) Reichenb. *46,
Cards.: Mwnt, GR 22/17.51. Stream. A. O. Chater, 1987.

PLANT RECORDS 185

102/2. NASTURTIUM MICROPHYLLUM (Boenn.) Reichenb. *77, Lanarks.: South of Strathaven,
GR 26/7.4. Burn. P. Macpherson, 1987, herb. P.M., det. T.C.G. Rich.

+102/6. RORIPPA AUSTRIACA (Crantz) Bess. 33, E. Gloucs.: Teddington Hands Roundabout,
Tewkesbury, GR 32/96.33. Roadside verge. R. J. Cooper, 1987. Only extant locality.

112/1. RESEDA LUTEOLA L. 98, Main Argyll: Bishop’s Glen, Dunoon, GR 26/16.76.
Disturbed ground. B. H. Thompson, 1987. 2nd record.

113/1. VIOLA ODORATA L. *93, N. Aberdeen: Insch, GR 38/63.28. Deciduous woodland. D.
Welch, 1987, ABD.

113/1 X 2. VIOLA ODORATA L. X V. HIRTA L. *80, Roxburghs.: West of Carham Station, GR
36/78.36. Limestone railway cutting. R. W. M. Corner, 1987, herb. R.W.M.C., det. D. M. Moore.

113/5. VIOLA REICHENBACHIANA Jord. ex Bor. *67, S. Northumb.: Devil’s Water near
Ordley, GR 35/94.59. Steep grassy bank. G. A. Swan, 1987. Gunnerton Crag, GR 35/91.75.
Limestone scrub. A. J. Richards, 1987. Both herb. G.A.S., det. D. M. Moore. 1st confirmed and
2nd record.

113/12 X 13. VIOLA TRICOLOR L. X V. ARVENSIS Murr. *80, Roxburghs.: Smailholm Craigs, St
Boswells, GR 36/63.34. Shallow soil over rock. M. Little, 1985, herb. R.W.M.Corner, det. D. R.
McKean.

115/5 X 6a. HYPERICUM PERFORATUM L. X H. MACULATUM Crantz subsp. MACULATUM 7 A fe
Lanarks.: Bothwell Castle, GR 26/68.59. Grassy area. J. H. Dickson, 1984, GL, det. N. K. B.
Robson.

123/14. SILENE ALBA (Mill.) E. H. L. Krause *103, Mid Ebudes: Near Bishop’s House, Iona,
GR 17/28.24. Waste ground. J. W. Clark, 1987. 1st confirmed Mull record.

7124/cor. LYCHNIS CORONARIA (L.) Desr. *70, Cumberland: Rockcliffe, Carlisle, GR 35/
35.61. Scrub on sandstone river cliff. R. E. Groom, 1979, still present in 1987, LANC.

127/1. DIANTHUS ARMERIA L. 11, S. Hants.: Avon Forest Park, St Leonards, GR 41/12.01.
Sandy acid grassland. Mrs F. Woodhead, 1986. Only extant locality.

127/4. DIANTHUS PLUMARIUS L. *73, Kirkcudbrights.: Below Port Mary House, GR 25/75.45.
Shingle on shore. C. D. Preston & O. M. Stewart, 1986, E.

127/8. DIANTHUS DELTOIDES L. +11, S. Hants.: Avon Forest Park, St Leonards, GR 41/12.01.
Sandy grass heath. R. P. Bowman, 1986, herb. R.P.B., det. S. Hooper. Only extant locality.

129/1. SAPONARIA OFFICINALIS L. +77, Lanarks.: Bellahouston, GR 26/56.63. Grassy waste
ground. P. Macpherson, 1987, herb. P.M. 2nd record. +99, Dunbarton: Near Dog’s Loan,
Dunbarton, GR 26/40.74. Sandy shore. E. Brock, 1981. 2nd record.

131/10. CERASTIUM DIFFUSUM Pers. +*64, Mid-W. Yorks.: Adel, Leeds, GR 44/27.39.
Garden weed. K. Bradbury, 1987, det. P. P. Abbott. Embsay Station, GR 44/00.53. Platform and
trackside. E. Crackles & J. Duncan, 1987. 1st and 2nd records. +*79, Selkirks.: Galashiels, GR
36/47.37. Old railway track. J. A. Murray, 1986, det. D. R. McKean. +80, Roxburghs.:
Galashiels, GR 36/47.37. Old railway track. J. A. Murray, 1986, det. D. R. McKean. 2nd record.

131/12. CERASTIUM SEMIDECANDRUM L. *79, Selkirks.: Old Redhead, GR 36/44.37. Rocky
roadside bank. R. W. M. Corner, 1987, herb. R.W.M.C.

132/1. MyosoToN AQuaATicuM (L.) Moench 59, S. Lancs.: Lostock, GR 34/66.08. Wet
ground. V. Gordon, 1987. Only extant locality.

141/2. ARENARIA LEPTOCLADOS (Reichb.) Guss. 77, Lanarks.: Eastfield, Glasgow, GR 26
63.61. Bare ground. P. Macpherson, 1987. 2nd record.

4141/6. ARENARIA BALEARICA L. *99, Dunbarton: Helensburgh, GR 26/29.83. Garden wall.
A.McG. Stirling & A. Rutherford, 1985. Present since 1950.

186 PLANT RECORDS

147/1. ILLECEBRUM VERTICILLATUM L. +59, S. Lancs.: Fazakerley, GR 33/38.97. Old railway
lines. B. K. Byrne, 1987. 2nd record.

148/1. SCLERANTHUS ANNUUS L. 7, N. Wilts.: Near The Column, Savernake Forest, GR 41/
23.64. Cornfield. D. Green & J. Newton, 1985. Only extant locality.

149/1a. MONTIA FONTANA L. subsp. FONTANA *52, Anglesey: Twyn-y-Parc, Bodorgan, GR
23/36.64. Coastal turf near gullery. C. D. Preston, 1986, NMW, det. S. M. Walters.

149/1b. MonrtTIA FONTANA L. subsp. CHONDROSPERMA (Fenzl) Walters *80, Roxburghs.: Near
Crailing Hall, Oxnam, GR 36/70.21. Sandy soil over basalt. South end of Broadlee Loch, GR 36/
40.09. Peaty flush. Both R.W.M. Corner, 1986, herb. R.W.M.C., det. S. M. Walters. Ist and 2nd
records.

149/1c. MONTIA FONTANA L. subsp. AMPORITANA Sennen 38, Warks.: Sutton Park, GR 42/
09.96. Marshy ground. J. C. Bowra, 1986, WAR, det. S. M. Walters. 1st record since 1875.

149/1d. MONTIA FONTANA L. subsp. VARIABILIS Walters 52, Anglesey: Llyn Coron, GR 23/
37.69. Close-grazed lakeside turf. C. D. Preston, 1986, NMW, det. S. M. Walters. 2nd localized
record. *81, Berwicks.: Fans, GR 36/61.41. Sandy bank. M. E. Braithwaite & M. Mutch,
1987, herb. M.E.B., det. S. M. Walters.

+DISPHYMA CRASSIFOLIUM (L.) L. Bolus *52, Anglesey: North of Bwa Du, Rhoscolyn, GR
23/26.76. Coastal cliff. C. D. Preston, 1986, photographs in CGE. Originally planted, first seen
1976, now thoroughly naturalized. 1st Welsh record.

4153/4. AMARANTHUS LIVIDUS L. 38, Warks.: Guy’s Cliffe, Warwick, GR 42/28.66. Waste
ground. J. C. Bowra, 1985, WAR, det. T. B. Ryves. Still present in 1987. 1st record since 1874.

154/3. CHENOPODIUM VULVARIA L. 70, Cumberland: Eden Bridge, Lazonby, GR 35/55.40.
Landscaped picnic area. R. E. Groom, 1984, LANC, det. J. M. Mullin. 1st record this century.

154/11. CHENOPODIUM MURALE L. 70, Cumberland: Great Salkeld, GR 35/55.36. Garden.
R. W. M. Corner, 1982, LANC, det. J. M. Mullin. 1st record this century.

+154/16. CHENOPODIUM GLAUCUM L. *70, Cumberland: Eden Bridge, Lazonby, GR 35/55.40.
Landscaped picnic area. R. E. Groom, 1984, LANC, det. J. M. Mullin.

[156/1 x 2. ATRIPLEX LITTORALIS L. X A. PATULA L. 95, Moray: Delete record published in
Watsonia, 12: 350 (1979), specimen at E redetermined as A. littoralis L. X A. prostrata Boucher ex
DC. by P. M. Taschereau. |

156/4. ATRIPLEX GLABRIUSCULA Edmondst. 59, S. Lancs.: Hightown, GR 34/29.02. High-tide
mark. V. Gordon, 1987. Only extant locality.

156/5. ATRIPLEX LACINIATA L. 59, S. Lancs.: Hightown, GR 34/29.02. High-tide mark.
V. Gordon, 1987. Only extant locality.

156/lon. ATRIPLEX LONGIPES Drejer *68, Cheviot: Near Beal Point, GR 46/08.42. Under
Scirpus maritimus in salt-marsh. G. A. Swan, 1983, herb. G.A.S., det. P. M. Taschereau.

+KOCHIA SCOPARIA (L.) Schrader *70, Cumberland: Eden Bridge, Lazonby, GR 35/55.40.
Landscaped picnic area. R. E. Groom, 1984, LANC, det. J. M. Mullin.

7168/8. GERANIUM MACRORRHIZUM L. *99, Dunbarton: Craigrownie, GR 26/22.81. Pathside.
A.McG. Stirling & A. Rutherford, 1985.

168/14. GERANIUM PUSILLUM L. *75, Ayrs.: Maidenhead Bay, GR 26/21.08. Sandy waste
ground near shore. A.McG. Stirling, 1986, E.

7170/4. OXALIS EUROPAEA Jordan 52, Anglesey: Henblas, GR 23/42.72. Garden weed. BSBI
meeting, 1987, NMW, det. C. D. Preston & R. H. Roberts. 2nd record. *99, Dunbarton:
South King Street, Helensburgh, GR 26/30.82. Garden path. A. Rutherford, 1983.

7170/exi. OXALIS EXILIS A. Cunn. *80, Roxburghs.: Kelso Abbey, GR 36/72.33. Lawn.

PLANT RECORDS 187

Greycrook, St Boswells, GR 36/60.30. Garden weed. Both M. E. Braithwaite, 1987. 1st and 2nd
records. *81, Berwicks.: Rhymers Tower, Earlston, GR 36/57.38. Lawn. M. E. Braithwaite &
R. W. M. Corner, 1987, herb. M.E.B.

192/1. TRIFOLIUM ORNITHOPODIOIDES L. *26, W. Suffolk: Red Lodge, Freckenham, GR 52/
69.70. Sandy waste ground. D. Dupree, 1953 & P. H. Oswald, 1987, CGE.

192/10. TRIFOLIUM STRIATUM L. +59, S. Lancs.: Near Lydiate, GR 34/34.04. Laneside.
V. Gordon & G. Sloman, 1987. Only extant locality.

192/occ. TRIFOLIUM OCCIDENTALE D. E. Coombe *41, Glam.: Tear’s Point & Mewslade, GR
21/41.87. Cliff-top turf. J. Dunn, 1987, conf. D. E. Coombe. Ist Welsh record.

7192/pan. TRIFOLIUM PANNONICUM Jacq. 59, S. Lancs.: Melling, GR 34/37.00. Railway
embankment. V. Gordon, 1987. 2nd record.

7193/1 car. ANTHYLLIS VULNERARIA L. subsp. CARPATICA (Pant.) Nyman var. PSEUDOVULNERARIA
(Sag.) Cullen *73, Kirkcudbrights.: East of Castle Douglas, GR 25/78.62. Rough ground.
O. M. Stewart, 1986, E, det. J. R. Akeroyd.

200/3. ASTRAGALUS GLYCYPHYLLOS L. *70, Cumberland: West of Culgaith, GR 35/59.29.
Basic bank. R. W. M. Corner, 1987, LANC.

206/12. VICIA LUTEA L. +38, Warks.: Near Brandon Marsh N.R., GR 42/38.75. Raised grassy
bank. J. C. Bowra, 1987, WAR, det. P. J. Copson. 2nd record.

+207/6. LATHYRUS SYLVESTRIS L. 39, Staffs.: Burton-on-Trent, GR 43/23.24. Grassy bank.
B. R. Fowler, 1976. 1st post-1930 record.

+207/gra. LATHYRUS GRANDIFLORUS Sibth. & Sm. *39, Staffs.: Pennant Road, Rowley Regis,
GR 32/96.87. Rubble. M. Doulton, 1987, herb. B.R. Fowler.

+209/1 < 2. SPIRAEA SALICIFOLIA L. X S. DOUGLAS Hook. *59,S. Lancs.: Trawden, GR 34/
91.39. Hedge. V. Gordon, 1987, det. A. J. Silverside. *70, Cumberland: Lees Hill,
Kirkcambeck, GR 35/55.67. E. Rhone & M. R. Smith, 1983, LANC, det. A. J. Silverside.

4209/1 x< alb. SPIRAEA SALICIFOLIA L. X S. ALBA Duroi *99, Dunbarton: Near Rosneath
Castle, GR 26/25.82. Rough marshy ground. A.McG. Stirling & A. Rutherford, 1979, det. A. J.
Silverside. Vast, long-established colony.

+209/2. SPIRAEA DOUGLASII Hook. 59, S. Lancs.: Between Chew Moor and Lostock Junction,
GR 34/67.08. Waste ground. V. Gordon, 1987, det. A. J. Silverside. 2nd record.

+209/2 < alb. SPIRAEA DOUGLASII Hook. X S. ALBA Duroi *59, S. Lancs.: Chadwicks Farm
Lane, Abram, GR 34/61.00. Hedge. V. Gordon, 1987, det. A. J. Silverside.

+209/alb. SPIRAEA ALBA Duroi 73, Kirkcudbrights.: Maxwelltown, GR 25/97.75. Waste
ground. O. M. Stewart, 1986, E, det. A. J. Silverside. 2nd record.

+ARUNCUS DIOICUS (Walter) Fernald *99, Dunbarton: Near Ardencaple Farm,
Helensburgh, GR 26/28.83. Waste ground. A. Rutherford, 1977.

211/9 X 11/129. RuBus cagsius L. X R. ULMIFOLIUS Schott *38, Warks.: Newbold Pacey,
GR 42/29.57. Churchyard hedgerow. J. W. Partridge, 1987, det. A. Newton.

211/11/22. RUBUS WARRENII Sudre *70, Cumberland: West of Fletchertown, GR 35/20.42.
Hedge. G. Halliday & A. Newton, 1986, LANC.

211/11/226. RUBUS SUBTERCANENS W. C. R. Wats. *99, Dunbarton: Shore of R. Clyde,
Dalmuir, GR 26/47.71. Waste ground. A.McG. Stirling & A. Newton, 1980, herb A.McG.S., det
A. N. Ist Scottish record.

211/11/arm. RUBUS ARMIPOTENS Barton ex A. Newton +*75, Ayrs.: Skelmorlie, GR 26/19.67.
Roadside. A. Newton & A.McG. Stirling, 1987, E, det. A. N. 1st Scottish record.

188 PLANT RECORDS

211/11/blo. RuBUS BLOxAMII Edees *7, N. Wilts.: Near Grand Avenue, Savernake, GR 41/
23.65. Glade. A. Newton & D. Allen, 1986, herb. R. D. Randall.

211/11/lan. RuBUS LANATICAULIS Edees & A. Newton *75, Ayrs.: West of Dalquarran
Castle, Dailly, GR 26/26.02. Wood border. A. Newton & A.McG. Stirling, 1987, E, det. A.N. 1st
Scottish record.

