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JOURNAL AND PROCEEDINGS OF
THE BOTANICAL SOCIETY
OF THE BRITISH ISLES

VOLUME 8

EDITED BY
E. F. GREENWOOD, G. HALLIDAY,
M. C. F. PROCTOR AND N. K. B. ROBSON

1970-71

PRINTED FOR THE SOCIETY BY
Willmer Brothers Limited, Birkenhead

PUBLISHED AND SOLD BY THE BOTANICAL SOCIETY OF THE BRITISH ISLES
C/O DEPARTMENT OF BOTANY, BRITISH MUSEUM (NATURAL HISTORY), LONDON SW7 5BD

DATES OF PUBLICATION

Part 1, pp. 1-96, 10TH January 1970.
Part 2, pp. 97-194, 3rp August 1970.
Part 3, pp. 195-344, 6TH April 1971.
Part 4, pp. 345-447, 3rp August 1971.

CONTENTS

THE JOURNALS OF THE BOTANICAL SOCIETY OF THE BRITISH ISLES

THE DISTRIBUTION OF Dryopteris assimilis S. WALKER IN BRITAIN. By
J. A. CRABBE, A. C. JERMY AND S. WALKER ,

THE STATUS OF Puccinellia pseudodistans (CREP.) JANSEN & WACHTER
IN GREAT BRITAIN. By B. M. G. JONES AND L. E. NEWTON

A BIOSYSTEMATIC AND ECOLOGICAL STUDY OF Salicornia IN THE DEE
ESTUARY. By P. W. BALL AND K. G. BROWN

Euphrasia brevipila AND E. borealis IN THE BRITISH ISLES. By P. F. YEO
_ SHORT NOTES

THE IRISH VICE-COUNTIES. By D. A. WEBB

PLANT RECORDS. Compiled by E. C. WALLACE

BOOK REVIEWS

OBITUARIES ...

REPORTS

NOTES ON THE HAWKWEEDS (Hieracium sensu lato) OF WESTERN SCOT-
LAND. By A. G. KENNETH AND A. McG. STIRLING

A REVISION OF SOME OF THE TAXONOMIC CHARACTERS OF ee
australe FEE. By R. H. ROBERTS ee ;

‘SrupiEs IN Potentilla anserina L. By D. J. OCKENDON AND S. M.
WALTERS ue “es sia me ne a

PLANT RECORDS FROM MULL AND THE ADJACENT SMALL ISLANDS. 2. -
J. F. M. CANNON AND E. B. BANGERTER

Exotic Amelanchier SPECIES NATURALISED IN EUROPE AND THEIR
OCCURRENCE IN GREAT BRITAIN. By F.-G. SCHROEDER

SHORT NOTES
PLANT RECORDS
BOOK REVIEWS
OBITUARY
REPORTS

PRESIDENTIAL ADDRESS 1970: BOTANICAL CONSERVATION IN BRITAIN,
PAST, PRESENT AND FUTURE. By E. MILNE-REDHEAD

PAGES
2

3-15

17-26

27-40
41-44
45-49

a
53-61
63-68
69-74
75-95

97-120

. 121-134

... 135-144

145-153

. 155-162
. 163-164
. 165-167
. 168-180
. 181-182
. 183-194

. 195-203

INDEX TO WATSONIA VOLUME 8
TAXONOMY OF THE Linum perenne GROUP IN EUROPE. By D. J. OCKENDON 205-235
THE Ranunculus acris L. COMPLEX IN EUROPE. By S. M. CoLeEs ... 237-261

STUDIES ON BRITISH POPPIES 4. SOME ASPECTS OF VARIABILITY IN THE
BRITISH SPECIES OF Papaver AND THEIR RELATION TO BREEDING

MECHANISMS AND ECOLOGY. By S. ROGERS ey eS: ... 263-276
A CYTOTAXONOMIC STUDY OF Lamiastrum galeobdolon (L.) EHREND. &

POLATSCHEK IN BRITAIN. By S. WEGMULLER ae fe ... 277-288
FREQUENCIES OF PIN AND THRUM PLANTS IN A WILD POPULATION OF THE

COWSLIP, Primula veris L. By D. R. LEES ... we ae ... 289-291
SHORT NOTES a See ie sa ues Ei me 2s 293-301
PLANT RECORDS .... Bet oe is. “ ne _ .. 802-312
BOOK REVIEWS oe ae su $e sic oi, 7 ... 313-322
REPORTS ... as eee ae ae Mi A ee ... 323-338
PANEL OF REFEREES AND SPECIALISTS a: ae2 a, ode ... 339-344
STUDIES ON Cytisus scoparius (L.) LINK WITH PARTICULAR REFERENCE

TO THE PROSTRATE FORMS. By J. J. B. GILL AND S. WALKER ... 345-356
BOTANICAL KEYS GENERATED BY COMPUTER. By R. J. PANKHURST __.... 35 7—368
SIX BRAMBLES (Rubi) FROM THE NORTH MIDLANDS. By A. NEwron ... 369-377
STUDIES ON GLOUCESTERSHIRE POPULATIONS OF A//ium eee (BIEB.)

G. Don. By D. M. BARLING Be at ... 379-384
OBSERVATIONS ON Phyteuma tenerum R. SCHULZ IN ENGLAND. By M.

KOVANDA oy a a oak ine aoe sat ... 385-389
THE MANCHESTER POPLAR. By C. A. STACE se Ae ses ... 391-393
SHORT NOTES ae: a: - ia ay: Fe be ... 395-402
PLANT RECORDS, ye: - sie o ces a ine set ... 403-411
BOOK REVIEWS ee ae oe Lise aN bf oa ... 412-422
OBITUARY ... Ri an vot mt ee: ne a ... 423-424
REPORTS... Ms oe ae cee ae te a ... 425-433
THE VICE-COUNTY GUIDE. Compiled by F. H. PERRING... ae? ... 435-447

INDEX sais we cus ae ie ir ex hes ..-» 451-469

iCa

f the Botan

ceed

0

ings o

Saher,

4

.

Bes

nna?

ab '

$

ok

nt

oh

Botanical Society of the British Isles

Patron: Her Majesty Queen Elizabeth the Queen Mother

Applications for membership should be addressed to the Hon. General Secretary:
c/o Department of Botany, British Museum (Natural History), Cromwell Road, London,
S.W.7, from whom copies of the Society’s Prospectus may be obtained or from the Hon.

Treasurer at 12/14 Wigmore Street, London, W.1.

Officers for 1969-70

Elected at the Annual General Meeting, 3rd May 1969
President, E. Milne-Redhead

Vice-Presidents, Prof. J. G. Hawkes, Prof. D. H. Valentine, J. E. Lousley, Dr F. H.
Perring

Honorary General Secretary, Dr I. K. Ferguson
Honorary Treasurer, J. C. Gardiner

Honorary Editors, E. F. Greenwood, Dr G. Halliday, Dr M. C. F. Proctor, Dr
N. K. B. Robson

Honorary Meetings Secretary, Mrs M. Briggs

Honorary Membership Secretary, Mrs J. G. Dony

All enquiries concerning sales of, or advertising space in, the Society’s publications
should be addressed to D. H. Kent, 75 Adelaide Road, West Ealing, London, W.13.

Watsonia, 8, 1-2 (1970). 1

The Journals of the Botanical Society of the British Isles

This is the first issue of a journal formed from the amalgamation of the Society’s two previous
journals, Watsonia and Proceedings of the Botanical Society of the British Isles, but continuing
under the former title and with continuity of volume number. The new journal will be published
twice a year, in January and July, and there will be four parts per volume. Abstracts from
Literature, formerly published in Proceedings, will appear separately and annually as BSBI
Abstacts. Editor

J. G. Dony President, 1967-9

In the early years of the Society, which dates back to 1836, its journal consisted of
an annual report of plants collected during the year as the Botanical Exchange
Club and Society of the British Isles (the name used by the Society for very many
years). Slim reports of a few pages increased in size as critical botanists, critical in
more senses than one, made longer notes on each other’s gatherings and deter-
minations. In the course of time the exchange reports took second place to
longer supplementary items which included a wide variety ranging from purely
domestic matters, such as lists of members and accounts of meetings, to valuable
papers of a taxonomic and topographical nature.

In 1947 the name of the Society was changed to its present one, by which
time the exchange of specimens had diminished to such an extent that its end
was certain and near. The next major change in the journals came in 1949 with
the separation of the domestic items from the rest. This was done in a Year Book
which was published for four years during which period the exchange reports
lapsed. At the same time a new journal, Watsonia, was launched under the
distinguished editorship of E. F. Warburg who was to remain its editor for
fourteen years. It was originally intended that this should contain Plant Notes,
Plant Records, Book Reviews, Obituaries and Abstracts from Literature, as well
as papers on the taxonomy and distribution of British plants. It was, in short,
to reflect all those facets of the study of the British flora which must for ever
be associated with the pioneer work of H. C. Watson and with which the Society
was mainly concerned.

Botanical studies were, however, going through a period of transition which
could be met only by another change in the Society’s journals. In 1953 the page
size of Watsonia was increased to allow for larger diagrams and tables, so
necessary in papers on the cytogenetics and experimental taxonomy of British
plants, the number of which was steadily increasing. Indeed, Watsonia was at
that time mainly intended for such papers. In the same year Proceedings of the
Botanical Society of the British Isles, an entirely new journal, was launched with
D. H. Kent as its editor. His skilled editorship was to last for twelve years. It was
intended that Proceedings should contain all that had previously been included
in the Year Book as well as such items as Plant Notes, Plant Records, Obituaries
and Abstracts from Literature. In addition it would contain papers dealing with
plant distribution. It seemed a very clear division of the material the Society
would need to publish and for a time it worked exceedingly well.

2 J. G. Dony

In recent years the study of the British flora has once more changed as no
doubt it will change again. Cytotaxonomy is no longer favoured to the extent
that it was twenty years ago with the result that a clear-cut division of papers is
no longer so easily achieved. Papers have appeared in one journal which could
with equal justification have been included in the other. If there had ever been a
claim, which is doubtful, that Watsonia was a medium for the professional
botanist and Proceedings for the amateur, it has long since passed. Indeed, some
of the more valuable scientific papers have come from the pens of amateurs
whilst lighter and more discursive articles have not infrequently been contributed
by professionals. A further problem arises from the recent increase in Abstracts
from Literature, which are of great value to some readers but are only of
passing interest to others.

The editors of the two journals and the Publications Committee of the
Society have given considerable thought to the future of the Society’s periodical
publications and have reported that they can see no justification for continuing
to have two periodicals. The needs of today could be served more effectively with
only one journal, an amalgamation of the present two, to be called Watsonia
and subtitled Journal and Proceedings of the Botanical Society of the British
Isles. To this change the Council of the Society agreed. The new journal will
contain all that has previously been included in the two previous ones and will
appear twice a year. In addition Abstracts from Literature will be published
under a separate cover annually and in this form may prove to be even more
useful to those members and subscribers who now value it.

As all contributions will now appear in one journal, it is hoped that an
increasing number of amateur botanists will be encouraged to be more ambitious
in the projects that they undertake. Their intimate knowledge of growing plants
is an essential complement to the laboratory techniques of research workers. Such
collaboration between amateur and professional botanists has been a unique
feature of the history of the Society and has led to the British flora being the
best understood in the world.

March, 1969.

Watsonia, 8, 3-15 (1970). 3

The Distribution of Dryopteris assimilis S. Walker
in Britain

By J. A. CRABBE*, A. C. JERMY* and S. WALKER**

* British Museum (Natural History), London.
*, Department of Genetics, LiverpoolUniversity.

ABSTRACT

The Dryopteris dilatata complex in Britain consists of two allotetraploids, D. dilatata (Hoffm.)
A. Gray and D. carthusiana (Vill.) H. P. Fuchs, and an ancestral parental diploid D. assimilis
S. Walker. The latter has hitherto been confused with D. dilatata sensu stricto and this paper
discusses the variation of the characters used to separate D. assimilis and gives the known
distribution in Britain. In particular the variation of the perispore is discussed in{the light of
studies made on the scanning electron microscope.

INTRODUCTION

The presence in Europe (in Britain, Norway and Switzerland) of a diploid
cytotype of the Dryopteris dilatata complex was first pointed out by Manton
(1950). Following cytogenetic studies (Walker 1955, 1961), it was proposed that
this taxon be regarded as a species, Dryopteris assimilis S. Walker (1961: 607),
based on Lastrea dilatata var. alpina T. Moore, which, as long ago as 1845,
had been recognised as a distinct variety. D. dilatata (Hoffm.) A. Gray and
D. carthusiana (Vill.) H. P. Fuchs (D. lanceolatocristata (Hoffm.) Alston,
in Dandy 1958) are both tetraploid species. A survey of material of these three
species in the major British and Irish herbaria (by J.A.C. and A.C.J.) showed that
D. assimillis has a wider distribution than hitherto thought and led to further
collecting of living material, which has been examined cytologically by S.W.
Diploid plants attributable to D. assimilis can now be reported from 23 localities;
Roberts (1965) recorded the presence of diploids in Snowdonia, N. Wales.

Since 1961 several papers have dealt with the morphology of and variation
within D. assimilis (Walker & Jermy 1964; Dopp & GAtzi 1964; Nannfeldt
1966; Widén, Sarvela & Ahti 1967; Simon & Vida 1966). Chromatographic
investigation by Hegnauer (1961), Wieffering, Fikenscher & Hegnauer (1965),
and Widén & Sorsa (1966) into the intraspecific variation of phenolic and other
compounds is proving to have interesting results. The discussion below, of
characters found useful in this survey, stresses the similarities and differences
between the three species mentioned above; it is summarised in Table 1, p. 6.

Material from the following herbaria has been studied: British Museum
(Natural History) (BM); Cambridge University (CGE); National Museum of
Ireland, Dublin (DBN); Royal Botanic Garden, Edinburgh (E); Glasgow
University (GL); Royal Botanic Gardens, Kew (K); Manchester Museum
(MANCH); National Museum of Wales, Cardiff (NMW); Oxford University
(OXF); and those of J. E. Lousley and E. C. Wallace.

4 J. A. CRABBE, A. C. JERMY AND S. WALKER

SPORE ORNAMENTATION AND CHROMOSOME NUMBER

We examined about 1500 specimens for this survey and found that the most
reliable identification characters are spore ornamentation and chromosome
number and these have therefore been stressed in the list of identifications
below. We feel that with a little experience certain vegetative characters can be
correlated with this spore type and these and the variation within them will be
briefly discussed.

The sculpturing of the perispore of D. assimilis differs from that of D. dilatata
and D. carthusiana in density and shape and size of projections. In D. dilatata,
50 per cent. or more of the surface is covered with subconical blunt spines, the
majority of which are touching and coalescing towards their bases (see Plate 1a).
The differences between the spores of D. carthusiana and D. dilatata are some-
what more subtle; those of the former are possibly more rugose and the pro-
jections shorter in relation to their width, many forming rounded papillae
rather than spines (see Plate 1b). The perispore of D. assimilis is 10-25 per cent.
covered with more finely tapered, blunt spines which only rarely are close
enough to coalesce at their bases (see Plate 1c and d).

Of the 102 herbarium specimens of D. assimilis with mature spores studied,
only two populations showed a spore-type which could not immediately be
placed in the assimilis range. One of these was from Borrowdale (collected by
Mr F. Jackson in 1963) and the spores were more rugose and had extremely few
spines; this first gathering was tentatively identified (Jermy 1968), on spore
and gross morphology, as a hybrid involving the D. dilatata aggregate and
D. aemula (Ait.) Kuntze, a diploid not hitherto regarded as a member of this
complex. A further search in the field failed to find this problem plant but similar
plants from the same locality proved to be diploid with regular meiosis. These
do not show aemula characters but are morphologically within the range of
D. assimilis and are regarded as that species in this paper. However, this popula-
tion may have been isolated for a long time and the spore morphology is not
typical (see Plate le); it needs further investigation. It is possible that
D. assimilis is elsewhere in upland sites in the Lake District; the specimen at
Kew collected at ‘Keswick’ in 1859 by Wright could well be from the same
Borrowdale locality, or at least the same valley.

The second is the plant recently found by Mr J. W. Dyce near Cullen, Banff-
shire (see below) and as yet its chromosome number is unknown. The spines
on the perispore are denser and more robust and bear a resemblance to those of
D. carthusiana (see Plate If).

When an investigation into the finer structure of the perispore (perine) using
the scanning electron microscope was begun, it was realised that immature
stages could well be interpreted as different spore types. A preliminary investiga-
tion using ultra-thin sections through developing sporangia suggested that the
perispore is laid down within a relatively short time and that the sporopollenin
is built up on a predetermined pattern; the density of the projections at least
would be obvious, therefore, from the outset of perispore formation (Pettitt,
J.M., personal comm., 1968). The degree of rugosity could well be an artifact
accentuated on drying, when it would then reflect the thickness of the various
walls of the spore. It is concluded that differences seen in the assimilis type of
spore are real and not arrested stages of the dilatata type.

It is said above that the chromosome number (n = 41) is a reliable character.

PLATE 1. Perispore surface in the Dryopteris dilatata complex (all x 6000). (a) D. dilatata [Corley,

Argyll]; (b) D. carthusiana [Crabbe & Jermy 11790, Sussex]; (c) D. assimilis [Jermy 6173,

Argyll]; (d) D. assimilis [Corley, Strome Ferry, W. Ross]; (e) D. assimilis (Jackson, Borrowdale,
Cumberland]; (f) D. assimilis [Dyce, Cullen, Banffshire].

PLATE 2. Spores and median pinnae of species of the Dryopteris dilatata complex. (a) D. car-
thusiana; (b) D. dilatata sensu stricto; (c) D. assimilis.

—— SSeS SS

THE DISTRIBUTION OF DRYOPTERIS ASSIMILIS IN BRITAIN =)

We are aware, however, that other diploid taxa, possibly originating from the
same ancestral stock, may well be very close morphologically to D. assimilis.
Without an investigation into their breeding behaviour these may not obviously
disclose themselves.

RANGE OF VARIATION IN SPECIMENS WITH AN ASSIMILIS SPORE-TYPE

RHIZOME

The rhizome is basically an erect system which falls into a semi-prostrate
condition in old specimens. Large D. assimilis plants show a more open-branched
rootstock than the compact head of two or three crowns seen in a well-established
D. dilatata. The side branches of the rhizome of D. assimilis may be distinctly
creeping thus having a growth-form similar to the hybrid D. carthusiana x dila-
tata. In both D. dilatata and D. assimilis the leaves arise close to each other, thus
forming a closed ‘shuttlecock’ crown; in D. carthusiana, the leaves are more
widely spaced along a creeping rhizome and never form a closed ‘shuttlecock’
crown. Widén et al. (1967) describe internal secretory hairs in the intercellular
spaces of the rhizome (and petiole) which are smaller (60-80 um) but more
frequent in D. assimilis than the other two species where they average 80-
100 um long.

SCALES ON RHIZOME AND STIPE

All the scales around the growing apex of all three species are triangular-
lanceolate; those on the stipe base are similar, often grading into very narrow,
hair-like scales. Also found on the stipe are more ovate scales with acuminate
tips and in D. assimilis these are frequently bullate (blistered or balloon-like), a
condition not seen on the other two species. The colour of the scales of D. assi-
milis is very variable, both concolorous forms and those with a darker centre
being found. Pale concolorous scales are found always in D. carthusiana
and on the stipes of juvenile leaves produced on adventitious side branches in
D. dilatata (small plants of D. dilatata from high altitudes occasionally have
pale scales also). Large carthusiana-like plants showing a darker median stripe
in their scales usually suggest hybridity with dilatata. Some populations of
D. assimilis have concolorous scales of a distinct red-brown colour not seen
in the other two species.

FROND HABIT, TEXTURE AND COLOUR

D. assimilis shows the erectness of habit characteristic of D. carthusiana;
this stiffness is reflected in the pinnae and lamina itself and the blade is flat from
the very beginning. In D. dilatata the whole frond reposes at an angle of 60° or
less and the blade throughout is more flaccid. On the ultimate segments, the
lamina tissue itself is reflexed from the midrib (cf. Plate 2a, b and c), a condition
more marked in young fronds but nevertheless obvious throughout the life of
the frond.

The colour of the lamina of D. assimilis is a mid- to yellow-green which dries
to a yellow-brown green in the herbarium. To some extent D. carthusiana has
this same character; D. dilatata, on the other hand, is a darker, more bluish
(olive) green which remains dense in the dried state. Obviously the resulting
colour of dried specimens varies to some extent with the method and speed of
drying but that of fresh material is not affected by shade or similar habitat
conditions.

6 J. A. CRABBE, A. C. JERMY AND S. WALKER

FROND SHAPE AND SIZE

The ratio of length of stipe to length of lamina ranges from 2:3 to 1:1 in
both D. carthusiana and D. assimilis; in D. dilatata it is more often 1:2.
D. assimilis from woodland sites can attain 1 m in height, as can D. carthusiana
in similar situations, but D. dilatata reaches up to 1-6 m. The range of height
of frond is greatest in D. assimilis, some smaller specimens 10-15 cm high
retaining their low stature after three years in cultivation; D. dilatata nay also
have small fronds at high altitudes and in exposed places.

The shape of the lamina in D. dilatata varies from ovate- triangibe to
lanceolate-triangular; in D. carthusiana the frond is narrower, almost linear-
lanceolate in outline. D. assimilis ranges between the two but more typical
specimens are linear-lanceolate; some small specimens and juvenile leaves tend
to be ovate or even triangular. The shape of the lowest pinna has-been used as a
diagnostic character for D. assimilis (Walker 1955, Clapham, Tutin & Warburg
1962, Heywood 1964, Simon & Vida 1966); unfortunately this exaggerated
lowest basiscopic pinnule, often half as long as the pinna itself, emphasising a
triangular outline, can be found in D. dilatata. The record of D. assimilis for
Kerry (v.c. H2) by Vida (1966, p. 281) is based solely on this character (pers.
comm.) and as no other Irish specimens have come to light in our search, this
record must be taken as extremely doubtful. On the other hand, however, the
shape of the median pinnae does point to a distinction and, as Plate 2 shows,
D. assimilis has a smaller length/breadth ratio than either of the other two.

The degree to which a fern leaf is dissected is difficult to define because two
axes of development are concerned; it is, however, so often the basis of the
general appearance of a species.

TABLE 1. COMPARISON OF CHARACTERS

D. assimilis

Rhizome

erect to semi-prostrate;
frond bases forming open
shuttlecock

Scales

concolorous, pale or red-
brown, or with darker
median stripe; some
ballooned scales on stipe

Frond shape|habit

erect, ovate-lanceolate to
linear-lanceolate, small
fronds ovate-triangular;
flat; teeth often turned up
Frond dissection
sinus open, cut +
tule; teeth obtuse

to cos-

Frond colour
yellow-green

D. dilatata

erect or rarely semi-pros-
trate; frond bases forming
dense shuttlecock

witha darker median stripe
(concolorous only on ad-
ventitious shoots or at high
altitudes); no ballooned
scales

lax, ovate-triangular, rarely

becoming lanceolate; con-
vex; teeth turned down

sinus closed, cut 4 or 2/3 to
costule; teeth obtuse

olive-green

D. carthusiana

creeping; frond bases more
distant, not forming dis-
tinct shuttlecock

concolorous and pale;
ballooned scales absent

erect, (linear-) lanceolate;
flat; teeth turned up

sinus closed, cut + half-
way to costule; teeth acute

yellow-green

THE DISTRIBUTION OF DRYOPTERIS ASSIMILIS IN BRITAIN q|

In D. assimilis the cutting, especially towards the tips of the pinnae, is deep
and the segments are distinct; in fact some fronds are so finely dissected that,
when the upper half only is seen, the plant may be taken for Athyrium filix-
femina. In both D. dilatata and D. carthusiana, the sinus between the segments
rarely reaches more than half-way to the midrib (see Plate 2). Furthermore this
open pattern is accentuated in D. assimilis by the sinus being wide, whereas in
the other two species it is closed and the lamina edges often touch.

GENERAL COMMENTS ON THE VARIATION

Dryopteris assimilis shows then considerable variation in scale colour, frond
shape and the cutting of pinnae but there is insufficient correlation between
these characters to warrant any further formal taxonomic separation. Widén
et al. (1967, p. 5) describe a form from S. Finland with finely dissected pinnae
and with distinct red-brown scales, which they compare with D. dilatata var.
pseudo-spinulosa Rosendahl (1916, p. 327), but stress that on morphological and
chemical grounds it ‘is of no taxonomic importance’. This form is present in
W. Scotland also, e.g. Jermy 6176, where n = 41.

Thomas Moore described many varieties of Lastrea dilatata (Moore 1855;
1857), one of which (var. alpina) is, of course, the type of D. assimilis. Another
variety whose gross morphology and spores suggest D. assimilis is L. dilatata
var. ordeanae described from a lowland site at Kilmory, Argyll. It has a broader,
finely dissected frond with dark-centred scales and is characteristic of the
D. assimilis found around 1000 m in the Austrian Tyrol. Other specimens in
the Moore herbarium at Kew, from Bute and labelled ‘L.d. var. micromera
T. Moore’ are also D. assimilis. The type of var. micromera, however, is from
Devon and other specimens so named are from SW. Wales; this southern
element has spores of the D. dilatata type and needs further investigation. The
well-worked site of Ben Lawers has produced not only the type of D. assimilis
but also L.d. var. fraseri, a form with dark-centred scales and an ovate-triangular
frond, which is extremely (and abnormally?) finely cut; it has the assimilis
spore-type.

Two forms that stand out in the field, in cultivation and in the herbarium
have not, as far as can be seen, been named. One is a plant with crowded
(i.e. overlapping), ascending pinnae and pale reddish-brown scales; one such
population of very small plants from Glen Spean (Stirling & Walkinshaw
1 & 2) that keep their small stature in cultivation has been cytologically in-
vestigated and found to be diploid. Similar, but larger plants have been detected
in the herbarium from other parts of Scotland. The other is a plant collected in
fairly dry, acid, dwarf heath near Cullen, Banffshire by J. W. Dyce, now in
cultivation. It has distinct creeping rhizomes and pale scales, an ovate-triangular
frond with very narrow pinnae and oblong, truncate pinnules (cf. the form known
as D. dilatata var. collina: see Moore 1855, tab. 26A). The spines on the peri-
spore are somewhat denser than in normal D. assimilis (see Plate 1f) and the
plant may turn out to be a form of D. carthusiana; it certainly needs further
investigation.

DETAILED DISTRIBUTION OF D. ASSIMILIS

The distribution of the specimens seen is summarised on the map (Fig. 1); the
data is listed below in vice-county order and, within each vice-county, it follows

8 J. A. CRABBE, A. C. JERMY AND S. WALKER

the numerical order of the National Grid co-ordinates. The overall distribution
in Britain is that of an ‘arctic-alpine’ as defined by Matthews (1955); it follows
closely the distribution of such species as Carex bigelowii Torr. ex Schwein.,
Silene acaulis (L.) Jacq. and Saxifraga nivalis L. and, in England, that of
Woodsia ilvensis (L.) R. Br. In Europe it is mainly northern in distribution,
reaching 71° N. in Norway; in central and southern Europe it is confined to the
mountain ranges.

DISTRIBUTION
of
DRYOPTERIS
ASSIMILIS
S. Walker
(Gy onogee hy
examined

e@ herbarium
specimens

al

Ry

— a)

0 1 2 3 4 5 6

Ficure 1. Distribution of Dryopteris assimilis S. Walker in the British Isles.

THE DISTRIBUTION OF DRYOPTERIS ASSIMILIS IN BRITAIN 9

Its presence, together with other arctic-alpines, on the higher mountains of
Britain suggests it may have been a member of our Late-glacial flora. Its ecology
on the other hand is that of a woodland plant, a habitat which it often frequents
at lower altitudes (e.g. at almost sea level in western Scotland). Did our moun-
tain form spread from central European woodlands because of its ability to find
a niche amongst the bolders of a stable moraine? If so, why did it not return
to the lowlands in some quantity to the habitats where we now find D. dilatata?
We need to know more of the ecology of D. assimilis and to decide whether in
fact there are two ecological races (and diploid cytotypes).

SPECIMENS OF DR YOPTERIS ASSIMILIS IDENTIFIED BY THE AUTHORS

(a) confirmed by examination of chromosomes (by S.W.)