211/11/nor. RuBUS NoRVICENSIS A. L. Bull & Edees *7, N. Wilts.: Near Grand Avenue,
Savernake, GR 41/23.65. Glade. A. L. Bull, 1986, herb. R.D. Randall, det. A. Newton.

212/13 * 15. POTENTILLA X MIXTA Nolte ex Reichb. *103, Mid Ebudes: Craigdarroch House,
Arinagour, Coll, GR 17/22.57. Grass by wall. J. W. Clark, 1986, E, det. D. R. McKean.

218/2. AGRIMONIA PROCERA Wallr. 25, E. Suffolk: Nacton, GR 62/20.40. Woodside track, on
sandy soil. Suffolk Trust for Nature Conservation Recording Team, 1986. 2nd record.

220/2. ALCHEMILLA CONJUNCTA Bab. +*39, Staffs.: Coseley, GR 32/94.93. Limestone slope.
B. R. Fowler, 1987, CGE, det. S. M. Walters. +*77, Lanarks.: Busby, GR 26/57.56. Waste
ground. P. Macpherson, 1987, herb. P. M.

220/3/3. ALCHEMILLA FILICAULIS Buser subsp. FILICAULIS 68, Cheviot: Hobberlaw, GR 46/
17.11. Old limestone quarry. G. A. & M. Shaw, 1987, herb. G.A.S., det. S. M. Walters. 2nd
record.

$220/3/12. ALCHEMILLA MOLLIS (Buser) Rothm. *11, S. Hants.: Plain Heath, Holmsley, GR
40/22.99. Short turf. R. P. Bowman, 1985, herb. R. P. B., det. S. M. Walters. *99, Dunbarton:
Near Rhu Station, GR 26/27.84. Rough ground by roadside. A. McG. Stirling & A. Rutherford,
1976.

+223/2. SANGUISORBA MINOR Scop. subsp. MURICATA Briq. 83, Midlothian: Arthur’s Seat, GR
36/27.72. Roadside. N. F. Stewart, 1983. 1st record since 1906.

+226/3. PRUNUS CERASIFERA Ehrh. *77, Lanarks.: Between Glasgow and Renfrew, GR 26/
51.67. Edge of woodland. P. Macpherson & E. Teasdale, 1986, herb. P. M., det. R. D. Meikle.

+227/2. COTONEASTER SIMONSII Baker *39, Staffs.: Brocton, GR 33/96.18. Woodland margin.
B. R. Fowler, 1987.

+229/pru. CRATAEGUS PRUNIFOLIA (Lam.) Pers. *39, Staffs.: Broadhurst Green, GR 33/98.15.
Roadside verge. B. R. Fowler, 1974.

232/1 X 5/1. SORBUS AUCUPARIA L. X S. ARIA (L.) Crantz *39, Staffs.: Newborough, GR 43/
12.22. Roadside woodland. M. B. Jennings, 1987, herb. B.R. Fowler.

+232/6/lat. SORBUS LATIFOLIA (L.) Crantz sensu stricto *39, Staffs.: Brown’s Coppice E. of
Hanbury, GR 43/18.28. Deciduous woodland. B. R. Fowler, 1987, det. P. J. M. Nethercott.

232/7. SORBUS TORMINALIS (L.) Crantz 64, Mid-W. Yorks.: Studley Royal, GR 44/28.69.
Magnesian limestone cliff. P. P. Abbott, 1987. 2nd record.

+237/hel. CRASSULA HELMSII (T. Kirk) Cockayne *33, E. Gloucs.: Arle Court, Cheltenham,
GR 32/91.21. Muddy edge of lake. S. C. Holland & D. S. Dudley-Smith, 1987, conf. F. H.
Dawson. *39, Staffs.: Brocton, GR 33/97.19. Flooded disused gravel pit. B. R. Fowler, 1987,
herb. B.R.F. Known here for 10 years. *46, Cards.: Ynys-las, GR 22/69.93. Pool in dune slack.
M. Wainwright, 1986.

+239/4. SAXIFRAGA CF. UMBROSA L. *99 Dunbarton: Clachan Woods, Rosneath, GR 26/
25.83. Semi-natural woodland. A.McG. Stirling & A. Rutherford, 1983, det. C. D. Brickell.

4239/4 x 5. SAXIFRAGA UMBROSA L. X S. SPATHULARIS Brot. *99, Dunbarton: Bank of
Gielston Burn, GR 26/34.77. Steep, wet rocks. A.McG. Stirling & A. Rutherford, 1985, det. C. D.
Brickell.

4239/7. SAXIFRAGA CYMBALARIA L. *69, Westmorland: Winderwath, Culgaith, GR 35/59.29.
Garden weed for many years. A. Wilson, 1986, LANC.

PLANT RECORDS 189

+240/1. TELLIMA GRANDIFLORA (Pursh) Douglas ex Lindley *99, Dunbarton: Rhu, GR 26/
26.84. Marshy ground near stream. A. Rutherford, 1984. Known here since 1940s.

}+PELTIPHYLLUM PELTATUM (Torr. ex Benth.) Engler *99, Dunbarton: Duhill, Helensburgh,
GR 26/29.83. Rough ground near seepage. A. Rutherford, 1978, still present in 1984.

+RODGERSIA PODOPHYLLA A. Gray *99, Dunbarton: Bank of R. Leven near Dumbarton Golf
Course, GR 26/39.76. Well established under willows. B. McNeil, 1986, det. C. D. Brickell.

4244/1. PHILADELPHUS CORONARIUS L. *69, Westmorland: South of Barbon, GR 34/62.81.
Edge of plantation. C. E. Wild, 1987, LANC.

249/2. LYTHRUM HYSSOPIFOLIA L. 13, W. Sussex: Thorney Island, GR 41/74.03. Depression in
arable field. D. J. P. Miller, 1987, BM. 1st record since 1853. *34, W. Gloucs.: Swan Lake,
Slimbridge, GR 32/72.04. Marshy edge of lake. B. Stewart & N. Woodward, 1987.

252/1. HIPPOPHAE RHAMNOIDES L. +93, N. Aberdeen: Cruden Bay, GR 48/09.36. Sand. D.
Welch, 1987. 1st post-1930 record; probably a relic of introduced plants reported by Trail in 1904.

254/1 x 3. EPpPILOBIUM HIRSUTUM L. X E. MONTANUM L. *70, Cumberland: West of
Boltonfellend, GR 35/47.68. Roadside ditch. G. Halliday, 1986, LANC.

254/4. EPILOBIUM LANCEOLATUM Seb. & Mauri 7, N. Wilts.: Old Totterdown, GR 41/13.71.
Scrub. K. Payne, 1986, herb. D. Green. 2nd record.

254/5. EPILOBIUM ROSEUM Schreber *69, Westmorland: Brathay Hall, Ambleside, GR 34/
36.03. G. Halliday, 1986, LANC. 1st seen here by A. J. Richards in late 1970s.

254/7. EPILOBIUM TETRAGONUM L. *59, S. Lancs.: Victoria Park, St Helens, GR 33/50.96.
D. Jolly, 1987. 1st localized record.

254/12. EPILOBIUM ALSINIFOLIUM Vill. *93, N. Aberdeen: Creag an Sgor, GR 38/37.21.
Flushed shaly earth. D.Welch, 1987, ABD.

+256/2. OENOTHERA ERYTHROSEPALA Borbas *75, Ayrs.: Troon, GR 26/33.29. Sandy waste
ground by railway. A.McG. Stirling, 1987.

+256/3. OENOTHERA STRICTA Ledeb. ex Link *29, Cambs.: Dernford Fen, GR 52/47.50.
Sedge fen. G. M. S. Easy, 1987, herb.G.M.S.E.

+256/cam. OENOTHERA CAMBRICA Rostanski 4, N. Devon: East-the-Water, Bideford, GR 21/
45.26. Disused railway. W. H. Tucker, 1986. 2nd record.

262/2. CALLITRICHE PLATYCARPA Kiitz. 73, Kirkcudbrights.: Carton Bridge, GR 25/83.66.
Slow, deep burn. O. M. Stewart, 1986, E. 1st post-1930 record.

264/1. THESIUM HUMIFUSUM DC. 26, W. Suffolk: Bury Hill, Moulton, GR 52/66.64. Chalk
grassland. E. M. Hyde, 1987. 2nd extant locality.

+268/hib. HEDERA HIBERNICA (Kirchner) Bean ‘Hibernica’ *99, Dunbarton: Torwood Road,
Rhu, GR 26/27.83. Verge. A.McG. Stirling & A. Rutherford, 1971.

+268/col. HEDERA COLCHICA (C. Koch) C. Koch *99, Dunbarton: Greenfield, Garelochhead,
GR 26/24.90. Wooded shore. A. Rutherford, 1970.

7271/1. ASTRANTIA MAJOR L. 39, Staffs.: Fradswell, GR 33/99.31. Shady churchyard. E. S.
Edees, 1976. 1st post-1930 record.

285/4. APIUM INUNDATUM (L.) Reichb. f. 7, N. Wilts.: Durley, Savernake Forest, GR 41/
23.64. Pond. A. Summers, 1986, herb. D. Green. Only extant locality.

288/1. CICUTA VIROSA L. *81, Berwicks.: Everett Moss, GR 36/60.43. Fen. M. E. & P. F.
Braithwaite, 1987, herb. M.E.B. Previously reported site (Yetholm Loch) is in v.c. 80.

300/7. OENANTHE FLUVIATILIS (Bab.) Coleman 7, N. Wilts.: R. Churn, Cricklade North
Meadow N.N.R., GR 41/09.94. K. Payne, 1986. Only extant site.

190 PLANT RECORDS

311/1 X +2. HERACLEUM SPHONDYLIUM L. X H. MANTEGAZZIANUM Somm. & Levier wah,
Man: Kionlough, Bride, GR 24/45.99. Wet hollow with willow scrub. M. Devereau & L. S.
Garrad, 1987.

319/2. EUPHORBIA LATHYRIS L. +*73, Kirkcudbrights.: Carsethorn, GR 25/99.59. Shore.
O. M. Stewart, 1987.

319/7. EUPHORBIA PLATYPHYLLOS L. *39, Staffs.: Terrace Street, Blackheath, GR 32/96.86.
Rubble. L. Goodby, 1987, herb. B. R. Fowler.

+319/16. EUPHORBIA CYPARISSIAS L. 70, Cumberland: Beck Farm, Wheyrigg, GR 35/19.48.
Rough trackside. M. Porter, 1987, LANC. 1st post-1930 record.

320/1/4 POLYGONUM ARENASTRUM Boreau *77, Lanarks.: Cathkin, Glasgow, GR 26/62.57.
Side of reservoir. A.McG. Stirling & P. Macpherson, 1983, herb. P.M. Shieldhall, Glasgow, GR
26/53.65. Waste ground. P. Macpherson & E. Teasdale, 1986. 1st and 2nd records.

320/2. POLYGONUM OXYSPERMUM C. A. Meyer & Bunge ex Ledeb. subsp. ram (Bab.) D. A. Webb
& Chater 25, E. Suffolk: Felixstowe, GR 62/29.33. Shingle beach. J. Heath, 1987, IPS. 1st
post-1930 record.

320/13. POLYGONUM MITE Schrank 12, N. Hants.: Bramshill Common, GR 41/74.62. Lake-
shore. C. R. Hall, 1986. Only extant locality. 25, E. Suffolk: Iken Wood, GR 62/39.56. Ditch.
I. M. Vaughan, 1985, herb. E. M. & M. A. Hyde, det. J. R. Akeroyd. 2nd extant locality.

[320/16. POLYGONUM DUMETORUM L. 38, Warks.: Delete records published in Watsonia, 13:
335 (1981), specimens have been redetermined as P. aubertii Louis Henry by J. R. Akeroyd.]

+320/20. POLYGONUM SACHALINENSE F. Schmidt *77, Lanarks.: Castlemilk, GR 26/60.59.
Wooded river valley. P. Macpherson, 1986, herb. P. M.

4320/22. POLYGONUM CAMPANULATUM Hooker fil. *99, Dunbarton: Cardross, GR 26/34.77.
Streamside. A.McG. Stirling, 1972, still present in 1985.

325/1/3. RUMEX TENUIFOLIUS (Wallr.) Love 79, Selkirks.: E. side of Haze Law, Craik Forest,
GR 36/30.11. Forestry track. R. W. M. Corner, 1987, herb. R.W.M.C. 2nd record.

325/11 X 12. RUMEx crispus L. X R. OBTUSIFOLIUs L. *25, E. Suffolk: Landguard Common,
Felixstowe, GR 62/28.32. Soil bank. A. Copping, 1981.

325/12 X 14. RUMEX oBTuSIFOLIUS L. X R. SANGUINEUS L. *64, Mid-W. Yorks.: Bishop
Wood near Selby, GR 44/55.34. P. P. Abbott, 1987, herb. P.P.A., det. W. A. Sledge.

325/15. RUMEX CONGLOMERATUS Murray 83, Midlothian: Duddingston Loch, GR 36/28.72.
Marshy loch edge. J. Sinclair, 1946, E, det. D. R. McKean. 1st post-1930 record.

+327/1. SOLEIROLIA SOLEIROLIT (Req.) Dandy *99, Dunbarton: East Montrose Street,
Helensburgh, GR 26/30.32. Garden wall. A. Rutherford, 1980.

343/5. SALIX TRIANDRA L. *77, Lanarks.: ““Cunningar Loop’, Rutherglen, GR 26/62.62.
River-bank. A. J. Silverside & A. McG. Stirling, 1981.

343/5 X 9. SALIX TRIANDRA L. X S. VIMINALIS L. *69, Westmorland: Temple Sowerby Moss,
GR 35/61.27. R. W. M. Corner, 1985, LANC, det. R. D.Meikle. *77, Lanarks.: ““Cunningar
Loop’’, Rutherglen, GR 26/62.62. River-bank. A. McG. Stirling & A. J. Silverside, 1981, herb.
A.McG.S., det. R. D. Meikle as S. X mollissima Hoffm. ex Elwert var. undulata (Ehrh.)
Wimm. ;

343/9 X 11. SALIX VIMINALIS L. X S. CAPREA L. *77, Lanarks.: Kinning Park, Glasgow, GR
26/56.63. Old railway line. P. Macpherson, 1987, herb. P.M., det. R. D. Meikle. *99
Dunbarton: Loch Lomond, Balloch Park, GR 26/38.72. Shingle shore. A.McG. Stirling, 1986,
herb. A.McG.S., det. R. D. Meikle.

343/9 X 13. SALIX VIMINALIS L. X S. AURITA L. *99, Dunbarton: R. Leven, Dunbarton, GR
26/39.76. Swampy river-bank. A.McG. Stirling, 1984, herb. A.McG.S., det. R. D. Meikle.

PLANT RECORDS 191

343/12 < 14. SALIX CINEREA L. X S. MYRSINIFOLIA Salisb. *77, Lanarks.: Gorbals, Glasgow,
GR 26/58.64. Roadside. P. Macpherson, 1987, herb. P.M., det. R. D. Meikle.
343/12 X 15. SALIX CINEREA L. X S. PHYLICIFOLIA L. *99, Dunbarton: St Germans Loch,

Bearsden, GR 26/54.71. Loch margin. A.McG. Stirling, 1984, herb. A.McG.S., det. R. D. Meikle.

+343/ude. SALIX UDENENSIS Trautv. & Mey. *70, Cumberland: Viol Moor, GR 35/59.40.
R. W. M. Corner, 1985, LANC, det. R. D. Meikle.

351/1. GAULTHERIA SHALLON Pursh *80, Roxburghs.: Kaim Burn, Edgerston, GR 36/68.12.
Woodland. R. W. M. Corner, 1986, herb. R.W.M.C.