V.C. 70 CUMBERLAND: Holmrook, c. 200 ft., 34/0799, H. V. Corley, 1967 (BM);
Borrowdale, W slopes of Glaramara, 2000 ft., 35/2512, F. Jackson 2 & 4, 1966 (BM).

V.C. 88 MID PERTH: Ben an Eachan, c. 2500 ft., 27/5638, A. C. Jermy 3363,
Sept. 1962 (BM); Meall na Tarmachan, c. 2500 ft., 27/5838, S. Walker 63103, 1963
(BM); Ben Lawers, c. 3000 ft., 27/6341, S. Walker 63114, 1963 (BM).

V.C. 92 SOUTH ABERDEEN: above Loch Muick, c. 2000 ft., 37/2989, S. Walker
65001, 1965 (BM).

V.C. 96 EASTERNESS: Beinn Braeriach, c. 2500 ft., 28/9500, S. Walker 66002,
1966 (BM); Coire an Lochain, c. 3000 ft., 28/9803, S. Walker 66003, 1966 (BM);
Faicaill a’ Choire Chais, c. 3000 ft., 28/9904, S. Walker 66004, 1966 (BM).

V.C. 97 WESTERNESS: Stob Coire an Easain, Glen Spean, c. 3000 ft., 27/2372,
A. McG. Stirling & D. A. Walkinshaw I & 2, 24 Sept. 1966 (BM); Stronanime, Ardgour,
17/9583, H. V. Corley 674 & 678, 1967 (BM); Beinn na Lap, 27/3769, H. V. Corley 14,
1966 (BM).

V.C. 98 MAIN ARGYLL: Beinn Fraochaidh, 1600-1750 ft., 27/0251, A. C. Jermy
6176, 27 Aug. 1966 (BM); Coire Dubh, Beinn Fraochaidh, 27/0252, H. V. Corley,
1965 (BM); Allt Coire Ghlias, c. 3000 ft., 27/1028, S. Walker 65010, 1965 (BM);
Bidean nan Bian, c. 1750 & 2250 ft., 27/1354, S. Walker 65005 & 65006, 1965 (BM);
Beinn an Lochain, 27/2208, H. V. Corley, 1965 (BM); Beinn Laoigh, c. 1900 ft.,
27/2626, H. V. Corley, 1965 (BM); Beinn Udlaidh, c. 1900 ft., 27/2732, H. V. Corley 9,
1966 (BM); Coire Daimh, 27/2732, H. V. Corley 6, 1966 (BM); Beinn Dothaidh,
c. 3000 ft., 27/3240, S. Walker 63113, 1963 (BM); Coire an Dothaidh, c. 2250 ft.,
27/3240, S. Walker 63112, 1963 (BM).

V.C. 105 WEST ROSS: 2 miles W of Strome Ferry, 18/8032, S. Walker 63109, 1963
(BM).

V.C. 111 ORKNEY: Naversdale, Orphir, N30/3304, E. M. Bullard, Aug. 1966 (BM.

(b) determined by examination of spores (except those marked * where ripe spores
were not found) or by other morphological criteria

V.C. 48 MERIONETH: Moelwyn Bach, Ffestiniog, 23/6644, R. E. Hughes, Aug.
1967 (NMW)*.

V.C. 49 CAERNARVON: Yr Eifl, 23/3544, R. H. Roberts, 22 July 1967 (NMW);
Cribbyn, 23/65, H. A. Hyde, July 1925 (NMW)*; Cwm Brynog, 2000 ft., 23/6054,
J. H. Lewis, 29 July 1875 (MANCH); Cwm Glas, 23/6155, J. E. Griffith (NMW);
Cwm Glas Mawr, 23/6155, R. E. Hughes, June 1967 (NMW)*; Cwm Glas Bach,
23/6156, R. E. Hughes, June 1967 (NMW)*; Cwm Llafar, 23/6565, J. E. Griffith,
July 1893 (NMW)*.

10 J. A. CRABBE, A. C. JERMY AND S. WALKER

V.C. 65/66 NORTH-WEST YORK/DURHAM: Teesdale, 35/82, Mitchinson,
1860 (K).

V.C. 68 CHEVIOTLAND: Hulne Parks, 46/1912, W. Richardson, Aug. 1868 (BM)*.

V.C. 69 WESTMORLAND: High Cup Nick, Murton, 35/7426, A. H. G. Alston
4293, 13 July 1939 (BM)*.

V.C. 70 CUMBERLAND: Borrowdale, W slopes of Glaramara, 35/2512, A. C.
Jermy 6424-30, 23 June 1968 (BM); Keswick, 35/2723, Wright, 1859 (K).

V.C. 72 DUMFRIES: Moffat, 36/0805, J. Anderson, 1861 (K); Midlaw Burn,
Moffatdale, 1500 ft., 36/1205, E. S. Marshall, 23 July 1907 (BM, CGE, E)*, 12 Oct.
1907 (K)*; Hartfell, near Moffat, 36/1113, P. N. Fraser, 21 Aug. 1861 (EK, K); Carrifran
Glen, 36/1514, P. N. Fraser, 25 July 1860 (E); Loch Skene, 36/1716, P. N. Fraser,
15 Sept. 1860 (E), 24 July 1866 (K)*.

V.C. 78 PEEBLES: Mossfennan, 36/1136, P. N. Fraser, 13 July 1860 (E).

V.C. 85 FIFE: wood in estate, Balmuto, 36/2289 (introduced ?) J. T. Boswell-Syme,
June 1876 (BM); Kirkaldy, 36/2791, W. Stewart (K).

V.C. 86 STIRLING: Ben Lomond, 3100 ft., 27/3302, B. W. Ribbons, 23 Oct. 1965
(GL); North Corrie, Ben Lomond, 3000 ft., 27/3602, R. Mackechnie, 20 Aug. 1932,
(Herb. Lousley; Herb. Wailace).

V.C. 87 WEST PERTH: Meall Mor nan Eag, 27/3418, 1650 ft., H. V. Corley 126,
27 May 1968 (BM)*; Glen Falloch, c. 2400 ft., 27/3622, E. S. Marshall, 25 July 1890
(CGE)*.

V.C. 88 MID PERTH: Beinn Oss, 27/2825, F. M. Webb, 5 Aug. 1879 (E); An
Binnein, Ben More, 3300 ft., 27/4322, F. J. Hanbury, 26 July 1889 (BM)*, E. S.
Marshall, 26 July 1889 (CGE); Killin, 27/5732, E. Liddell, 1871 (K); Glen Lyon, 27/5646,
P. N. Fraser, July 1885 (E)*; Ben Lawers, 3000 ft., 27/6341, A. H. G. Alston, ex Univ.
Botanic Garden, Leeds, 1952 (BM), J. H. Balfour (BM)*, J. T. Boswell-Syme, Aug. 1851
(BM), Aug. 1866 (E), Cleminshaw, July 1905 (CGE)*, J. N. Frankland, Aug. 1949 (BM),
P. N. Fraser, 5 Aug. 1856 (E), J. Hardy, Aug. 1865 (CGE), C. W. Hope, 5 Aug. 1856 (E),
T. Moore, 1855, [LECTOTYPE of Lastrea dilatata var. alpina T. Moore and HOLO-
TYPE of Dryopteris assimilis S. Walker] (K), Wilson, Sept. 1855 (NMW); Ben Lawers
E corrie, 27/6341, A. J. Wilmott, 21 July 1928 (BM), R. David, 25 July 1964 (CGE)*;
Ben Lawers SW corrie, 2000 ft., 27/6341, J. E. Lousley, 12 Aug. 1932 (Herb. Lousley)* ;
An Stuc above Lochan nan Cat, 3000 ft., 27/6341, A. C. Jermy 1884 bis (BM)*,
E. S. Todd 25 & 26, (OXF); Meall Garbh, 3000 ft., 27/6441, E. S. Marshall, 22 Aug.
1887 (CGE); Aberfeldy, 27/8548, W. H. Forbes, 8 Oct. 1892 (OXF).

V.C. 89 EAST PERTH: Beinn Pharnagain, on boulder scree, 1750 ft., 27/4562,
K. Trewren 3, 19 Sept. 1965 (BM); S side of Ben Vrackie, 2600 ft., 27/9563, J. E. Lousley,
18 Aug. 1966 (Herb. Lousley); below Corrie Cas ea gallach, Ben-y-Glow, 2000 ft.,
27/9873, P. Halligey, 8 July 1964 (BM)*.

V.C. 90 FORFAR: Glas Maol (Forfarside), 37/1676, E. F. Linton, 6 Aug. 1883,
(BM)*; Glen Clova, Corrie Winter, 37/2774, W. A. Shoolbred, 30 June 1904 (NMW)*;
Loch Brandy, E side, 37/3375, H. & J. Groves, 25 July 1878 (BM); Corrie Fee, Clova,
2500 ft., 37/2575, A. C. Jermy 3405, 22 July 1964 (BM)*; Glen Clova, 37/2575,
E. F. Linton, 21 Aug. 1884 (OXF), Wollaston, 1873 (K).

+ The specimen recorded as D. assimilis from Knock Ore Gill (V.C.69) by A. Eddy &
D. Welch (Proc. bot. Soc. Br. Isl., 6: 326 (1967)) is D. dilatata.

THE DISTRIBUTION OF DRYOPTERIS ASSIMILIS IN BRITAIN 11

V.C. 92 SOUTH ABERDEEN: in the higher parts of the Grampians, 27/88,
H. C. Watson (BM)*; Cairn Toul, Coire Lochan Maine, 3000 ft., 27/9697, E. C.
Wallace & R. Mackechnie, 5 July 1934 (Herb. Lousley*; Herb. Wallace): Pools of
Dee, Lairig Ghru, Cairngorms, 28/9503, W. Edgar Evans, 31 July 1907 (E)*; Derry
Cairn Gorm, near Braemar, 37/0198, F. J. Hanbury, 23 July 1885 (BM)* ; Beinna Bhuird,
3000 ft., 37/0898, Greville, 1831 (E); Braemar, Castleton, 37/1491, A. Croall 167,
August 1854 (CGE, K); Morone, Braemar, 37/1288, C. Bailey, 6 Sept. 1864 (MANCH);
Glen Callater (highland above Loch Kander), 37/1880, J. Fraser, Sept. 1885
(MANCH)*; Glen Callater, Braemar, 37/1883, A. Croall, 1855 (K); J. F. Duthie,
August 1874 (BM)*; rocks below Lochnagar cliffs, 37/2485, E. C. Wallace, 8 Sept.
1934 (NMW)*.

V.C. 93 NORTH ABERDEEN: 3 miles S of Keith, near Pitgour Farm, 38/4445,
M. McCallum Webster 7891, 14 Aug. 1962 (BM).

V.C. 94 BANFF: Cabrachan, 38/3826, D. Clarke, 1864 (K); Crannoch Hill, Cullen,
c. 200 ft., 38/5266, J. W. Dyce, Sept. 1966, (BM).

V.C. 96 EASTERNESS: north-facing slopes of Coire Garblack, 27/8794, Proctor &
Goodway G3/82, June 1953 (CGE)*; Braeriach, 2800 ft., 27/9599, J. A. Wheldon,
July 1909 (CGE, OXF); Corrie an Lochan, Braeriach, 3300 ft., 27/9599, E. S. Marshall,
12 Aug. 1898 (BM)*; above Corrie an Lochan, 3500 ft., 38/0004, J. E. Lousley, 1 July
1953 (Herb. Lousley)*.

V.C. 97 WESTERNESS: Ben Nevis, 2750 ft., 27/1671, M. S. Campbell, 16 July 1937
(BM)*, K. Trewren, 22 Sept. 1965 (BM), Herb. Terras, 1887 (E); Ben Alder, 27/4971,
E. C. Wallace, 10 July 1946 (Herb. Wallace)*; S side of Coire Ardair, Crage Meagaidh,
2200 ft., 27/4387, H. V. Corley 124, 30 Aug. 1967 (BM); Coire nan gall, L. Laggan,
27/4886, F. J. Hanbury, 28 July 1916 (BM, E)*.

V.C. 98 MAIN ARGYLL: Kilmory, 16/78, A. Brown, 1862 (K); Ormay Woods,
17/8101, J. W. Dyce 6, Sept. 1968 (BM); above L. Craignish, 200 ft., 17/8204, H. V.
Corley, May 1968 (BM)*; Ben More, Cowal, 26/1090, A. G. Kenneth, 7 Apr. 1966
(Herb. Wallace); Tailor’s Leap, Glen Nant, 27/0128, H. J. Bruty, Sept. 1968 (BM);
Beinn Fraochaidh, 1600-1750 ft., 27/0251, A.C. Jermy 6173, 27 Aug. 1966 (BM, E);
Beinn nan Aighenan, 3000 ft., 27/1441, H. McAllister, May 1961 (BM); Sgurr an
Ulaidh, 27/1153, H. V. Corley 10, Apr. 1966 (BM); Fionn Glenn, Glencoe, 27/1557,
R. Mackechnie (Herb. Wallace)*; Ben Donich, near Lochgoyl Head, 2500 ft., 27/2204,
A. G. Kenneth, 19 Nov. 1966 (BM); Ben Laoigh, 3000 ft., 27/2626, J. W. Dyce, 1967
(BM), C. I. & N. Y. Sandwith, 21 July 1936 (K)*; Clach Leathad, 3 miles SW of
Kingshouse Inn, 27/2449, E. S. Marshall (CGE)*; Beinn Dhorain, 3000 ft., 27/3238,
G. C. Druce, Aug. 1898 (OXF)*.

V.C. 99 DUNBARTON: Ben Vane, 2800 ft., 27/2709, J. E. Lousley, 27 July 1966
(Herb. Lousley)* ; Ben Vorlich, 27/2912, P. N. Fraser, 10 Aug. 1864 (E).

V.C. 100 CLYDE ISLES: Arran, 16/26, J. Anderson, 1857 (K); head of Glen Dubh,
Arran, 16/9833, J. W. Dyce 5, Sept. 1968 (BM); Bute, 26/06, E. Liddell, 1869 (K)*.

V.C. 101 KINTYRE: Ballenach, Lochgilphead, 16/8491, J. Anderson, 1860 (K),
E. Greatorex, 1861 (K); Erine, Lochfyneside, 16/8576, A. G. Kenneth, Oct. 1966 (BM);
Cam Loch, near Lochgilphead, 16/8286, H. V. Corley, May 1966 (BM)*; Ardrishaig,
Loch Fyne, 16/8585, F. Griffith, 1858 (K).

V.C. 104 NORTH EBUDES: Hallaig, Raasay, 18/5436, M. McCallum Webster,
20 June 1957 (BM)*.

12 J. A. CRABBE, A. C. JERMY AND S. WALKER

V.C. 105 WEST ROSS: SW side of A’Chailleach, 28/1371, A. G. Kenneth, 19 July
1967 (BM)*; Meall a Chrasgaidh, Fannich Forest, 28/1873, E. C. Wallace, 22 July
1948 (Herb. Wallace)*, A. G. Kenneth, 4 July 1967 (BM)*.

V.C. 106 EAST ROSS: Coire Dombain, Carn Eige, Glen Cannich, BLES Ee. Cs
Wallace, 18 July 1947 (Herb. Wallace).

V.C. 107 EAST SUTHERLAND: Ben More Assynt, 2300 ft., 29/3120, F. J. Hanbury,
12 July 1890 (BM)*, E. S. Marshall, 12 July 1891 (CGE).

V.C. 108 WEST SUTHERLAND: Achmelvich, 250 ft., 29/0524, J. Anthony, 25 July
1955, (E); Ben Stack, 29/2642, A. G. Kenneth, 8 July 1967 (BM)*; NW. end of Foinaven,
29/3149, A. G. Kenneth, 10 July 1967 (BM)*; Cranstackie, 29/3555, A. G. Kenneth &
E. M. Bullard, 13 July 1967 (BM)*; Ben Hope, 29/4749, A. G. Kenneth, 18 July 1966
(BM)*.

V.C. 109 CAITHNESS: Summit of Morven, 39/0028, F. J. Hanbury, 18 July 1887
(BM)*.

V.C. 110 OUTER HEBRIDES: South Uist, 08/71 to 74, A. Somerville, July 1888
(BM)*; near Loch Maddy, North Uist, 08/9367, W. A. Shoolbred, 12 July 1894
(BM, CGE, NMW); Cleisham, Harris, 19/1507, W. S. Duncan, August 1889 (BM)*;
north rocks, Sgaoth Ard, North Harris, 19/1603, J. W. Campbell, 1 Aug. 1939 (BM)*.

V.C. 111 ORKNEY: Hoy: Enegass, SE side of meadow of the Kame, N30/1904,
H. H. Johnston 4733 (E)*; Mainland: hill between Sowa Dee and Stany Knowe,
Sandwick, 310 ft., N30/2214, H. H. Johnston 2594 (E, K)*; Naversdale, Orphir,
N30/3304, J. T. Boswell, Aug. 1875 (BM, CGE, E, NMW), 4. H. Johnston, 7 Sept.
1878 (E); quarries E of Loch of Stenness, N30/3304, J. T. Boswell, Aug. 1875 (BM, E);
Russa Dale, Stenness, 300 ft., N30/3309, &. M. Bullard, 24 Aug. 1966 (BM).

V.C. 112 ZETLAND: Bressay, N41/4940, R. Tate, 26 June 1865 (CGE)*.

HYBRIDS

Triploid hybrids have been detected in populations where D. assimilis and
D. dilatata grow together in S. Aberdeen, v.-c. 92, Easterness, v.-c. 96, Argyll,
v.-c. 98 and West Ross, v.-c. 105 (all coll. S. Walker); and in Kintyre, v.-c. 101
(coll. A. G. Kenneth). They are intermediate between the parents, having the
dark-centred scales and dissection of D. dilatata and the flatness of lamina and
erectness of D. assimilis; the sporangia are mostly abortive. Cytological
investigation shows that chromosome pairing at meiosis in these hybrids
agreed with that of synthesised hybrids (Walker 1955), giving approximately
equal numbers of bivalents and univalents. Wild hybrids between these species
have been reported and cytologically examined previously from Bavaria
(Walker 1955) and Switzerland (Dépp & Gatzi 1964). No wild hybrids between
D. assimilis and D. carthusiana have been recorded for Britain at the time of
writing. Other plants with abortive spores which on morphological grounds
have some features characteristic of D. aemula (Ait.) O. Kuntze (see Jermy
1968) may turn out to be hybrids between that species and D. assimilis or
D. dilatata; such hybrids have to be proved cytologically and must remain
dubious until they are further studied.

ACKNOWLEDGMENTS
We would like to record our thanks to the curators of the herbaria mentioned

THE DISTRIBUTION OF DRYOPTERIS ASSIMILIS IN BRITAIN 13

above for loaning material to the British Museum for study. We would further
thank the many people who have sent material in, especially Miss Elaine
Bullard, H. V. Corley, J. W. Dyce, F. Jackson, A. G. Kenneth and R. H.
Roberts, who have sent living material for cytological investigation. For
technical help with the scanning electron microscope, we thank Miss Patricia
Sims of the Botany Department and Brian Martin and his colleagues of the
Electron Microscope Unit, British Museum (Natural History); we are grateful
to Cambridge Scientific Instruments Ltd and to Miss J. Killingworth for their
help in the early stages of this investigation.

REFERENCES

CLAPHAM, A. R., TuTIN, T. G. & WarBuRG, E. F. (1962). Flora of the British Isles, 2nd ed.
Cambridge.

Danpy, J. E. (1958). List of British Vascular Plants. London.

Dopp, W. & GAtTzi, W. (1964). Der Bastard zwischen tetraploider und diploider Dryopteris
dilatata. Ber. schweiz. bot. Ges., 74: 45-53.

HEGNAUER, R. (1961). Uber infraspezifische chemische Variabilitat bei einigen Dryopteris-
Arten. Pharmaceutica Acta Helvetiae, 36: 21-29.

HeEYwoop, V. H. (1964). Dryopteris in Flora Europaea, vol. 1, pp. 20-22., Cambridge.

JermMy, A. C. (1968). Two new hybrids involving Dryopteris aemula. Br. Fern Gaz., 10: 9-12.

Manton, I. (1950). Problems of cytology and evolution in the Pteridophyta. Cambridge.

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

Moore, T. (1855). The ferns of Great Britain and Ireland, nature-printed folio ed. London.

Moore, T. (1857). The handbook of British ferns, etc., 3rd ed. London.

NANNFELDT, J. A. (1966). Dryopteris dilatata och Dr. assimilis i Sverige. Botaniska Notiser, 119:
136-152.

RoBeErTs, R. H. (1965). Dryopteris assimilis S. Walker in Snowdonia. Nature Wales, 9: 162-164.

ROSENDAHL, H. V. (1916). Bidrag till sveriges ormbunksflora. III. Svensk bot. Tidskr., 10:
312-339.

Simon, T. & VIDA, G. (1966). Neue Angaben zur Verbreitung der Dryopteris assimilis S. Walker
in Europa. Annls Univ. Scient. bpest. Rolando Eodtvés, Sect. biol., 8: 275-284.

WALKER, S. (1955). Cytogenetic studies in the Dryopteris spinulosa complex. I. Watsonia, 3: ©
193-209.

WALKER, S. (1961). Cytogenetic studies in the Dryopteris spinulosa complex. II. Am. J. Bot.,
48: 607-614.

WALKER, S. & JERMY, A. C. (1964). Dryopteris assimilis S. Walker in Britain. Br. Fern Gaz., 9:
137-140.

WIDEN, C. J., SARVELA, J. & AuTi, T. (1967). The Dryopteris spinulosa complex in Finland.
Act. bot. fenn., 77: 1-24.

WIDEN, C. J. & Sorsa, V. (1966). A chromatographic and cytological study of the Dryopteris
spinulosa complex in Finland. Hereditas, 56: 377-381.

WIEFFERING, J. H., FIKENSCHER, L. H. & HEGNAUER, R. (1965). Chemotaxonomische Unter-
suchungen mit Dryopteris-Arten—6. Dryopteris spinulosa Komplex. Pharmaceutisch
Weekblad, 100: 737-754.

14 A. C. JERMY AND B. S. MARTIN

APPENDIX: A NOTE ON THE PREPARATION OF
MATERIAL FOR THE SCANNING
ELECTRON MICROSCOPE

By A. C. JERMY and B. S. MARTIN*
* British Museum (Nat. Hist.) Electron Microscope Unit

MOUNTING MATERIAL

Whenever possible, large quantities of spores were shaken on to a clean sheet of tracing,
greaseproof, or similar high glazed paper and the aluminium stub, covered with the
appropriate mountant, was inverted and pressed gently on to the spore-mass. Where
spore material was scarce, a sorus was moistened with water to which had been added
a drop of 0-1 per cent solution of ‘Alcopol 0’ wetting agent and the selected undehisced
sporangium lifted to the prepared stub. On drying, dehiscence was provoked with
the tip of a needle. Latterly the mountant used was double-sided Sellotape because,
(i) it is easy to apply; (ii) it remains sticky for a considerable time; and (iii) tape plus
coated specimens can be removed from stub at the end of the investigation and
stored for future reference. One disadvantage is that, on drying, the Sellotape latex
may crack and form a distracting background to the specimen. There is also evidence to
indicate that double-sided Sellotape increases the amount of contamination in the

Position 2 en

Revolving platten
with specimen
stubs ve

Support with
motorised drive

Filament 1

Filament 2

: Base of vacuum chamber ;
i m

cae

Filaments connected.in parallel to 10 v. supply

FiGurE 2. Sectional diagram showing the positions of filaments and specimen holders in the
coating unit vacuum chamber.

APPENDIX 15

microscope column. Other mountants, e.g. “Turtox’ clear mountant (water soluble),
‘Durofix’ nitrocellulose cement (benzene soluble) and ‘Araldite’ epoxy resin were used
with equally good effect but did not have all the advantages mentioned above.

COATING THE SPECIMEN

The stubs were coated with gold-palladium or gold metal alone; this latter has become
the standard for botanical material investigated at the British Museum. The metal
wire was evaporated in a vacuum of 10-4 Torr in an Edwards 12E6 coating unit. Approx-
imately 4cm of 36 SWG gold wire were ‘wound around the V-notch '!of each of two
filaments (consisting of approximately 5cm of 25 SWG tungsten wire); these were
simultaneously gently heated, sufficient to melt the gold and then the current was
increased to c 35 amps for evaporation. (The filaments were connected, in parallel, toa

THE DISTRIBUTION OF DRYOPTERIS ASSIMILIS IN BRITAIN

10 volt supply). At all times both the filaments and the gold wire were handled with
‘Velin’ tissue to prevent grease contamination. The stubs were fixed to a platen which
revolved about 5 revolutions per second at such distances and angles from the filament
as to allow even deposition of metal (see Fig. 2). A second evaporation of a similar
amount of gold was then carried out, the angle of incidence of the specimen platen to the
filaments having been altered to position 2 (see Fig. 2).

It is essential that a continuous metal coat covers both stub and spores, and to this
end an improved platen has been designed which gives the stubs an additional planetary
rotation during coating. It is estimated that the total thickness of metal deposited may
be as much as 30 nm from the four filaments. It was found best to coat speciments the
same day as observations were being made and specimens studied over a period of six
months were re-coated as many as three or four times with no obscuring of detail at the
magnifications used (up to x 10,000).

The scanning microscope used was the Cambridge Scientific Instruments Limited
‘Stereoscan’ using an accelerator voltage of 20K V. Photographs were made on Ilford
HP3 film developed in Microdol-X.

Watsonia, 8, 17-26 (1970). 17

The status of

Puccinellia pseudodistans (Crep.) Jansen & Wachter
in Great Britain

By B. M. G. JONES{ and L. E. NEWTON*
Queen Mary College, University of London

ABSTRACT

Puccinellia pseudodistans is described in literature as intermediate between P. distans and
P. fasciculata. It was found to be cytologically and morphologically distinct from P. distans,
but in most characters is similar to P. fasciculata. Experimental work suggests that it is con-
specific with P. fasciculata, the two taxa representing environmentally induced phenotypes with
no genetic distinction. P. fasciculata is the earlier name for the species.

INTRODUCTION

Puccinellia pseudodistans was first described in 1865 by the Belgian botanist
Francois Crépin. Crépin’s material was from the western Mediterranean and
was published as Glyceria pseudodistans Jansen and Wachter, who found the
plant in Spain and in Holland, transferred the species to Parlatore’s genus
Puccinellia in 1935. The first published record of its occurrence in Britain was in
1936, in a paper by Lousley, who had collected specimens at Seasalter, Kent in
1930 and at Grain, Kent in 1935. In the British Herbarium at the British
Museum (Natural History) (BM) there is a specimen collected at Grain, Kent
in 1893 as Glyceria borreri (= Puccinellia fasciculata) and which is annotated
by Lousley ‘earliest British specimen seen’ (i.e. of P. pseudodistans).

Tutin (1962) lists five species of Puccinellia in the British flora. Whilst the other
species are described fully, P. pseudodistans, which is described as ‘more or less
intermediate between P. distans and P. fasciculata’, is dismissed rather briefly.
Tutin also reports that ‘Sterile plants intermediate between P. distans and
P. fasciculata and probably hybrids between the two, occur occasionally where
these species grow together’. Hubbard (1954) also treats P. pseudodistans with
brevity in a work which is of value generally for its detailed accounts of British
grasses. These scant references seem to imply that the authors do not consider
P. pseudodistans to have the same taxonomic status as the other species, and
Tutin has said elsewhere (Tutin 1955) that he regards it as an ‘uncertain entity’.

In published descriptions of the three species P. pseudodistans, P. distans and
P. fasciculata the principal differences are as shown in Table 1 & Plate 1. These
differences were examined critically by investigations on wild populations and
by cultivation experiments. Material was collected from various localities in
South-East England, mostly from Essex and Kent (Newton 1965).

INFLORESCENCE FORM
Four of the characters given in Table 1 concern the form of the inflorescence.
In the field it was found that these characters were quite distinctive, and with a
few exceptions there was no difficulty in determining collected material.

+Present address: Department of Botany, Royal Holloway College, University of London.

* Present address: Department of Biological Sciences, University of Science & Technology,
Kumasi, Ghana.

B

18 B. M. G. JONES AND L. E. NEWTON

Plants of P. fasciculata were always found in very dry habitats, whereas the
other two species always occurred in wet situations. In cultivation, P. fasciculata
responded to a rich soil and abundant water by growing much more vigorously.

TABLE 1. DIAGNOSTIC CHARACTERS REPORTED IN PUBLISHED

P. distans

Panicles symmetrical.

Panicle branches all bare
at base.

Panicle branches fully
deflexed at maturity.

Panicle branches swollen
at base.

Nerves of lemma not
reaching the tips

Anther length:

DESCRIPTIONS.

P. pseudodistans

Panicles symmetrical.

Only longer panicle
branches bare at base.

Panicle branches spreading
but not defiexed.

Panicle branches not
swollen at base.

Middle nerve of lemma
minutely projecting at
tip.

Anther length:
0:-4-0:5 mm

P. fasciculata

Panicles unilateral.

Only longer panicle
branches bare at base.

Panicie branches
obliquely erect.

Panicle branches not
swollen at base.

Middle nerve of lemma
usually minutely
projecting at tip.

Anther length:
0-6-1:0 mm

0:5-1:0 mm

When this species flowered for the first time in cultivation, the inflorescences were
observed to be different in general form from those remaining from the last
flowering season in the wild. The characters of the new inflorescences were
similar to those of P. pseudodistans. One of these plants was then transferred to a
soil-filled earthenware pipe in order to simulate the dry conditions of the habitat,
which was usually a dry bank. The base of the pipe was set in a pan of soil and
apart from an occasional overhead spray, water was given only through the pan.
The panicles produced after transfer to these conditions, especially the later
ones, were similar to those on the original collected plant (Fig. 1).

Further cultivation experiments were then prepared to investigate the plasticity
of the inflorescence characters. Three clones of each of the three species were
used. Three sets of material were planted in four-inch clay pots, and a fourth
set was planted in pipes as described above. All were kept in a greenhouse. One
pot of each clone was kept in a mist unit; a second was placed on a tray of sand
which was permanently moist; a third was placed on an open staging of wooden
slats and watered only once a week. The pipes were placed on the open staging;
they were watered from above once a week, and the soil in the pans was watered
every day. The experiment was set up in March so that the plants would have a
full growing season before flowering in late summer.

Plants in the mist unit produced similar results to those which were on the
trays of damp sand. The plants which were in pots and watered only once a
week showed poor growth, and several died in hotter weather. Plants in the
pipes were healthy, though less vigorous than the well-watered plants.

PLATE |.

Herbarium specimens of Puccinellia

a. P. distans: Swalecliffe, Kent. Wet flush on the side of the sea-wall. 2” = 42
(L. E. Newton No. 53).

b. P. fasciculata: Grain, Kent. Dry ground on sea-wall. 2” = 28 (L. E. Newton
No. 43).

c. P. pseudodistans: Erith marshes, Kent. Muddy waste-heaps by roadside. 2” = 28
(L. E. Newton No. 36).

d. P. pseudodistans: Jaywick, Essex. Mud at side of lane behind sea-wall. (L. E,
Newton No. 58)

i
i
a oT y fi
ni - .

: F i
va (tn 1
if ; bee

i
i
-

PUCCINELLIA PSEUDODISTANS IN BRITAIN 19

FiGuRE 1. The effect of returning a plant of P. fasciculata to drought cultivation. The in-

florescence on the left was developed when water supply was abundant; the inflorescences in

the centre and on the right were formed when water supply was limited. (Drawn from a
photograph in Newton 1965).

Differences of cultivation conditions produced little variation in the in-
florescence characters of P. distans. The four panicle characters given for this
species in Table 1 were manifest in all conditions of cultivation. P. pseudodistans
and P. fasciculata were similar to one another in their different response to the
different conditions. In drought cultivation all plants of both ‘species’ produced
short unilateral panicles with all branches bearing spikelets down to the base.
In wet cultivation they all produced long symmetrical panicles, the longer
branches of which were bare at the base. None had branches with swollen
bases or becoming deflexed.

MORPHOLOGY OF LEMMAS

Fresh flowers from living plants were dissected and examined under a low
power binocular microscope. Floral bracts were flattened for observation by
spreading them with needles on strips of transparent cellulose tape. No appreci-
able difference was observed between flowers at different positions in a spikelet.

Lemmas have rounded apices in P. distans and are subacute in the other two

20 B. M. G. JONES AND L. E. NEWTON

species. The hyaline margin, which is characteristic of all floral bracts in this
genus, was much more prominent at the lemma tips in P. distans than in the
other species (Fig. 2). In P. distans the nerves all stopped well short of the tip

FiGureE 2. Lemmas of P. distans (left), and P. fasciculata (right) compared with a lemma of
P. pseudodistans (centre). (The stippled areas are green, the clear areas membraneous;
all x 20).

of the lemmas, never extending into the hyaline region. In the other species the
central nerve always extended to the apex, as did the region of chlorenchyma, but
was never observed to be excurrent. In all three species, as also in the other two
British species of Puccinellia, the tips of all floral bracts were found to be finely
denticulate (Fig. 3). This is easily visible with the x 40 power of the binocular
microscope but is scarcely visible with a x 10 hand lens. This character appears
not to have been reported before. When an unflattened lemma is examined with
a hand lens the curled denticulate apex can easily give the impression of being
finely mucronate.

Ficure 3. Apex of lemma in P. distans (x 120). (Drawn from a photograph in Newton, 1965.)

PUCCINELLIA PSEUDODISTANS IN BRITAIN p2)|

ANTHER SIZE

Lousley’s (1936) measurements for P. pseudodistans are quoted by Tutin
(1962), but he and Hubbard (1954) give different ranges for P. distans:

P. distans P. pseudodistans P. fasciculata
Tutin 0-5-0:75 mm 0-4-0:5 mm 0-75-1:0 mm
Hubbard 0-8-1:(0 mm — 0-6 -1:(0 mm

Anther length was determined from flowers collected in the field and also from
flowers produced by plants in the cultivation experiment. In all cases the
measurements were made on exserted but undehisced anthers, and were
obtained with a low power microscope and camera lucida. The results (Newton
1965) reveal so much overlapping of ranges for the three species as to indicate
that anther length is not a reliable diagnostic character for individual plants.
Observations upon anthers from plants used in the cultivation experiments
indicate that water supply has some effect on anther length (Fig. 4).

— X R— S8—W
} P.distans.
——— X e—— 8-—D

— X ——. 30—W
P. fasciculata.
—+ xX He— 30-D

— xr— 1-—W
P.pseudodistans.
————4 X »-———_ 1-D

———— + X H— 12-—W
P.pseudodistans.
———————— | ———————— 12

a SSS
0.4 0.5 0.6 0.7 0.8 0.9

millimetres.

W = ‘wet cultivation’ D = ‘dry cultivation’
The numbers are reference numbers of the clones.

Ficure 4. Anther length of P. distans, P. fasciculata and P. pseudodistans at the conclusion of
the cultivation experiment. (Means, twice SE and ranges are shown).

LEAF EPIDERMIS

In his work on the Greenland species of Puccinellia, Sorensen (1953) found
that characters of the leaf epidermis were of taxonomic value. He found that
if the examination was made at a similar position on each plant, the variability of
the characters observed was no greater than that of the floral parts, and he was
able to prepare an identification key for the Greenland species based entirely
upon the structure of the leaf epidermis. None of the three British species under
discussion here occurs in Greenland.

Metcalfe (1960) examined only P. fasciculata and P. maritima in his anatomical
survey of the Gramineae. His descriptions for these two species differ in only
one character, viz. the presence of ‘macro-hairs’ on P. fasciculata and their
absence from P. maritima.

22 B. M. G. JONES AND L. E. NEWTON

In a paper concerning the use of epidermal characters in taxonomy, Tutin
(1955) compared the leaf epidermis of P. distans with that of P. pseudodistans,
which he had previously regarded as an uncertain taxon. No conclusion is
stated in the paper, but differences between the two are described and the text
implies that the author regarded these two taxa as distinct species. The principal
differences given by Tutin are shown below.

P. distans P. pseudodistans
Epidermal cells Relatively short and broad Long and narrow
Stoma length 30-35 um 30-35 um
Stomatal pit Somewhat cruciform Oval
Broader than long About twice as broad as
long
Stomatal flap | Overlaps guard cells obliquely Overlaps guard cells sym-
metrically

Longer than in P. distans

P.distans Dry —_—_<—<_<—_{ X }—___—_

(Ref. No. 8) Wet —_————4— X -——

P.fasciculata Dry SS ee

(Ref. No. 7) Wet SEER eed

P.pseudodistans Dry —————————

(Ref. No. 1) Wet et eee

P.maritima Wet yt FX

(Ref. No. 3)

P.rupestris Wet 1x?

(Ref. No. 21)
eee eS Se eee ee ee ee,
26 30 34 38 42 46 50

(um)

FiGcure 5. Guard cell lengths from culm leaves of Puccinellia species in cultivation experiments.
(Means, twice SE and ranges are shown).

Observations were made on the uppermost mature culm leaves of plants in the
cultivation experiment at least six months after the experiment was set up. The
leaves were boiled in ethyl alcohol for a few minutes and then in water. Small
portions from the middle of the leaf were mounted in lactophenol for examina-
tion.

The size of the ‘long-cells’ was not found to be a useful diagnostic feature, and
it is clear from the literature that it is not reliable. Tutin described the cells of
P. distans as ‘relatively short and broad’, whilst Sorensen describes the pidermis
of this species as ‘narrow-celled’. The ranges of stoma length were found to be

PUCCINELLIA PSEUDODISTANS IN BRITAIN 23

somewhat wider than those given by Tutin, and were of no diagnostic value
(Fig. 5). The shape of the stomatal pit varies considerably on any one leaf
according to the nature and position of adjacent cells, and it is not always easy to
apply the terms ‘cruciform’ and ‘oval’ as used by Tutin. The way in which the
stomatal flap overlaps the guard cells was not a consistent character. Some
distinct differences were observed, however. Papillae were fewer on P. distans
than on the other two. The cell walls were markedly sinuous on P. distans,
whereas those of the other two species were only slightly rippled. The cell walls
in P. fasciculata and P. pseudodistans had a conspicuous pitted appearance which
was not seen in P. distans. These observations support the apparent view of Tutin
that P. pseudodistans and P. distans are morphologically distinct, but go further in
showing a similarity between P. pseudodistans and P. fasciculata.

TABLE 2. NEW CHROMOSOME COUNTS ON NATIVE BRITISH
PUCCINELLIA SPECIES.

Species Origin Somatic chromosome number

P. distans N. of Leysdown, Kent 2n = 42
Swalecliffe, Kent DAD
Reculver, Kent 2n = 42
Jaywick, Essex 2ne— 42
P. fasciculata Allhallows, Kent 28
Grain, Kent 2 28
Shellness, Kent Dip IS
Wakering Stairs, Essex Zi 28
Jaywick, Essex 2ne— 25
P. pseudodistans Erith Marshes, Kent 2i— 28
Shellness, Kent 2n = 28
Wakering Stairs, Essex 2n— 28
Jaywick, Essex 2n-— 28

Between Burnham-on-Crouch and
Creeksea, Essex 228
P. rupestris Swalecliffe, Kent 2n = 42
Reculver, Kent 2n = 42

CYTOLOGY

Previous chromosome counts for British material of these species are as follows:

P. distans Dn 42 Lovis, in Tutin, 1955.
P. fasciculata Di 2S Rutland, 1941.
P. pseudodistans Dis Lovis, in Tutin, 1955.

Root tips from plants used in the present study were given a pretreatment in a
0.002 M solution of 8-hydroxyquinoline for three hours at 10°C, fixed for one
hour in 45 per cent. acetic acid, hydrolysed for 8 minutes in N/1 HCI at 60°C.,
stained in leucobasic fuchsin, and squashed in acetic carmine. The results are
given in Table 2. In all cases the counts obtained confirmed the earlier counts
quoted above.

24 B. M. G. JONES AND L. E. NEWTON
FLORAL BIOLOGY AND HYBRIDISATION

Most Puccinellias begin to flower in June, but the main flowering period is
July-August. Flowers on the plants in cultivation began to open at about
5.30 a.m. Anthers and stigmas started to emerge within an hour, and dehiscence
of the anthers was completed in most cases by about 8 a.m.

In P. distans the anthers protrude prominently in the manner expected of an
anemophilous plant. In the other two species, however, the anthers were
hardly clear of the lemma and palea when dehiscence occurred. Stigmas protrude
at the same time as the anthers and it seems most likely that self-pollination
occurs in these cases, with the pollen dropping on to the the stigmas of the
same floret or other florets in the same spikelet. Bagged inflorescences on each
species produced abundant seeds which germinated freely soon afterwards.
Several flowers on each species were emasculated immediately before anthesis
and bagged; none of these set seed.

Several plants were collected from wild populations as possible interspecific
hybrids because they were not easily determined on inflorescence characters.
At one locality (Jaywick, Essex), plants answering the descriptions of all three
species were found growing together in a small area, with some plants whose
characters were somewhat intermediate. All these plants were found to have a
pollen fertility of over 80 per cent. Those readily determined as P. distans had a
chromosome number of 2” = 42; all other plants in this collection were
Dig) == Ds),

ECOLOGY AND DISTRIBUTION
The Puccinellias are generally referred to in British literature as salt-marsh
plants, and they are listed under ‘Salt-marshes and Coastal mud flats’ in
Hubbard’s habitat lists (1954). Early field excursions in search of material soon
showed that this habitat description is misleading. Only P. maritima was found
on inter-tidal salt-marshes; the other species were always found well above
high water level.

P. distans and P. pseudodistans occur mostly in wet, muddy situations. In
contrast, P. fasciculata is found in dry places, as also is P. rupestris (Jones &
Newton 1965). Analysis of soil and mud samples from some localities showed
that P. distans and P. pseudodistans occur in poorly drained soils and P. fascicu-
lata grows in well drained soils. All are salt tolerant, but none is an obligate
halophyte as all survived with fresh water cultivation for up to three years.
Seeds germinated readily when supplied only with fresh water. It is interesting
to note that at the locality where all three species were found together, referred
to in the previous section, P. distans and P. pseudodistans were in silty soils in
the drip-zone of rain water from the top of a wall against which they grew,
whilst the P. fasciculata plants were growing in sandy shingle in a drier situation
further from the wall.

These plants were never found to be dominant in a mixed community, and
usually there was little or no competition from other plants. Results of a
competition experiment with the eminently successful weed Poa annua, in
which plants were grown together in pans, suggest that these species
cannot withstand competition. Field observations on collecting excursions
suggest that they appear mainly as pioneer plants on disturbed ground, suc-
cumbing later when other plants invade the area. Coastal erosion, common in

PUCCINELLIA PSEUDODISTANS IN BRITAIN DS

south-east England where these species are mostly found, and human activity
would provide the temporary habitats in which these relatively uncommon
species are able to grow.

DISCUSSION

Although P. pseudodistans is described in literature as intermediate between
P. distans and P. fasciculata, when they are encountered in the field all three
taxa normally present no difficulties in identification. They are readily deter-
mined on the basis of inflorescence form alone, and all the character differences
for the panicle given in Table 1 were found to be reliably diagnostic.

P. distans and P. pseudodistans are similar in their habitat preference, and the
panicle differences were still evident on plants in varying conditions of the
cultivation experiment, in spite of the greater plasticity of P. pseudodistans. A
detailed comparison (Newton 1965) showed that the two taxa are cytologically
and morphologically distinct.

As well as having distinctive panicle forms, P. fasciculata and P. pseudodistans
normally occur in different habitat conditions. In morphological characters
other than the panicle form, however, they were found to be remarkably
similar. In the cultivation experiments the two panicle forms were shown to be
extremes of a phenotypic plasticity which was parallel in the two taxa. The plasti-
city was induced by variation in water supply, and characters appearing in the
cultivation experiment corresponded to those found in similar water supply
conditions in habitat. When grown in similar cultivation conditions the two
taxa could not be distinguished. This leads to the conclusion that the two taxa
as found in the field represent environmentally induced phenotypes of one
species.

Continental material has not been investigated extensively, but some plants
collected by one of us (L.E.N.) as P. pseudodistans on the Mediterranean coast
showed a similar response to the cultivation experiment as that described for
the British material. It seems likely, therefore, that Continental plants cor-
responding to Crépin’s original material will be found to have the same status
as the British material.

The small size of the anthers in P. fasciculata and P. pseudodistans might
account for reports of sterile plants being found. There are two main reasons
why a particular plant may have smaller anthers than in related species, viz.
pollen abortion, often an indication of hybridity, and the occurrence of regular
self-pollination. Both in cultivated plants and in plants sampled in the field
there was no indication that small anthers were associated with reduction in
fertility. Whilst both Hubbard (1954) and Tutin (1962) say that sterile plants
occur in the wild, neither states how it was determined that the plants were
sterile. Herbarium specimens in BM labelled as hybrids and having small
anthers were found to have no indication of pollen sterility. Use of small anther
size as an index of pollen sterility is clearly misleading.

That self-pollination may account for the small anthers is suggested by other
observations. It was noted earlier that at anthesis the anthers protrude very
little from between the bracts and are not well exposed for wind-pollination.
Both taxa are described in literature as perennial, and they lived for three years
in cultivation. In the wild, however, they often behave as annuals; dead plants
were frequently found at the end of the flowering season, mainly in places where
the substrate had dried out. It is known that annual plants in north temperate

26 B. M. G. JONES AND L. E. NEWTON

regions are almost exclusively self-pollinated (Stebbins, 1960). It was found that
the Puccinellias (except P. maritima) are confined to situations where dis-
turbance creates a fresh substrate whose salinity inhibits rapid colonisation by
other plants. Self-pollination is probably a selective adaptation for survival in
this unstable ecological niche.

TAXONOMIC CONCLUSIONS

It is concluded that P. pseudodistans and P. fasciculata are ecads of one species.
However, as they appear in the field and herbarium as strikingly different
phenotypes with few intermediates, it seems useful to distinguish between them
at the level of forma. The nomenclature of the taxa investigated is as follows:

PUCCINELLIA DISTANS (L.) Parl., Fl. Ital., 1: 367 (1848).
Poa distans L., Mant., 1: 32 (1767).
Glyceria distans (L.) Wahlenb., Fl. Upsal: 36 (1820).
Sclerochloa distans (L.) Bab., Man. Brit.: 1., 370 (1843).
PUCCINELLIA FASCICULATA (Torr.) Bickn., Bull. Torr. Bot. Club, 35: 197 (1908).
forma FASCICULATA
Poa fasciculata Torr., Fl. N. Am.: 107 (1824).
Glyceria borreri Bab., Suppl. Engl. Bot., t.: 2797 (1837).
forma pseudodistans (Crép.) Newton & Jones, comb. nov.
Glyceria pseudodistans Crép., Nouv. Rem. Glyceria Gr. Heleochloa,
15 (1865).
Puccinellia pseudodistans (Crép.) Jans. et Wacht., Nedl. Kruidk. Archief.:
14: 10 (1935).

Voucher specimens of material used in this investigation have been lodged
with the British Museum (Natural History) (BM).
This work forms part of the material submitted by L. E. Newton for the degree
of M.Sc. in the University of London.

REFERENCES

HUuBBARD, C. E. (1954). Grasses. Harmondsworth.

Jones, B. M. G. & NEwrTon, L. E. (1965). Confusion of Puccinellia rupestris (With.) Fernald &
Weatherby and Poa annua L. in the field. Proc. bot. Soc. Br. Isl., 6: 171-172.

Lous.Ley, J. E. (1936). Puccinellia pseudo-distans (Crépin) Jans. & Wacht. in Britain. J. Bot.,
Lond., 74: 260-266.

METCALEE, C. R. (1960). Anatomy of the Monocotyledons, vol. 1. Oxford.

Newton, L. E. (1965). Taxonomic Studies in the British Species of Puccinellia. M.Sc. thesis,
University of London.

RUTLAND, J. P. (1941). The Merton Catalogue, A List of Chromosome Numbers of British
Plants. Suppl. No. 1. New Phytol., 40: 210-214.

SORENSEN, Th. (1953). A Revision of the Greenland Species of Puccinellia Parl. Meddr Gronland,
136(3): 1-179.

STEBBINS, G. L. (1960). Variation and Evolution in Plants. New York.

TuTIN, T. G. (1955). Species problems in plants with reduced floral structure. B.S.B.JI. Conf.
Rept. 4: 21-26.

TuTin, T. G. (1962). Puccinellia Parl. in CLAPHAM, A. R., TUTIN, T. G. & WARBURG, E. F.,
Flora of the British Isles, 2nd ed., pp.1133-1135. Cambridge.

Watsonia, 8, 27-40 (1970). 27

A Biosystematic and Ecological Study of Salicornia in
the Dee Estuary

By P. W. BALL and K. G. BROWN
The Hartley Botanical Laboratories, The University, Liverpool

ABSTRACT

Populations of Salicornia in the salt marshes of the Dee Estuary (Cheshire, England) are
composed of Salicornia europaea L. (2n = 18) and S. dolichostachya Moss (2n = 36) growing
intermingled. It is shown that nearly all individuals can be identified by means of a number of
morphological features, but anther-size is the most reliable character. S. dolichostachya was
the commoner species in the most open habitats. It was found that the radicle of seedlings of
S. dolichostachya elongated more rapidly than that of S. europaea and it is suggested that this
may to some extent explain the greater frequency of S. dolichostachya in the open habitats.
S. europaea was more frequent than S. dolichostachya in areas where there was a relatively high
cover of Puccinellia maritima (Huds.) Parl. and Spartina ‘townsendii’, and it was found that
many individuals of S. dolichostachya died without producing seed in these areas. It would
seem that S. europaea is better able to withstand competition from perennial grasses than is
S. dolichostachya.

INTRODUCTION

The genus Salicornia has always presented considerable taxonomic difficulty
and, despite extensive investigations in a number of European countries in
recent years, there is no general agreement as to the number of species which
occur in Europe. Whilst it is usually possible to recognise a number of taxa in a
given locality, difficulties often arise when attempts are made to correlate the
taxa from different localities. Because of these taxonomic difficulties most
ecological data have been recorded under the generic name only, or under a
collective specific name. In those cases where attempts were made to be more
precise, the information is of rather uncertain value as the criteria used in the
past to recognise species have been shown to be very unsatisfactory (Ball &
Tutin 1959, Ball 1960). The observations presented here are an attempt to
combine a morphological study with a preliminary autecological investigation.
Because of the necessity to examine each individual with some care, the quantity
of material used was relatively small. This was partly due to the time needed to
record the data, and partly to a desire not to deplete the numbers of Salicornia
plants in the areas investigated, since each sample examined had to be removed
from the marsh.

LIFE HISTORY

The life-history of Salicornia is very similar to that of many other annual
Chenopodiaceae. Seeds usually germinate in April or May and the plants grow
vegetatively until July or August, when fertile segments are produced, and
flowering commences soon after this (see Ball & Tutin (1959) for a detailed
morphological description). By the end of August to mid-September the plants
have ceased to grow and they have usually completed flowering by the end of
September, although a few flowers can often be found until well into October.
Seed reaches maturity from about mid-September onwards when the plants
begin to die. Seed is usually shed by falling out of the dead or dying parent plant,

28 P. W. BALL AND K. G. BROWN

but it is not unusual to find that the seeds have not been shed and have germina-
ted in situ the following spring, by which time the parent has often fallen over
and been buried.

MORPHOLOGICAL INVESTIGATIONS

The salt marshes of the Dee estuary in Cheshire are of recent origin and are
still expanding rapidly. The area that has been examined in some detail is that
between Parkgate and Neston (Grid ref. SJ 2777). This marsh is known to have
developed since about 1930 and it now extends for up to 1 km beyond the
seawall. Salicornia occurs mainly towards the lower edge of the marsh and in
many places is colonizing open sand or muddy sand. A small number of plants
were examined in 1955 and the population has been examined each year since
then by one of us (P.W.B.). This population is composed of two species, identified
as S. europaea L. (2n = 18) and S. dolichostachya Moss (2n = 36) by Ball &
Tutin (1959). Table 1 summarizes the morphological differences between the

TABLE 1. COMPARISON OF SALICORNIA EUROPAEA AND S. DOLICHOSTACH YA
(AFTER BALL & TUTIN 1959).

Character S. europaea S. dolichostachya
Colour Often red or purple Green or yellow, rarely reddish
Angle between mainstem __c. 45°, branches straight 45-90°, branches curving
and base of uppermost upwards
branches
Terminal spike (2-)5-9(-16) fertile segments (7-)12-25(—30) fertile segments
Fertile segments
Length (2:2-)2-5-4(-5:5) mm 3-6 mm
Diameter at base 24:5 mm 3-6 mm
Maximum diameter (2:5-)3-5-6 mm 3-6 mm
Shape A, rarely B (Fig. 2) C, rarely B (Fig. 2)
Central flower 1-7-2:5 x 1:7-2:5 mm 2:44:0 x 1:9-2:°9 mm
Lateral flowers 1-6-2:0 x 1-3-1-8 mm 1-8-3-0 x 1-5—2-:3 mm
Stamens Usually 1, exserted or not, 1-2, always exserted, not de-
usually dehiscing before ex- hiscing before exsertion
sertion
Anther size 0-3-0:5 mm (0-5—)0-6-0-9 mm
Pollen grain diameter (20—)24—28(-31) um (27—)31-34(-42) pm
Stomata (length of guard (20-)24-30(-33)yum (27-)29-36(-42) um
cells)
Seed size 1-0-1-7 mm 1-5-2:3 mm
Chromosome number A= as

two species, and it can be seen from this that no single character apart from the
chromosome number gives an absolute distinction between them. The chromo-
some numbers were determined on a small number of plants collected in 1955
and 1956, and in 1957 a large sample was collected in order to obtain further
information about the variation that occurred. Figure 1 illustrates some of
the results obtained from this sample. The two characters selected—the number
of fertile segments in the terminal spike and the ratio ‘maximum width of the
fertile segment/width at the base of the fertile segment’—give a clear division
into two taxa. The ratio chosen corresponds approximately to the fertile
segment shapes used in the 1966 study (Fig. 2). Plants with a ratio of 1-17 or

A BIOSYSTEMATIC AND ECOLOGICAL STUDY OF SALICORNIA 29

Fertile segment
ratio

=

1:50

e ae
‘ @ =2n=18
1-40 338 4 =2n=36
oltgi2'g
pa foanin® $84
2S
wee a
1:2 3 i :
on eo * ge
1.10 & o 23 e,
2
ry @® @@
i “3 e “beg 4 No. of fertile segments
8 e in terminal spike

1-00

FicureE 1. Scatter diagram of sample collected in 1957 and individuals of known chromosome
number collected in 1955-6.

“() (A al

A(a) B(A) C(X)

FiGureE 2. Fertile segment shapes used in the 1966 study.

more correspond to Shape A, those with a ratio of 1.12 or less correspond to
Shape C, the intermediate ones being similar to Shape B. The plants whose
chromosome numbers were determined in 1955 and 1956 have also been included
on this diagram, and it can be seen that the difference in chromosome number is
correlated with the morphological separation. In the revision by Ball & Tutin
(1959) it was emphasized that damaged and depauperate plants often presented
considerable difficulties and they were excluded from these samples.