358/4. VACCINIUM OXxYCOCccos L. 2, E. Cornwall: N.W. of Garrow Downs, GR 20/13.79.
Sphagnum bog. K. Bryson, T. J. Dingle & R. J. Murphy, 1987. 2nd record.

370/2. LYSIMACHIA NUMMULARIA L. *93, N. Aberdeen: Insch, GR 38/63.28. Moist deciduous
woodland. D. Welch, 1987, ABD.

371/1. TRIENTALIS EUROPAEA L. 103, Mid Ebudes: Near Loch Spelve, Mull, GR 17/66.24.
Area of bracken and heather. A. Gardner, 1987, photograph lodged with J. W. Clark. 2nd record.

372/1. ANAGALLIS TENELLA (L.) L. 7, N. Wilts.: Pike Corner, Ashton Keynes, GR 41/03.93.
Meadow ditch. D. Hornby, 1985, det. D. Green. 2nd post-1930 record.

388/1. POLEMONIUM CAERULEUM L. 68, Cheviot: S. bank of Ridlees Burn, GR 36/84.06. Steep
N.W.-facing andesite crag. G. A. & M. Swan, 1987. 2nd native locality.

7392/3. SYMPHYTUM ORIENTALE L. *99, Dunbarton: Luss churchyard, GR 26/35.92. Old wall.
A.McG. Stirling & A. Rutherford, 1978.

392/6. SYMPHYTUM TUBEROSUM L. +*7, N. Wilts.: Allington, GR 31/89.75. Roadside.
J. Presland, 1986, herb. D. Green. +*29, Cambs.: Harston, GR 52/43.50. Wooded ride.
G.M.S. Easy, 1987, herb G.M.S.E.

+392/ibe. SYMPHYTUM IBIRICUM Steven *99, Dunbarton: Below Cannisburn Hospital, GR 26/
54.70. Deciduous wood. A.McG. Stirling, 1985.
+BRUNNERA MACROPHYLLA (Adams) I. M. Johnston *69, Westmorland: Gossrig Hall,

Cliburn, GR 35/60.24. Well naturalized in grounds. R. W. M. Corner, 1984, LANC. TSB
Kirkcudbrights.: Knockbrex, GR 25/58.49. Ditch-bank. O. M. Stewart, 1986.

7393/1. BORAGO OFFICINALIS L. *73, Kirkcudbrights.: Carsethorn, GR 25/99.59. Sand on
shore. O. M. Stewart, 1987. 103, Mid Ebudes: Iona, GR 17/28.24. Waste ground. J. W. Clark,
1987. 1st Mull record.

400/1 x 4. MyosotTis scorPioipES L. X M. CAESPITOSA K. F. Schultz *70, Cumberland:
Parkside, Frizington, GR 35/02.15. Marshy field. C. C. Haworth, 1986, LANC, det. P. M. Benoit.

+AMSINCKIA INTERMEDIA Fischer & C. A. Meyer 38, Warks.: Bradnock’s Marsh, Barston,
GR 42/22.78. Waste ground. P. Noakes, 1984, det. P. J. Copson. 2nd record.

+PLAGIOBOTHRYS SCOULERI (Hooker & Arnott) I. M. Johnston *11, S. Hants.: Setley, GR
40/30.99. Locally abundant on silt and fine gravel in shallow flood hollows in levelled tip areas.
A. Hale, 1982, herb. Lady A. Brewis. Beaulieu Airfield, GR 41/35.00. Bare mud and gravel in
shallow flood hollow. R. P. Bowman, 1987, BM. 1st and 2nd records, both det. A. O. Chater.

1413/4. SOLANUM SARRACHOIDES Sendtn. 38, Warks.: Wasperton, GR 42/27.58. Waste
ground. J. W. Partridge, 1987, WAR, det. J. C. Bowra. 2nd record.

7414/1. SALPICHROA ORIGANIFOLIA (Lam.) Baill. 25, E. Suffolk: Ipswich, GR 62/17.45. Waste
ground. M. N. Sandford, 1987. 2nd record.

+416/3. VERBASCUM PHLOMOIDES L. *26, W. Suffolk: West Stow, GR 52/82.70. Roadside
verge. P. G. Lawson, 1986.

424/3. SCROPHULARIA UMBROSA Dumort. 38, Warks.: Moreton Morrell, GR 42/32.56. Ditch-
side. J. W. Partridge, 1987, WAR, det. P. J. Copson. Ist record since 1903.

192 PLANT RECORDS

+424/5. SCROPHULARIA VERNALIS L. *39, Staffs.: Terrace Street, Blackheath, GR 32/96.86.
Rubble. L. Goodby, 1987, herb. B. R. Fowler.

+425/1 X cup. MIMULUs GUTTATUS DC. X M. cuprEUS Dombrain *99, Dunbarton: Drumfad
Farm House, Glen Fruin, GR 26/31.84. Wet, peaty area. A.McG. Stirling & A. Rutherford, 1985,
det. A. J. Silverside.

+425/3. MIMULUS MOSCHATUS Dougl. ex Lindl. 79, Selkirks.: Lewenshope Burn, Yarrow,
GR 36/37.31. Bog by burn. D. J. Methven, 1987, herb. R. W. M. Corner. 1st record since 1917.

426/1. LIMOSELLA AQUATICA L. 32, Northants.: Pitsford Reservoir, GR 42/78.70. Muddy
reservoir margin. B. Adams, 1987. 1st post-1930 record.

1430/14. VERONICA PEREGRINA L. 77, Lanarks.: Bellahouston, Glasgow, GR 26/55.63.
Nursery weed. P. Macpherson, 1987, herb. P. M. 2nd record.

431/ X ama. HEBE X AMABILIS (Cheesem.) Cockayne & Allan *99, Dunbarton: Lennox
Drive, Helensburgh, GR 26/29.83. Dry waste ground. A.McG. Stirling & A. Rutherford, 1983.
Regenerating.

435/1/16. EUPHRASIA PSEUDOKERNERI Pugs]. 26, W. Suffolk: Bury Hill, Moulton, GR 52/
65.64. D. E. Coombe, 1986, det. P. F. Yeo. 2nd post-1930 record.

435/1/18. EUPHRASIA ARCTICA Lange ex Rostrup subsp. BOREALIS (Towns.) P. F. Yeo iS:
Hants.: Hookheath Farm, Lye Heath, Southwick, GR 41/64.08. Fen meadow. R. P. Bowman,
1985, herb. R.P.B., det. A. J. Silverside. Only extant locality.

440/8. OROBANCHE MINOR Sm. *75, Ayrs.: Ardrossan, GR 26/23.42. Disused railway track.
G. Mitchell, 1987, E.

462/1. LAMIUM AMPLEXICAULE L. 103, Mid Ebudes: Arnabost, GR 17/20.60. Sandpit rubbish
dump. J. W. Clark, 1987, E. 1st Coll record.

466/1. NEPETA CATARIA L. *7, N. Wilts.: Long Dean, GR 31/84.75. Hedgebank. D. Green,
1983. Little Cote Estate, GR 41/29.70. B. Phillips, 1985. Only extant localities.

469/2. SCUTELLARIA MINOR Hudson 67, S. Northumb.: Near Black Cleugh, GR 36/90.03.
Damp stream-bank. G. A. & M. Swan, 1987. Only extant locality.

+470/1. TEUCRIUM CHAMAEDRYS L. *77, Lanarks.: Busby, GR 26/58.56. Roadside wall.
P. Macpherson, 1986, herb. P.M., det. R. M. Harley.

472/1 int. PLANTAGO MAJOR L. subsp. INTERMEDIA (DC.) Arcangeli *73, Kirkcudbrights.: Port
Mary House, GR 25/75.45. Phragmites marsh on shore. O. M. Stewart, 1986, E.

1475/5. CAMPANULA PERSICIFOLIA L. *39, Staffs.: Gibbet Lane, Kinver, GR 32/86.83.
Roadside verge. C. B. Westall, 1985, still present in 1987. 70, Cumberland: By R. Caldew,
Buckabank, GR 35/37.48. Woodland. R. E. Groom, 1985, LANC. Ist post-1930 record.

+t475/pos. CAMPANULA POSCHARSKYANA Degen *46, Cards.: 1.5 km west of Sarnau, GR 22/
30.50. Roadside hedgebank. A. O. Chater, 1984.

+PRATIA ANGULATA Hooker fil. *80, Roxburghs.: Wilton Lodge Park, Hawick, GR 36/49. 14.
Lawn. M. E. Braithwaite, 1972, herb. R. W. M. Corner. Still present in 1987.

485/6. GALIUM PUMILUM Murray 8, S. Wilts.: Slay Down, Larkhill Artillery Ranges, GR 41/
09.49. Chalk grassland. R. D. Porley, 1986, det. M. J. Wiggington. 2nd record. *12,_N.
Hants.: Ladle Hill, GR 41/47.56. F. Rose, 1987, herb. Lady A. Brewis.

+492/1. LEYCESTERIA FORMOSA Wallich *99 Dunbarton: Rhu Point, GR 26/26.83. Roadside
verge. A. Rutherford, 1966.

494/2. VALERIANELLA CARINATA Lois. *77, Lanarks.: Bellahouston, Glasgow, GR 26/55.63.
Abundant nursery weed. P. Macpherson, 1987, herb. P. M.

494/5. VALERIANELLA DENTATA (L.) Poll. 64, Mid-W. Yorks.: Newthorpe, GR 44/45.32.

PLANT RECORDS 193

Edge of pea field. Leeds Naturalists Club Excursion, 1987, det. P. P. Abbott, 2nd localized post-
1930 record.

4503/1. GALINSOGA PARVIFLORA Cav. 4, N. Devon: Braunton, GR 21/46.35. Arable field.
W. H. Tucker, 1987. 2nd record.

$507/2. DORONICUM PLANTAGINEUM L. 80, Roxburghs.: Abbotsford, GR 36/50.33. Wooded
river-banks. M. E. Braithwaite, 1987, herb. M.E.B. 1st record since 1873. Doorpool Loch, GR
36/61.10. Woodland edge. R. W. M. Corner, 1987, herb. R.W.M.C. 2nd extant locality.

+509/2. PETASITES ALBUS (L.) Gaertner *67, S. Northumb.: Swin Burn near Swinburne
Castle, GR 35/93.75. Wooded stream-bank. G. A. Swan, 1985, herb. G.A.S. Extensively
naturalized.

4512/1. INULA HELENIUM L. 77, Lanarks.: Barncluith, GR 26/73.54. Waste ground. A.McG.
Stirling & A. Rutherford, 1986. 2nd record.

517/1. ANTENNARIA DIOICA (L.) Gaertner 33, E. Gloucs.: Cleeve Hill, GR 32/99.26. One
patch in short turf. D. S. Dudley-Smith & S. C. Holland, 1987. Only extant locality.

4519/8. ASTER LANCEOLATUS Willd. *67, S. Northumb.: R. Wansbeck W. of Morpeth, GR
45/18.86. River-bank. G. A. Swan, 1984, herb. G.A.S. *70, Cumberland: R. Duddon, Duddon
Bridge, GR 34/19.88. River-bank. G. Halliday, 1975, LANC. *99, Dunbarton: Near
Helensburgh Upper Station, GR 26/29.83. Embankment. A. McG. Stirling & A. Rutherford,
1985. All records det. P. F. Yeo.

+522/1. CONYZA CANADENSIS (L.) Cronq. 70, Cumberland: Denton Holme Industrial Estate,
Carlisle, GR 35/39.55. Old railway sidings. R. E. Groom, 1987, LANC. 2nd record.

+526/1. ANTHEMIS TINCTORIA L. *73, Kirkcudbrights.: Castle Douglas, GR 25/75.62. Waste
ground. O. M. Stewart, 1987, E.

+533/4. TANACETUM PARTHENIUM (L.) Schultz Bip. 103, Mid Ebudes: The Lodge, Arinagour,
GR 17/21.57. Garden weed. J. W. Clark, 1987. 1st Coll record.

4536/1. ECHINOPS SPHAEROCEPHALUS L. *69, Westmorland: Far Arnside, GR 34/45.76.
Roadside. C. E. Wild, 1986, LANC. *70, Cumberland: Longwath, Moorhouse, GR 35/32.56.
Roadside grassland. R. E. Groom, 1986, LANC.

540/3 < 4. CIRSIUM PALUSTRE (L.) Scop. X C. ARVENSE (L.) Scop. *77, Lanarks.: Dalton, GR
26/67.58. Partly overgrown track. J. H. Dickson, 1987, GL.

554/1. LACTUCA SERRIOLA L. *46, Cards.: Cardigan, GR 22/17.45. Roadside. A. O. Chater.
A. P. Fowles & C. D. Preston, 1987.

554/2. LACTUCA VIROSA L. 81, Berwicks.: Blount Bank, New Ladykirk, GR 36/90.47.
Wooded bank. M. E. Braithwaite, 1987, herb. M.E.B. 2nd extant locality.

558/1/4. HIERACIUM ALPINUM L. *105, W. Ross: An Teallach, GR 29/07.85. Rocky slopes and
detritus. A. G. Kenneth, 1984, herb. D. J. Tennant. *107, E. Sutherland: Ben Hee, GR 29/
42.33. Detritus near summit. A. G. Kenneth, 1986, herb. D. J. Tennant.

558/1/28. HIERACIUM CENTRIPETALE F. J. Hanb. *73, Kirkcudbrights.: Meikle Millyea, GR
25/51.83. Rocks in gully. D. J. McCosh, 1986, E.

558/1/32. HIERACIUM DASYTHRIX (E. F. Linton) Pugs. *73, Kirkcudbrights.: Cairnbaber, GR
25/48.76. Rocks. D. J. McCosh, 1986, herb. D. J. McC., det. P. D. Sell.

558/1/38. HIERACIUM AMPLIATUM (W. R. Linton) A. Ley *73, Kirkcudbrights.: Cairnbaber,
GR 25/48.76. D. J. McCosh, 1986, herb. D. J. McC., det. P. D. Sell.

558/1/40. HIERACIUM LANGWELLENSE F. J. Hanb. *73, Kirkcudbrights.: Cairnbaber, GR 25/
48.76. Rocks by burn. D. J. McCosh, 1986, herb. D.J.McC., det. P. D. Sell.

558/1/46. HIERACIUM STENOPHOLIDIUM (Dahlst.) Omang *69, Westmorland: Longsleddale
above Sadgill, GR 35/48.07. Dry rocks by beck. G. Halliday, 1986, LANC, det. P. D. Sell.

194 PLANT RECORDS

558/1/56. HIERACIUM SUBRUDE (Arv.-Touv.) Arv.-Touv. *103, Mid Ebudes: Totamore, Coll,
GR 17/17.56. Rock cleft. K. A. Cassels, 1986, E, det. A.McG. Stirling.

558/1/67. HIERACIUM SUBPLANIFOLIUM Pugsl. *69, Westmorland: Meathop quarry, GR 34/
43.79. Limestone cliff. J. M. Mullin, 1987, BM, det. P. D. Sell.

588/1/92. HIERACIUM SCOTOSTICTUM Hy]. *70, Cumberland: R. Eden, E. of Low House Farm,
Holmwrangle, GR 35/51.48. By wooded path. G. Halliday, 1985, LANC, det. P. D. Sell.

588/1/94. HIERACIUM DURICEPS F. J. Hanb. *77, Lanarks.: Hillshie Burn, Camps Reservoir,
GR 36/03.21. Rocks by waterfall. D. J. McCosh, 1987, E. :

558/1/99. HIERACIUM GRANDIDENS Dahlst. *77, Lanarks.: Craigend Muir, Glasgow, GR 26/
65.68. Old wall. A. McG. Stirling & J. Mitchell, 1966, herb. A.McG.S. Gartloch, GR 26/67.67.
Old coal bing. A.McG. Stirling & J. H. Dickson, 1985. Ist and 2nd records.