In 1966 it was decided to make a comparative ecological study of the two
species. It was apparent from the previous investigations that they were growing
side by side in the lower parts of the marsh although the proportions of each
varied considerably from one habitat to another. If the results obtained were to
have any validity then it would be necessary to identify all individuals in any
condition. The first part of the 1966 investigation was therefore aimed at checking
the reliability of a variety of characters that might be used to identify the
individuals in this particular population. Two main areas (A and B) were
selected for sampling and small samples were also taken from two others

30 P. W. BALL AND K. G. BROWN

(C and D). Area A was a low mound, about 140 metres in diameter, on the
seaward side of the marsh. It was surrounded by broad shallow channels on all
sides, these having Puccinellia maritima—Spartina marsh on the opposite bank
on 3 sides and almost bare sand and mud on the fourth side. The vegetation on
the central part of the mound comprised a dense sward of Suaeda maritima,
with up to 2000 plants per sq. metre, and relatively few Salicornia plants. Around
the perimeter was a more open community of Salicornia and Suaeda maritima.
Area B was a transect taken from the lower edge of the marsh from bare sand
through an open Salicornia community into a closed Puccinellia maritima—
Spartina marsh. Area C was about half-way between the edge of the Puccinellia
maritima—Spartina marsh and the mean high water level, whilst Area D was at
about the maximum high water level with Festuca rubra as the dominant species.
Areas C and D contained only S. europaea, so although they were of interest
when examining the variation found in this species they were not used for the
later comparative studies. For the detailed investigations a 2-metre quadrat in
the middle of a transect through Area A was chosen and a sample of 50 plants
taken. Another sample was taken from the perimeter of A, and others were
obtained by means of random 0-5 metre quadrats. Similar samples were taken
from Area B. Small collections were also made from Areas C and D. The
previous studies of Salicornia in this locality had indicated that only a limited
number of characters were likely to be of value in separating the two species so
only the following data were recorded: (1) height of plant (ii) total number of
branches, or total number of fertile segments (111) number of fertile segments in
the terminal spike (not recorded if damaged) (iv) length of the 2nd—4th fertile
segments in the terminal or a major lateral spike (v) shape of the fertile segments
(A, B or C, Fig. 2) (vi) anther length (when available) (vii) length of guard
cells of stomata (epidermis taken from the sterile segment at the base of the
terminal or a major lateral spike). Figures. 3-7 were prepared from this data.

9
Anther 1966 (All samples)

length (mms)

7

08+ 2 @ ® e
ee
a e ee ee e © ee
® @ oe
0-6 ecst «68 000 Se e
r e

0-5 ed e &
O4+ © eSeeceed @ 6

ee e rv)
0:3 gee cos © 0

Stomata
19 20 21 22 23 24 25 26 27 28 size

FIGuRE 3. For legend see p. 31

A BIOSYSTEMATIC AND ECOLOGICAL STUDY OF SALICORNIA 31

FIGURES 3-7. Pictorial scatter diagrams of samples collected in 1966. Stoma size is measured in
arbitrary units.
+ = 2n = 36 | = 2n = 18
A = Fertile segment shape A (Fig. 2)
A = Fertile segment shape B (Fig. 2)
x = Fertile segment shape C (Fig. 2)

1966 (All samples)
Anther length (mms)

0-9
0-8 xX ef
A
0-7 4 A & x x g x
: Ae Se X x
06 ISS ee x
x x
K
x
x
a“ ray
A
6:4 ahkhabshane ana A
Aa A A
ae
4 a aA aA A
‘ : Bd
0-3 bs A A a A A
A
No. of
fertile) ccoments
in termina
5 10 15 20 epike
FIGURE 4. For legend see above
1966 (All samples)
Ht. of plant (cms)
35
39
25
20
15
10 No. of
fertile segments
in terminal
5 10 15 20 spike

FiGure 5. For legend see above

32 P. W. BALL AND K. G. BROWN

1966 (Sample A45)

Anther
length (mms)

0:8:

Ht. of plant (cms)
10 15 20 25 30 35 40

Ficure 6. For legend see p. 31

1966 (All samples)

No. of
fertile segments
in terminal
spike
20
x
X XXy XX
x XX x
15 x x x
x
x Kx
x A

A 7a a& x x

ee a A ax
ya
Assh A 4 x
AA A A A

Ab Adbsda dA

5 A A&bADAAAA
7a. AAS AAAAAAA
rN AA A
Stomata
19 20 21 22 23 24 25 26 27 28 size

FIGURE 7. For legend see p. 31

It will be seen from these diagrams that the height of plant is not correlated
with anther size or fertile segment shape, whilst anther size, stomatal size and fer-
tile segment shape are all strongly positively correlated. There is a weaker
positive correlation between these 3 characters and the number of fertile
segments in the terminal spike. There is also some correlation between height

A BIOSYSTEMATIC AND ECOLOGICAL STUDY OF SALICORNIA 33

of plant and the number of fertile segments in the terminal spike although the
pattern of variation is different for each species.

Figures 4 and 7 confirm that, by means of a combination of fertile segment
shape and the number of fertile segments in the terminal spike, it is possible to
identify the majority of individuals in the field. Where any doubt arises this can
almost always be resolved by anther length or stomatal size, or a combination of
these two measurements. Where anthers were available it was found that the
distinction was almost complete, S. dolichostachya having anthers (0:-45—)0-5 mm.
long or more, whereas S. europaea had anthers 0-4 mm long or less.

Transect A

Over 200
150
No. of
Salicornia 100
individuals
70
50
40
30 a)

% cover
80

Suaeda
maritima

2m Quadrats

Ficure 8. % cover of Suaeda maritima through Transect A and the frequency of Salicornia
dolichostachya (O) and S. europaea (A).

Cc

34 P. W. BALL AND K. G. BROWN
ECOLOGICAL INVESTIGATIONS

Once it had been demonstrated that it was possible to identify the vast majority
of individuals in this locality, an attempt was made to compare the
detailed distribution of the two species in the areas studied, and to compare
their performances in terms of dry weight and seed production. Transects
2 metres wide were taken through Areas A and B; that through A was 120
metres long, running roughly at right angles to the shoreline, and that through
B was 22 metres long. (In A the first 21 2-metre quadrats were examined, but
after this only every 3rd quadrat was examined; in B every quadrat but one
was examined.) The percentage cover of each species was recorded, but for
Salicornia the total number of individuals of each species was also noted. The
results obtained are illustrated in Fig. 8.

In Area A, Salicornia dolichostachya occurred throughout. In the part of the
transect with large areas of bare ground (quadrats 1-16, 62-end), this and
Suaeda maritima were the only species present. The density of Salicornia and
Suaeda increased in quadrats 17-21, S. dolichostachya still being the commonest
species, but S. europaea appearing more frequently. In the central region
(quadrats 36-48) S. europaea became more frequent, both in number of indi-
viduals and as a proportion of the total population of Salicornia. In those
quadrats with very high (c. 90 per cent.) Suaeda cover the number of individuals
of Salicornia was extremely low and they were often difficult to find.

Transect B

*% cover

Puccinellia maritima
and Spartina

100
80
60
40

20

No. of Salicornia
individuals

over 200

2m Quadrats

Lay reat Sewanee Ole Om ia. aS 27/9 oral Oneal

FIGURE 9. Combined % cover of Puccinellia maritima and Spartina through Transect B
and the frequency of Salicornia dolichostachya (O) and S. europaea (A).

A BIOSYSTEMATIC AND ECOLOGICAL STUDY OF SALICORNIA S)3)

- Whilst it is impossible to draw any definite conclusions from these data a
number of points should be noted. Firstly, S. dolichostachya occurred in most
parts of the areas sampled; secondly, S. europaea did not occur where there was
less than 40% Suaeda cover; thirdly, the largest number of Salicornia individuals
occurred in those areas where the Suaeda cover was between 40 and 70%
(in only one case up to 85 per cent.); fourthly, with Suaeda cover in excess of
85%, Salicornia virtually disappeared.

Transect B gave a somewhat similar picture, but with some slight variations
(Fig. 9). On the open sand and mud the number of individuals was extremely
low, there being only 5 plants of S. dolichostachya and 2 of S. europaea in the
first three 2-metre quadrats. A larger sample collected nearby for morphological
examination comprised 20 individuals of S. dolichostachya and 4 of S. europaea.
In the quadrats where Puccinellia maritima and Spartina first appeared, the
number of Salicornia individuals increased considerably, but where the com-
bined Puccinellia-Spartina cover exceeded 50 per cent. the number declined.
S. dolichostachya showed a more rapid decline than S. europaea, and it was
noticed that in mid-September many individuals of S. dolichostachya in quadrats
6-8 were in a very poor condition and appeared to be dying without producing
seed. The last quadrat of this transect contained only S. europaea.

Areas C and D were isolated from the main populations of Salicornia and
both contained only S. europaea. In both cases the area containing Salicornia
did not greatly exceed one 2m quadrat so only the one could be examined in
each. In Area C it was found that Puccinellia maritima was the dominant
species with 80 per cent. cover; in Area D Festuca rubra was the dominant
species with 75 per cent. cover.

The above information was obtained in mid-September when many Salicornia
plants were still in flower and very few contained ripe seed. During October some
attempt was made to investigate the variation in size and seed production of the
two species in different parts of these habitats. Further samples were obtained by
means of random 0:5-metre quadrats. All individuals were collected from each
quadrat and an estimate of the seed production obtained by counting the total
number of fertile segments (Tables 2 and 3). Previously it had been shown that

TABLE 2. VARIATION OF MEAN NO. OF FERTILE SEGMENTS PER
SALICORNIA PLANT WITH % COVER OF SUAEDA MARITIMA IN AREA A.

% COVver Mean no. of fertile segments per plant
Suaeda maritima S. europaea S. dolichostachya

10 676 967

15 640 118

20 — 394

45 176 83

50 37:5 DS)

50 40 49

70 58 53

there was no significant difference between the two species in the number of
seeds in each fertile segment, the mean ranging from 2-8 to 3-8 seeds per segment.
When these samples were taken it was found that all of the S. dolichostachya in
areas with more than some 50 per cent. Spartina-Puccinellia cover had died,

36 P. W. BALL AND K. G. BROWN

TABLE 3. VARIATION OF MEAN NO. OF FERTILE SEGMENTS PER
SALICORNIA PLANT WITH % COVER OF PUCCINELLIA MARITIMA AND
SPARTINA IN AREAS B AND C AND WITH FESTUCA RUBRA IN AREA D.

% Cover Mean no. of fertile segments per plant
Puccinellia maritima S. europaea S. dolichostachya
and Spartina
0 245 544
0 — 451
30 187 47
40 27 65
75 (Festuca rubra) (D) 53 _
80 (C) SV —

and it seemed doubtful whether they had produced any seed. Thus, in contrast to
S. europaea, under these conditions S. dolichostachya appeared to fail com-
pletely.

The difference in distribution of the two species of Salicornia raised a number
of problems. On the open sand and mud the individuals of both species were
extremely large and well-grown and produced a large quantity of seed. The
observations by Wiehe (1935) in the Dovey estuary, Cardiganshire, indicated that
in a habitat where only Salicornia occurs the density of the individuals was
closely related to the frequency of submersion by the tide. This was related to the
effect of movement and drag on seedlings and young plants, the critical period
being 2—3 days undisturbed by tides. From the photographs published by Wiehe
it appears that what is probably S. dolichostachya predominated in the area
where there was frequent disturbance, while both S. dolichostachya and
S. europaea (or another diploid species) occurred in the dense Salicornia
community.

This suggested that germination rate and the initial rate of growth of the
seedlings might explain the difference in frequency between S. dolichostachya
and S. europaea on the open sand in the Dee estuary.

GERMINATION EXPERIMENTS

Ten plants of each species were selected at random and stored in polythene
bags at 2°C for a short period. The spikes were then broken up into small
pieces and divided into six parts. These were given different treatments prior to
germination as follows:

(a) Frozen for 7 days in a sealed container.

(b) Stored at 10°C for 7 days in a sealed container.

(c) Soaked in 100 ml of distilled water for 7 days at 10°C.

(d) Soaked in 100 ml of 50% filtered seawater (made up with distilled water)
at 10°C for 7 days.

(e) Soaked in 100 ml of 100% filtered seawater at 10°C for 7 days.

(f) Seeds extracted from the segment, dried at 25°C and then kept at 10°C
for 7 days in a sealed container.

After treatment 25 seeds were placed in vermiculite soaked in distilled water
in petri dishes, there being four replicates of each. Daily records were kept of the
number of seeds that had germinated. The results obtained are shownin Figure 10.

A BIOSYSTEMATIC AND ECOLOGICAL STUDY OF SALICORNIA 37

S. dolichostachya
“% Germination

100

ps

S. europaea

100

Days after
6 12 16 SOWING

Ficure 10. Germination of Salicornia seeds kept under various conditions.

Kept at 10°C (Treatments (b) and (c) combined)
Soaked in 100% seawater

Soaked in 50% seawater

Extracted and dried at 25°C

Frozen for 7 days

OP pe x
HTT Te a

In general, the treatments do not affect the rate of germination or the total
percentage germination of either species over a period of 16 days. The only
exception to this was treatment (f) which with S. europaea gave a higher germina-
tion rate and a higher percentage total germination than was obtained for the
other samples of S. europaea. A surprising feature of these results was the
consistently lower rate of germination and the lower total germination of
S. europaea as compared with S. dolichostachya, except for treatment (f) in
which S. europaea was almost identical with S. dolichostachya. These experiments
were performed in the late autumn of 1966 and it was too late to obtain further
seed samples to check these results in the spring of 1967. Plants were therefore
collected from Area A in the autumn of 1967 and stored in polythene bags at
2°C until the spring of 1968 before any further experiments were carried out.
These experiments, which were primarily concerned with measuring the rate
of growth of the newly germinated seedlings, gave much quicker and almost

38 P. W. BALL AND K. G. BROWN

identical rates of germination (87 per cent. for S. europaea and 90 per cent.
for S. dolichostachya after 4 days) for both species. The most likely explanation
for the results obtained in the autumn of 1966 is that the 1966 seeds of
S. europaea were not fully mature. It has been noted by a number of authors,
e.g. Ball & Tutin (1959), that S. dolichostachya flowers slightly earlier than
S. europaea, and it is likely therefore that the seeds of S. dolichostachya also
mature earlier.

GROWTH OF SEEDLINGS

A number of plants of both species were collected from Area A in October
1967 and stored in polythene bags at 2°C until the end of February 1968. By
this time the fleshy parts of the plants had decayed so it was not possible to mix
the broken spikes at random. As far as possible they were cut into pieces
containing only one fertile segment and 10 pieces were placed in vermiculite
soaked in distilled water in petri dishes. 6 replicates of each species were prepared,
and they were placed in a heated greenhouse kept at about 21°C. To avoid
confusion only one seedling from each segment was allowed to grow. The day
on which a seed germinated was recorded and 4 days later the lengths of the
plumule and radicle were recorded separately. The results obtained are shown
in Fig. 11. Whilst there is a small difference in the mean length of the plumule of

Plumule Radicle
No. of
Individuals

| S. europaea

S. dolichostachya

.

5
5

Ficure 11. Frequency distribution of the length of the plumule and radicle of Salicornia
seedlings 4 days after germination.

S. europaea and S. dolichostachya (8-9 mm against 9-5 mm), the total range of
variation found in the two species is almost identical. The difference in mean
radicle length is much more marked (3:7 mm against 4-9 mm). The results

A BIOSYSTEMATIC AND ECOLOGICAL STUDY OF SALICORNIA 39

obtained from both species were distorted by the fact that in many seedlings
the radicle failed to elongate at all or it elongated by a small amount and then
shrivelled. Most radicles 2 mm long or less were shrivelled, and so too were
some longer radicles. No satisfactory explanation of this phenomenon could
be found. Whilst it often occurred in instances where the radicle was exposed to
the light, this was not always so. The seedlings were always in direct contact
with wet vermiculite and the humidity in the petri dishes was high, there being
droplets of water permanently on the lid. In a few cases in which the radicle had
shrivelled, adventitious roots were beginning to appear at the base of the stem.
However, the results obtained show clearly that in S. dolichostachya seedlings
the radicle elongates more rapidly than in S. europaea, both in absolute length
and as a proportion of the total length of the seedlings.

DISCUSSION

It is not proposed to discuss in detail the taxonomic situation in Salicornia in the
Dee estuary. The main purpose of the systematic part of the present investigation
was to show that two species occurred in this locality and that most individuals
could be reliably identified. Similar populations were described by K6nig
(1939, 1960) from N. W. Germany and by Nannfeldt (1954) from S. Sweden.
However, taxonomic conclusions drawn from these populations may not be
valid for localities in south and east England, so further discussion is best left
until populations from this area have been described.

The results of the ecological study largely confirmed the conclusions from the
previous more casual observations. On completely open ground it appears that
plants of the diploid S. europaea are less able to establish themselves than the
tetraploid S. dolichostachya. However, once established in this habitat, S. euro-
paea is almost as successful as S. dolichostachya in terms of the seed production
of individual plants. With the increasing stability afforded by plants of other
species and by the somewhat flatter ground found in the centre of Area A and
in the middle and upper parts of Area B, plants of S. europaea are able to
establish themselves more readily. The results obtained in the seedling growth
experiments indicate that the radicle of seedlings of S. dolichostachya grows
more rapidly than that of S. europaea. In the habitats most exposed to tidal
disturbance this could be expected to give S. dolichostachya a marked advantage
over S. europaea. The observations by Wiehe (1935) strongly support the view
that establishment of the seedlings is the critical factor in determining whether
Salicornia can grow in a habitat that is frequently covered by the tide. It is
unlikely that the same tidal level will be critical in each locality. Many factors
such as the stability of the surface substrate, the slope of the land and the rate at
which the water flows over a particular area are likely to play an important part
in determining the critical point.

The behaviour of both species of Salicornia when growing in competition with
other species is not absolutely clear. A very slight difference in tidal level could
substantially affect the availability of water during parts of the summer and
early autumn and it was not possible to exclude this factor. In Area A, where
Salicornia was competing with Suaeda maritima, another, but more robust,
annual, both Salicornia species seemed able to survive and to produce seed at
approximately the same rate per plant. In area B, where competition was with
perennial grasses, S. europaea appeared to be rather more successful than

40 P. W. BALL AND K. G. BROWN

S. dolichostachya. However, in this case it is not possible to ignore the possible
effects of tidal level, although there did not appear to be any significant
difference between the heights of quadrats 6 and 11. The failure to find any
S. dolichostachya plants in the areas similar to quadrats 8-11 of transect B in
October, combined with the fact that in September many of the small plants
of S. dolichostachya from this area appeared to be dying without producing seed
suggests that, for reasons not determined, this species is less able to compete
successfully with perennials than is S. europaea. If only the plants that come to
maturity are considered, seed production per individual does not show any
marked differences in those areas where both species occur in competition with
other species. However, no individuals of S. dolichostachya were found with less
than a total of 6 fertile segments and only 4 (out of 147) with 10 or fewer
fertile segments. The corresponding figures for S. europaea were 18 and
43 (out of 237) (Fig. 12). This would appear to suggest that the minimum
vegetative growth requirement for S. dolichostachya to successfully flower and
fruit is greater than that for S. europaea.

1966 (All samples)

Cumulative 100 pe Bere a
70 a Pas

90
80
70

60

No. of fertile segments

+4 +_+—_—
5 10 15 20 30 40 50 75 100 150 200 300 400 500 750 10002000 per plant

Fi GuReE 12. Total number of fertile segments in each plant expressed as a cumulative %.
O = S. dolichostachya A = S. europaea

ACKNOWLEDGMENTS

We wish to thank Dr C. D. K. Cook and Dr T. McNeilly for their advice and
helpful criticism, and for a considerable amount of practical and technical help.

REFERENCES

BALL, P. W. (1960). The experimental taxonomy of Salicornia. Ph.D. thesis, University of
Leicester.

BALL, P. W. & TuTIN, T. G. (1959). Notes on annual species of Salicornia. Watsonia, 4: 193-205.

Konic, D. (1939). Die Kromosomenverhaltnisse der deutschen Salicornien. Planta, 29:
361-375.

Konic, D. (1960). Beitrage zur Kenntnis der deutschen Salicornien. Mitt. Fl.-Soziol. Arb.,
8: 5-58.

NANNFELDT, J. A. (1954). Nagot om slaktet Salicornia i Sverige. Svensk Bot. Tidskr., 49: 97-109.

WIEHE, P. O. (1935). A quantitative study of the influence of tide upon populations of Salicornia.
J. Ecol., 23: 323-333.

Watsonia, 8, 41-44 (1970). 41

Euphrasia brevipila and E. borealis i the British Isles

P. F. YEO
University Botanic Garden, Cambridge

ABSTRACT

The correct name at specific rank for British and Irish plants known until now as Euphrasia
brevipila is E. borealis (Towns.) Wettst. Plants from Orkney and Shetland known as E. borealis,
E. borealis var. speciosa and E. borealis var. zetlandica are referable to E. arctica Lange ex
Rostrup, which is endemic to the Faeroes, Orkney and Shetland. Plants from the rest of
the British Isles referred by Pugsley to E. borealis are correctly placed, though plants corr-
esponding to the north English form regarded by him as typical of E. borealis are not as
widespread as he indicated.

The publication in 1930 of H. W. Pugsley’s Revision of the British Euphrasiae
brought about a major clarification of the taxonomy of the British Eyebrights,
and since that time British students of the genus have endeavoured to apply
Pugsley’s findings. Wettstein (1896) had recognized both E. borealis (Towns.)
Wettst. and E. brevipila Burnat & Gremli as British, distinguishing E. borealis
from E. brevipila by its lack of short glandular hairs on the leaves. Pugsley
(1930: 519) accepted this distinction but amplified it, saying that E. brevipila
differed from E. borealis “in its rather narrower and more finely toothed leaves,
which are lighter in colour and nearly always more or less glandular; and its
flowers are lilac instead of white. On an average, its habit of growth approximates
less closely to the early-summer type than that of E. borealis . . .” In fact, Pugsley
(I.c., p. 517) considered that there had been some confusion in the material of
E. borealis that Wettstein had cited and asserted that the most characteristic
gathering cited in Wettstein’s original description was Townsend’s from
Borrowdale (dated 1884 according to the citation by Pugsley, l.c., p. 515). Pugsley
stated that E. borealis was “‘widely distributed in Scotland, less so in England
and Ireland”’, and that “‘In the Orkneys and Shetlands it appears to be particularly
abundant and variable’.

In the course of my studies I have had no difficulty in recognizing plants
corresponding with Pugsley’s E. borealis. The characters I have used have
included also the usually more elliptic capsule, as compared with E. brevipila,
but, on the other hand, it has not been possible to accept the flower-colour
character. Further, it has to be accepted, as was done by Pugsley (1930: 518),
that absence of glandular hairs is not diagnostic of E. borealis vis a vis
E. brevipila. Where 1 have had difficulty in following Pugsley is with regard to
distribution. Clearly, it has seemed to me, the stronghold of E. borealis is the
English Lake District and the northern Pennines, where it seems largely to
replace E. brevipila. From here it spreads north into the Scottish border counties
and, increasingly sparingly, south to the Peak District and Charnwood Forest
(Leicestershire); it has also been collected as far south as Hitchen in Hertford-
shire where it is, no doubt, long since extinct. I found that the plants from
Orkney and Shetland referred to E£. borealis and its vars. zetlandica Pugsl. and
speciosa Pugsl. were also clearly recognizable. But in between the Scottish border

42 P. F. YEO

counties and the Pentland Firth, and in Wales, I have hardly ever, if at all, seen
specimens corresponding with either the Lake District form or the Orkney and
Shetland forms of E. borealis. From the north coast of Scotland itself I have seen
specimens which I formerly thought might be included in E. borealis, but I now
think they are young and luxuriant plants of a form of E. nemorosa that occurs
on the coasts of north and north-west Scotland. I have never seen borealis-like
plants from Ireland. The specimens cited by Pugsley (1930) from Tiree, v.-c. 103,
(Macvicar, BM) are much branched, while those of Marshall’s which he cites
from Brora, v.-c. 107, are marked by him in BM as “? hybrid’, so neither
corresponds with the north English plant. Otherwise, I have seen only Towns-
end’s Braemar specimens from among the material cited by Pugsley from
Scotland apart from Orkney and Shetland, and Pugsley himself says that some
of the Braemar material is uncharacteristic. In fact, Pugsley (1930) records
E. borealis from only seven vice-counties from the mainland and Western Isles
of Scotland, and from seven vice-counties from the very much smaller area of
northern England. Thus, although Pugsley’s outline of distribution quoted
above gives the impression that E. borealis is found mainly in Scotland, his
vice-county records clearly show a concentration of the species in N. England.

Regarding the taxonomy of E. borealis, I have gradually come to the view that
the Orkney and Shetland plants are not the same as the north English-south
Scottish ones from which they are, in my view, spatially disjunct. Further, I have
doubted the practicability of recognizing the plants from the latter area as
specifically distinct, and I finally abandoned this attempt, having in recent years
determined such specimens as E. brevipila. The variation which has in any case
to be allowed in E. brevipila is so great that it seems pointless to try to treat the
rather minor departures from it represented by E. borealis as having specific rank.
However, the geographical significance of this variation should not be lost sight
of and may eventually receive taxonomic recognition at some infraspecific level.
During work on North American eyebrights in recent years, Mr P. D. Sell and
I had to investigate the name Euphrasia arctica Lange ex Rostrup, and we
concluded that the type came from the Faeroes, and that it represented the same
taxon as the “EF. borealis” of Orkney and Shetland.

The conclusions outlined so far have been rather tentatively embodied in the
notes on E. borealis in the Critical Supplement to the Atlas of the British Flora
(Perring & Sell, 1968). (The distribution map given there, however, includes
records authenticated by Pugsley and the late E. F. Warburg, as well as by
myself). It will be seen that if these conclusions are fully accepted the Orkney
and Shetland plants will become E. arctica and the rest will be merged with
E. brevipila, a name which has priority over E. borealis. The name E£. borealis
would then disappear into synonymy. This situation is also implicit in determina-
tions that I have made in recent years, and the purpose of this note is to prevent
these conclusions being acted upon. This is because further relevant factors
have come to light, as a result of my beginning work on an account of Euphrasia
for Flora Europaea. In doing this work I have learned that Wettstein was apt to
accept pairs of species differing only in the presence or absence of glandular
hairs, and have found that if this character is ignored all the original material of
E. brevipila from Haut Valais in Switzerland can easily be accepted as a slightly
precocious, subalpine state of E. stricta Lehm. The E. brevipila of the British
Isles, on the other hand, is not conspecific with E. stricta; it usually has larger
capsules than E. stricta and has the basal pairs of teeth of the lower floral leaves

EUPHRASIA BREVIPILA AND E. BOREALIS IN THE BRITISH ISLES 43

patent, instead of forwardly directed (antrorse) as in E. stricta. The name E.
brevipila must therefore be abandoned for British plants, and the earliest name
available at specific rank for British ““E. brevipila’’ is then E. borealis. Thus the
British “E. brevipila”’ and “‘E. borealis” (excluding specimens referableto E. arctica)
are to be united not under the former name but under the latter. It may be that in
Flora Europaea E. borealis will be reduced below the rank of species, but
pending a final decision on this it seems best to go over straight away to using
the name E. borealis. From now on it will be necessary to show on determinations
of some Continental specimens that EF. brevipila is a synonym of E. stricta, and it
is therefore important to make it clear that I do not regard British “‘E. brevipila”’
as synonymous with E. stricta.

For the benefit of readers concerned with Continental species, it should be
mentioned that I accept that members of the F. borealis group occur in Scan-
dinavia and eastern Europe.

Mr Sell and I have dealt with the typification of E. arctica and E. brevipila
in Our manuscript paper on American Euphrasia. The nomenclature and
typification of E. borealis is set out below.

E. borealis (Townsend) Wettstein, Mon. Gatt. Euphr. 108 (1896) (““Townsend
pro var. E. Rostkovianae in sched., in H. Schinz, H. Townsend’’).

E. rostkoviana Hayne forma borealis Townsend in Rep. Botl. Exch. Club Br.
Isl., 1: 307 (1891 for 1890).
Lectotype: E. rostkoviana Hayne f. borealis m.s. Grassy border of field nr.
the Manse, Braemar. Aug. 25. —90. Frederick Townsend (CGE).

E. brevipila auct. non Gremli, quoad pl. Britannicae et Hibernicae.

The lectotype of E. rostkoviana f. borealis was chosen by Mr Sell and me in
1968. Ideally it should have been chosen from Townsend’s own herbarium, but
Mr Sell has been unable to locate this despite repeated enquiries. Of the four
plants on the lectotype sheet all have the bracts setose and in two plants they
are also freely glandular. They are characteristic specimens of the plant normally
known in the British Isles as E. brevipila!

The original description of E. rostkoviana f. borealis appears in the Report of
the Distributor for 1890 but has apparently been overlooked by later authors; the
entire protologue reads as follows:

“‘Euphrasia rostkoviana, Hayne, f. borealis, m.s. Grassy border of field near the
Manse, Braemar, 25th Aug., 1890. E. rostkoviana f. borealis, has an included
corolla tube, and the whole plant is eglandular, or when glands are present,
they are very shortly stalked. It is probably a common form in Scotland.—
Fredk. Townsend”’.

It is not known whether Wettstein saw this printed matter attached to the
herbarium sheets he studied, but since he cited the type of E. rostkoviana
forma borealis and used the epithet borealis at specific rank, the basionym of
E. borealis must be E. rostkoviana f. borealis, its authority “(Towns.) Wettst.”’,
and its type the Braemar specimen. It is thus impermissible to typify E. borealis
except by the Braemar gathering of 25 Aug. 1890, and Pugsley’s treatment of
the Borrowdale form as typical of the species cannot be accepted as a lecto-
typification. In fact Wettstein used the Braemar gathering for both of his
illustrations (Taf. III, figs. 147-153; Taf. XI, fig. 7), and described it as the
“Original-Exemplar’’, that is, the type, in the legend on p. 308 to the photographic
illustration.

44 P. F. YEO

Plants described as E. borealis var. speciosa Pugsl. may be included in
E. arctica without separate recognition; they differ only in having rather larger
corollas than usual. E. borealis var. zetlandica Pugsl., on the other hand, is
probably derived by crossing between E. arctica and E. confusa; it could well be
retained as a variety of the former but I feel it is still too soon to make the
required new combination.

REFERENCES

PERRING, F. H. & SELL, P. D., ed. (1968). Critical Supplement to the Atlas of the British Flora.
London.

PuGsLey, H. W. (1930). A revision of the British Euphrasiae. J. Linn. Soc. (Bot.), 48: 467-544.

WETTSTEIN, R. VON (1896). Monographie der Gattung Euphrasia. Leipzig.

Watsonia, 8, 45-49 (1970). 45

Short Notes

97. CARDAMINE—Pollen.

Observations have been made on the pollen of Cardamine spp. including C. hirsuta,
C. impatiens, C. matthiola, C. resedifolia (2n = 16), C. pratensis (2n = 30), C. flexuosa
(2n = 32), and C. raphanifolia (2n = 46). Measurements of pollen diameter showed
that this character was not very useful in distinguishing the diploids from the poly-
ploids. Light microscope observations of the pollen surface showed a reticulate pattern
but no obvious distinguishing features.

Further investigations of the surface features were carried out using the Stereoscan
electron microscope. Low ( x 5000) and high ( x 15000) power photomicrographs were
exhibited at the 1968 Exhibition Meeting showing the extine pores and features
of the germinal pores. In the region of the germinal pores the reticulate pattern broke
down and, especially in C. matthiola, numerous papillae were seen. Measurements of
the extine pore diameters were found to be significantly different in all the species
examined. There were no significant correlations between extine pore size and breeding
system (C. matthiola, C. pratensis, and C. raphanifolia are self-sterile while the other
species are self-fertile) and between pore size and the ability to hybridise (C. flexuosa
with C. hirsuta, C. pratensis with C. raphanifolia). In contrast, there is a strong correla-
tion between extine pore size and chromosome number. The mean pore size varied
from0-:15 » m (C. resedifolia) to 0:21 u m (C. hirsuta) in the diploids to 0:24 » min
C. pratensis, 0:26 » min C. flexuosa and 0:42 » min C. raphanifolia.

Other species in the Cruciferae will be examined to determine if this correlation can
be made over a number of genera and if pollen extine pore size is useful in distinguish-
ing species.

R. P. ELuis & B. M. G. JONES.

122/2. ELATINE HYDROPIPER L.—New to Scotland.

In the West of Scotland the summer of 1968 was unusually dry and Loch Lomond was
reduced to prolonged low levels, exposing extensive areas of sand bars and mud flats
around the River Endrick mouth and its low-lying marshy hinterland. Among several
uncommon plants found on the freshly exposed surfaces was Elatine hydropiper L.
During July and August this plant was recorded in two localities on the mainland part
of the Loch Lomond National Nature Reserve, Dunbartonshire (Vice-county 99), and
in the marshes south of Balmaha, Stirlingshire (Vice-county 86). In each case it was
found in open communities on either undisturbed, bare sandy mud, or surrounding
agricultural watering ponds on the mud churned up by cattle. Since 1965 The Nature
Conservancy has maintained daily recordings of the levels of Loch Lomond from
which it has been possible to determine the height and period of inundation/emersion
of each colony of E. hydropiper. The annual rise and fall of the loch is between
5-7 feet with a summer average level of 24-5 ft O.D. The modal loch level, i.e. the level
occupying the highest percentage of time throughout the year, is 25:5 ft O.D. By
referring to the loch levels it was found that, with one exception which was not ascertain-
able, all colonies of E. hydropiper were growing at about 24-0 ft O.D., 1-5 ft below the
modal loch level. Taking the average for the years 1958-1967, sites at this level would
be exposed for 22 days per year. However in 1968 the number was 43, almost twice as
long, providing highly favourable conditions for germination. It is worth noting that
Mr D. McClintock recorded Elatine hexandra (Lapierre) DC. at Balmaha in July 1953
following a similar dry period and low loch level. This species was also noted by the

46 SHORT NOTES

present authors in the Endrick mouth area in the summer of 1968.

Sir Edward Salisbury has pointed out that E. hexandra is intolerant of competition
except with other small annual species. In general the same rule seems to apply to
E. hydropiper. In both the English sites from which the plant has been reported since
1960, the recorders note that it was either the only plant colonising the mud or its
nearest competitor was 1 metre away. At Loch Lomond, although the plant cover was
more dense than this, it was still extremely sparse. The most commonly associated
species were Peplis portula L., Callitriche stagnalis Scop. and Eleocharis acicularis (L.)
Roem. & Schult., all notably slender or weak-growing species.

In Sweden, Lohammar (1965) has produced comparable lists of species colonising
inundated muds on lake shores. At Lake Morjarvstraésket he records Eleocharis
acicularis, Subularia sp., Limosella sp., Crassula sp., Elatine hydropiper and Callitriche
verna; at Lake Jadersjon, on the dried-up lake floor, he notes Limosella sp., Subularia
sp., Elatine hydropiper and Callitriche verna. Similarly Pekkari (1965) working on the
aquatic vegetation of Swedish rivers found Subularia aquatica, Callitriche verna and
Elatine hydropiper growing with other small annuals in the shallow muddy bottoms of
broads, where there was less competition. The only perennials present were Eleocharis
acicularis, Isoetes echinospora and Ranunculus reptans.

At Loch Lomond annual fluctuations in water level seem to be important in com-
bining with wave action to maintain a strand of bare sand and mud, readily colonised
by weak annual species. In Sweden, although water levels differ in the way they
control the habitat, the effect in maintaining bare soil is much the same. Lohammar
(1965) states “The kind of substrate and annual rhythm in water level fluctuation are
prerequisites for this peculiar vegetation. During the period of low water in winter
frost penetrates down into these bottoms, and horizontal layers of ice cleave the
bottom material and cause it to expand and heave. On such an unstable substrate
Phragmites and other tall rhizomatose perennial plants never have the possibilities of
forming closed communities leaving the space free for small or weak competitors’.
When open conditions change, E. hydropiper would be expected to disappear in the
face of competition from more vigorously growing species. Praeger (1934) suggested
that even the irregular cleaning of the sides of the Newry Canal (Northern Ireland),
was sufficient to maintain a suitable habitat for large colonies of the plant there.

The nearest sites to Loch Lomond, in which E. hydropiper has been seen since 1930,
are in NE. Ireland, where it is recorded from eight 10 km squares covering four
Vice-counties, Louth, Armagh, Down and Antrim (Atlas of the British Flora, 1962).
Apart from NE. Ireland the only post-1930 records are from Surrey, Anglesey,
W. Sussex and Worcester and only in the last two has the plant been seen since 1960.
In both of these the habitat was either shallowly submerged sand or mud exposed by
fluctuating water levels. It is probable that, in Britain, the rarity of the plant may be
more apparent than real considering the difficulty of detection in seasons other than
those which are unusually dry.

We are grateful to Miss Noelle Hamilton (Biological Records Centre) and Messrs
F. Fincher and O. Buckle, County Recorders for Worcester and W. Sussex, for their
help in checking past and present records.

REFERENCES

LOHAMMAR, G. (1965). The Vegetation of Swedish Lakes, in The Plant Cover of Sweden. Acta
phytogeogr. suec., 50: 28-47.

PEKKARI, S. (1965). Notes on Aquatic Vegetation, in The Plant Cover of Sweden. Acta phyto-
geogr. suec., 50: 209-214.

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

PRAEGER, R. L. (1934). Fifteen miles of Elatine hydropiper. Ir. Nat. J., 5: 102-104.

SALISBURY, E. J. (1967). On the reproduction and biology of Elatine hexandra (Lapierre) DC.
(Elatinaceae); a typical species of exposed mud. Kew Bull., 2: 139-147.

E. T. IDLE, J. MITCHELL & A. McG. STIRLING.

SHORT NOTES 47

168/s. GERANIUM SUBMOLLE Steud. (or G. core-core Steud.)—In the Channel Isles.

The precise name of this, probably South American, plant cannot be settled because
the type specimens are still unobtainable. It is, however, most certainly not the New
Zealand G. retrorsum L’Herit. Its history in the Channel Isles is as follows:

1926. Paradis/Fort Doyle, Guernsey. T. J. Fogitt, (BM). Never seen there since.

1938. Railway crossing, Whitegates, Alderney. A. B. & A. K. Jackson & H. K. Airy
Shaw, (K).

1957. Railway line between naval quarries and Platte Saline, Alderney. Discovered
independently there by Dr H. J. M. Bowen, D. McClintock and Mrs B. H. S.
Russell, and still plentiful.

1963. Essex House and Reuthe’s Valley, Alderney. D. McClintock.

1966. Rue des Pres, Jersey. Mrs F. le Sueur.

1968. La Garenne, Guernsey (at the opposite end of L’Ancresse Common from
Paradis). Dr H. J. M. Bowen.

I know of no records for this plant anywhere else in the British Isles. Specimens from

the first and last localities were shown at the Exhibition Meeting in November 1968.

REFERENCES

KENT, D. H. (1959). Geranium retrorsum L’Herit., in Plant Notes. Proc. bot. Soc. Br. Isl.,
3: 284.

LousLey, J. E. (1962). Geranium retrorsum L’Hérit., in Plant Notes. Proc. bot. Soc. Br. Isl., 4:
413-414.

TOWNSEND, C. C. (1961). Geranium microphyllum Hook. f. as an adventive plant in Britain.
Watsonia, 5: 43-44.

TOWNSEND, C. C. (1964). More on the introduced Bordon and Alderney Geraniums. Proc.
bot. Soc. Br. [sl., 5: 224-226.

D. MCCLINTOCK.

506/4, 5 & 8. SENECIO SQUALIDUS L., S. VULGARIS L. and S. CAMBRENSIS ROSSER.

The status of the radiate varieties of Senecio vulgaris in Britain has recently been
discussed by D. E. Allen, who suggests the separation of these varieties into var.
denticulatus (O. F. Muell.) Hyland., a maritime plant, and var. hibernicus Syme, an in-
land plant that has spread considerably in recent times. However, reference to herbarium
and other records establishes a pattern where S. vulgaris var. hibernicus tends to occur
only after S. squalidus has been recorded in the same locality.

In 1966, a short-rayed plant closely resembling Rosser’s S. cambrensis (2n = 60),
the allohexaploid from SS. vulgaris (2n = 40) x S. squalidus (2n = 20), was found in
east London. Progeny from this plant segregated for characters of S. vulgaris and
S. squalidus, and all proved to be more or less tetraploid (2n = (36—) 40 (-44)). The
majority of these segregates resembled the parent plant in ray dimensions, and a
small number showed either rays of full size, as in S. squalidus, or were nearly or
completely rayless, as in S. vulgaris. Self-fertilised rayless plants produced rayless
offspring. The morphology of these is within the range of variation shown by S. vulgaris,
but they differ in their greater vigour, slightly larger seeds and pollen, a degree of seed
and pollen sterility, and the formation of a few multivalents at pollen mother cell
meiosis. Intermediate-rayed tetraploids have been crossed with S. vulgaris, and have
produced segregate offspring, some of which, again, closely resemble S. vulgaris.
It therefore appears probable that repeated back-crossing of these plants into
S. vulgaris could result in plants containing genetically determined characters of
S. squalidus, and that these characters (e.g. the ray character, and leaf shape) might be
sufficiently “diluted” in the population to appear as single gene variants. There have
been a number of reports of “‘short-rayed S. squalidus plants’ and of “‘giant radiate
groundsels” in the literature, and in herbarium records, but with no possibility of
analysing these plants chromosomally, it would be presumptive to conclude that

48 SHORT NOTES

these are plants of the type described above. We would, however, like to suggest that
the spasmodic reports of these anomalous plants indicate that they are not part of
stable populations, and hence might be indicators of transitory stages in the rapid
introgression of S. squalidus genes into S. vulgaris.

There are several possibilities as to the origin of these segregating tetraploids. These
will be reported on when we have more information, but observations made on the
progeny of the triploid (2x = 30) S. x londinensis Lousley (S. viscosus, 2n = 40
x JS. squalidus, 2n = 20) have indicated that plants with tetraploid or higher chromo-
some numbers can be produced. These have so far shown high sterility, in contrast to
the naturally occuring tetraploid from S. vulgaris and S. squalidus reported above.

This investigation has also raised the rather intriguing question of what happens to
the large-rayed tetraploid segregates. These bear a close resemblance to ordinary
diploid S. squalidus, but because of their increased chromosome number will not form
part of the same breeding population. They also differ in being self-compatible, which
would allow isolated individuals to reproduce themselves. Also in this context, it is of
interest to note that S. cambrensis, which appears from the herbarium records to have
arisen in Wales at least 40 years ago, has segregated to the extent that pure breeding
rayless forms can be seen there. Theoretically, self-maintaining populations of large-
rayed S. cambrensis segregates could have also been produced. This is speculative, but
could it be that tetraploid and hexaploid, as well as diploid races of “‘S. squalidus’’ occur,
as yet undetected, in the British Isles? These races, as well as those of S. cambrensis
resembling S. vulgaris, could easily have been mistaken for autopolyploids, with any
slight morphological difference from the “‘parent species” being attributed to evolution
in the reproductively isolated autopolyploid. Polyplotypes of other species have often
been described as autopolyploids because of their close morphological correspondence
to a presumed parental diploid, only to be re-evaluated as “‘segmental allopolyploids”
(Stebbins, 1947) on a more complete analysis. We hope that the present investigation
will show clearly that the concept of segregation following allopolyploidy (inter-
genomic segregation, Jones 1967) can offer another possible explanation of this
phenomenon.

REFERENCES

ALLEN, D. E. (1967). The Taxonomy and Nomenclature of the Radiate Variants of Senecio
vulgaris L. Watsonia, 6: 280-282.

Jones, B. M. G. (1967). The evolutionary advantages of polyploidy. Mimeographed copy of
lecture to the British Association at Leeds, September 1967.

Rosser, E. M. (1955). A new British Species of Senecio. Watsonia, 3: 228-232.

STEBBINS, G. L. (1947). Types of polyploidy: their classification and significance. Adv. Genet.,
1: 403-429.

P. Crisp & B. M. G. JONES.

558/-. HIERACIUM—Two Hieracia Sect. Alpestria from the British mainland.

At the time of Sell & West’s (1965) revision, Hieracium mirandum Sell & C. West
was known only from a single gathering in 1962 and H. carpathicum Besser, in Britain,
from four gatherings between 1889 and 1956. Further herbarium material of H. miran-
dum has since been discovered, collected by A. Ley at Ribblehead in 1902 and by
E. C. Wallace from Masson Hill, Bonsall in 1946. Bearing in mind the remarkable
persistence of hawkweeds in restricted localities, we and Dr C. West in 1967 searched
for both species without success, for H. mirandum around Masson Hill and for
H. carpathicum up the Lochsie Burn from the Spittal of Glenshee. In 1968, however, we
succeeded in finding both species. H. mirandum was growing in moderate quantity in
the Ribblehead area, where it was in grave danger from grazing animals and most
plants had the capitula nibbled off; H. carpathicum was on the bank of the Shee water a

SHORT NOTES 49

few miles below the Spittal, where it appeared to be in no danger unless from acquisi-
tive botanists. It was apparently absent between the Spittal and this new locality, but we
did not explore further downstream. By contrast H. magniceps Sell & C. West, hitherto
known only from a limited region near the Spittal, was frequent beside the river for
some miles further downstream.

REFERENCE

SELL, P. D. & WEsT, C. (1965). A revision of the British species of Hieracium section Alpestria
(Fries) F. N. Williams. Watsonia, 6: 85-105.

J. N. Mitts and J. R. J. MILLs.

Watsonia, 8, 51 (1970). 5]

The Irish Vice-Counties

Doubts having recently arisen as to the correct interpretation of the boundaries
of the biological vice-counties of Ireland, the Praeger Committee of the Royal Irish
Academy has considered the matter and recommends as follows:

The names, limits and extent of the biological vice-counties of Ireland should be
those set out in the map which forms the frontispiece to Irish Topographical Botany
by R. L. Praeger (published by the Royal Irish Academy in 1901 as Volume 23 (3rd
Series, Vol. 7) of its Proceedings).

In making this recommendation the committee has been influenced by the following
considerations:

(1) the map referred to, though on a small scale, is of admirable quality and clarity,
and permits the assignment to its vice-county of any locality with a margin of un-
certainty of less than half a mile;

(2) the text of Irish Topographical Botany forms by far the largest single unit of
biological records on a vice-county basis, and Praeger’s prestige and authority were
such that recorders of other groups of organisms were content to accept his judgement
in this matter;

(3) the decision is in general harmony with the policy followed in Great Britain of
“freezing”? county-boundaries in the form in which they existed when the recording
system was first instituted and ignoring subsequent administrative changes. Consider-
able changes in the Irish county-boundaries were made in 1898, but only one of these
(the transfer from Galway to Clare of a substantial area on the west side of Lough
Derg) was recognized by Praeger in his map or his text.

I hope to submit shortly to the Royal Irish Academy for publication in its Proceedings
a more detailed statement about those aspects of the vice-county-boundaries which
require clarification.

D. A. WEBB.
School of Botany, Trinity College, Dublin.

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Watsonia, 8, 53-61 (1970). 53

Plant Records
Compiled by E. C. Wallace

‘Plant Records’ are arranged in the order given in the List of British Vascular Plants
(L.B.V.P) by J. E. Dandy (1958).

Records where no date is given are for the year 1968.

The following signs are used:

§ before the L.B.V.P. number: to indicate that the paragraph contains information necessitating
a correction to an annotated copy of the Comital Flora.

{ before the L.B.V.P. 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.

* before the record: to indicate a new vice-county record, not published previous to this
issue in the Proceedings. Some of these records may, however, appear as dots on maps of
the particular species in the Atlas of the British Flora.

t before the record: to indicate a record additional to an annotated copy of the Comital Flora,
but published elsewhere prior to this issue of Watsonia.

[ ] enclosing a record: to indicate doubt as to the validity of the record, either of identification
or locality.

It will be useful if National Grid Co-ordinates, made as accurate as is thought advisable,
are added to all records. These will not be published, but the original cards containing them
will be filed and may be made available for reference.

§1/2. LYCOPODIELLA INUNDATA (L.) Holub (Lycopodium inundatum L.). *110,
Outer Hebrides; Loch an Tairbh Duinn, near Tarbert, Harris, F. BURNIER, comm.
J. OUNSTED.

1/3. LYCOPODIUM ANNOTINUM L. 98, Argyll; Meall a Bhuiridh, north-facing
slopes near Cam Ghleann, in small quantity, c. 2000 ft., A. G. KENNETH.

§3/1. ISOETES LACUSTRIS L. *106, E. Ross; Loch Beannacharain, Strath Conon,
M. McC. WEBSTER.
§4/10. EQUISETUM TELMATEIA Ehrh. *72, Dumfries; Tarras Water, near Cronks-

bank, locally abundant in ditches by roadside, 350 ft., R. W. M. CorNER, comm.
H. MILNE-REDHEAD. *82, Haddington; colony on bank of Oldhamstocks Water, near
Oldhamstocks, E. P. BEATTIE.

§7/2. HYMENOPHYLLUM WILSONIL Hook. *79, Selkirk; sheltered vertical rocks
above river, south side river Ettrick, below junction with Range/Cleuch Burn, very local,
colony covering 1-2 square feet, 1100 ft., R. W. M. CoRNER, comm. H. MILNE-
REDHEAD. 88, Mid Perth; wooded banks on rock, Glenlednock, near the Deil’s
Cauldron, 1967, G. P. WILSON, comm. A. W. ROBSON.

15/2. ASPLENIUM BILLOTI F. W. Schultz (A. obovatum auct.). 48, Merioneth;
crevices of rocks near the sea at Aberdovey in two places, the first records for the
south of the vice-county, P. M. BENorr and Mrs K. M. STEVENS.

15/S5a. ASPLENIUM TRICHOMANES L. subsp. TRICHOMANES. 46, Cardigan; rock
crevices on the east side of the river Rheidol in the gorge near Parson’s Bridge, P. M,
BENOIT. 47, Montgomery; a few plants in a crevice of an acid rock on the roadside at
Esgair Geiliog, on the Montgomeryshire side of the river Dovey, P. M. Benoit (1965,
Nature Wales, 9, 219). The subsp. quadrivalens D. E. Meyer is the usual subspecies in
Cardigan and Montgomery, P. M. BENOIT.

§15/8. ASPLENIUM SEPTENTRIONALE (L.) Hoffm. 192, S. Aberdeen; rocks at east
end of pass of Ballater. cf. Dickie Botanist’s Guide (1860), A. J. SOUTER.

54 PLANT RECORDS

§16/1. CETERACH OFFICINARUM DC. *84, Linlithgow; Hopetown Estate, one
plant on wall, E. P. BEATTIE.
§19/1. CySTOPTERIS FRAGILIS (L.) Bernh. *29, Cambridge; concrete platform of

disused Old North Road station, S. M. WALTERS.

21/6. DRYOPTERIS CARTHUSIANA (Vill) H. P. Fuchs (D._ lanceolatocristata
(Hoffm.) Alston). 94, Banff; rough ground, Culvie Hall, 1967, A. J. Souter.

21/8. DRYOPTERIS AEMULA (Ait.) O. Kuntze. 48, Merioneth; Cwm Cynfal, near
Llan Ffestiniog, about 30 plants, 1966, F. RosE & P. M. BENorr (1966, Nature Wales, 10,
87); river Ysgethin, Talybont, at least 2 clumps, P. M. BENoIT & Mrs K. M. STEVENS.

§21/a. DRYOPTERIS ASSIMILIS S. Walker. *90, Forfar; Caenlochan Glen, Glenisla;
*94, Banff; Coire Raibeirt, Cairn Gorm, U. K. DUNCAN, both conf. A. C. JERmMy.

§24/1. THELYPTERIS LIMBOSPERMA (AIll.) H. P. Fuchs (7. oreopteris (Ehrh.)
Slosson.) *82, Haddington; near Innerwick, 1964, E. P. BEATTIE.

24/4. THELYPTERIS DRYOPTERIS (L.) Slosson. 82, Haddington; in small quantity
in Bladdering Cleach, Oldhamstocks, 1967, E. P. BEATTIE.

§46/21. RANUNCULUS TRICHOPHYLLUS Chaix. tH6, Waterford; Carrickavrantry
Lake, 1967, I. K. Fercuson, det. C. D. K. Cook (1968, Ir. Nat. J., 16, 94).

46/23. RANUNCULUS BAUDOTI Godr. 95, Moray; brackish pool near Findhorn
estuary north of Invererne, 1954, M. McC. WEBSTER.

48/1. MyosuRUS MINIMUS L. 55, Leicester; colony of about 100 plants growing in a
hollow against a field gateway between Barrow and Mountsorrel. They were growing on
a deposit of flood silt. A search at similar spots in the neighbourhood failed to reveal
any more. Probably not hitherto recorded from the vice-county in this century, but
recorded from several localities in the last century (1968, Heritage, 30, 15).

§49/1. AQUILEGIA VULGARIS L. *+94, Banff; several plants established on
shingle of river Avon below Delnabo, Tomintoul; garden origin, M. McC. WEBSTER.

§57/2. CERATOPHYLLUM SUBMERSUM L. *29, Cambridge; Horseshoe Pond,
Madingley Park. Two variants occur, var. submersum and var. haynaldianum (Borbas)
G. Beck, R. C. CHAPMAN.

§58/3. PAPAVER LECOQI Lamotte. *+101, Kintyre; Knapdale, road block south
of Ardrishaig, casual, 1967, A. G. KENNETH.

58/4. PAPAVER HYBRIDUM L. 103, Mid Ebudes; potato patch in garden, Baugh,
Isle of Tiree, coll. proprietor of guest-house, comm. U. K. DUNCAN.

§65/3. CORYDALIS CLAVICULATA (L.) DC. *82, Haddington; in small quantity
in Bolton Muir Wood near Gifford, E. P. BEATTIE.
88/5 x1. COCHLEARIA DANICA L. X OFFICINALIS L. 48, Merioneth; shingle on

the shore at Llanaber, near Barmouth, with both parents, and morphologically
intermediate between them, pollen partially sterile, matching a synthesised hybrid of
this parentage raised by the recorder, 1967, P. M. BENOIT.

89/1. SUBULARIA AQUATICA L. 106, E. Ross; shores of Loch Beannacharain,
Strath Conon, M. McC. WEBSTER.

§94/4. DRABA MURALIS L. *84, Linlithgow; abundant on verges and paths at
an old mill near East Philipstown, E. P. BEATTIE.

+97/7. CARDAMINE RAPHANIFOLIA Pourret (C. Jatifolia Vahl, non Lej.). 33,
E. Glos.; about 50 plants in a withy holt on river Eye, a tributary of river Dikler near
Lower Slaughter, reported by R. C. L. HowrttT, and localised by Mrs DUDLEY-SMITH
and Miss D. E. DE VESIAN, det. B. M. G. JONES.

§98/3. BARBAREA INTERMEDIA Bor. 88, Mid Perth; railway embankment,
Glenfarg, frequent, 1965, A. W. Rosson. {H6, Waterford; damp ground near the
road at Newtown Cove, south of Tramore, 1962, I. K. FERGusoN, det. P. W. BALL
(1968, Ir. Nat. J., 16, 95).

PLANT RECORDS 35

113/9b. VIOLA PALUSTRIS L. subsp. JURESSI (Neves) P. Fourn. 48, Merioneth;
plants of V. palustris having petioles with conspicuous spreading hairs, at Coed
Cymerau national nature reserve, near Ffestiniog, provide the confirmation for v.c. 48
noted as lacking in the Critical Supplement to the Atlas of the British Flora, P. M.
BENOIT.

4113/10. VIOLA CORNUTA L. 94, Banff; several clumps by roadside near Inver-
charach, M. McC. WEBSTER.
4115/4. HYPERICUM CALYCINUM L. 84, Linlithgow; abundant in Hopetown Estate

for many years, E. P. BEATTIE.

§122/1. ELATINE HEXANDRA (Lapierre) DC. 48, Merioneth; sparingly by the
river Dysynni, Brynerug, doubtless the spot where it was collected by H. E. Fox in
1878 (OXF), P. M. BENoir and Mrs K. M. STEVENS; in abundance in shallow water
at edge of Bala Lake, P. M. BENoiT, J. G. & C. M. Dony. *98, Argyll; Loch Awe,
Inverliever Bay, on dried-out mud in some quantity, A. G. KENNETH.

123/2. SILENE MARITIMA With. 88, Mid Perth; cliff ledge, Corrychamaig, Glen
Lochay, 1967, A. W. ROBSON.

§131/10. CERASTIUM DIFFUSUM Pers. (C. atrovirens Bab.) +755, Leicester; a few
plants on ballast at Thorpe Satchville and fairly common along railway track,
H. B.[RADSHAW]: only doubtfully recorded before in Leicestershire (1968, Heritage,
30, 16).

133/1b. STELLARIA NEMORUM L. subsp. GLOCHIDISPERMA Murb. 48, Merioneth;
woods with rich humus at Brithdir and at Arthog, P. M. BENoiT, conf. P. S. GREEN.

136/10. SAGINA NODOSA (L.) Fenzl. 84, Linlithgow; marshy area, Blackness
Castle, 1967, E. P. BEATTIE. 94, Banff; bog, Glenlivet, M. McC. WEBSTER.