558/1/210. HIERACIUM STRICTIFORME (Zahn) Roffey *69, Westmorland: Stock Gill, Amble-
side, GR 35/3.0. S. H. Haslam, 1843, KDL, det. P. D. Sell.

558/1/atr. HIERACIUM ATRATICEPS (Pugsl.) P. D. Sell & C. West *88, Mid Perth: N. E. Coire,
Ben More, GR 27/44.24. Rock ledge. D. J. Tennant, 1978, herb. D.J.T., conf. P. D. Sell.

558/1/lar. HIERACIUM LARIGENSE (Pugsl.) P. D. Sell & C. West *92,S. Aberdeen: An Garbh
Choire, Cairn Toul, GR 27/95.98. Rocky slope. D. J. Tennant, 1973, herb. D.J.T., conf. P. D.
Sell.

558/1/not. HIERACIUM NOTABILE P. D. Sell & C. West *97, Westerness: Coire Ardair, Creag
Meagaidh, Loch Laggan, GR 27/43.88. Rock ledges. D. J. Tennant, 1979, herb. D.J.T., conf.
F De Sells

558/2/1 mic. PILOSELLA OFFICINARUM F. W. Schultz & Schultz Bip. subsp. MICRADENIA (Naegeli &
Peter) P. D. Sell & C. West 103, Mid Ebudes: Near Arnabost, GR 17/20.60. Grassy outcrops.
J. W. Clark, 1987, det. A.McG. Stirling. 1st Coll record.

+558/2/10 PILOSELLA PRAEALTUM (Vill. ex Gochnat) F. W. Schultz & Schultz Bip. subsp. ARVORUM
(Naegeli & Peter) P. D. Sell & C. West *8, S. Wilts.: Trowbridge, GR 31/85.58. Disused
railway siding. J. Presland, 1982, herb. D. Green, det. P. D. Sell.

+559/2. CREPIS VESICARIA L. subsp. HAENSELERI (Boiss. ex DC.) P. D. Sell *99, Dunbarton:
Forth & Clyde Canal, Kelvindale, GR 26/55.69. Canal tow-path. A.McG. Stirling, 1986, GL.

+570/3. ELODEA NUTTALLI (Planch.) St. John *83, Midlothian: Auchendinny, GR 36/25.61.
Pond. D. R. McKean & A. G. Miller, 1987, E, det. D. R. McK. *99, Dunbarton: Forth &
Clyde Canal between Bowling and Old Kilpatrick, GR 26/46.72. Abundant in canal. BSBI
Meeting, 1987, E, det. C. D. Preston, N. F. Stewart & A.McG. Stirling.

577/13. POTAMOGETON PUSILLUS L. 99, Dunbarton: Net Bay, Loch Lomond N.N.R., GR 26/
43.88. Pool near loch shore. A.McG. Stirling & J. Mitchell, 1987. 2nd record.

581/1. NAJAS FLEXILIS (Willd.) Rostk. & Schmidt 103, Mid Ebudes: Loch Ballyhaugh, Coll,
GR 17/17.58. Mesotrophic loch. A. Walker & H. J. Noltie, 1987, E. 2nd record.

589/2. POLYGONATUM ODORATUM (Miller) Druce +*98, Main Argyll: Achafour, by bridge
across Ardyne Burn, GR 26/10.70. Woodland edge. B. H. Thompson, 1987.

600/1 xX +2. HYACINTHOIDES NON-SCRIPTA (L.) Chouard ex Rothm. xX H. HISPANICA (Miller)
Rothm. *99, Dunbarton: Daligan, GR 26/32.84. Roadside wood. A.McG. Stirling & A.
Rutherford, 1979.

+600/2. HYACINTHOIDES HISPANICA (Miller) Rothm. *77, Lanarks.: Braehead, GR 26/51.67.
Woodland. P. Macpherson & E. Teasdale, 1986, herb. P. M.

+601/arm. MUSCARI ARMENIACUM Leichtlin ex Baker *77, Lanarks.: Between Glasgow and
Renfrew, GR 26/51.67. Trackside bank. P. Macpherson, 1987, herb. P.M., det. E. J. Clement.

605/4. JUNCUS COMPRESSUS Jacq. 11, S. Hants.: Ham Common, Sopley, GR 40/14.98. Short

PLANT RECORDS 195

grassland on flood plain near river. C. Chatters, 1986, conf. R. P. Bowman, 1987. Only extant
locality.

605/fol. JuNcus FoLiosus Desf. *12, N. Hants.: Fleet Pond, GR 41/82.54. Sandy ground at
water’s edge. A. R. G. Mundell, 1981, herb. A. Brewis, det. T. A. Cope. 14, E. Sussex:
Cooden Beach Golf Club, Bexhill, GR 51/70.06. Spring running into ditch. Lady R. FitzGerald &
L. B. Burt, 1987, herb. R. F., det. T. A. Cope. 1st post-1930 record. *46, Cards.: Nant-llwyd,
Soar y Mynydd, GR 22/78.52. Marshy track. A. O. Chater & D. Davies, 1987, NMW, det. C. A.
Stace.

7607/11. ALLIUM PARADOXUM (Bieb.) G. Don *39, Staffs.: Abbots Bromley, GR 43/08.24.
Roadside verge. I. J. Hopkins, 1983. 70, Cumberland: Near Blennerhasset, GR 35/16.40.
Roadside. C. C. Haworth, 1986, LANC. 2nd record.

+607/uni. ALLIUM UNIFOLIUM Kellogg *99, Dunbarton: Behind Cardross Park, GR 26/34.77.
Damp, basic soil in deciduous wood. A.McG. Stirling & A. Rutherford, 1985, det. C. D. Brickell.

+616/sib. IRIS SIBIRICA L. *77, Lanarks.: Ibrox, GR 26/55.64. P. Macpherson, 1986, herb.
P. M., det A. C. Leslie.

+618/2. CROCUS PURPUREUS Weston *77, Lanarks.: Craigton, GR 26/53.64. Old graveyard.
P. Macpherson, 1985, herb. P. M., det. A. C. Leslie.

4618/5. Crocus FLAVUS Weston *70, Cumberland: Deanscales, GR 35/09.26. Roadside
verge. N. Botham, 1987, LANC.

+CURTONUS PANICULATUS (Klatt) N. E. Brown *99, Dunbarton: Ardencaple Castle, GR 26/
28.83. A. Rutherford, 1975.

624/3. CEPHALANTHERA RUBRA (L.) Rich. *12, N. Hants.: K. Turner, 1986, herb. Hampshire
County Museum Service.

625/4. EPIPACTIS LEPTOCHILA (Godfery) Godfery 11, S. Hants.: Hedgemoor Copse, W.
Tytherley, GR 41/26.31. Under beech near edge of wood. V. A. Williams, 1987, conf. R. P.
Bowman, photograph in herb. R.P.B. Only extant locality.

636/1 Xx 643/1. GYMNADENIA CONOPSEA (L.) R.Br. X DAcTYLORHIZA FUCHSII (Druce)
So6 *14, E. Sussex: Coburn Bottom near Lewes, GR 51/43.09. N.-facing chalk downland.
D. C. Lang, 1987. *39, Staffs.: Ecton, GR 43/09.58. N.-facing roadside bank on limestone.
B. R. Fowler, 1987, det. R. H. Roberts.

7648/1. LYSICHITON AMERICANUM Hultén & St John *98, Main Argyil: R. Add at Kilmichael
Glen, GR 16/8.9. Riverside. A. G. Kenneth, 1986. Has spread downstream 6 km from origin at
Kirnan gardens.

7649/2. ARUM ITALICUM Mill. subsp. ITALICUM 46, Cards.: Croeslan, Llandysul, GR 22/38.44.
Roadside hedgebank. G. Harrison, 1986. 2nd record.

+650/min. LEMNA MINUSCULA Herter *11, S. Hants.: The Hatches, Moortown, GR 41/14.03.
Mill stream. G. Halliday, 1984, LANC. R. Test, Horsebridge, GR 41/34.30. Stream. J. A. Moore,
1984. 1st and 2nd records, both det. A. C. Leslie.

654/1. ERIOPHORUM ANGUSTIFOLIUM Honck. 8, S. Wilts.: Pains Bridge, Pewsey, GR 41/16.60.
Water meadows. D. Green & R. Randall, 1983. 2nd extant locality.

654/2. ERIOPHORUM GRACILE Roth 11, S. Hants.: Widden Bottom, Sway, GR 40/28.99.
Valley bog. R. P. Bowman, 1987. 2nd extant locality.

654/4. EERIOPHORUM VAGINATUM L. *29, Cambs.: Ninewells, Thriplow, GR _ 52/4.4.
N. Maynard, 1842, CIECRO, det. D. E. Coombe. Chippenham Fen, GR 52/6.6. A. Shrubbs,
undated but probably c. 1890, CGE. 1st and 2nd records.

655/9. SCIRPUS LACUSTRIS L. subsp. TABERNAEMONTANI (C. C. Gmelin) Syme 7, N. Wilts.:
Cotswold Water Park, Ashton Keynes, GR 41/01.93. Gravel pit. D. Green, 1986, herb. D.G. Only
extant locality.

196 PLANT RECORDS

656/3. ELEOCHARIS QUINQUEFLORA (F. X. Hartmann) Schwarz 7, N. Wilts.: Pike Corner,
Ashton Keynes, GR 41/03.93. Ditch in damp meadow. D. Hornby & D. Wells, 1986, herb.
D. Green. Ist record since 1885. *55b, Rutland: Shacklewell Hollow, GR 43/97.07. Marsh.
D. Drewett, 1987. 1st v.c. 55 record since 1922.

656/6. ELEOCHARIS UNIGLUMIS (Link) Schult. *8,S. Wilts.: Seend Head, GR 31/92.59. Water
meadow. F. Rush, 1985, det. D. Green.

657/1. BLYSMUS COMPRESSUs (L.) Panz. ex Link 81, Berwicks.: Fangrist Burn, GR 36/69.49.
Burn-side. M. E. & P. F. Braithwaite, 1987, herb. M.E.B. 2nd extant locality.

658/1. CYPERUS LONGUS L. +*7,N. Wilts.: Kemble, GR 31/99.97. Streamside marsh. M. A.
R. & C. Kitchen, 1985. Holt, GR 31/86.61. Path. E. Curtis, 1986. 1st and 2nd records.

661/1. CLADIUM MARIScus (L.) Pohl +*7, N. Wilts.: Cotswold Water Park, Somerford
Keynes, GR 41/01.94. Edge of gravel pit. D. Green & S. C. Holland, 1986. 11, S. Hants.:
Gilkicker Point, Gosport, GR 40/60.97. Hollow in shingle behind beach. R. P. Bowman, 1986,
herb. R.P.B. 2nd extant record. *46, Cards.: Near Ciliau Aeron, GR 22/4.5. Fen. A. P.
Fowles, 1987, NMW.

663/4. CAREX HOSTIANA DC. 7, N. Wilts.: Pike Corner, Ashton Keynes, GR 41/03.93. Damp
meadow. D. Green & D. Hornby, 1986. 2nd extant locality.

663/15. CAREX PSEUDOCYPERUS L. *46, Cards.: Near Ciliau Aeron, GR 22/4.5. Fen. A. O.
Chater & A. P. Fowles, 1987.

663/16. CAREX ROSTRATA Stokes 8, S. Wilts.: Jones’ Mill, Pewsey, GR 41/16.61. Water
meadow. D. Green, 1984, DZS. Only extant locality.

663/17 xX 21. CAREX VESICARIA L. X C. ACUTIFORMIS Ehrh. *11, S. Hants.: Ebblake,
Ringwood Forest, GR 41/10.06. Acidic, wet carr. R. P. Bowman, 1986, BM, det. A. O. Chater &
R. W. David. Ist British record.

663/18. CAREX X GRAHAMII Boott *97, Westerness: Beinn na Socaich, Glen Spean, GR 27/
23.74. Wet flush. D. J. Tennant, 1980, BM, det. A. C. Jermy.

663/20. CAREX RIPARIA Curtis +*79, Selkirks: Howlands plantation, Yair Forest, GR 36/
45.32. Dominant in two old ponds. C. O. Badenoch, 1986, herb. R.W.M. Corner.

663/20 X 33. CAREX RIPARIA Curtis X C. LASIOCARPA Ehrh. *29, Cambs.: Grunty Fen, GR
52/4.7. Herb. C. M. Lemann, 1833, CGE, det. E. Nelmes, conf. R. W. David.

663/40. CAREX ORNITHOPODA Willd. *70, Cumberland: By R. Eden S. of Armathwaite, GR
35/50.44. R. W. M. Corner, 1986, LANC.

663/46. CAREX ELATA All. *33, E. Gloucs.: Cerney Wick, GR 41/07.94. Fen. S. C. Holland &
R. W. David, 1987, herb. S.C.H., det R.W.D.

663/47. CAREX ACUTA L. 79, Selkirks.: R. Ettrick, Thirlestane, GR 36/28.15. Fen. C. O.
Badenoch, 1987, herb. R.W.M. Corner. 2nd record.

663/54. CAREX PANICULATA L. 103, Mid Ebudes: Arnabost, GR 17/20.60. Marsh. J. W.
Clark, 1987, E. 1st Coll record.

663/54 x 56. CAREX PANICULATA L. X C. DIANDRA Schrank *98, Main Argyll: Loch Baile
a’Ghobhainn, Lismore, GR 17/86.43. A. McG. Stirling & J. Mitchell, 1984, E, det. A. O. Chater
& R. W. David. 2nd Scottish record.

663/54 x 71. CAREX PANICULATA L. X C. REMOTA L. *39, Staffs.: Sher Brook Valley, GR 33/
98.18. Streamside. D. J. Tinston, 1987, BM, det. A. O. Chater.

663/55. CAREX APPROPINQUATA Schumach. 79, Selkirks.: Back Loch, Alemoor, GR 36/35.15.
Mire. R. W. M. Corner, 1986, herb. R.W.M.C. 2nd record.

663/67. CAREX SPICATA Hudson 77, Lanarks.: Craigton, GR 26/54.64. Grassy area.
P. Macpherson, 1986, det. A. O. Chater. 2nd record.

PLANT RECORDS 197

663/68. CAREX MURICATA L. subsp. LAMPROCARPA Celak. 79, Selkirks.: A7 road by Big Wood,
S. of Selkirk, GR 36/47.25. Foot of roadside cutting. P. F. Braithwaite, 1987. 2nd extant record.

663/69. CAREX ELONGATA L. 25, E. Suffolk: Reydon, GR 62/4.7. Woodland. F. W. Simpson,
1987, det. A. O. Chater. Only extant locality.

669/4. GLYCERIA MAXIMA (Hartm.) Holmberg 81, Berwicks.: Banks of R. Tweed, Fishwick,
GR 36/91.49. M. E. Braithwaite & M. Mutch, 1987, herb. M.E.B. Only extant locality.

+670/dif. FESTUCA DIFFUSA Dumort. 77, Lanarks.: Shieldhall, GR 26/53.65. Waste ground.
P. Macpherson, 1986, herb. P. M., det. P. J. O. Trist. 2nd record.

670/1 X 671/1. FESTUCA PRATENSIS Hudson X LOLIUM PERENNE L. *79_ Selkirks.: East
Buccleuch Farm, Ettrick, GR 36/32.14. Track verge. R. W. M. Corner, 1987, herb. R.W.M.C.

4676/15. PoA CHAIXII Vill. 38, Warks.: Leamington Spa, GR 42/31.65. Canal bank. J. C.
Bowra, 1987, herb. J.C.B., det. P. J. O. Trist. 2nd record. *103, Mid Ebudes: Torosay Castle,
Mull, GR 17/72.35. Edge of pond. J. W. Clark, 1987, E, det. H. J. Noltie.

677/1. CATABROSA AQUATICA (L.) Beauv. 81, Berwicks.: Everett Moss, GR 36/60.43. Muddy
pool. M. E. & P. F. Braithwaite, 1987, herb. M.E.B. 2nd extant locality. 99, Dunbarton:
Hunter’s Burn near Loch Bowie, GR 26/41.75. Muddy stream bank. A.McG. Stirling & 4
Rutherford, 1987, E. Ist record since 1892.

+683/4. BROMUS INERMIS Leyss. 77, Lanarks.: Kelvindale, GR 26/55.69. Roadside. A. J.
Silverside, 1977, still present 1987, herb. P. Macpherson. Ist record since 1920.

683/5. BROMUS STERILIS L. 93, N. Aberdeen: Fingask, GR 38/77.26. Old railway. D. Welch,
1987, ABD. Ist post-1930 record.

1683/7. BROMUS DIANDRUS Roth *46, Cards.: Penyrangor, Aberystwyth, GR 22/58.80.
Rocky slope. A. O. Chater, 1982, NMW, det. P. J. O. Trist.

4683/9. BROMUS TECTORUM L. 11, S. Hants.: Dibden Bay, GR 41/41.09. Sparse turf on
reclaimed land. R. P. Bowman, 1987, herb. R.P.B., det. P. J. O. Trist as var. hirsutus Regel. 1st
post-1930 record.

683/10 x 13. BROMUS HORDEACEUS L. X B. LEPIDUS Holmberg *7, N. Wilts.: Pewsey Down
N.N.R., GR 31/90.64. Downland. K. Payne, 1984, det. P. J. O. Trist.

683/11. BROMUS FERRONII Mabille *93, N. Aberdeen: Aberdour, GR 38/89.65. Sea cliff. D.
Welch, 1987, ABD, det. P. J. O. Trist.

4683/18. BROMUS SECALINUS L. 26, W. Suffolk: Hermitage Farm, Clare, GR 52/77.46. Wheat
field. F. Edmonds, 1987, det. P. J. O. Trist. 2nd post-1930 record.

4683/20. BROMUS CARINATUS Hooker & Arnott *39, Staffs.: Wom Brook near Common
Lane, Wombourne, GR 32/87.92. Bank of brook. C. B. Westall, 1987, herb. B.R. Fowler.

685/3 x 4. ELYMUS REPENS (L.) Gould X E. pYCNANTHUS (Godron) Melderis *11, S. Hants.:
Eling, GR 41/36.12. Edge of creek. A. C. Leslie, 1986, herb. A.C.L.

686/1. LEYMUS ARENARIUS (L.) Hochst. +*38, Warks.: Winson Green, GR 42/04.88. Sandy
waste ground by canal. J. W. Partridge, 1987, WAR, det. P. J. O. Trist.

689/1. KOELERIA MACRANTHA (Ledeb.) Schultes 99, Dunbarton: Kilpatrick Braes, GR 26/
46.73. Dry turf near basalt rock outcrops. A.McG. Stirling, 1987, E. Ist record since 1891.

698/1. CORYNEPHORUS CANESCENS (L.) Beauv. +*39, Staffs.: 1 mile S.W. of Kinver, GR 32/
83.82. Loose sand on S.E.-facing heathy slope. B. R. Fowler, 1977, BM, det. E. J. Clement. Still
present in 1986.

700/2. CALAMAGROSTIS CANESCENS (Weber) Roth *72, Dumfriess.: Old course of Kinnel
Water, Lochmaben, GR 35/09.83. Winter-flooded silty sand. E. Kungu & M. E. R. Martin, 1987,
K, det. T. A. Cope.

198 PLANT RECORDS

702/1. APERA SPICA-VENTI (L.) Beauv. 11, S. Hants.: Fryern Court, Fordingbridge, GR 41/
14.16. Border between fields. J. Ounsted, 1987. Only extant locality.

702/2. APERA INTERRUPTA (L.) BEAUV. *39, Staffs.: Bilston, GR 32/94.96. Disused railway
track. B. R. Fowler, 1976, herb. B.R.F., det. J. P. M. Brenan. Pensnett, GR 32/91.88. Spoilheap.
I. C. Trueman, 1986. 1st and 2nd records.

703/1. POLYPOGON MONSPELIENSIS (L.) Desf. 725, E. Suffolk: Beccles, GR 62/42.91. Field
entrance. T. Abrehart, 1987, det. E. Beaumont, G. W. Maybury & P. J. O. Trist, 2nd record.

+713/aqu. PHALARIS AQUATICA L. *7, N. Wilts.: Manor Farm, Rockley, GR 41/72.15.
Arable headland. M. Ponting, 1986, DZS, det. R. Payne.

Watsonia, 17, 199-207 (1988) 199

Book Reviews

Jupiter Botanicus. Robert Brown of the British Museum. D. J. Mabberley. Pp. 500, with 24 colour
plates and 64 text figures. J. Cramer, Braunschweig. 1985. Price £43.00 (ISBN 3—7682-1408-7).

It may seem strange to be reviewing a book three years after its emergence from the press — and
five even after the date of its preface. The reason for doing so in this case is that publication was
attended by a quite exceptional piece of misfortune, as a result of which scarcely any copies were
sent out at the time for review. That was not a fair measure of the volume’s value and importance,
and belatedly steps have been taken to repair the omission.

At the same time it is an open secret that the manuscript was a long time finding a publisher in
the first place. And one can see why that was: to judge from the finished text and from hints
dropped in the acknowledgements, it bore all too much resemblance to too many doctoral theses,
in that so much effort was expended on the research that seemingly little was left over for
producing a narrative that was easy, let alone enticing, to read. Biography is a particularly well-
established genre, and it was surely courting danger not to make more attempt to meet the
expectations that the public (or at any rate publishers) have of titles in that category.

Which is a pity, because Robert Brown is certainly a figure of the first rank in botanical history
(whether or not one accepts the author’s claim that he was “‘Britain’s greatest botanist’); a full-
scale study of his life and work has long been needed, and Dr Mabberley has fulfilled the scholarly
side of his task with exemplary care and thoroughness. The problem seems to have been that he fell
in love with his material and could not bring himself to discard enough of it to produce an account
of the necessary selectiveness. Just about everything one could possibly want to know about Brown
and his times is consequently to be found in his pages. It is indeed, as he terms it himself, a
“source-book’’, and one that will be consulted, and with confidence, by generations of historians
and taxonomists to come.

Much of the volume is necessarily devoted to Brown’s work on the Australian flora and his
pioneering researches in a variety of botanical fields apart from taxonomy. There is necessarily
much, too, on his years successively in the employ of the Linnean Society (1805-22), Sir Joseph
Banks (1810-27) and the British Museum, where he was responsible for heading a new botanical
department from 1827 till his death in 1858 in his 85th year. B.S.B.I. members, however, are likely
to find the two opening chapters of more particular interest, for in these Brown’s little-known
contributions to field botany in these islands are described.

It was while at Edinburgh University, as a 17-year-old medical student, that Brown apparently
embarked upon botany seriously, starting a herbarium and exploring his native county of Angus in
the company of the more experienced George Don. In 1791 their partnership was speedily
rewarded with a singular piece of beginner’s luck: the addition of Scirpus hudsonianus to the
British flora in its only-ever-known station, the Moss of Restenneth. Three years after that, Brown
on his own found a new grass, on which Smith was to bestow the name Alopecurus alpinus (though
at first he mistakenly gave the credit for its discovery to Don). As an ensign in the Fifeshire
Fencibles Brown then found himself posted to Ulster. There he was the first to recognise Sagina
maritima as a distinct species and to collect several mosses that are now known to have been new to
science. This work on bryophytes brought him into touch with James Dickson, the fellow Scot and
Covent Garden nurseryman then compiling the fourth of his cryptogamic Fasciculi, and to that
Brown became a major, if unacknowledged, contributor.

A special word of praise must be reserved for the series of superb watercolours by the Bauers
reproduced in a special section at the end of the book. Over half of these, mainly of Australian
plants and animals, are enjoying here their public début, after years of lying hidden away in the
magnificent iconographical collections of the British Museum (Natural History).

D. E. ALLEN

200 BOOK REVIEWS

The National Trust book of wild flower gardening. John Stevens. Pp. 192. Profusely illustrated with
colour photographs, additional colour drawings and b. & w. marginal thumb-nail sketches. Dorling
Kindersley Publishers Ltd., London, in association with the National Trust. 1987. Price £14.95
(ISBN 0-86318-219-4).

The concept of growing wild flowers in gardens has dramatically increased in popularity in the last
decade, with a corresponding public awareness of the loss of wild flowers from native habitats. This
book aims at encouraging the creation of a piece of countryside in urban and garden settings, as
part compensation for the fast-disappearing countryside in the wild.

The attractively produced book has good colour photographs of more than 100 growing plants
featured in the “Creating wild flower gardens” section. These are arranged for six types of garden:
sunny, shady, semi-shady, water, rock and seaside, each species with photograph, description and
cultivation instructions. The grouping is supplemented by eleven double spreads of wildflower
snippets photographed on a parchment-coioured background.

A 30-page section entitled ‘“Technique of wildflower gardening” includes notes on seed
dispersal, collection and sowing of the seeds, general planning, cultivation and looking after the
wildflower garden. A further 30-page section, entitled ‘“‘Wildflower catalogue and useful
information’, lists plants for gardens of different aspects with cultivation notes for 156 of these. A
section on useful addresses (B.S.B.I. not included) and an 8-page index complete the book. 3¢
National Trust Wildflower Gardens are listed, actively promoted by the Trust as part of its
conservation role. John Sales, Chief Gardens Adviser, states in his foreword that the wildflower
areas also enhance the overall design of these gardens.

This book should entice even tidy gardeners to consider wildflower cultivation. John Stevens was
a pioneer in growing wild flowers for seed to meet the current need to supply this “‘quiet revolution
taking place in our gardens’’, and he offers valuable practical suggestions on their cultivation from
his experience. Growing plants increases our knowledge, and it is the author’s hope that to grow
wild flowers will also increase the gardener’s appreciation of these in the wild.

M. Briccs

A check-list of mycorrhiza in the British flora. J. L. & E. L. Harley. New Phytologist supplement
105 (2). Pp. 102. Available from the Executive Editor, The New Phytologist, Department of
Botany, The University, Sheffield S10 2TN. Price £5.00.

Contrary to the assumptions of most botanists (and many mycologists), mycorrhizal associations
between plants and fungi are very widespread. In the British flora, there are few species which
have not been reported to have some sort of association with fungi, apart from annual plants which
presumably have too short a life cycle for the association to be feasible.

The work includes an introduction which details the various types of mycorrhizal association,
and in very basic terms the sorts of fungi that are involved, the check-list itself, with some notes on
its organization, and no less than 723 references.

The list is arranged according to the Excursion Flora of the British Isles (3rd ed., 1981).
Information on the type of mycorrhizal association, references to literature, and to the Biological
Flora of the British Isles (Journal of Ecology (1941), et seq.), and a section of miscellaneous notes,
are included. A distinction is made between studies on British material, and European studies on
Species occurring in Britain.

As the compiler of another large check-list, 1 am well aware that the amount of effort expended
on such a project is out of all proportion to the number of printed pages produced. The authors
should be congratulated on producing such an extensive and informative list. However, I have a
few minor criticisms, which should not be allowed to detract from the favourable impression
gained of the work as a whole.

The list would have been more compact if a tabular format had not been chosen. Plants whose
associations have not been investigated are tabulated to no real advantage, though the blank
columns may encourage other workers to investigate these species. References to the Biological

BOOK REVIEWS 201

Flora could usefully have been incorporated into the general reference column. More detail might
have been given of the fungal component of the mycorrhiza, though I am aware that in many cases
it is difficult to be sure of the identity of the symbiotic partner.

This check-list should be of interest to botanists and mycologists alike, and will be as useful to
workers in other parts of northern and western Europe as to those from Britain. At the very
modest price quoted, it can be warmly recommended.

P. F. CANNON

Checklist of European pteridophytes. L. N. Derrick, A. C. Jermy & A. M. Paul. Sommerfeltia, 6:
i-xx, 1-94 (1987). Botanical Garden and Museum, University of Oslo, Trondheimsveien 23B, N—
0562 Oslo 5, Norway. Price NOK 70.00 (ISBN 82-—74-002-0; ISSN 0800-6865). Obtainable from J.
W. Dyce, Hilltop, 46 Sedley Rise, Loughton, Essex IG10 1LT.

This list is the first major updated output to be produced directly from the database of the
European Science Foundation’s Taxonomic, Floristic and Biosystematic Documentation System.
It is essentially an updating, amplification and correction of the relevant data from Flora Europaea,
Vol. 1 (T. G. Tutin et al., 1964). Whereas the latter contained 244 accepted pteridophyte taxa, the
present list contains 329, the increase being to a considerable extent due to the inclusion of more
hybrids. The synonymy is vastly more extensive than that in Flora Europaea, and a total of 1259
synonyms are listed, all with their places of publication. Country by country distributions are given
for all taxa at and below the rank of species, as well as a brief indication of world distribution. The
arrangement of families, genera, species, subspecies (although the typical one comes first) and
hybrids is alphabetical, and the system of Lellinger is followed for the circumscription of families.
Only one variety (Athyrium distentifolium var. flexile) is recognised, but two new varietal
combinations made in the introduction are surprisingly not included in the list. There seems to be a
high degree of accuracy in the citations, and the nomenclature has clearly been very thoroughly
checked. In a rare misprint, Asplenium scolopendrium subsp. anti-jovis is given for Britain instead
of Bulgaria. So far as British and Irish taxa are concerned, apart from hybrids, the main changes
from the last authoritative account, C. N. Page, The Ferns of Britain and Ireland (1982), are that
Ceterach and Phyllitis are again sunk into Asplenium, the plant we have been calling Dryopteris
affinis subsp. stilluppensis has become subsp. cambrensis Fraser-Jenkins (and subsp. robusta is
sunk into subsp. borreri), and Polypodium australe Fée has again become P. cambricum L. There
are six pages of valuable taxonomic and nomenclatural notes in the introduction.

The list is printed from camera-ready copy taken directly from the database. This has resulted in
a layout and typography that are confusing and difficult to follow. Apologies are made for the lack
of any diacritical signs (which of course makes the list difficult to quote from unless one is in a
position to check from source what accents are required). Equally unfortunate, though, is the lack
of any variation in type-face except for upper and lower case, and differentiation between accepted
names and synonyms has to rely entirely on a rather poor system of indentation. The plant names
are written to include the authors not only of the names but of the books in which they were
published (if different), e.g. CRYPTOGRAMMA CRISPA (L.) R.BR. INHOOKER - Gen. Fil.
t. 115 B (1842). It is much to be hoped that future publications of this sort will not continue to
offset the advantages of quick and easy updating by using such unhelpful and inaccurate
typography. The synonymy is also not as user-friendly as it might be; and although some synonyms
are explained away, others, for example Aspidium distans Viv. (1825), under Dryopteris affinis
(Lowe) Fraser-Jenkins, based on Nephrodium affine Lowe (1838), are not, and the reader would
have to repeat the work which the authors of the list have presumably already done to discover why
the earliest epithet, distans Viv., is not the one to be adopted.

This list will immediately become the standard checklist for Europe and will be of great interest
even for those only interested in the 110 or so British and Irish taxa that it covers. With its detailed
synonymy and thorough checking it should also help to stabilise the notoriously unstable
nomenclature of this group of plants.