§137/1. MINUARTIA VERNA (L.) Hiern. *87, W. Perth; Dumyat, dry slopes,
1883; not at all plentiful, ‘F.R.C.’ (E.). *89, E. Perth; Lochton, 1883 (E.). *103, Mid
Ebudes; Ben More, Mull, mountain ledge, 1956, J. SUTHERLAND (Hb. Edinb.), all det.
or conf. and comm. G. HALLIDAY.

4141/6. ARENARIA BALEARICA L. 88, Mid Perth; flat sandstone block by edge of
river Almond between mouth of Francesfieldburn and that of the Miltonburn, 1967;
base of dead tree trunk, south bank of river Almond, Blackwoodlands, 1967, A. W.
ROBSON.

143/4. SPERGULARIA MEDIA (L.) C. Presl. 84, Linlithgow; salt marsh, Blackness,
1967, E. P. BEATTIE.

149/1d. MONTIA FONTANA L. subsp. VARIABILIS Walters. 96b, Nairn; flush at
foot of 25 ft raised beach, Culbin, north of Mariston Farm, M. McC. WEBSTER.
101, Kintyre; Knapdale, Isle of Danna, 1967, A. G. KENNETH, det. S. M. WALTERS
(as subsp. fontana).

§154/3. CHENOPODIUM VULVARIA L. *+81, Berwick; several plants by tip,
Chirnside Paper Mill, O. Stewart & M. McC. WEBSTER.

163/2. MALVA SYLVESTRIS L. 101, Kintyre; Keil rocks near Southend, perhaps
only casual, A. G. KENNETH.

164/1. LAVATERA ARBOREA L. 82, Haddington; abundant on the island of
Craigleith as well as on the Bass Rock, E. P. BEATTIE.

168/1. GERANIUM PRATENSE L. +84, Linlithgow; roadside from Whitequarries

_ to Abercorn, and elsewhere in the vice-county, E. P. BEATTIE.
7168/3. GERANIUM ENDRESSII Gay. $2, Haddington; a small patch on a grassy

verge near Blance, E. P. BEATTIE. 109, Caithness; a large clump near cottages between
Reay and Fouss, M. McC. WEBSTER.

§173/3. ACER CAMPESTRE L. ttH6, Waterford; woods at Newtown Cove
south of Tramore, 1964, I. K. FERGUSON (1968 , /r.Nat. J., 16, 95).

56 PLANT RECORDS

§185/1. GENISTA TINCTORIA L. *84, Linlithgow; small colony on towpath of
Union Canal between Winchburgh and Broxburn, E. P. BEATTIE.

*191/2. MELILOTUS OFFICINALIS (L.) Pall. 48, Merioneth; railway bank west of
Aberdovey Station, 1968, P. M. BENoiIr & Mrs K. M. STEVENS. A species not given
in Benoit & Richards’s Contribution to a Flora of Merioneth (1963).

207/1. LATHYRUS APHACA L. 24, Bucks.; road embankment near Ashley Green,
apparently spreading, J. ROGERSON.
4208/1. PHYSOCARPUS OPULIFOLIUS (L.) Maxim. 82, Haddington; several bushes

in Saltoun Wood, 1967, E. P. BEATTIE.

212/12. POTENTILLA CRANTZI (Crantz) G. Beck ex Fritsch. 88, Mid Perth;
rocky gorge, eastern flanks of Stob an Aonich Mhair, Talla, Bheithe, 1965, A. W.
ROBSON.

~216/2. GEUM MACROPHYLLUM Willd. 87, W. Perth; uncut grass triangle at
entrance to Blairdrummond Park, 1965; second vice-county record, A. W. ROBSON.

220/3(9). ALCHEMILLA GLOMERULANS Buser. 79, Selkirk; locally common in wet
meadowland at confluence of Ettrick and Timal Waters, R. W. M. CoRNER, comm.
H. MILNE-REDHEAD.

220/3 (12). ALCHEMILLA MOLLIS (Buser) Rothm. 88, Mid Perth; dominant on a
quarry floor, Craigend, Moncrieffe Hill, 1965, A. W. ROBSON.

225/1. ROSA ARVENSIS Huds. 88, Mid Perth; roadside verge, Lynedoeh drive-
way, 1967, Miss J. STEWART, det. and comm. A. W. ROBSON.

~225/5. ROSA RUGOSA Thunb. 82, Haddington; one large bush, Thornton loch,
E. P. BEATTIE.

239/3. SAXIFRAGA HIRCULUS L. 94, Banff; Upper Cabrach, three small adjacent
groups; annual count of flowers produced 1965-68 was from 20 to 40, A. J. SOUTER.

239/4. SAXIFRAGA UMBROSA L. 88, Mid Perth; rocky banks, Buchanty Spout,
south side of river Almond, 1967, R. CORNER.

239/15. SAXIFRAGA HYPNOIDES L. 83, Edinb.; Linn Dean Water, a large colony,
E. P. BEATTIE.

$241/1. TOLMIEA MENZIESII (Pursh) Torr. & Gray. 88, Mid Perth; muddy banks
of the Farg, in deep shade, Glenfarg, 1965, A. W. Rosson; under willow and alder
wood on sandy soil on small island at confluence of Turretburn and river Earn, 1966,
G. P. WILSON, comm. A. W. Rosson. 89, E. Perth; edge of backwater of river Tummel,
under trees, Moulinlam; by stream under trees, Quarrymill Den, Scone, 1967, A. W.
ROBSON.

§247/1. DROSERA ROTUNDIFOLIA L. *84, Linlithgow; abundant at Easter Inch
Moss, Blackburn, 1967, E. P. BEATTIE.
247/3. DROSERA INTERMEDIA Hayne. 49, Caern.; Cors Graianog, on bare wet

peat mud on rocky slope, C. A. SINKER, comm. R. H. ROBERTS.

254/5. EPILOBIUM ROSEUM Schreb. 95, Moray; distillery yard, Carron, M. McC.
WEBSTER.

262/5. CALLITRICHE HERMAPHRODITICA L. 96, Easterness; pool on moor one
mile north-west of Glen Mazeran Lodge, M. McC. WEBSTER.

4271/1. ASTRANTIA MAJOR L. 88, Mid Perth; fully naturalised at Falls of
Edinumple, J. N. MILLs.

304/1. MEUM ATHAMANTICUM Jacq. 48, Merioneth; in quantity on the edges of
meadows at Garneddwen, between Bala and Dolgellau, rediscovery of a site reported
by John Ray in 1662, 1967, W. M. Conpry.

§319/11. EUPHORBIA EXIGUA L. *t95, Moray; distillery yard, Knockando, a
single plant, G. SHEPHERD, comm. M. McC. WEBSTER.

PLANT RECORDS 57

§320/5. POLYGONUM VIVIPARUM L. *79, Selkirk; locally common on wet rocks
and in flushes above banks of Tima Water below Meerlees, 850 ft, R. W. M. Corner,
comm. H. MILNE-REDHEAD.

320/14. PoLYGONUM MINUS Huds. 48, Merioneth; marsh by Bala Lake, sparingly,
providing confirmation in the original Merioneth station noted as desirable by D. A.
Jones in Rep. botl. Soc. Exch. Club Br. Isl., 5, 674 (1920), P. M. Benoit, J. G. & C. M.
Dony.

+320/22. POLYGONUM CAMPANULATUM Hook. f. 38, Warwick; a single large
colony in Sutton Park, Miss D. A. CADBURY & R. C. READETT.

325/11 x 8. RUMEX X PROPINQUUS Aresch. 82, Haddington; by roadside near
Pilmuir, V. LEATHER & E. P. BEATTIE, det. J. E. LOUSLEy.

325/14. RUMEX SANGUINEUS L. 49, Caern.; grounds of Bryn Gwynant Youth
Hostel, shady drive, 1967, R. M. BURTON, comm. R. H. ROBERTs.

§335/2. BETULA PUBESCENS Ehrh. subsp. PUBESCENS. *§2, Haddington; in all the
Lammermuir Deans, 1967, E. P. BEATTIE.
4351/1. GAULTHERIA SHALLON Pursh. 88, Mid Perth; coniferous woodland,

Balgowan Woods, Findo Gask, 1966; wood partly felled in 1967, and unable to refind
plant in 1968, A. W. ROBSON.

358/3. VACCINIUM ULIGINOSUM L. 72, Dumfries; peat bog, Archie Grain, above
Langshahburn, Eskdalemuir, very local, scattered in area of c. 12 square yards with
Vaccinium myrtillus, R. W. M. CORNER, comm. H. MILNE-REDHEAD.

§358/4. VACcINIUM oxycoccos L. *84, Linlithgow; Tailend Moss, near Bathgate,
E. P. BEATTIE.

360/1. ORTHILIA SECUNDA (L.) House. 88, Mid Perth; by rocky banks of stream,
Auchnafree, Glenalmond, 1966, A. W. Rogson. 89, E. Perth; rocky banks, Glen Bruar,
gorge west of Meall nan Lath, 1966, A. W. RoBSOoN.

§364/2. EMPETRUM HERMAPHRODITUM Hagerup. *101, Kintyre; Cruach Lusoch,
west face, c. 1300 ft, in small quantity, 1968, A. G. KENNETH.
§385/3. GENTIANELLA AMARELLA (L.) Borner. *84, Linlithgow; in some quantity

in Dalmeny Estate, E. P. BEATTIE.

1392/3. SYMPHYTUM ORIENTALE L. 95, Moray; bank by Forres tip, Mrs
CHRISTIE, comm. M. McC. WEBSTER.

$394/1. TRACHYSTEMON ORIENTALIS (L.) G. Don. 82, Haddington; small colony
by the Gifford Water in Yester Estate, 1967, E. P. BEATTIE.

§400/3. MyYOSOTIS STOLONIFERA Gay (M. brevifolia C. E. Salmon). *67, North-
umberland, S.; in S. Tynedale at Faugh Cleugh on the Thinhope Burn, 1350 ft (conf.
A. E. WADE); by a tributary of the Ayle Burn, 1250 ft; Ayle Common, 1625 ft; in
W. Allendale by a small waterfall near Smallburns, 1450 ft; in E. Allendale on Shield-
bank Sike, 1450 ft (conf. A. E. WADE); Ellershope Burn, 1460 ft, G. A. & M. Swan.

4406/2. CALYSTEGIA PULCHRA Brummitt & Heywood (C. dahurica auct., non (Herbert)
G. Don). 84, Linlithgow; verge of Kinneil Wood near Bo’ness, 1966, E. P. BEATTIE.

410/1. ATROPA BELLA-DONNA L. +83, Edinb.; in some quantity in Warriston
Cemetery, E. P. BEATTIE.

422/2. KICKXIA ELATINE (L.) Dumort. +48, Merioneth; Aberdovey, on path up
to the Trefeddian Hotel, about 13 plants, 1963, E. TEAGUE (1964, Nature Wales, 9, 64),
and garden weed in several places about the town, Mrs K. M. STEVENS, a species not
included in Benoit & Richards’s Contribution to a Flora of Merioneth (1963).

7423/1. CYMBALARIA MURALIS Gaertn., Mey. & Schreb. 95, Moray; wall at
Earlsmill, Darnaway, M. McC. WEBSTER.

7424/2. CyMBALARIA PALLIDA (Ten.) Wettst. 83, Edinb.; wall in Princes Street
Gardens, Edinburgh, V. LEATHER & E. P. BEATTIE.

58 PLANT RECORDS

7425/1 x 2. MIMULUS GUTTATUS DC. x M. LUTEUS L. 13, W. Sussex; Fulking,
1961, J. T. H. KniGut, conf. R. H. ROBERTS.

426/2. LIMOSELLA SUBULATA Ives. 48, Merioneth; in some quantity in a wet
place on Morfa Harlech, a third Merioneth station for one of Britain’s most local
species, P. M. BENoIT (1966, Nature Wales, 10, 45).

1430/14. VERONICA PEREGRINA L. 16, W. Kent; vegetable garden, Tonbridge,
S. L. MELVILLE, comm. E. B. BANGERTER.

§435/1 (5). EUPHRASIA FOULAENSIS Townsend ex Wettst. *94, Banff.; grassy sea
cliffs, Strathlene, M. McC. WEBSTER, det. P. F. YEO.
§435/1 (11). EUPHRASIA CAMBRICA Pugsl. *48, Merioneth; cliffs in Cwm Aran,

Cader Idris, at 2000 ft, 1965, P. M. BENolrT, det. P. F. YEo.

§435/1 (15). EUPHRASIA CONFUSA Pugs]. *94, Banff; fixed shingle by the burn
Glenlivet, M. McC. WEBSTER, det. P. F. YEo.

439/1. LATHRAEA SQUAMARIA L. 88, Mid Perth; mixed wood, mainly sycamore,
Ardeonig, immediately east of Manse grounds, 1966, Miss V. M. THomM, comm.
A. W. ROBSON.

440/4. OROBANCHE ALBA Steph. ex Willd. 65, N.-W. York;scars above Carperby,
J. D. Lovis; first record for over 40 years, C. M. Ros (1969, Naturalist, 1969, 24).
88, Mid Perth; one plant among wood shavings, Methven station yard, 1967, Miss
J. STEWART, det. D. J. HAMBLER, comm. A. W. ROBSON.

442/2. UTRICULARIA NEGLECTA Lehm. 89, E. Perth; small pool beside the path
along east flank of Creag Bhreac, 1966, A. W. ROBSON.

442/3. UTRICULARIA INTERMEDIA Hayne. 49, Caern.; open pools in old cuttings
in peat bog, Cors Graianog, with Menyanthes, etc., C. A. SINKER, comm. R. H.
ROBERTS.

446/1. LyCOPUS EUROPAEUS L. 84, Linlithgow; a few plants on canal bank near
Linlithgow golf course, 1965—: 86, Stirling; a few plants on canal bank near Polmont,
E. P. BEATTIE.

474/1. WAHLENBERGIA HEDERACEA (L.) Reichb. 42, Brecon; among boulders
beside stream, Glen Haffes, 1965, J. N. MILLS.

7483/1. ASPERULA TAURINA L. 88, Mid Perth; banks of Turret burn, Crieff,
completely naturalised over a wide area under trees on sand and shingle, 1966, G. P.
WILSON, comm. A. W. ROBSON.

485/2. GALIUM BOREALE L. 48, Merioneth; volcanic rock in Cwm Pryser, near
Trawsfynydd, close to where Thomas Ruddy reported it at about the end of the last
century, WEST WALES NATURALISTS’ TRUST, MERIONETH BRANCH (1964, Nature Wales,
9, 73).

485/3 x 4. GALIUM X POMERANICUM Retz. 88, Mid Perth; roadside verge, Dall,
Rannoch, one plant, 1966, A. W. RoBSOoN.
485/10. GALIUM ULIGINOSUM L. 96b, Nairn; rare on the 25 ft raised beach north

of Maviston Farm, Culbin, M. McC. WEBSTER.

§495/3. VALERIANA DIOICA L. *89, E. Perth; marshes, Loch Cluny, local,
A. W. ROBSON.

506/6 x 7. SENECIO xX VISCIDULUS Scheele. 83, Edinb.; Arthur’s Seat, E. P.
BEATTIE.

7507/2. DORONICUM PLANTAGINEUM L. 89, E. Perth; abundant in woodland,
particularly under sycamore and ash, Binn Hill, A. W. ROBSON.

§7512/1. INULA HELENIUM L. *93, N. Aberdeen; by streamlet at Raemurrack—
road from Davidston to Cairnie, A. J. SouTER. 96, Easterness; south bank of Beauly
River below Lovat Bridge, J. N. MILLS.

PLANT RECORDS 59

§513/1. PULICARIA DYSENTERICA (L.) Bernh. *106, E. Ross; roadside by Glen-
marksie, near Loch Luichart dam, a well-established colony, U. K. DUNCAN.

§515/2. GNAPHALIUM NORVEGICUM Gunn. *105, W. Ross; many plants in a
gully, Cluanie Forest, A. McG. STIRLING & E. C. WALLACE.

540/7. CIRSIUM HETEROPHYLLUM (L.) Hill. 48, Merioneth; in quantity in wet
pasture and Corylus scrub near Trawsfynydd, the first record for the vice-county this
century, P. M. BENOIT.

§551/2. PICRIS HIERACIOIDES L. *+48, Merioneth; side of disused railway line
Barmouth Junction, 3 plants when first noted in 1967, increased to c. 40 in 1968,
clearly introduced in this artificial habitat in acidic country, P. M. BENOIT.

555/1. MYCELIS MURALIS (L.) Dumort. 93, N. Aberdeen; roadside wall, Drum-
blair, near Forgue, 1966, A. J. SOUTER.

§556/1. SONCHUS PALUSTRIS L. *11, S. Hants. ; estuary, Beaulieu, Mrs GOODHART,
det. D. MCCLINTOCK, comm. A. BERENS.
577/6 x 1. POTAMOGETON XSPARGANIFOLIUS Laest. ex Fr. 103, Mid Ebudes;

submerged near margin of loch, Loch Riaghain, Isle of Tiree, U. K. DUNCAN, det.
J. E. DANDY.

§577/12. POTAMOGETON RUTILUS Wolfg. 1103, Mid Ebudes; submerged, Loch an
Eilein, Heylipoll, Isle of Tiree, U. K. DuNCcAN, det. J. E. DANDY.

§594/1. FRITILLARIA MELEAGRIS L. t750, Denbigh; on a wooded bank by the
river Dee, Overton-on-Dee, 1944, Major P. BoLLom (K), see also Turrill, W.B., Roy.
Hort. Soc. Lily Year Book 1951-52, 108 (1952), comm. D. H. KENT.

597/1. GAGEA LUTEA (L.) Ker-Gawl. 88, Mid Perth; on river sand under trees,
banks of Turret burn, Crieff, 1966, A. W. ROBSON.

§603/1. PARIS QUADRIFOLIA L. +106, E. Ross; among rocks in wood by small
stream, north side of Loch Achilty near Contin, Mrs J. APPLEYARD, comm. U. K.
DUNCAN.

628/2. LISTERA CORDATA (L.) R. Br. 109, Caithness; moorland, Dunnet Head,
M. McC. WEBSTER.

630/1. GOODYERA REPENS (L.) R. Br. 88, Mid Perth; coniferous woodland,
Balgowan Woods, Findo Gask, 1966; wood partly felled in 1967, and unable to
rediscover plant in 1968, G. P. WILSON, comm. A. W. ROBSON.

§633/1. CORALLORHIZA TRIFIDA Chatel. *80, Roxburgh; willow carr, Blackpool
Moss, 800 ft, 1956; willow carr, Branxholme, Wesser Loch, 900 ft, 1965, R. W. M.
CORNER.

§644/1. ACERAS ANTHROPOPHORUM (L.) Ait. f. *38, Warwick; a few plants in
rough grassland on calcareous soil at Ufton Fields, D. J. JEFFRAY.

§7646/1. ACORUS CALAMUS L. *82, Haddington; in small quantity in Pressmen-
nan Lake near Stanton, E. P. BEATTIE.

§650/2. LEMNA TRISULCA L. *84, Linlithgow; abundant in canal, E. P. BEATTIE.

652/3. SPARGANIUM ANGUSTIFOLIUM Michx. 88, Mid Perth; artificial pond,
Dalnaspidal, mouth of Allt Dubhaig, 1967, A. W. RoBSon.

652/4. SPARGANIUM MINIMUM Wallr. 48, Merioneth; in quantity, fruiting in a
pool near Garth Angharad, Arthog, apparently the first certain Merioneth record of
this species as distinct from S. angustifolium which is frequent in the mountain lakes,
P. M. BENOIT.

§655/3. ScIRPUS MARITIMUS L. *94, Banff; small colony in brackish pool, East
Bay of Portsoy, towards East Head, A. J. SOUTER.

656/1. ELEOCHARIS PARVULA (Roem. & Schult.) Link ex Bluff, Nees & Schau. 48,

Merioneth; pools in saltmarsh on the Mawddach estuary near Penmaenpool,

a second station in the vice-county, barren as elsewhere in Wales, P. M. BENOIT.

60 PLANT RECORDS

656/4. ELEOCHARIS MULTICAULIS (Sm.) Sm. 49, Caern.; Cors Graianog, in old
peat cuttings, C. A. SINKER, comm. R. H. ROBERTS.
§656/6. ELEOCHARIS UNIGLUMIS (Link) Schult. 161, S.-E. York; in two marshes

between Wansford and Driffield, E. CRACKLEs, det. S. M. WALTERS (1969, Naturalist,
1969, 21).

§663/4. CAREX HOSTIANA DC. *82, Haddington; Aberlady Nature Reserve,
1967, E. P. BEATTIE.

§663/7. CAREX LEPIDOCARPA ‘Tausch. *82, Haddington; Aberlady Nature
Reserve, 1967, E. P. BEATTIE.

§663/23. CAREX STRIGOSA Huds. *49, Caern.; Penrhyn Park, near Bangor,
M. J. WIGGINTON, comm. R. H. ROBERTS.

663/28. CAREX LIMOSA L. 88, Mid Perth; Sphagnum bog, river Quaich near
Loch Freuchie, A. W. ROBSON.

663/36. CAREX CARYOPHYLLEA Latourr. 96, Easterness; bank by river near Glen
Mazeran Lodge, M. McC. WEBSTER.

§663/48. CAREX AQUATILIS Wahlenb. *83, Edinb.; Thriepmuir Reservoir near
Balerno, 1967, P. MYERSCOUGH, comm. E. P. BEATTIE. 109, Caithness; a large clump by
the Reay burn east of the golf course; mouth of Forss river, A. BERENS and M. McC.
WEBSTER.

663/56. CAREX DIANDRA Schrank. 93, N. Aberdeen; marsh north-west of Cairnie
Junction on border of v.c. 94, A. J. SouTER. 98, Argyll; Kerrera Island, near Ballie-
more, in a small marshy area, 1967, A. G. KENNETH, det. E. C. WALLACE.

§663/81. CAREX DIOICA L. *§82, Haddington; Aberlady Nature Reserve, 1967,
E. P. BEATTIE.

669/3. GLYCERIA DECLINATA Bréb. 88, Mid Perth; stream, Dunning golf course,
1966, A. W. ROBSON.

672/5. VULPIA AMBIGUA (LeGall) More. 4, N. Devon; Braunton Burrows,
1967, M. M. SAYER, conf. M. McC. WEBSTER.

673/c. PUCCINELLIA CAPILLARIS (Lilj.) Jansen. 94, Banff; stony path by the sea,

Strathlene: 108, W. Sutherland; among rocks by the sea, Port Skerra, Melvich,
M. McC. WEBSTER, det. C. E. HUBBARD.

§674/2. CATAPODIUM MARINUM (L.) C. E. Hubbard. *§82, Haddington; wall at
Barns Ness, E. P. BEATTIE.

4676/14. POA PALUSTRIS L. 89, E. Perth; riverside, Inchyre, 1966, A. W.
ROBSON.

1676/15. Poa CHAIxiI Vill. 89, E. Perth; avenue under ancient trees, Dunkeld
House policies, two small colonies, A. W. ROBSON.

683/3. BROMUS BENEKENII (Lange) Trimen. 33, E. Glos. Pope’s Wood near
Painswick, 1932: 34, W. Glos.; above river Wye at English Bicknor, 1966, E. MILNE-
REDHEAD.

§685/2. AGROPYRON DONIANUM F. B. White. *94, Banff; roadside cliff, Craig-
halkie, Tomintoul, 1965, A. J. Sourer, det. A. MELDERIS.

§686/1. ELYMUS ARENARIUS L. *83, Edinb.; shore at Cramond, E. P. BEATTIE.

§693/1. HELICTOTRICHON PRATENSE (L.) Pilg. *82, Haddington; shore at
Barns Ness, near Dunbar, E. P. BEATTIE.

696/4. DESCHAMPSIA SETACEA (Huds.) Hack. 108, W. Sutherland; pool by
bridge over river Naver, Altnahara, M. McC. WEBSTER.

700/1. CALAMAGROSTIS EPIGEJOS (L.) Roth. 104, N. Ebudes; rocky gully,
Tokavaig, Sleat, Isle of Skye, E. C. WALLACE & A. McG. STIRLING.

PLANT RECORDS 61

§701/4. AGROSTIS GIGANTEA Roth. *101, Kintyre; Knapdale, Isles of Danna;
south of Crear, near Kilberry, 1965, A. G. KENNETH, det. C. E. HUBBARD.

§709/1. MILIUM EFFUSUM L. 106, E. Ross; small rock outcrop by small stream
in wood, Teandore, Black Isle, U. K. DUNCAN.
§711/1. HIlEROCHLOE ODORATA (L.) Beauv. *79, Selkirk; boggy depression

beside stream, parish of Roberton, 875 ft, 1965, R. W. M. CorNER, comm. H. MILNE-
REDHEAD.

§7713/2. PHALARIS CANARIENSIS L. tH6, Waterford; grass verges and tip
south of Waterford on Tramore Road, 1967, I. K. FERGUSON (1968, Ir. Nat. J., 16, 97).

|

Watsonia, 8, 63-68 (1970). 63

Book Reviews

Abbreviated Titles of Biological Journals. Third edition, compiled by P. C. Williams.
Pp. viii + 47, interleaved. Biological Council, London. 1968. Price 12s. 6d.

This booklet has been produced for the convenience of contributors to, and editors
of, journals covering a wide field of biology and medicine, with the aim of guiding
them quickly to accepted abbreviations of titles of journals most likely to be referred to.
It is essentially an alphabetical listing of the full titles of the 1420 journals most
frequently cited in 45 periodical publications of societies affiliated to the Biological
Council (therefore with a strong British bias), together with their corresponding
abbreviations given in the fourth edition (1963-1965) of the World List of Scientific
Periodicals. As a concession to those who prefer the American Standards Institute
abbreviations to those of the British World List, the American version is also indicated
in brackets if it differs from the World List abbreviation. The list is interleaved with
blank pages to allow additional entries to be made.

To those not involved in such problems it may come as a surprise to learn that
abbreviation of journal titles has been a matter for prolonged argument and inter-
national debate. It does not seem to matter particularly to most people whether one
cites Am. J. Bot., Amer. J. Bot., Amer. Journ. Bot. or the full American Journal of Botany,
so long as the meaning is clear. However, librarians and computer feeders seem to be
unanimous in their wish to impose a uniform system of abbreviations on the whole
world, which might seem a laudable object were it not for the fact that they cannot agree
among themselves as to which system should be adopted. Some of the problems are
lucidly and realistically discussed in the Foreword to this booklet by the compiler.
Here the main question is seen to be whether one should settle for the British (World
List) system or the American system, and as neither one nor the other has any con-
clusive argument in its favour the present booklet is a compromise between the two.
But having myself been brought up in the tradition of taxonomic botany I find both
these systems frequently exasperating from both practical and aesthetic points of view.
The long-established conventions used in a vast body of literature in taxonomic botany
are at variance with these more recently devised systems in many of the principles
involved. In the last five years, extensive lists of journal abbreviations have been
published in this country in Flora Europaea (vol. 1, 1964; vol. 2, 1968), in the Index to
European Taxonomic Literature (annually since 1966) and in D. H. Kent’s Index to
Botanical Monographs (1967), while in America the Hunt Botanical Library has
produced its colossal Botanico-Periodicum-Huntianum (1968), reviewed elsewhere in
this journal, and in Russia T. Zaikonnikova has produced a shorter list in Nov. Sist.
Vyss. Rast. (Leningrad) (1968). Although each of these differs from the others in minor
details they all reflect the long tradition of taxonomic botany and differ extensively from
the World List and A.S.I. abbreviations. The problem of selecting one universally
acceptable system of abbreviations is thus far from being even a single choice between
two alternatives, and at the moment international standardization seems to be only a
vain hope. We insular British should at any rate note the warning given in the Foreword
of this book that some of the abbreviations we happily recommend make words far
from acceptable in polite company in France.