A. O. CHATER

202 BOOK REVIEWS

Iconographia palynologica Pteridophytorum Italiae. E. Ferrarini, J, Ciampolini, R. E. G. Pichi
Sermolli & D. Marchetti. Extracted from Webbia 40 (1). Pp. 202, with 71 b. & w. plates. Firenze.
1986. Price not stated (ISSN 0083-7792).

This large and useful volume contains much valuable information. All Italian taxa are included,
with their spores depicted as 550 micrographs. The description of the spores is greatly aided by the
provision of a very useful illustrated and cross-referenced glossary of the terms used within.
Chromosome numbers, genomic formula, notes on reproduction and the total geographic range
(with distribution maps cited) are given. Note that the spore sizes given are derived from spores
observed in air, and also that the photographs, although generally good, are like my own SEMs —
sometimes a little less than sharp.

A key is given to the genera, species and subspecies of all Italian Pteridophytes, based on their
spore characters. This will prove valuable to many, including palaeobotanists.

This collaborative work by four authors has been competently co-ordinated by Pichi Sermolli in
his usual thorough-going manner. The work is a precursor to his book on Italian ferns, which is in
preparation now.

P. J. EDWARDS

The Brightest Jewel. A history of the National Botanical Gardens, Glasnevin, Dublin. E. C. Nelson
& E. M. McCracken with original watercolours by Wendy F. Walsh. Pp. 275, with 1 colour
photograph, 15 colour plates and 150 b. & w. illustrations. Boethius Press, Kilkenny, Ireland.
1987. Price £26.00 (ISBN 0-86314—083-1).

The National Botanic Gardens at Glasnevin have a distinguished record in the history of
horticulture world wide. Established in 1795 they were at one time the second largest botanic
garden in the British Empire. This volume traces the Gardens from their formation through the
peak of their international fame in the 19th century up to the present day.

From 1838 David Moore and later his son Frederick, who retired in 1922, established contacts at
home and abroad and built up rich and extensive collections of plants. One of their particular
enthusiasms was tropical orchids, but Glasnevin is also famous for raising many new cultivars of
Nerine, Lachenalia and a host of other plants.

The work not only represents the definitive history of Glasnevin but also provides considerable
insight into the history of botanic gardens, plant collecting and horticulture of the 19th and 20th
century. A particularly noteworthy feature is the pairing of early illustrations and photographs with
recent counterparts. How little the Yew Walk has changed in 80 years is remarkable.

The planning and construction of the Curvilinear Range, contemporary with the Palm House at
Kew, shows that the Victorian period suffered its share of problems and frustrations.

The work is admirably researched and written in a most readable and enjoyable style.

I. K. FERGUSON

The Euphorbiales. Chemistry, taxonomy & economic botany. Edited by S. L. Jury, T. Reynolds,
D. F. Cutler & F. J. Evans. Pp. iv + 326, with black & white illustrations. Academic Press,
London. 1987. Price £15 (ISBN 0—12-3924804) softback.

This volume is the published proceedings of a joint symposium organised by the Linnean Society of
London and the Phytochemical Society of Europe and has been reprinted from Bot. J. Linn. Soc.,
94: 1-326 (1987). Of the 17 papers, ten are on various aspects of chemistry, six are taxonomic in
some way and one deals with ethnobotany. Of the taxonomic papers, B.S.B.I. members will
probably be most interested in the review of the classification and relationships of the Euphorbiales
by G. L. Webster, and the discussion by A. Radcliffe-Smith of families segregated from the

BOOK REVIEWS 203

Euphorbiaceae. Surprisingly for a symposium of this nature, there is only one paper which applies
chemical evidence directly to the solution of a taxonomic problem — and this deals with Euphorbia
esula and its relatives in North America. Many of the other chemical papers discuss aspects of the
carcinogenic properties of diterpene esters and their derivatives, compounds widespread in the
Euphorbiaceae. Accepting the pronounced chemical bias to the symposium, this volume is well-
written and well-presented. I doubt, however, whether it contains sufficient of interest to B.S.B.I.
members to make it a good buy for them.

R. J. GORNALL

Coevolution and systematics. Edited by A. R. Stone & D. L. Hawksworth. The Systematics
Association Special Volume No. 32. Pp. xii + 147, with 30 text figures. Oxford University Press,
Oxford. 1986. Price £22.50 (ISBN 0—19-857703-6).

All evolution is coevolution to some extent, there being a continuous spectrum from the most
tenuous to the most intimate of coevolutionary interactions. It is with the extreme of the latter, the
relationship between parasite and host, characterized by reciprocal evolutionary change in interac-
ting species involving partial coordination of non-mixing gene pools, that this volume is concerned.
The papers are largely addressed to the question of the extent to which ‘Fahrenholtz’s rule’ and its
corollaries, which essentially state that parasites and hosts have congruent phylogenies, really do
apply in practice. The problem is considered both generally and in relation to special groups of hosts,
such as Nothofagus and Australian marsupials, and parasites, such as aphids and lice.

The overall conclusion is that Fahrenholtz’s and similar rules are invalid as generalities and may
produce erroneous hypotheses of relationship; groups of parasites associated with a holophyletic
group of hosts may or may not be holophyletic, as the case may be. The conclusions reached are
pertinent ta the consideration of other intimate and biologically significant coevolutionary
phenomena, such as endosymbiosis, mycorrhiza, lichenization and gut biota. Hypotheses of pattern
should always be developed independently of those of process.

C. JEFFREY

The flowering plants and ferns of North Lancashire. L. A. & P. D. Livermore. Pp. iv+148, with 735
distribution maps. L. A. & P. D. Livermore, 8 Durham Avenue, Lancaster, LA1 4ED. 1987. Price
£4.95 + £1.00 postage & packing (ISBN 0—9512644—0-0).

This publication covers the northern part of Watsonian vice-county 60, West Lancashire, specifically
the administrative area of Lancaster District Council. To the best of my knowledge no account of the
flora of this area, or of the vice-county as a whole, has been published since Wheldon & Wilson’s The
flora of West Lancashire appeared in 1907 (reprinted 1978). The recent publication is a praiseworthy
attempt to fill the gap and may arguably be considered a model for other district floras.

The survey area covers 220 square miles (c.570 km”), involving some 180 tetrads, with fieldwork in
the main carried out during the last six years. The area of the botanical survey is considered in terms
of Wheldon & Wilson’s Flora, but no attempt is made to give a detailed account of the geography and
geology (on cost grounds); rather the reader is referred to the earlier work.

A short account compares the changes in flora since 1907, and the mapping system by the standard
tetrad method is described. There follows a brief synopsis of the area’s habitat types with their
associated flora. This is a very rich district ranging from the Lancashire coast to the Pennine
moorland, and from acidic mosslands to the Morecambe Bay limestone.

The main list of the flora includes hybrids as well as plants of garden or similar origin and is
followed by 732 distribution maps. Critical groups are well covered with the exception of Hieracia,
though, given the difficulty in obtaining identifications of this genus, this is hardly a criticism of the
authors. However, although the referees for the more critical groups are acknowledged, no

204 BOOK REVIEWS

reference is made to which of the records they identified. Neither is there any mention of where the
voucher specimens are housed. Inevitably in any work such as this there are odd omissions,
Eriophorum latifolium for one; and is Epilobium tetragonum quite so common?

Yet, one can only praise this publication and the hard work that has so obviously gone into its
production. It provides a valuable account of the area’s flora and will undoubtedly become a
standard datum from which to assess the floristic changes in future years. It is a must for field
botanists in that part of England or anyone intending to spend any time in this rewarding part of
the country. The authors’ attempt to keep the price so low is commendable, but I for one would
prefer to pay more and have something a little more robustly bound.

P. JEPSON

Biosystematics in the Nordic Flora. Edited by Bengt & Lena Jonsell. Acta Universitatis
Upsaliensis/Symbolae Botanicae Upsaliensis 27 (2). Pp. 256. Distributor: Almqvist & Wiksell
International, Stockholm — New York [1986]. 1987. Price SEK 160 (ISBN 91-554-1941-0).

This is an account of the Proceedings of a symposium held at the Royal Swedish Academy of
Sciences on the occasion of the centenary of the Bergius Botanic Garden, Stockholm University,
Sweden, on August 27th-29th, 1985. The symposium brought together a large gathering of
botanists, many of whom were active in biosystematic studies in the Nordic flora, and the 25 papers
presented cover a wide range of topics, including much to interest British and Irish botanists.
There are excellent accounts by two prominent English taxonomists, Professor Clive Stace on
“Hybridization and plant taxonomy” and Dr Max Walters on “‘Alchemilla: a challenge to the
biosystematists”, while critical groups are well represented with studies on the Campanula
rotundifolia complex, the Carex flava group, Cochlearia and Euphrasia. Research on the
Gramineae is covered by papers on the genera Anthoxanthum, Deschampsia and Festuca. ‘‘Man’s
influence on the establishment and composition of the national flora” by M. Ryberg. provides an
account of the role of man as a dispersal agent in the Swedish flora and of the effect of human
influence on natural and semi-natural plant communities; it contains much data relevant to these
islands. A further paper by R. Svensson & M. Wigren, entitled ‘A changing flora — a matter of
human concern’, describes the decline of many cornfield weeds and the changing pattern of village
ruderals due to modern grain-cleaning methods, improved manure management and better
husbandry. This is a useful publication, similar in many ways to some of the B.S.B.I. Conference
Reports. It should be acquired by all those with an interest in the flora of northern Europe.

D. H. KENT

Planting a Bible garden. F. N. Hepper. Pp. vii + 160, with 73 line drawings and 41 colour photos.
Her Majesty’s Stationery Office, London. 1987. £6.95 paper-back. (ISBN 0—11—250011-0).

Nigel Hepper has a close interest in this subject — he is the author of HMSO’s Bible plants at Kew
and has himself laid out Bible gardens. The result is, for those inclined that way, all that could be
wished for — discussions on the frequent extreme uncertainties of just what plant is intended by the
various references and translations, and how to grow them or substitutes. For many are impossible
out of doors outside the Holy Land, and some even there.

Personally, I would find such an odd assortment of plants unsatisfactory, above all when the best
that can be done is quite often to have a similar, or not so similar, species — ideas towards this are
listed under each sort described, e.g. the 1867 yellow British hybrid Cytisus X praecox for the white
Retama, or Iris pseudacorus for Acorus. Mr Hepper makes the best of these difficulties, and one
wonders why (so far as I know) only two churchyards include any of these as such, as many are easy
enough to grow. The book is a tour de force by him, all the drawings, all the photos and all the text
being his.

D. McCLINTOCK

BOOK REVIEWS 205

The complete book of British berries. D. C. Lang. Pp. 223, with 108 colour photographs and 103
line drawings. Threshold Books, London. 1987. Price £19.95 (ISBN 0-901366-34-X).

In this book, David Lang confines his attention to those fruits which can loosely be called berries.
The author’s claim to deal with all the true fleshy fruits found as natives or widespread
introductions in Britain is fully justified, with 96 species covered, including some, such as Fuchsia
magellanica, of which fruit illustrations are rare.

The introductory text has chapters explaining about the different kinds of fleshy fruits and where
to find them, plant poisons and how to treat them, a species list entitled ‘“Can we eat them’’, and a
second list of the species covered in the book with known or potentially poisonous ones marked.
There are a number of minor but alarming discrepancies here. Depending on which list is
consulted, Rowan for example, is either good to eat or potentially poisonous, a distinction of some
importance to the eater. Nor is the text any more reassuring, concluding with the statement that
“the process of boiling involved in making jelly or jam could well destroy these (poisonous)
properties”. Only ‘“‘could well’? I think I will stick to strawberry jam!

At the end of the book are a rather serendipitous glossary, an extensive bibliography and a
usefully cross-referenced index. The middle of the book is devoted to descriptions of the species,
arranged in the taxonomic order used in Flora Europaea (not the one most familiar to British
botanists). In addition to the text, which describes leaves and flowers as well as fruits, and includes
notes on ecology and distribution, each entry is accompanied by a colour photograph of the fruit
and a line drawing, usually of a flowering stem, flower or fruit. The photographs are excellent,
sharply focused, and with true colours, and are the best feature of the book. The line drawings are
rather crude, adding little to the book. For comments on the history and uses of the various fruits
the author relies heavily on those oft-quoted herbalists, Culpeper and Gerard. I found Mr Lang’s
own comments on the toxicology, symptoms and treatment of the poisonous species of more
interest.

This particular book falls into an odd category, more than a coffee table glossy, less than a field
guide. The author conceived his book as an aid for those with ‘“‘minimal botanical knowledge to
identify plants in fruit’. The lack of any kind of key to the species is a major drawback, and I fear
Mr Lang’s intended readers may find using his book for this purpose a frustrating exercise.

J. R. PREss

The plant-book. A portable dictionary of the higher plants. D. J. Mabberley. Pp. xii+706.
Cambridge University Press, Cambridge. 1987. Price £20.00 (ISBN 0—521—34060-8).

When Willis’s Dictionary of the flowering plants and ferns was revised and enlarged by H. K. Airy
Shaw for the 7th edition (1966), he confined the entries to generic and family names and those of
higher ranks, omitting much that had been treated by Willis himself in the earlier editions (1896—
1931). The missing parts (common names, uses, etc.) were subsequently included by F. N. Howes
in A dictionary of useful and everyday plants (1974), whilst Shaw’s 8th edition of the Dictionary
proper (1973) was in the same format as the 7th. Many botanists, however, while appreciating the
increased and more up-to-date coverage of the works of Shaw and Howes, lamented the absence of
a single-volume comprehensive small-size dictionary; and it is this lacuna that David Mabberley
has attempted to fill.

Mabberley’s book is certainly small (23 x 12.5 x 4 cm); and it is more comprehensive than
Shaw’s in that it contains a large number of common names from many regions of the world and a
considerable amount of economic information in addition to the purely taxonomic and
distributional data. It is not — nor could it be — completely comprehensive. The less usual botanical
synonyms, for example, have been omitted; and I failed to find one or two modern common
names, e.g. Gallant Soldiers (Galinsoga parviflora). On the whole, however, I should regard the
use of the word ‘comprehensive’ as well justified; and the wide coverage of literature has made it
reasonably up-to-date. Shaw’s volumes include summaries of the classifications of both Bentham &
Hooker and Engler & Prantl; but the text is full of his own ideas of family and generic

206 BOOK REVIEWS

relationships, only some of which have gained acceptance. Mabberley has ‘“‘tried not to innovate”’
and has adopted Cronquist’s system (1981) as standard; and it is that system that is summarized at
the end of this volume. His own views, therefore, are seldom revealed; but they do sometimes
appear, e.g. when he uses the sign ‘~’ (‘‘sometimes included in, has recently been included in,
should be included in or is very close to’’) before a generic name. The references to recent or
standard monographs or revisions, however, are a very welcome innovation and do compensate, to
some extent, for the absence of the subfamilial and tribal positions of the genera that have been a
feature of ‘Willis’. Was it necessary, though, to include a nom. nov. (p. 371) for a genus of orchids
in such a work?

The format, in general, is acceptable and helpful. Would that it were always possible, however,
to distinguish which author’s names are abbreviated. This can be done only for those of generic
synonyms, not for those whose generic names are accepted. All in all, however, this really is a
book that should be on the bookshelf of every botanist, not to mention other biologists who are
concerned with vascular plants.

N. K. B. RoBson

The origins of angiosperms and their biological consequences. E. M. Friis, W. G. Chaloner & P. R.
Crane. Pp. x+358, with 44 text figures and 15 tables. Cambridge University Press, Cambridge.
1987. Price £27.50 (ISBN 0—521-—32357-6).