Perhaps the main objection to the World List abbreviations is to their use of con-
tractions (particularly those without a full-stop after them) as distinct from abbrevia-
tions. Even people relatively familiar with the idea may be stumped by such contrac-
tions as Archo, Bd, Fd, Fdn, Fm, Reprium and Wld for Archivio, Board, Food, Federation,
Farm, Repertorium and World (not Wild), while Jb., Jbr., Mschr., Rc., Zbl. and dt. for
Jahrbiicher, Jahresbericht, Monatschrift, Rendiconti, Zentralblatt and deutsch may take

64 BOOK REVIEWS

even more getting used to. Abbreviation seems to have become an end in itself, and
intelligibility is sacrificed for no apparent gain. If one must distinguish between
Annales and Annalen why cannot one write them in full instead of the cumbersome
Annls and Annin? The space saved is immaterial and even a computer can cope with
the two extra letters with no difficulty. The decapitalisation of initial letters is of no
practical importance but is an added aggravation, particularly in proper names as in
Proc. bot. Soc. Br. Isl. And in view of this last abbreviation it is difficult to see why
bot. becomes botl (no full-stop!) in Rep. bot! Soc. Exch. Club Br. Isl. More important,
however, is the use of a now incorrect title such as Repertorium novarum specierum regni
vegetabilis, the original title of what changed to Feddes Repertorium (which is not given)
in 1943. Similarly, in the American equivalents given, the adoption of the titles
Allgemeine botanische Zeitung (which is in fact the subtitle) and Jahrbiicher fiir
Systematik, Pflanzengeschichte und Pflanzengeographie for Flora (Jena) and Botanischer
Jahrbiicher respectively, is to be greatly deplored. It is clear, of course, that these faults
cannot be attributed to the compiler of the book reviewed here but have originated in
the primary works from which this list is taken.

For those who want a handy guide to the World List or A.S.I. abbreviations of the
commoner biological and medical journals used in this country this booklet is a well-
compiled, well-produced and useful tool. The present reviewer, however, and many
others concerned in botanical bibliography, will regard many of the abbreviations
offered here with great misgivings. If one standard list is to be adopted by taxonomic
botanists then the Botanico-periodicum-Huntianum seems to be the most acceptable
on account of its much more sensible abbreviations and excellent comprehensiveness.

R. K. BRUMMITT.

The Use of Biological Literature. R. T. Bottle and H. V. Wyatt. Pp. ix + 286.
Butterworths, London. 1966. Price £2. 18s. Od.

This is the first book to attempt the formidable task of providing a guide to the use
of the literature of the various aspects of pure and applied biology, a field with numer-
ous interdisciplinary connections. The editors have adopted a form of presentation of
the material that makes it readable and every working biologist should read it. On the
other hand, this method detracts from its use as a reference work. This fault would
have been lessened if all chapters had been provided with an Appendix of reference
works like that provided by Hambler for Chapter 7.

The editors were the first to organise a course on the use of biological literature at
Bradford University, a course, however, rather biased towards biochemistry and
pharmacy, and of comparatively little value to the systematist. The editors inform us
that this volume is intended for use in planning courses for graduates and under-
graduates and they have accordingly appended sample questions for use in practical
work. I hope this volume will stimulate the provision of such courses in the use of
biological literature, both for those still at university and also for those who have
already started their research careers. No written guide, however good, is a substitute
for supervised practical work.

It is obvious that the several contributors had a more detailed knowledge of some
aspects of the subject that they were covering than they had for others. It would have
enhanced the value of this book if the editors had submitted their treatments to other
experts for comment and addition of important omissions, thereby producing a more
even treatment of the whole field. In general, I consider that agriculture and ecology
should have been more adequately covered and that the more important guides to
palaeontology, palaeozoology, palaeobotany and anthropology should have been,
included.

The editors have allowed their contributors to cover their subject in their own way.
Hambler’s coverage of botany is therefore quite different from that of Archer for zoology.
Hambler’s title ““Taxonomy, treatises and museums” bears little relationship to its
content. He largely restricts his coverage to volumes relating to the British flora and

BOOK REVIEWS 65

states that he is omitting the taxonomist’s guides. Why? The title of the book implies
it is a complete guide to biological literature and are there no budding taxonomists
among the graduates and undergraduates at our Universities? Surely they need to
be trained in the use of their particular tools, as much as other biologists! It is no
doubt because of this limitation that the following important taxonomic references
have been omitted. They are the encyclopaedic treatments of the plant world—
Engler’s Das Pflanzenreich and Die natiirlichen Pflanzenfamilien; De Candolle’s
Prodromus systematis naturalis regni vegetabilis; Bentham and Hooker’s Genera
plantarum, etc. Obviously for the same reason, Hambler does not mention the Index
Nominum Genericorum and the important microfiche reproductions of several famous
herbaria including that of the most famous of all systematic botanists, Linnaeus.
The Bibliography of Agriculture is not referred to in this chapter, in spite of the fact
that a number of British taxonomic botanists consider that it is the best guide to date
to taxonomic literature. Surely such journals as the American Fern Journal, Revue
algologique, Revue bryologique et lichénologique and Revue mycologique, which contain
abstracts covering the literature on the lower plant groups, should have been included
in this chapter and also in the chapter on Abstracts, Indexes, etc. Just’s Botanisches
Jahresberichte and Botanisches Zentralblatt are not mentioned by Hambler but are
given in the chapter on Abstracts, etc., where these services are stated erroneously to
be the current German ones; they both “‘died’”’ during the war. The chapter on Abstracts
etc., does not mention the German service which is currently being published,
Excerpta botanica; on the other hand this is given by Hambler. This is a good example
of the lack of editorial control which is unfortunately evident elsewhere in this volume.
Hambler, to my surprise, does not include in his appendix of reference works such
important general titles as Tansley’s The British Islands and their Vegetation, Good’s
Geography of the flowering plants; Engler’s Die Vegetation der Erde and Kerner &
Oliver’s The natural history of plants. Nor does he include the very important guides to
literature: Pritzel’s Thesaurus Literature Botanicae and the library catalogues of the
British Museum (Nat. Hist.), Royal Botanic Gardens, Kew, Linnean Society and Royal
Horticultural Society.

No two historians would write the same brief guide to the development of plant
science. I personally regret the following omissions from the chapter on history and
biography: the very important role played by scientific societies and their publications,
especially that of our own Royal Society; planned natural history exploration, initiated
in this country by Sir Joseph Banks, which in later years gave opportunity to travel to
such famous naturalists as Charles Darwin and has led to the superb collections of
natural history specimens at such institutions as the British Museum (Nat. Hist.) and
the Royal Botanic Gardens, Kew; finally the outstanding contribution to the advance-
ment of systematic botany and zoology made by Carl Linnaeus. In the appendix of
references to this chapter I was surprised to find that Britten and Boulger’s A Biogra-
phical Index to deceased British and Irish botanists was omitted.

Dr Wyatt in his epilogue provides a stimulating discussion of the need, the value and
the methods of communication. This, I feel, would have been better placed as a
prologue. Finally, I am sorry that the editors decided to limit the scope of the book to
The Use of Biological Literature. 1 hope they will therefore consider widening the cover-
age in future editions, and also, by the assistance of a wider range of contributors,
produce a more comprehensive and balanced treatment of the subject.

PHYLLIS I. EDWARDS.
The Friends of John Gerard (1545-1612), Surgeon and Botanist. Robert H. Jeffers.

Pp. 99. The Herb Grower Press, Falls Village, Connecticut. 1967. Available in Britain
from Daniel Lloyd, 4 Hillcrest Avenue, Chertsey, Surrey. Price £1. 10s. Od.

Mr Jeffers, like many others before him, has fallen under the spell of John Gerard
and his Herball. This new book is an attempt to defend Gerard against well-founded

E

66 BOOK REVIEWS

criticism of his scientific standards and conduct as an author, and with this in mind the
events of his life are set out in chronological sequence. It is, however, unlikely that
many readers will be persuaded to modify their acceptance of balanced and scholars’
judgments of Gerard’s virtues and failings such as those set out in C. E. Raven,
English Naturalists from Neckam to Ray (1947).

The importance of Mr Jeffers’ book is that it is almost the first attempt to make use
of local record offices for botanical research. It is a reminder that even at this late
stage in collecting information about early naturalists, the methods and sources
employed by local historians offer scope for further discoveries. Mr Jeffers has pro-
duced new facts about Gerard and his contemporaries (his field extended far beyond
Gerard’s “‘friends’’) by consulting sources such as A London Subsidy Roll, 1589, which
were not available to earlier writers. Unfortunately the absence of page references
makes it difficult to check most of his statements and to ascertain which of them are
new. Failure to include an index also reduces the value of the book.

No reasons are given for the acceptance of the unlikely record of Veronica spicata
from Barnes (p. 34). This also appears in How’s Phytologia, being based on Veronica
recta minima, and has long being regarded as an undoubted error (cf. Boulger in
Salmon, C. E. (1931) Flora of Surrey, p. 43. London). Mr Jeffers goes on to state that
Gerard found Veronica spicata at Barnes independent of Stephen Bredwell.

Mr Jeffers has embarked on a promising line of research. It is to be hoped that he
will continue his work in record offices and publish it with full references so that it can
be put to maximum use.

J. E. LousLey.

Flora Europaea. Volume 2. Rosaceae to Umbelliferae. Edited by T. G. Tutin, V. H.
Heywood, N. A. Burges, D. M. Moore, D. H. Valentine, S. M. Walters and D. A.
Webb, with the assistance of P. W. Ball, A. O. Chater and I. K. Ferguson. Pp. xxvii +
455 with 5 maps. Cambridge University Press, London. 1968. Price £7. 7s. Od.

As the scope and aims and the origin of Flora Europaea have already been explained
in detail in this journal (cf. Watsonia, 6: 319, (1967)), this review will welcome only the
second volume. This will be cheered by all botanists who have used the first one with
success. They will feel as satisfied in using the new one, which appears only 4 years
later. The editorial committee, now increased by D. M. Moore (Reading), and the
51 contributors have done a remarkable work, as some large families are included:
Rosaceae, Leguminosae, Umbelliferae. The volume covers the second part of Rosales,
the Geraniales, Rutales, Sapindales, Celastrales, Rhamnales, Malvales, Thymelaeales,
Guttiferales, Violales, Cactales, Myrtales and Umbelliflorae, and includes the Cucurbi-
tales as well.

In the apogamous genera the leading species are described and keyed and their
minor relatives mentioned beneath them without description but with full citation.
Rubus has been evaluated by Y. Heslop-Harrison, Alchemilla by S. M. Walters and
B. Pawlowski and Sorbus by E. F. Warburg and Z. E. Karpati. It may be worth while
to mention the larger genera in this volume: Astragalus, worked out by A. O. Chater,
Vicia and Lathyrus by P. W. Ball, Trifolium by D. E. Coombe, Hypericum by N. K. B.
Robson and Viola by D. H. Valentine, H. Merxmiiller and A. Schmidt.—All informa-
tion about characters, habitat and distribution is given in the same approved way as
before; and so are the maps, the family key and the general chapters that have been
repeated from the first volume, partly in a more concise form, and completed by
the few necessary additions.

One essential advantage of this work is that it elucidates the taxonomic and geo-
graphical connections and graduations of taxa that appear too much isolated in local
floras. May it be used by many people, and may its good progress continue.

F. MARKGRAF.

BOOK REVIEWS 67

Humming Birds and their Flowers. K. A. Grant and V. Grant. Pp. 115. Columbia
University Press, New York and London. 1968. Price £7. 17s. Od.

Verne Grant started as a biosystematist with a strong grounding in evolutionary
genetics and produced some valuable scientific books before entering the field of
floral ecology. He and his wife proceeded to make observations in the field and syn-
thesized information from all angles, first for the Polemoniaceae as a family, now for a
different cross-section of the ecosystem.

Their joint investigations into the interactions between flowers and humming birds
(mainly in the western U.S.A.), already published, deserve to be reproduced in the
present colourful compilation and presented to a broader public willing to digest fully
the scientific material.

The plates (75 colour photographs, mainly of birds visiting flowers) are attractive and
also illustrative of the importance of ornithophily in the U.S.A. (41 proven and 86
probable cases), though in most places (as far as Alaska) the birds are migratory
components of the ecosystem. The genus Castilleja predominates, as basically bird-
adapted. Penstemon and Aquilegia are of secondary importance, being bird-adapted in
some species only. One has seen sharper photographs, but those presented have a
special flavour, the catching of purely natural conditions.

The adaptive evolutionary changes in the bird-flowers are analysed, along with the
taxonomy, distribution, anatomy and ethology of their bird-partners. The lack of
flower-constancy in humming birds is recognised as leading to mixed visits. The
consequent visits to many flowers of one plant are also described. Possible regulations,
ensuring cross-pollination nevertheless, are mentioned briefly.

The prevalence of convergent red coloration is causally analysed. The possibilities
of (i) mere preference for red or (11) better perception by birds and better contrast with
foliage (cf. red berries for fructivorous birds) are dismissed, on the grounds that there
are some counter-indications and also (not entirely convincing) that in tropical regions
with resident flower birds, non-red flowers are frequently visited. The authors favour a
synecological explanation, viz. that the migrant birds in the north require a common
signal, indicating “‘their’’ species of flowers, which may occur widely scattered among
others and are preferred for their quantity of nectar, which is greater than in insect-
flowers. Other colours are already associated with insects (butterfly-red is apparently
negligible). In this way the flowers, too, obtain specificity and a way out in the competi-
tion for pollinators.

The Grants found, after quantitative elaboration of their observations, that mountain
regions are seasonal coilecting grounds for birds and are thus also regions where
ornithophiles from diverse plant genera congregate. One might make the objection
that the sympatric development of many ornithophiles in one genus remains un-
explained, on account of the lack of flower-constancy and of other speciating factors
in the visitors. The authors answer that even in Castilleja the many species are rarely
sympatric and that (especially in the tropics) length of bill and tube and also differences
in the biotope provide stronger causes of speciation.

Conclusion: The book is a modern, causal approach to biosystematics and floral
ecology, important to evolutionists, both botanists and zoologists, and, moreover,
highly enjoyable for all general naturalists.

L. VAN DER PIL.

Welsh Ferns, Clubmosses, Quillworts and Horsetails. H. A. Hyde and A. E. Wade,
fifth edition by S. G. Harrison. Pp. xii + 178 with 14 plates and 85 text figures.
National Museum of Wales. Cardiff. 1969. Price £1. Ss. Od.

Welsh Ferns was first published in 1940 and quickly established its reputation as the
best descriptive systematic account of British ferns, for despite its title all British
species were included. The call for three new editions during the succeeding 22 years
was proof of its soundness and utility. The present edition, revised, expanded and

68 BOOK REVIEWS

rearranged by the new Keeper of Botany at Cardiff, differs more from previous editions
than any of these did from one another.

Welsh Ferns has now become Welsh Pteridophytes by inclusion of all species of
Lycopodium, Selaginella, Isoetes and Equisetum. These cover 21 species the descriptive
and distributional treatment of which conforms with that of the ferns. The ferns proper
have been rearranged according to Pichi-Sermolli’s system of classification. These
changes, together with the necessarily expanded introduction and key to genera, plus
the inclusion of much new matter covering additional species, subspecies and hybrids,
revised Welsh records and textual emendations and additions, have necessitated
resetting the pages throughout the book.

The descriptions are models of clarity and the overall standard of presentation of
information is so consistently high that few points call for comment other than praise.
The usual data covering distribution in Wales and beyond have been accidentally
omitted after Polypodium vulgare. If four variants of Equisetum arvense qualify for
inclusion, why not also the striking E. palustre var. polystachyum which is plentiful at
Newborough Warren? Recent critical work on Asplenium trichomanes and Asplenium
hybrids is incorporated in the account of this genus and Dryopteris assimilis is added to
the list of Dryopteris species. But how long will it be before Athyrium flexile receives its
just recognition as a distinct species, a status unanimously accorded to it by all who
have seen the living plant but rejected by those who know it only as herbarium
specimens? In cultivation at Leeds it retained all its peculiarities unaltered for many
years. The negative evidence of a chromosome complement identical with that of
A. distentifolium has been largely responsible for denying to it its rightful status.

Fifteen text figures have been supplied to cover the additional groups included in
this edition and two more plates have also been added. The drawings combine artistic
merit with diagnostic utility, though Fig. 21 will require replacing in the next edition
for it depicts the large and small leaves of Selaginella kraussiana in the opposite
relationship to one another to those they actually occupy. Is it not time also that the
dusky and too familiar figures illustrating the life-history of the Male Fern, which have
done service for close on a century, were now replaced by drawings more in harmony
with those which illuminate the rest of the book? These are however minor criticisms
of a volume which is likely to remain for a long time to come the standard work on
British Pteridophytes. Mr Harrison deserves congratulations and thanks for adding
still further to the value of a book of proved worth.

W. A. SLEDGE.

The Identification of Flowering Plant Families. P. H. Davis and J. Cullen. Pp. 122 with
6 text figures. Oliver and Boyd, Ltd., Edinburgh and London. 1965. Price 12s. 6d.

This book is divided into 10 parts: Preface, introduction, usage of terms, examining
the plant, using the key, key to the groups, arrangement and description of families,
further identification, glossary, and index. A list of abbreviations used is on the inside
of the back cover.

As indicated in the preface, the keys were written for use in the North Temperate
regions. Mexico, Florida, most of India, and subtropical China are excluded. I and my
students have used the keys extensively in two parts of the United States, central
California and western Montana, for two seasons. The keys work very well for plants
of these regions with very few exceptions. For instance, some western North American
species of Hydrophyllaceae have 1-celled gyncecia and hence cannot be determined
readily in the key.

The section entitled ‘“‘Usage of terms’’ is excellent and the discussion of the terms
hypogynous, perigynous, and epigynous is the best I know of in English. Other sections
are useful and well done. This book should receive wide usage in North America, as
well as in Europe.

JOHN H. THOMAS.

Watsonia, 8, 69-74 (1970). 69

Obituaries

KATHLEEN BEVER BLACKBURN

It was with great regret that her friends learned of the death of Dr Kathleen Bever
Blackburn in August 1968.

Dr Blackburn graduated with honours in Botany at Bedford College, London, in
1913. From 1914 to 1918 she was Lecturer in Botany at Southlands Training College,
Battersea, where she took her M.Sc. in 1916 for a thesis on plant anatomy. She was
awarded the D.Sc. of London University in 1924 for her contributions to the investiga-
tions of sex in plants and in 1930 was awarded the Trail Medal of the Linnean Society.
In 1918, she was appointed to a lectureship at Armstrong College (later King’s College.)
Newcastle-on-Tyne, and she remained there for the rest of her academic career. She
became Reader in Cytology in 1947 and retired in 1957.

Perhaps her most important contributions to science were those which she made in
collaboration with the late Professor J. W. Heslop Harrison. They were both remarkable
people; and they combined a deep interest in flowering plants with an awareness of the
importance of genetics and cytology in the interpretation of evolution and the clarifica-
tion of taxonomy. In the period 1921-24 they produced papers on the cytotaxonomy of
the Salicaceae, and the genus Rosa, which were amongst the first of their kind, and
which laid the foundation for the tremendous advances in cytotaxonomy and bio-
systematics in later years. The cytological work in these studies was all done by
Dr Blackburn. Incidentally, they produced the first reports of sex chromosomes in
flowering plants (in Populus), and Dr Blackburn produced convincing evidence a little
later in Silene dioica and S. alba. She continued to study the cytotaxonomy of the
Caryophyllaceae; and she also inspired a long series of post-graduate students to
undertake work of the same kind; her reputation was international. Towards the end
of her career, she became interested in pollen analysis, and in this field, too, she made
important contributions, carrying out investigations of peats both in the Hebrides and
in the north of England.

This steady output of distinguished work was combined with a heavy teaching load.
In particular, she was a pioneer in the teaching of practical plant cytology, and her
course was famous. She spared no pains to make her classes as full and exact as
possible, and her technique was excellent. She was in demand as a University tutor;
and she was respected and liked by her pupils and, indeed, by all who came into
contact with her.

To naturalists in the north-east of England she gave devoted service. Herself a skilled
field botanist, she explored the countryside, often to collect material for her classes or
for her research work. She was secretary of the Northern Naturalists’ Union from
1940 to 1955, and served on the management committee of the Hancock Museum.

When I went to Durham in 1945, Dr Blackburn characteristically took time and
trouble to introduce me to local people both inside and outside the University. She
was a charming person, ever ready to help when it was needed, but never making a fuss.
It was tragic that soon after her retirement she became seriously ill, and was unable
to travel or, eventually, to work. Had she kept her health, she would have rejoiced in
the vigorous growth of the Northumberland and Durham Naturalists’ Trust and its
activities in the counties which she had known and served so well.

PRINCIPAL PUBLICATIONS

1921
(With J. W. H. Harrison) The status of the British Rose forms as determined by their
cytological behaviour. Ann. bot., 35: 159-187.

70 OBITUARIES

1923
Sex chromosomes in plants. Nature, 112: 687-688.

1924
The cytological aspect of the determination of sex in the dioecious forms of Lychnis.
J. exp. Biol., 1: 413-430.
(With J. W. H. Harrison) Genetical and cytological studies in hybrid roses. 1. The
origin of a fertile hexaploid form in the pimpinellifoliae-villosae crosses. J. exp. Biol.,
1: 557-570.
(With J. W. H. Harrison) A preliminary account of the chromosomes and chromosome
behaviour in the Salicaceae. Ann. bot., 38: 361-378.

1925
Chromosomes and classification in the genus Rosa. Am. Nat., 59: 200-205.

1927
Chromosomes and their relation to Rose problems. Am. Rose A., 1925: 54-58.
Polyploidy within a species (Silene ciliata). Nature, 120 : 157-158.

1928
Chromosome number in Silene and the neighbouring genera. Zeitschr. Vererb.,
Suppl., 1: 439-446.

1929
On the occurrence of sex chromosomes in flowering plants, with some suggestions as to
their origin. Proc. Int. Cong. Plant Sci., 1: 299-306.

1930
Polyploidy within the species. Internat] Cong. Bot., Sth, Cambridge. Abs. Commun.
Pp. 145-146.
(With J. J. Boult) The status of the genus Saponaria and its near allies considered in
the light of their cytology. Proc. Univ. Durham phil. Soc., 8: 260-266.

1933
Notes on the chromosomes of the duckweeds (Lemnaceae) introducing the question of
chromosome size. Proc. Univ. Durham phil. Soc., 9: 84-90.

1934
Wasting disease of Zostera marina. Nature, 134: 738.

1936
Notes on Valerians. Vasculum, 22: 52-55.

1938
On the occurrence of a hermaphrodite plant of Empetrum nigrum L. J. bot., 76: 306—
307.

1939
The Crowberry. Vasculum, 25: 12-24.
The Limosella plants of Glamorgan. Part 2. Chromosomes and species. J. bot., 77:
67-71.

1946
On a peat from the Island of Barra, Outer Hebrides. New Phytol., 45: 44-49.

1949
Chromosomes and classification in Rosa. In Wilmott, A. J. (ed.), British Flowering
Plants and Modern Systematic Methods, pp. 53-57. London.

1952
The dating of a deposit containing an elk skeleton found at Neasham, near Darlington,
County Durham. New Phytol., 51: 364-377.

1953
Notes on modern research methods in taxonomy. Rep. Trans. Soc. Guernés., 15: 169-170.
(With A. W. Adams) Cytology in Herniaria. Proc. bot. Soc. Br. Isl., 1: 380.

1957
(With J. K. Morton) The incidence of polyploidy in the Caryophyllaceae of Britain and

Portugal. New Phytol., 56: 344-351.
D. H. VALENTINE.

OBITUARIES 71

HUMPHREY GILBERT-CARTER
(1884-1969)

Address delivered at the Memorial Service in Trinity College Chapel, Cambridge,
Ist March, 1969.

In 1909 there entered Trinity College a young man of 25 who, having gained a medical
qualification in the University of Edinburgh, and taken subsequent clinical study in
Marburg, had relinquished medicine for botany, and had come to work as an ad-
vanced student under C. E. Moss, then Curator of the Herbarium in the Cambridge
Botany School. It was a decision of moment both for Humphrey Gilbert-Carter and
for botany in Cambridge, for, after serving for the years between 1913 and 1921 as
economic botanist to the Botanical Survey of India, he was recalled to the newly-
created Directorship of the University Botanic Garden, a post which he held, in associa-
tion first with curatorship of the Herbarium, and later with a University Lectureship,
until retirement in 1950.

I little realised in 1921, as a callow and intense student beginning Part II of the
Tripos, that Humphrey and I were being launched side by side into a period of close
and rewarding friendship ended only now.

In the Botanic Garden hewas essentially the scholar-director rather than the executive
director. His Guide to the University Botanic Garden and his Descriptive labels for
Botanic Gardens, both published within three years of his appointment, were lucid,
erudite and original. These qualities, together with his intense capacity for friendliness,
attracted from all quarters a general sympathy for and interest in the Garden, not
least from Reginald Cory, who having already subsidised the Guide and the Director’s
House, finally left to the University his munificent Cory Bequest.

Humphrey was charged at his appointment “‘to teach Systematic Botany in the
Garden’ and indeed he always exhibited a strong aversion from formal lecture-room
courses designed for force-feeding data into notebooks and minds, infinitely preferring
the role of informal teacher, setting alight the imagination of his pupils so that they
hastened off to do for themselves the work of garnering and consideration. Who will
dispute that this ancient practice is in fact the very heart of education?

Most of ail it was in his frequent and gladly attended field excursions that Humphrey
displayed his outstanding quality as a teacher. In the words of his pupil and successor
to the Curatorship of the Herbarium, Dr S. M. Walters, ‘““He practised naturally and
without strain that fundamental technique of the gifted teacher: his pupils were instructed
in the delightful game of learning by an enthusiast who could not prevent his interest
in people, or his love of botany, from shining through, however dull the day or tedious
the text-book.”’ Those who, like himself, had the perfectly retentive mind of the taxono-
mist, would follow at his heels, gathering now and then an explicit exposé of a taxono-
mic problem to which the text-books yielded no clue whatever: these gleanings came to
light or had their undisclosed importance in the work of taxonomists and naturalists
for decades afterwards.

Above all, his former students will recall the heart-warming friendliness of the
Sunday tea-parties that he and Dorine gave at Cory Lodge, ably abetted by their
beautiful and unselfconscious daughters, a function so generally popular that double-
ranking round the table was the rule. The ice was broken by Humphrey’s mischievous
medical-school jocularity and the prim were often disconcerted to find, deposited in
their lap, a shaggy and affectionate Old English sheep-dog whose warm breath was
alleged to have strong therapeutic value. ‘Grace’ and ‘Grasshopper’, ‘Lightning’ and
“The Little One’... how could students thus addressed feel out of the warmth of the
family circle that enlarged so much as the years went by and that has so many of its
members come here today?

Humphrey Gilbert-Carter was a man of formidable accomplishments and wide
knowledge. His own disavowal of any claim to be a scientist sprang largely from a

TP HUMPHREY GILBERT-CARTER

modesty that undervalued his own capacity and from his own high personal integrity
of thought. It was these qualities that enabled Humphrey without effort to establish
and retain the friendship of many great scholars in this and other Universities. Only
one class of friendship would I prefer to this: I mean the quick and unreserved friend-
ship of small children and young people, folk who respond instinctively to gentleness
and assurance: Humphrey also had this abundantly.

Humphrey had tremendous facility for languages and the great Oxford lexico-
grapher, C. T. Onions, said of him that, outside the ranks of the professionals, Humphrey
was the best-informed philologist he had ever met. This facility was constantly at the
disposal of students and friends: there were German and Scandinavian reading classes
and sessions to initiate or revive contact with the beauties of the Georgics. He translated
a large part of the life-work of Raunkaier from the Danish, embellished his Guide to
the Garden with terms in Sanskrit, Arabic and Hindustani, and could take up con-
versation in their own language with the surprised lascars on a P. & O. liner. Small
wonder that his Glossary of the British Flora, which went through three editions, was
so authoritative and so welcome. We may recall with a smile what he wrote in Catkin-
bearing plants concerning the right use of words: “It is the duty of the teacher to
check the facility with which students learn words of whose meaning they are entirely
ignorant, a process comparable to swallowing flints, which are not only entirely
indigestible, but which because of their relentlessness will set up lasting internal
mischief.”

I shall, I trust, have made it clear that Humphrey Gilbert-Carter was a broad man.
He would undoubtedly have approved Bacon’s view that ““Travel in the younger sort
is a part of education; in the elder a part of experience.’’ He travelled widely himself
and always shunned a parochial attitude, aiming to bring Cambridge botany into close
touch with that of other countries. It is a habit of contact that has happily continued,
and the success of that current and massive undertaking, Flora Europaea owes no
small debt to it. Nor was Humphrey’s own approach to the practice of taxonomy in
the least narrow: he would have us smell and taste a plant, view it whole in the field
and have full regard to its favoured ecological situations and biological responses. He
exhibited sympathy with, though not participation in, the movement of younger
taxonomists, many his old students, into fields of experimentation and even into
enquiry as to the basic purposes and processes of taxonomy itself.