The contents of this fascinating and informative book were mostly presented as papers at the Third
International Congress of Systematic and Evolutionary Biology at the University of Sussex in
August 1985. They provide, overall, a clear and reasonably comprehensive picture of the current
‘orthodox’ view of the origin(s) of the angiosperms, although references are made by some
contributors to the divergent opinions of e.g. Meeuse, Axelrod and Burger (but not to those of
Melville).

An ‘Introduction to angiosperms’ by the editors provides a very useful summary of the facts and
opinions about their nature and evolutionary history, after which Doyle & Donoghue summarise
previous ideas about their relationships and give a detailed account of a cladistic analysis of the
groups concerned. From this it would appear that the angiosperms are the sister-group of the
Bennettitales + Pentoxylales + Gnetales, and that all these taxa form the sister-group of the
Caytoniales, which in turn links them to the Glossopteridales (a group of seed ferns). Parrish’s
discussion of Late Cretaceous and Early Tertiary geography and climate provides a helpful
framework into which the data of the other authors can be placed and indicates that large areas of
the earth were semi-arid (with steppe, taiga or Mediterranean climates) when the angiosperms
were apparently undergoing their major evolutionary divergence.

Of the other contributions, the chapter by Friis & Crepet on “Time of appearance of floral
features’ will be of great interest to many Watsonia readers, whilst Collinson & Hooker’s
description of the ‘Early Tertiary of southern England’ reveals a very different vegetation from
that encountered now on B.S.B.I. excursions, e.g. reedswamp with palms.

In general, I can thoroughly recommend this book for those who want to know the current
orthodox state of knowledge and opinion on the angiosperms, i.e. that they are monophyletic,
arose in or soon before the Early Cretaceous, were probably Magnolioid in form, had pinnate
venation in leaves and ‘carpels’, etc. Apart from Doyle & Donoghue’s preliminary discussion, you
will not find much about the views of those who disagree with this picture, although it is clear that
the simple flower of e.g. Chloranthaceae appeared very early in the evolutionary history of the
angiosperms. Would that palaeontologists took seriously Melville’s data on dichotomous venation
and gonophylls in the angiosperms and their relatives.

N. K. B. ROBSON

BOOK REVIEWS 207

Pasture-woodlands in lowland Britain. P. T. Harding & F. Rose. Pp. 89, with 8 colour plates.
Institute of Terrestrial Ecology, Monks Wood, Huntingdon. 1986. Price £5.00 plus £1.10 p. & p.
from Publications Section, I.T.E., Merlewood Research Station, Grange over Sands, Cumbria
LA11 6JU (ISBN 0—904282-91-0).

This attractively produced book is sub-titled “A review of their importance for wildlife
conservation’’. Many botanists will no doubt be familiar with Francis Rose’s work on the lichen
epiphytes of old trees, but the importance of ancient trees with dying and dead wood for saproxylic
invertebrates (particularly beetles) is perhaps less well-known.

The authors indicate that pasture-woodlands are better represented in lowland Britain than in
most of north-western Europe, with the New Forest being the most important of all.
Recommendations for their conservation and future research are given.

The numbers of lichens and insects associated with each tree are given. Oak is the best for both
groups (326 lichens, 423 insects and mites). It is of interest that sycamore is high up the list with 194
lichens — not far behind ash, beech and elms in this respect.

An important feature of the book is the tabulated schedule of all pasture woodlands with an
assessment of the conservation importance of each. There is also an extensive list of references.

R. C. STERN

208 Watsonia, 17, 208-216 (1988)
Obituaries

DOROTHY ADLINGTON CADBURY
3 (1892—1987)

Dorothy Cadbury was born on 14th October, 1892 and died on 21st August, 1987 at the age of 94.
Through much of her life she worked in Cadbury Bros. at Bournville and was a director for many
years, retiring from the firm in 1952. She was a Quaker and throughout her life maintained a strong
commitment to the Society of Friends.

Dorothy always said that her botanical interests started at an early age. As a child she had helped
her mother to find about 1,000 flowers, which were carefully coloured in her J/lustrations to the
Bentham & Hooker Flora of the British Isles. This activity was continued into her old age, and
nothing gave her more pleasure than finding another species which she could ‘paint into’ her book
of Illustrations.

She joined the Wild Flower Society in 1937, and each year sent in records beginning in the first
week in March or sometimes even earlier. She was Warwickshire Recorder for that Society for a
time and continued to send in records up to 1986. In 1939 she joined a group carrying out a
botanical survey of the Isle of Mull. Dorothy joined the B.S.B.I. in 1936 and remained a member
for the rest of her life. She took a lively interest in the Society’s meetings, particularly in the
November Conversazione.

From quite early in her career Dorothy Cadbury began to take a special interest in water plants,
though it was apparently by accident that she became a ‘Potamogetonist’. Apparently, she had sent
some pond weeds to a Wild Flower Society friend for name checking, and the friend had taken
them to the British Museum to be identified. The identifiers were Dandy and Taylor (later, Sir
George Taylor, who became the Director of the Royal Botanic Gardens, Kew). Sir George had a
great admiration for Dorothy’s Potamogeton work, and she was most gratified to be asked to
collect pondweeds officially for the British Museum.

During field excursions in Warwickshire, and indeed everywhere she went, we all knew that the
merest hint of an interesting pondweed would send her rushing into pond, canal or river fully
clothed and shod, without any hesitation, to collect her specimens.

One of the interesting outcomes of this work was the discovery of two rare hybrids, the first
named in her honour Potamogeton X cadburyae by Dandy and Taylor and collected in Seeswood
Pool, Nuneaton. According to these authors the hybrid is ‘‘a most surprising cross, but there is no
doubt about the parentage, as examination of the material will quickly show . . . Subsequent visits
to the Warwickshire locality have failed to reveal more of the plants’? (see Cadbury et al., A
computer-mapped Flora, a study of the county of Warwickshire: 218, 1971). The second hybrid was
named P. X pseudofriesii Dandy & Taylor and was collected in v.c. 14, near Buckenham
Ferry. She made some 400 Potamogeton collections for the British Museum, from many different
parts of the British Isles, of which 192 were from Warwickshire. These records included most of the
British species and natural hybrids.

Dorothy Cadbury did not neglect local natural history interests. She joined the Birmingham
Natural History Society in 1950 and remained a member until her death. One of her tasks for the
Society was to compile a complete list of the flowering plants of Edgbaston Park, a task which she
tackled with her usual enthusiasm and energy. She was very active, both on field excursions and in
giving talks during the winter sessions, particularly on her travels abroad, on wild flowers in
general and certainly on pondweeds in particular. Her enthusiasm and excitement during
excursions was infectious and was communicated to everyone within earshot. It was truly an
enjoyable and an energetic experience to be with Dorothy in the field!

I was privileged to work with her on the Revision of the Flora of Warwickshire for a period of
nearly 20 years. This was a joint project between the Birmingham Natural History Society
(B.N.H.S.) and the Botany Department of the University, together with help from the
Birmingham Museum and Art Gallery. The project was officially begun in January, 1950, the joint
recorders being Mr R. C. Readett (B.N.H.S.) and Mr Peter Green (University), with Dorothy

t

OBITUARIES 209

Cadbury as a committee member. Peter Green left in 1951 and I came to Birmingham in 1952.
After a number of changes to the Flora organization, Miss Cadbury, Mr Readett and I were
appointed joint recorders and editors with the assistance of Mr M. C. Clark.

Collation and verification of records was undertaken every Tuesday, year in and year out, by
Dorothy Cadbury together with various assistants, Elsa Pickvance taking part for most of the
period. In the evenings of each Tuesday, Dorothy together with Robert Readett and myself, would
identify ‘critical’ material sent in by collectors, make policy decisions from time to time and eat a
delicious Cadbury tea. This went on continuously, week by week, from 1954 to about 1970 — some
16 years. Throughout all this period Dorothy worked with an enthusiasm and determination that
was amazing. When the Computer-mapped Flora was published in 1971, Dorothy’s work
throughout the years was evident to all. This book stands as a tribute to her and to Robert Readett,
both gifted amateurs, who were able to teach me, as a professional, many lessons in field botany
and local flora work.

When I suggested to the Committee in 1964 that we ought to computerize our records and seek a
way to have the distribution of each species computer-mapped, I was surprised that without the
slightest hesitation Dorothy at once agreed to and enthusiastically supported this idea; I had
imagined that, since she grew up before the computer age, I should have some difficulty in
persuading her and Bob Readett to go along with this suggestion. On the contrary, they both took
to it with great enthusiasm.

After the publication of the Flora, into which Dorothy had put an energetic 20 years of her life, I
thought that she would wish to sit back and relax for a little while. I was completely mistaken; she
forged ahead with local natural history excursions and pond-dipping practically to the end of her
life.

Truly, here was a great personality whose friendship I greatly enjoyed. It was a privilege to have
known and worked with her. Always modest, but with whole-hearted devotion to field botany, she
is sadly missed by all who knew her.

J. G. HAWKES

JOHN HARRIS CHANDLER
(1909—1987)

B.S.B.I. members who knew him well will be sorry to learn of the death on 5th September 1987 of
John Chandler, a member since 1956. Though born at Markyate, near Luton, he had an intimate
knowledge of the countryside around Stamfora, where he had lived since the early 1950s. He was a
meticulous recorder who studied his own area in detail. During the Maps Scheme he took on a
number of 10km squares, and the Stamford square was one of the best recorded in England. He
was never afraid to tackle critical or difficult groups, particularly Rosa, Salix and Ulmus. His work
on these groups, especially the first, and in recording generally was of great significance in the
publication of three local Floras. Guy Messenger, author of the Flora of Rutland, writes that,
without John Chandler, that work would never have been begun, let alone finished. He tried to
persuade John to allow his name to appear on the title page, but, with his usual modesty and
generosity, he refused.

John Chandler also gave considerable help to Terry Wells in fieldwork for the forthcoming Flora
of Huntingdonshire and the Soke of Peterborough; his knowledge of the localities for many rare
plants in the Soke was invaluable and he was forever on the lookout for newcomers. It was he who
first noticed Elodea nuttallii in the Soke in 1974 and carefully traced its spread along the dykes and
river systems. His detailed study of Rosa, which fortunately occurred before the period of
widespread hedge removal, provides a historic record of the variety which existed around Stamford
and Peterborough before the fields were enlarged and the hedges destroyed. This detailed
approach is reflected in a letter he wrote to Joan Gibbons about the Flora of Lincolnshire: “I have
collected about 50 specimens so far, particularly in the Uffington and Holywell areas. It seems
better to work a small area in detail rather than odd bushes here and there. A small gravel pit at
Uffington has about a dozen bushes and at least half of that number are different — several ‘canina’

210 OBITUARIES

forms, a couple of ‘dumetorum’ varieties and ‘obtusifolia sclerophylla’.’’ This was his forte — a mind
and an eye for detail.

He was President of the Lincolnshire Naturalists’ Union (1965-66), and his Presidential
Address, which re-emphasised his love of his own locality, was ‘Uffington Gravel Pits — an
ecological study’. Uffington is c. 2 km east of Stamford.

In professional life John Chandler was a Customs and Excise Officer. This work stimulated his
interest in Revenue stamps, and, during the last years of his life when ill-health made fieldwork
more difficult, he was joint author of a book on The newspaper and almanac stamps of Great
Britain and Ireland; published in 1981, this is the definitive work on the subject. It demonstrates his
ability to compile and write; and, had he lived in the centre of a county instead of on the boundary
of three, he would undoubtedly have joined the ranks of B.S.B.I. members who have written
County Floras. Instead, three other authors have reason to be grateful for his unselfish assistance.

T. C. E. WELLS & I. WESTON

THOMAS GASKELL TUTIN, M.A., Sc.D., F.R.S.
(1908—1987)

Thomas Tutin, who died aged 79 on 7th October, 1987, was very widely renowned as an
extraordinarily productive and persistent author and compiler of Floras. His name will remain
familiar to plant taxonomists for centuries to come through his floristic achievements, two of which
stand out in excellence and importance.

Flora of the British Isles was published in 1952, with A. R. Clapham and E. F. Warburg as co-
authors, and, as ‘CTW’, quickly became the standard British reference work. The circumstances
surrounding the commencement of this work, and the early roles played by H. Gilbert-Carter and
A. G. Tansley, have been set in print by Clapham (1978), and for these and for many other
anecdotes and early reminiscences of Tom Tutin the reader is referred to that work, the existence
of which forbids too much repetition here. The impact of Flora of the British Isles cannot easily be
exaggerated. As a 13-year-old, it was the first ‘proper flora’ I ever used, but its degree of detail, its
level of authoritativeness, and its up-to-date nomenclature and classification were equally novel
and came as just as much a shock to those who were already experts in the study of British plants.
The reason for this is easy to see — it replaced, as the standard British Flora, C. C. Babington’s
Manual of British botany (1843), G. Bentham’s Handbook of the British flora (1858) and J. D.
Hooker’s The Students’ Flora of the British Islands (1870), all several times revised, but none very
adequately and none after 1924. The changes and new data that had accumulated up to 1952 were
predictably very numerous and very far-reaching. ‘CTW’ became the British botanists’ ‘bible’
almost overnight, and its sequence and nomenclature were soon adopted by the B.S.B.I. as its
basis for recording via J. E. Dandy’s List of British vascular plants(1958), which used the sequence
of ‘CTW’ “for obvious reasons of convenience’’.

The success of ‘CTW’ was largely the result of Tutin’s devotion to the overall editing and to the
incorporation of the efforts of his two co-authors, a devotion amply acknowledged by them on
page xviii of the first edition. Authorship of the various families was divided approximately equally
between the three (see pages 1507-1508 of the first edition). This position of pre-eminence of Flora
of the British Isles has been maintained to the present day by the appearance of a second edition in
1962 and a third in 1987, Tutin’s last work. Warburg having died in 1966, the families written by
him in the first two editions were revised for the third by D. M. Moore. The second and third
editions are, of course, improvements on the first, but, seen in relation to the amount and
availability of contemporary data, they probably represent a lesser achievement; in my opinion the
level of excellence of the first edition has not since been equalled in British Floras. The popularity
of ‘CTW’ was further established, and its use widened, by the preparation of a shortened version,
Excursion Flora of the British Isles, in 1959, with later editions in 1968 and 1981.

Much of the above concerns ‘CTW’ rather than Tom Tutin, but anyone who knew Tom soon
realised that the two were inextricably entwined. Moreover, Tom was a quiet, self-effacing man

OBITUARIES 211

who rarely volunteered information, let alone that of a personal nature, and in some ways it was
easier to understand the man via his work than directly.

The other major work with which Tom was involved was, of course, Flora Europaea (5 volumes,
1964-1980). The part Tom played as Chairman of the Editorial Committee, from its first formal
meeting at Leicester in 1956 until his death, has been referred to by Clapham (1978) and Webb
(1978), and an apt acknowledgement appears on page xv of Volume 5 (1980). A short history of
Flora Europaea also appears on pages xvii-xx of Volume 5. During the 1960s and 1970s Flora
Europaea gradually took over from Flora of the British Isles as Tom’s main botanical interest.
Indeed the influence of the former on the latter is made abundantly clear from a study of the
progressive editions of ‘CTW’ and of the Excursion Flora. The sequence of families in Flora
Europaea (taken from the twelfth edition of Syllabus der Pflanzenfamilien (1964)) was never
adopted in ‘CTW’, but the later editions were altered to conform with the generic and specific
sequences of the families, as was the classification in the majority of instances, and even the
descriptive data of later editions were emended in the light of those in Flora Europaea. Webb
(1978), in analysing the contents of Flora Europaea, pointed out that Tom wrote more accounts
(1307 species) than any other author, and concluded that “For individual achievement Tutin, . .
is in a class by himself; . . .”

Tutin’s contribution to botanical literature will also be known to all B.S.B.I. members through
two of our handbooks — British Sedges, produced in 1968 with A. C. Jermy (the second edition,
renamed Sedges of the British Isles, appeared in 1982, but Tutin played no part in its preparation),
and Umbellifers of the British Isles, produced in 1980. These books, the first two of the series, are
indispensable to students of sedges and umbellifers, and the first has become the best seller of any
of the Society’s publications. He wrote no major separate works on his favourite family, the
grasses, but he prepared the account of these for ‘CTW’ and was the general editor as well as a
major author for the family in Flora Europaea.

Tom Tutin was born, appropriately, at Kew on 21st April, 1908, where from an early age he
developed a keen interest in natural history, a devotion shared and fostered by his father, a
biochemist. Outside plants his favourite group became the Lepidoptera, and he retained an
interest in butterflies until his last years. His sighting of the season’s first Brimstone, or his
discovery on holiday of a good colony of Marsh Fritillaries, were to him items well worthy of
remark at the first opportunity to anyone who he thought shared his delight in these creatures.

In 1920 his family moved to the Bristol area, and from 1927 to 1930 he read Natural Sciences at
Cambridge University. There he met many people, both teachers and students, who were to shape
his future career. Perhaps his foremost contemporaries were P. W. Richards and E. F. Warburg,
but there were others just before and just after with whom he rubbed shoulders and who together
generated an exciting atmosphere from which emerged a probably unparalleled number of
distinguished botanists. In his second long vacation (1929) Tom and Warburg took part in an
expedition to Madeira and the Azores.

After graduation Tom took a series of short-term posts that alternated with his participation in a
number of foreign expeditions, notably to Spain and Morocco in 1931, British Guiana in 1933 and
Lake Titicaca in 1937. All these trips, especially that to British Guiana, deeply influenced his view
of plants and probably helped to determine the direction of his future career. Immediately after
graduation he remained at Cambridge to work on Greenland fossils collected by A. C. Seward. In
1933 he joined the Marine Biological Station at Plymouth to investigate the wasting disease of
Zostera, and after his return from Lake Titicaca he held posts at King’s College, London, the
University of Manchester, and, as part of the war effort, in the Geographical Section of the Naval
Intelligence Division of the Admiralty.

In these early years he published papers on topics as diverse as Greenland fossils, rot-holes in
trees, the Zostera parasite and fresh-water algae, as well as floristic and taxonomic notes on plants
from the Azores, S. America and Britain. His interest in aquatic plants, apparently commencing
with the Zostera work, flourished on his trip to Lake Titicaca, where he studied the larger algae
and developed ideas on the concept of the hydrosere succession. While at Manchester he spent a
good deal of time at Windermere, where he met Winifred Anne Pennington, whom he married in
1942. His wife, who answers variously to the names Winifred Pennington, Mrs T. G. Tutin, and
Professor W. A. Tutin, F.R.S., is a world authority on lake sediments. They had one son and three
daughters, one of whom, Caroline, has worked for the past 16 years on the threatened gorillas and

bt

212 OBITUARIES

chimpanzees in tropical Africa. Her regular letters to Tom over this period were a great delight to
him, and he often passed on from them tit-bits of general natural history interest.

Virtually all of Tom’s papers that appeared after 1942 concerned the floristics and taxonomy of
British and European flowering plants. In 1944 he was appointed Lecturer-in-charge of the
embryonic Department of Botany at University College, Leicester. He was appointed Professor of
Botany in 1947, holding this post through the granting of the charter to the University of Leicester
in 1957 until the foundation of the School of Biological Sciences there in 1967. During this period
he devoted much effort to the establishment of a thriving Department of Botany, especially
through the formation of a University Botanic Garden and Herbarium at Leicester, and to the
detailed study of the British Flora. The Botanic Garden was originally on the University campus,
but in 1947 was moved to an ideal 16-acre site two miles away at Oadby, opposite Leicester Race-
Course. The Herbarium was commenced in 1946 and now comprises about 120,000 specimens. At
first it reflected Tutin’s interest in the British flora, and contains his collections from most parts of
the British Isles, including many of the well-known localities of rare species. Those early
collections were augmented by duplicates from correspondents and friends such as J. E. Lousley
and E. F. Warburg, and by the collections of his chief technician, E. K. Horwood (son of A. R.
Horwood, co-author of The flora of Leicestershire and Rutland (1933)) and of one of his first
appointees, Miss A. P. Conolly. Parts of Tutin’s own herbarium (notably the grasses) were kept
separate, but they were all finally incorporated in the mid-1970s. Later, increasingly so as Flora
Europaea got underway, more Continental material was added (the names Horwood and Conolly
remain prominent here), and the valuable collections of Mrs B. M. Allen, O. Polunin and B. E.
Smythies were attracted by the determination service provided by Tom and A. O. Chater, who was
Flora Europaea Research Assistant at Leicester between 1960 and 1977. Their joint identification
sessions became a feature of the Leicester Herbarium for an hour or two after tea every afternoon,
and one soon learned that that period was not an appropriate time to interrupt with queries or
points relating to teaching or administrative matters.

In 1967 the Leicester scene was changed dramatically by the institution of the School of
Biological Sciences, which embraced five Departments (now seven). The greatly increased
administrative duties and inter-departmental liaison that this entailed were not to Tom’s liking
(principally, they ate into time that could be devoted to Flora Europaea), and Professor H. E.
Street was appointed from Swansea as Professor of Botany and Chairman of the School. Tom
became the first occupant of the newly created Chair of Plant Taxonomy. Upon his retirement
from this in 1973 he was awarded the title of Emeritus Professor, and until early 1985, when the
manuscript for the third edition of Flora of the British Isles was delivered to the press, he worked
almost daily in the herbarium on his various floristic projects.

Tom Tutin’s many floristic achievements were recognized by his award of a Gold Medal of the
Linnean Society of London in 1977, an Honorary Doctorate by Trinity College, Dublin in 1979,
and Fellowship of the Royal Society in 1982. He was President of the B.S.B.I. from 1957 to 1961,
following his Vice-Presidency from 1952-1956. Despite his name being a household word with
British botanists, I doubt whether he was well known personally by many of the present
membership of the B.S.B.I. He stopped attending our Annual Exhibition Meeting about ten years
ago because he felt he no longer knew many of the exhibitors and visitors. His own natural
reticence and the awe in which his juniors held him probably mitigated against the forging of new
friendships from casual personal meetings. However, once this initial difficulty was overcome, one
found a most warm-hearted and friendly person, who would most readily help with any query or
problem. He received many plants for identification; on the whole his replies were prompt,
accurate and brief. I believe that he never kept copies of his replies (certainly not since 1973, when
they were virtually all hand-written), and that he threw away the original letter of enquiry once it
was answered. Tom never used two words where one was sufficient, and had a pathological distaste
for confrontation or fruitless argument. He never in any way sought the limelight, and was always
surprised when anyone attempted to bring him into it. To celebrate his seventieth birthday, H. E.
Street had arranged for a book to be written by a number of his colleagues and former students
(Street 1978). Tom did not learn of this until a relatively late date, by which time Street had
unfortunately died of a heart attack. I was left to make the few remaining arrangements, and one
sentence in a brief note that Tom wrote to me shortly before we were due to gather for a surprise
celebratory dinner seems to me to sum up much of his character: “‘I find it incredible that anyone

OBITUARIES 713

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PLaTE 2. Thomas Gaskell Tutin, 1908-1987. Courtesy of Mrs T. G. Tutin. Taken early or mid 1960s.

214 OBITUARIES

should go to so much trouble over one so undeserving who, after all, has only done as far as
possible what he most wanted to do.”

His interests included observing plants and insects, listening to music and watching cricket. In his
younger days he enjoyed plant-hunting, gardening and playing the flute.

Tom Tutin was one of a dwindling number of distinguished botanists coming from the pre-war
Cambridge stable. In British plant taxonomy he started an era in 1952 with ‘CTW’, the many
obvious errors and gaps that it exposed being investigated by a much wider-ranging new generation
of research workers and amateurs. It generated a new interest in, and awareness of, the British
flora that coincided with the post-war expansion and the flourishing phase in the history of the
B.S.B.I. described by D. E. Allen (1986) as ‘“‘Full Steam Ahead.”’ His death equally marks the end
of an era, but we are left with tangible evidence of his achievements not only in his Floras and
Handbooks but also in the form of the enormous amount of diverse research and data-gathering
that these in turn generated. I know of two plants named after him: Atractylis tutinii Franco, Bot. J.
Linn. Soc., 71: 47 (1975) (endemic to Cabo de Gata, Spain); and Mentha x tutinii Pinto da Silva,
Cat. Pl. Vasc. Acores, p. 103 (1966). The latter has no description but refers back to a description
by J. Fraser in J. Bot., Lond., 70: 39 (1932), where the plant is described as M. piperita x M.
rotundifolia.

REFERENCES

ALLEN, D. E. (1986). The botanists. Winchester.

CLaPHaM, A. R. (1978). Thomas Gaskell Tutin, in Street, H. E., ed. Essays in plant taxonomy, pp. ix—xviii.
London, New York & San Francisco.

StreEET, H. E., ed. (1978). Essays in plant taxonomy. London, New York & San Francisco.

Wess, D. A. (1978). Flora Europaea — a retrospect. Taxon, 27: 3-14.

PUBLICATIONS OF T. G. TUTIN

This list is additional to that in Clapham (1978); it includes two omissions from the latter and all the
post-1976 publications that I have traced. In addition there are a few minor errors in Clapham’s
list. The most important are the page-numbers of the fifth article cited, which should read J. Bot.,
Lond. , 72: 306-314, 333-341; and that Aquilegia should be deleted from the entries for Flora
Europaea Volume 1.

1. 1964 A vegetacgao dos Agcores. Acoreana, 6: 8-32.

2. 1975 Apium, Bidens, Zostera & Poa, in Stace, C. A. ed. Hybridization and the flora of the
British Isles. London, New York & San Francisco.

3. 1978 (with C. A. Stace). A new species of Gaudinia from Spain. Bot. J. Linn. Soc., 76: 353-
356.

4. 1978 Short notes, in HEywoop, V. H., ed. Notulae Systematicae ad Floram Europaeam
spectantes, 20. Bot. J. Linn. Soc., 76: 361, 363, 365, 369.

5. 1980 (edited with V. H. Heywood et al.). Flora Europaea, Vol. 5. Cambridge. Author of
Zosteraceae and Gramineae pro parte.

6. 1980 Umbellifers of the British Isles. London.

7. 1981 (with A. R. Clapham [& E. F. Warburg]). Excursion Flora of the British Isles, 3rd ed.
Cambridge.

8. 1987 (with A. R. Clapham & D. M. Moore). Flora of the British Isles, 3rd ed. Cambridge.

C. A. STACE

OBITUARIES 215

DAVID HENRIQUES VALENTINE
(1912—1987)

Professor D. H. Valentine died in Manchester on 10th April, 1987. He was born in Salford on 16th
February, 1912, and was educated at Manchester Grammar School and St John’s College,
Cambridge. Initially involved with plant physiology, his research interests soon led him to consider
the relationships and distribution of flowering plants, a move to plant taxonomy which he pursued
for the rest of his life. In 1936 he was appointed Curator of the Herbarium at the University of
Cambridge and, in 1938, gained a Fellowship at St John’s College there. During World War II he
was employed by the Ministry of Food to carry out research into methods for dehydrating
vegetables. In 1945 he was appointed Head of the Department of Botany in the, then, Durham
Colleges in the University of Durham, first as Reader and later, in 1950, as Professor. In 1966 he
returned to his native Manchester (which, he always avowed, was schén), where he was Professor
of Botany until his retirement in 1979.

Valentine was a vigorous member of the Editorial Committee of Flora Europaea (which of his
colleagues can forget his insistent query at one meeting, “‘what the hell is NERC’’?, as the vogue
for such abbreviations moved towards the current unintelligibility), a member of the Council of the
Linnean Society of London (1968-71, 1976-80), President of the International Organization of
Plant Biosystematists (from 1974) and President of the Botanical Society of the British Isles from
1977 to 1979. In the late 1960s he was prominent in the group which undertook to assess the
potential effects of the dam at Cow Green in Upper Teesdale and was instrumental in the related
research programme.

His early studies on variation in Viola riviniana, during the 1940s, led Valentine into the then-
developing field of genecology. He soon expanded into the broader field of what would come to be
known as biosystematics, first by exploring the relationships between Viola riviniana and V.
reichenbachiana (later extending this to considerations of the cytogenetical relationships between
these and other violets) and then by his studies on Primula. He clarified the patterns and processes
of hybridization between British primulas, both in the field and in the experimental garden,
showing, amongst other things, the role of endosperm breakdown in accounting for the differential
success of reciprocal hybrids. In the field and in the laboratory Valentine’s interests extended to
Centaurea, Impatiens, Potentilla and Senecio. His Presidential Address to The Botanical Society of
the British Isles, in 1978 (Watsonia 12: 201-207, 1979), not only outlined the scope of his own
researches but also provided a most stimulating overview of the current British biosystematic
scene, in which amateur and professional members of our Society were making, or could make,
their mark.

Together with such contemporaries as Boécher, Gregor and Merxmiller, Valentine was
prominent in the European group which counterbalanced the North American, largely
Californian, school in the heyday of biosystematics during the 1950s and 1960s. His perceptive
studies of, above all, Viola and Primula, his introduction of the concepts of abrupt and gradual
speciation, and his continuing concern with the interrelationships between taxonomy and ecology,
have ensured his distinguished place in European botany. In addition to his considerable
contributions as editor and author for Flora Europaea, Valentine wrote over 30 scientific papers
and edited the volume resulting from the valuable international meeting on “Taxonomy,
Phytogeography and Evolution’, which he organized in Manchester in 1971.

Despite the research record outlined above, however, Valentine was, above all, a great teacher.
In 1951 I entered his Department as a callow undergraduate. Quite properly, ‘the Professor’ was
viewed with awe. It soon became evident that his kindly, though firm, guidance was leading to our
absorbing some of his knowledge and love of the British flora. His well-organized lectures and field
trips (always designed so that a glass of beer was at hand during the midday stop) were
instrumental in cultivating our appreciation of plant taxonomy. During the final examinations, how
we were helped to overcome our nervousness by the cherries and strawberries which he produced
during the intervals of the practicals!

I was the fourth, I believe, of his numerous graduate students. Unstintingly, he supported us
with ideas and the unpublished data from his work on Viola and Primula, with which to prosecute
our Ph.D. programmes. We were allowed into the tea-time discussions during which Valentine and
Dr Jack Crosby introduced us to the art of friendly, though gladiatorial, debate. Afternoon tea at

216 OBITUARIES

his home with Joan and their five children was always a delight and a welcome reminder of the
family life often missed by those of us living in college or digs. Nor was the conversation confined
to matters botanical — Valentine’s appetite for detective novels was a pleasurable, and much
discussed, revelation.

I am not sure how often a callow student becomes the friend and colleague of ‘the Professor’.
This was certainly the case between David Valentine and many of his students. His numerous
distinctions never caused him to lose his empathy with people, especially the young— the ‘common
touch’, as Kipling put it. We, his students, were privileged to have David Valentine as our mentor
and friend; we are grateful for his life.

D. M. Moore

(b)

Pate 3. David Henriques Valentine (1912-1987). a) Probably taken in the early 1950s; courtesy of Mrs H.
Parsons. b) Taken in 1967 at a Flora Europaea symposium in Seville; courtesy of D. M. Moore.

tin,

CAMBRIDGE

Flora of the British Isles

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A. R. CLAPHAM, the late T. G. TUTIN and D. M. MOORE

The third edition of this the standard British Flora will continue to be the definitive reference
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