Humphrey was one of the least self-seeking of men, rejecting personal ambition in
favour of devotion to the interests of his students, who have repaid him through the
years with a warmth of affection and gratitude rare even in this home of dedicated
teachers. He will be remembered not only in his own considerable published work but
in the larger volume that he inspired in his pupils. His modest view of his own memorial
can fittingly close my remarks. At his first meeting with the Old Botanic Garden
Syndicate in May, 1921, he recommended the planting of widely-set and well-grown
specimen trees, and urged .. . “Let us look ahead and think of those who will take our
places. Neither shall we ourselves die altogether if the coming generation of students
remember us when they study the great trees on the lawn, or the elders of the University
and town bless our memories as they rest beneath their shade.”

H. GopDwiIn.

I would like to add a few personal recollections of Humphrey Gilbert-Carter to
Professor Godwin’s excellent appreciation of him. I do so in the hope that they will be
of interest to those who did not know him; those who did will have many happy
memories of their own, but they may like to be reminded of the existence of Fragments
of a Botany School Excursion Pantomime (Tea Phytologist, 4/37r3, 8—9 (1950)).

I first met Humphrey at the beginning of the Long Vacation Term in 1928, at the
end of my first year in Cambridge, a year in which one thing had become clear to me:
that whatever I ultimately did, it would not be Botany. Long before the end of the
6-week term I had learnt that Botany is a special way of life linking the Arts and

HUMPHREY GILBERT-CARTER 73

Sciences. My earlier decision was soon reversed and I had no second thoughts about it.
Every excursion, whether a short one after tea to Coe Fen or the railway ballast pits
at Chesterton or a whole day at Wicken or on the Breckland, was not only full of
interest but also an adventure. Humphrey’s older friends sometimes came on these ex-
cursions and consequently our outlook was widened by meeting perhaps a Consul
from a remote part of Asia or a distinguished chemist. So, without our realizing it, he
made us aware of the interdependence of all kinds of knowledge and of the richness of
the world.

On short suburban excursions, usually between tea and opening time, we not only
looked at weeds and garden escapes but were shown the gardens of his friends or even
of total strangers. If Humphrey saw some unusual plant or exceptionally fine tree in
someone’s garden, he would go up to the door, knock, and after a short conversation
with the owner, be invited to bring his class in to see the garden.

He was always kind and considerate; the most severe reproof that I ever heard him
administer was ‘‘that was very incurious of you’’. During his period as President of the
Botanical Section of the Cambridge Natural History Society he encouraged the,
usually young and inexperienced, speakers by the way in which he opened the dis-
cussion. On one occasion when a paper had been given with which he completely
disagreed he started by saying “‘Of course there are two sides to everything except...”
(then there was a pause long enough for everyone to wonder what was coming)
.. . “Cambridge station’’. This, Iam sure, helped greatly to make the ensuing discussion
less acrimonious than it would otherwise have been.

Humphrey, though no mean performer on the ‘penny whistle’, was not really
interested in music. One summer evening after a concert to which we had been, Mrs
Gilbert-Carter asked some of us to go home with her for a cup of tea. The night was
fine, so naturally we strayed from the kitchen into the garden to look for the hedgehogs
which frequented it. Before long an unmistakable voice from the darkness above said,
“IT am observing the satellites of Jupiter; come and join me’’. We went upstairs and
found Humphrey with a small telescope, anxious to share his enjoyment with us.

On a winter afternoon shortly after the war we walked together through Trumping-
ton to Grantchester and, as we were about to come back across Grantchester Meadows,
he suggested that we might see if Tansley was in. He was, and on his own, so he asked
us to stay for tea. Over tea Tansley asked me to write a British Flora. I had never
thought of such an undertaking, but before we left plans had been drawn up and were
speedily put into action. So Humphrey not only instructed the three of us in field and
herbarium, but was a vital link in the chain which led to the production of the Flora.

At the lunch given to celebrate his 80th birthday he said “‘Our search is a search for
beauty; those who go into the field and observe, like those who ascend the spiral stair-
case whose treads are made of DNA, are seeking for the same thing’’. I have to quote
from memory and so may not have worded it as well as he did.

I count myself fortunate indeed in having had such a teacher and friend for over 40
years.

T. G. TUTIN.

BIOGRAPHICAL NOTE

Humphrey Gilbert-Carter was born on 19th October 1884, the second son of
Sir Gilbert Thomas Gilbert-Carter, K.C.M.G. He was educated at Tonbridge School
and the Universities of Edinburgh, Marburg and Cambridge. He was Economic
Botanist to the Botanical Survey of India from 1913 to 1921 and married in 1914,
having four daughters. From 1921 until his retirement in 1950 he was Director of the
Cambridge University Botanic Garden and, from 1930, University Lecturer in Botany.
He was an Honorary Associate of the Linnean Society of London. He died on January
4th, 1969.

74 BIOGRAPHICAL NOTE
BIBLIOGRAPHY

1913
Genera of British Plants. Cambridge.
1917
Report on the Industrial Section of the Indian Museum for the year 1916-17. Calcutta.
1921
(With D. N. Carter) Useful Plants of the District of Lakhimpur in Assam. Records of
the Botanical Survey of India, 6 (9): 353-420.
1922
A Guide to the University Botanic Garden. \st ed. Cambridge.
1924
Descriptive Labels for Botanic Gardens. Cambridge.
1930
Our Catkin-bearing Plants. 1st ed. Oxford, 2nd ed. Oxford (1932).
1934
Translations of C. C. Raunkiaer, The Life Forms of Plants and Statistical Plant
Geography.

1936
British Trees and Shrubs. Oxford.
1937
Translation of C. C. Raunkiaer, Plant Life Forms.
1947
A Guide to the University Botanic Garden. 2nd ed. Cambridge.
1950
Glossary of the British Flora. 1st ed. Cambridge, 2nd ed. Cambridge (1955), 3rd ed.

Cambridge (1964).
He also contributed the botanical entries to the Cambridge Italian Dictionary. General
Editor, Barbara Reynolds. Vol. 1, 1962; vol. 2, (in the press).

T. G. TUTIN.

Watsonia, 8, 75—95 (1970). 75

Reports

REPORT OF THE COUNCIL FOR 1968

This report and the audited accounts relate to the calendar year 1968. Figures in
brackets are the 1967 ones, for comparison.

The year 1968 was a notable one for the Society in four ways in particular: a long-term
programme of ‘network research’ was launched; British Sedges, the first volume of what
the Society hopes may be a series, met with a tremendously successful reception; the
Society entered into discussions with botanical societies of three other nations to
consider a proposal for some form of European federal structure; and the annual
subscription was raised. If it was a year in which steadily rising costs at length forced
the Society to face the inevitable and take essential corrective measures, it was a year
which also saw promising developments in several new directions.

It was a year, too, of sharp recovery in the recruitment of new members, after the
unusually poor results in this respect in 1967. The 128 who joined constituted, in fact, a
19 % increase over the year before. As usual, the South of England accounted for almost
two-thirds of this figure. One in every six, it is encouraging to note, was a Junior Member.
This rise, moreover, was not accompanied by any parallel rise in membership erosion:
89 (90) were lost by reason of death, resignation or the operation of Rule 28. The year
ended with a membership total of 1,733 (1,694), representing a net gain of 39 (18). The
result was sufficient to halt the steady decline in the annual rate of growth that had been
in evidence over the last four years.

This rise in membership, nevertheless, was not nearly sizeable enough to enable the
Society to postpone for any longer the raising of the annual subscription—which has
been held at the same level for no less than eleven years. Like most other bodies of its
type, the B.S.B.I. has no means of sidestepping the general inflation of costs, the effect of
which has been particularly marked on two of its indispensable staples: postage and
printing. On the recommendation of Council, accordingly, it was agreed at a Special
General Meeting on November 23 that the annual subscription for Ordinary and
Subscriber Members be raised by ten shillings, to £2, with effect from Ist January 1969.
While a breathing-space has now been gained, therefore, it is clear that economies
must be pursued and even greater efforts made to increase membership, if the Society is
to avoid a continuing depletion of its reserves.

As will be seen from the accounts which are appended, during the past year the
Society’s expenditure substantially exceeded its total income from subscriptions, from
dividends and interest, and from the sales of journals. The charge for the year for the
printing and distributing of our journals includes rather more than a normal year’s
charge at current prices for Proceedings but less than a full year’s costs for Watsonia.
For the first time the accounts show separately the expenditure incurred on storing and
distributing the back numbers of our journals, a share of these costs having been allocated
to the Publications Fund. Other expenditure contains little of an abnormal kind and
includes about £200 for the services of the Institute of Biology for various secretarial
purposes. A new item is the cost of printing and postage for the ‘network research’
programme launched in 1968. Dividends and interest received show a Satisfactory
increase, but this source of income cannot be maintained if our reserves are depleted by
expenditure in excess of annual income.

The Publication Fund shows that proceeds from the sales of British Sedges exceed the
expenditure incurred, after deducting the grant received from the Royal Society.
Altogether about 2,500 copies had been sold by the end of the year and as a steady
demand continues for this excellent handbook, it was decided to print off an additional

76 REPORTS

500 copies before the type was broken up; there will then be nearly 1,000 copies available
for the future. Other satisfactory items in the Publications Fund are the receipt of
£417 7s Od. in respect of royalties from the Critical Supplement to the Atlas of the
British Flora, further royalties from the Atlas itself and from Conference Report No. 9,
as well as initial royalties from D. H. Kent’s Index of Botanical Monographs.

THE MEETINGS COMMITTEE, after an exceptionally heavy year in 1967, in 1968 enjoyed
something of a respite. There were just two non-routine events to organise: a wine-and-
cheese party, in the evening following the Anual General Meeting, to celebrate the
publication of the Critical Supplement to the Atlas of the British Flora: and a further
Local Flora Writers’ Conference, this time at Aberystwyth, at which some 50 or so
members engaged in a stimulating exchange of views on the kinds of items that ought to
be included in any local Flora worthy of the name. The Society is much indebted to
Dr J. P. Savidge for his help in organising this useful and enjoyable weekend.

It was London’s turn this year for the Annual General Meeting, which duly took
place in the rooms of The Linnean Society in May. Fifty-eight members were present to
hear, as the prelude to this, the first Presidential Address, ‘The B.S.B.I. in a Changing
Britain’. The afternoon of the same day was given over to a series of talks introducing the
new programme of ‘network research’, and this was followed by an excursion to
Virginia Water, in Berkshire, on the Sunday.

The Committee also organised the customary Annual Exhibition Meeting in
November, held as usual in the Department of Botany, British Museum (Natural
History), by kind permission of the Keeper. Forty-three exhibits were staged, maintaining
the high total newly achieved in the previous year. There was the normal large attendance
of members and guests, of whom 65 stayed on for the now traditional evening Reception.

In addition to a visit to the South London Botanical Institute during the winter, four
field meetings were arranged—a return from the bare minimum to which the English
total was temporarily cut down as an experiment in 1967. These were at Bury St.
Edmunds, in Hampshire (for orchids), at Taunton, and in Co. Durham (for roses). The
meeting at Taunton doubled up as the South West Regional Meeting, at which this
Region duly elected its Representative for the vacancy on Council.

THE JUNIOR ACTIVITIES SUB-COMMITTEE arranged two field meetings in Britain in 1968—
in Cornwall and Perthshire—and another of its very popular meetings overseas, this time
in the Alps once again. Applications for this last proved unexpectedly numerous and
many, regrettably, had to be turned down because of the limited accommodation
available. In compensation, a further meeting on the Continent is being planned with
the least possible delay.

THE DEVELOPMENT AND RULES COMMITTEE had a very quiet year—so quiet, in fact, that
it has begun to have doubts about its continued usefulness. The major items considered
have been the present operation of the regional system in England and various matters
pertaining to publicity. The latter have led to the setting up of a small, specialist working
party to explore this whole aspect in fuller detail.

THE PUBLICATIONS COMMITTEE Was One of two standing committees to have a change of
secretary during the year, Mr Wanstall finally retiring after a record-breaking fourteen
years in office. It has been responsible for the production of two parts of Proceedings
and two parts of Watsonia (for which a new cover has also been designed). Plans for the
Society to publish further, companion volumes to British Sedges, dealing with other
difficult groups have been among other matters discussed.

Early in the year the viability of the Society’s much-appreciated abstracts service, at
any rate in its present form, was considered a sufficient matter for concern to warrant the
setting up of a joint working party with the Flora Europaea organisation to review its
character and position within a wider context. This body has so far carried out a wide-
ranging survey of the existing network of abstracting and indexing services in plant

REPORTS UW

taxonomy as a whole and is devoting study, more particularly, to the possibilities of
mechanised retrieval in this field. Should this prove feasible, the way may be opened to
integrating the Society’s service with others that run more or less in parallel, streamlining
its operation and—to some extent at least—safeguarding its continuity.

Mention should also be made here of the publication by Academic Press on the joint
behalf of the B.S.B.I. and the Linnean Society of the Report of the very successful 1967
Conference at the University of Liverpool. The thanks of the Society are due to Professor
Heywood for undertaking the very considerable task of editing this.

THE CONSERVATION COMMITTEE was the other standing committee to have a change of
secretary during the year. The Society is indebted to Mr Streeter for all his work on its
behalf in the past—and, in a sense, henceforward as well; for he will be continuing to
serve the Committee’s cause at one remove, in his capacity as chairman of the new
liaison committee of the Society for the Promotion of Nature Reserves. Three permanent
representatives of the B.S.B.I. Committee were also invited to sit on this useful new body,
which in future will act as a ‘clearing-house’ for the conservation concerns of a whole
range of societies such as ours and represent them in regular discussions with the
Nature Conservancy. Through this new machinery the Committee has already been
consulted on a proposed code governing introductions into reserves and of the pos-
sibility of producing for schools a list of plants that can be collected with the minimum
risk of harm. The Committee similarly contributed representatives, by special invitation,
to the County Trusts’ Conference at Canterbury and to a reception at St. James’s Palace
held in connection with “The Countryside in 1970’ Conference. It also gave advice on the
botanical implications to the new Inland Waterways Amenity Advisory Council. All the
while, throughout the year, the progress of the Wild Plant Protection Bill, in the framing
of which representatives of the Society have played a leading part from the very first,
was watched with understandable anxiety. In this connection it was, perhaps, ironic
that two of this year’s more worrying ‘threats’ were the digging up of Lady’s Slipper
Orchids in Yorkshire and the commercial exploitation of primroses in Devon.

THE RECORDS COMMITTEE was enlarged this year to include representatives of each of
the three Regional Committees. It has continued with its long-term task of improving
the Society’s whole system of record-collection, to this end supplying to further Recorders
who requested them sets of index drawers and cards. It has continued, too, to keep the
progress of the many local Floras known to be in preparation under close review and
provided specialist advice in this connection in one or two further instances. An impor-
tant new departure was the inauguration, at the end of the year, of a newsletter to
Recorders. This work in improving communications was furthered by the holding of
another Local Flora Writers’ Conference and by meetings that the Secretary was
fortunately able to have during the year with Recorders in both Scotland and Ireland. A
meeting of the Committee to debate the future basis for the Society’s recording con-
cluded with a recommendation to adhere to the Watsonian vice-county system, for a
variety of reasons, despite its acknowledged imperfections. In addition, it was agreed
that in future grid-references should accompany records published by the Society
(though, obviously, this will be done with discretion) and that efforts should be made to
press ahead with a census catalogue, so that a pocket register of up-to-date vice-county
distributions can be made generally available.

This Committee also took over responsibility during the year for the new programme
of ‘network research’. The initial responses to both of the first two enquiries have been
encouraging, well over 300 completed records card for the Symphytum survey having
been received by the end of October. Both enquiries will be continuing in 1969 and a
third, a Mistletoe survey, is planned for the coming winter season.

Of the three Regional Committees—
THE COMMITTEE FOR THE STUDY OF THE SCOTTISH FLORA arranged ten field meetings, of

78 REPORTS

which one had to be cancelled through lack of support. The nine which took place were
in Lanarkshire, Argyllshire, Upper Nithsdale, Lammermuir Deans, the Isle of Tiree, the
Isle of Mull, East Lothian and (two) Midlothian. In addition, the Annual Exhibition
Meeting was held jointly with the Andersonian Naturalists of Glasgow in the Department
of Botany, University of Glasgow, by kind permission of Professor P. W. Brian.

THE WELSH REGIONAL COMMITTEE also arranged its customary full programme of field
meetings. Those in 1968 were in the counties of Caernarvon, Pembroke, Denbigh,
Glamorgan, Montgomery and Brecon. The Annual General Meeting was at Gregynog
Hall, Montgomery, in September, and a number of exhibits were shown at this. The
Region also staged an exhibition at the annual conference of the Council for the
Protection of Rural Wales in Aberystwyth, which aroused considerable interest. The
Welsh Regional Bulletin did not appear during 1968, but an issue is to be circulated early
in 1969. Two editors have now been appointed and each will be responsible for a
January and August issue respectively.

THE IRISH REGIONAL COMMITTEE has now instituted a newsletter. Two extended field
meetings were held in under-worked counties (Co. Cavan and North Tipperary) and a
day meeting in the Boyne Valley. The Annual General Meeting and Exhibition took
place in late October at University College, Dublin.

WARBURG MEMORIAL FUND. By the end or the year, donations received in response to the
joint appeal with the British Bryological Society for the Warburg Memorial Fund
amounted to £474. This sum is considerably less than the target which it is hoped to
achieve. Members have recently been reminded of the appeal and it is hoped that those
who have not already made a donation to the fund will now send one to the Hon.
Treasurer. It will be remembered that the intention is to make an award out of the Fund,
annually or less frequently, to a young botanist for travel, preferably in the period
between leaving school and going to university. A joint committee with the British
Bryological Society is being established to formulate the rules for making awards out
of the Fund.

The Council thanks the many friends of the Society who have helped during the year
to further its progress and also all those members who have taken an active part in
organising its affairs, on committees, as representatives on other bodies, and in other
ways. In particular, thanks must be expressed to the authorities of University College,
London, the Nature Conservancy, the Linnean Society of London and the City of Perth
Museum and Art Gallery, for allowing the regular use of their premises for Council
or committee meetings; to the University College of Wales, Aberystwyth, for allowing the
Local Flora Writers’ Conference to be held there; and to the Trustees of the British
Museum (Natural History), for the use of their Lecture Hall and Conversazione Room
in connection with the Annual Exhibition Meeting. Finally, we thank our Honorary
Auditors, Messrs. Price Waterhouse & Co., for their continuing services.

J. G. Dony, President.
D. E. ALLEN, Hon. General Secretary.
11th March 1969.

BALANCE SHEET

REPORTS

AS AT 31st DECEMBER, 1968

79

1967 1967
General Fund: Investments:
Balance 1st 44% Defence
January 1968 ... £4,238 3 10 Bonds ... £500 O
Add: Premium on 5% Defence
conversion of Bonds ... . 2,000 0 0
Defence Bonds... 30 0 O 5% National
Development
4,268 3 10 Bonds ... 1,000 0 0
Less: Excess of 54% National
expenditure over Development
Income for year 785 11 10 Bonds 500 0O
£4,238 £3,482 12 0 6% British
3,437 Publications Fund . . 4,446 1 2 Savings Bonds... 1,000 0 0
40 Benevolent Fund ... ; 39 11 O 84% Local
2,409 Creditors for Printing and General Authority
Expenses ... 2,404 19 11 Bonds ... 1,000 0 0
108 Subscriptions Received in Advance 171 5 0 The Equities
Investment
Fund for
Charities—
750 units
(Market ea
£1,602) .. .. 1,429 13 9
£5,000 £7,429 13 9
Cash at Bank:
Deposit Account ... 2,500 0 0
Current Account .. 614 15 4
5,232 3,114 15 4
£10,232 £10,544 9 11] £10,232 £10,544 9 1
J. C. GARDINER,
Hon. Treasurer.
GENERAL INCOME AND EXPENDITURE ACCOUNT
for the year ended 31st December, 1968
1967 1967
Cost of Printing and Distributing: £2,372 Subscriptions received ... £2,483 19 1
Proceedings, 426 Dividends and interest received Sikes Ugh 98)
Vol. VII, Part 2 Proceeds from sales:
(balance) and provi- Proceedings EBIEXS. OF
sion for Part 3 yea oO SLO Watsonia ... 206°) AKO © 2!
Watsonia, Old B.E.C. Reports 30 3 10
Vol. VI, Part 6 508 ————._ 466 13 9
(balance) and prov- — Excess of expenditure over income
ision for Vol. VII, Charged to General Fund 785 11 10
Part 1 904 4 10
Storage and distri-
bution of back
numbers (less provi-
sion) t eae 94 2
LD 2 ———_—_—— £3,309 8 10
275 Printing and Distributing Annual
Report and Notices of Field
and Indoor Meetings .. 291) OF 2
35 Expenses of Council and Com-
mittee Meetings 66 16 0
100 Travelling Expenses of Officers,
Secretaries and Members at-
tending Meetings 100 13 0
443 General printing, stationery, post-
ages, telephone and petty ex-
penses ... Bb acts fe LOO Ze 9
137 Expenses of Regional Organisation,
including Card Indexes for
Recorders 7D) Wi
— Network Research printing and
postages 119 4
29 Field and Indoor “Meetings, ex-
penses less fees received. Gs 5)
20 Contribution to Wild Plant Pro-
tection Working Party —
155 Excess of income over expenditure
credited to General Fund _—
£3,306 £4e3 15) 1118) £35306 Se ee la

el Be

80 REPORTS
PUBLICATIONS FUND

Transactions in 1968

1967
1967
Cost of Printing, Advertising and Distributing: | £3,415 Balance from 1967 £3,437 0 4
£223 British Sedges SO 17/2 Sales:
Grant from Royal — British Sedges me 1-620 litiaed
Society ... sue 2SOMMOM SO 128 Conference Reports 60 6 9
——————— £1,252 17 2 14 Other publications 47 11 10
— Storage and Distribution of back —————._ 1,728 16 2
numbers sk fee Bae 60 0 O Royalties received:
9 Printing price-list of publications .... — 95 Atlas of the British
Balance to 1969: Flora’ BET 142 tS kG
General spo 2s 7 & — Critical Supplement to
Atlas Royalties less the Atlas sax OPAL Se
outgoings soo P40] NS) SG 17 Conference Report
3,437 ——————- 4, 446 1 2 Norore2.: aoe 619 0
— Index to Botanical
Monographs ... 26 10 4
—————_ 593 1 10
£3,669 £5,758 18 4] £3,669 £9575 Sone

TEESDALE DEBENCE COMMITTEE REPORS

The only transactions during 1968 on the Teesdale Defence Fund have been the receipt
of interest amounting to £82 5s. 9d. on the Deposit Account. The question of whether
or not the Fund can be accorded charitable status remains unresolved. Following legal
advice, further representations were made to the Charity Commissioners, but neither
they nor the Inland Revenue have yet been prepared to reverse their views. At the
present time we are awaiting the settling by Counsel of a formal submission to the
Commissioners following which the Commissioners will make their final determination;
this could be reversed only by appeal to the High Court. Although no monies have been
disbursed during the year liabilities in respect of legal fees amounting to about £150
have been incurred and will have to be discharged before the Fund can be wound up.

J. C. GARDINER, Hon. Treasurer.
11th March, 1969.

REPORTS $1

ANNUAL GENERAL MEETING, MAY 3rd, 1969

The Annual General Meeting of the Society was held at Merlewood Research
Station, Grange-over-Sands, Lancashire, on Saturday, 3rd May, 1969, at 12 noon.
Dr J. G. Dony (President) was in the Chair and 36 members were present.
The Minutes of the last Annual General Meeting were read and were adopted on the
motion of Mrs B. H. S. Russell, seconded by Mr E. F. Greenwood.

REPORT OF COUNCIL

The Report together with the Accounts for the year 1968 had been circulated to all
members. There being no queries, the Report was adopted on the motion of Miss
C. M. Rob, seconded by Mrs B. H. S. Russell.

ELECTION OF PRESIDENT

Mr E. Milne-Redhead had been nominated by Council. The Chairman said that
Mr Milne-Redhead had been a member of the Society for 35 years during which time
he had held many offices. It was Mr Milne-Redhead who had in fact introduced
Dr Dony himself to the Society. Unfortunately Mr Milne-Redhead was not able to be
present because the meeting coincided with Open Day at the Royal Botanic Gardens,
Kew. The Chairman proposed that Mr Milne-Redhead be elected and the proposal
was carried unanimously. Dr Dony continued in the Chair for the remainder of the
Meeting.

ELECTION OF OFFICERS

Following Mr Milne-Redhead’s election to the Presidency and Miss U. K. Duncan’s
wish not to be re-elected a Vice-President, Council had nominated Mr J. E. Lousley and
Dr F. H. Perring for election as Vice-Presidents, thus ensuring continuity of their
valuable work, and this, together with the re-election of Professor J. G. Hawkes, was
carried unanimously. Dr Dony said that Mr D. E. Allen, Honorary General Secretary,
wished to resign his office and his resignation had been received with much regret.
The Chairman spoke with praise of Mr Allen’s two very useful years of work as
General Secretary, during which time the Society had embarked on the Network
Research Programme and on a scheme to collaborate with other European botanical
societies. Mrs B. H. S. Russell proposed the Meeting’s warmest thanks to Mr Allen
for all his work, seconded by Miss C. M. Rob and followed by loud applause.

Dr I. K. Ferguson had been nominated to succeed Mr Allen. Dr Dony said that
Dr Ferguson’s work with Flora Europaea at the Department of Botany of the British
Museum and now his membership of the Kew staff were a fitting background for the
office. His election was proposed from the chair and carried unanimously.

Mr J. C. Gardiner had been nominated by Council for re-election as Honorary
Treasurer and he was re-elected unanimously.

Mr E. F. Greenwood, Dr M. C. F. Proctor and Dr N. K. B. Robson were unani-
mously re-elected as Honorary Editors. Dr G. Halliday had been nominated by Council
as a fourth Honorary Editor to take the place of Dr C. D. K. Cook who was no longer
working in this country. Dr Halliday’s election was carried. The Chairman voiced
the Society’s thanks to the hard-working Editors.

Mrs M. Briggs was proposed by the Chairman for re-election as Honorary Member-
ship Secretary following Council’s nomination and the meeting unanimously carried
the proposal.

AMENDMENTS TO THE SOCIETY'S RULES AND ELECTION OF AN HONORARY
MEMBERSHIP SECRETARY
The proposal to elect an Honorary Membership Secretary was discussed in con-

junction with the proposal to amend the Society’s rules. The following amendments
were approved.

$2 REPORTS

Rule 6: add ‘“‘Honorary Membership Secretary”

Rule 7: add ‘“(F) The Honorary Membership Secretary shall be responsible for the
Register of Members, and shall undertake such related work as may be
mecessary”’

Following Council’s nomination, Mrs J. G. Dony was elected and she will therefore

continue with the valuable work she has already done for many years.

The following amendment to Rule 11 was approved by the Meeting: “‘Notice of a
Regional Meeting shall be given at least two months in advance” (instead of three
months as at present) and “In the event of a nomination not being received four weeks
before the date of the meeting Council shall have the power to make a nomination’”’
(instead of eight weeks as at present).

ELECTION OF NEW MEMBERS TO COUNCIL

In accordance with Rule 10 Mr J. E. Lousley, Dr F. H. Perring and Mr P. J. Wanstall
retired. Three nominations had been received: Mr D. McClintock, Mr R. M. Burton
and Mr. H. G. Messenger and these were elected, their seniority for purposes of Rule 10
being decided (in the order shown above) by lots drawn by the Chairman.

ELECTION OF HONORARY MEMBERS

The election of Mrs C. M. R. Schwerdt and Sir Edward Salisbury as Honorary
Members was proposed by Council. Dr Dony in proposing that the Meeting approve
these nominations said Mrs Schwerdt was President of the Wild Flower Society and
had done much to promote interest in the British Flora. He went on to say Sir Edward
Salisbury, the ““Grand old man”’ of British Botany, had been a member of the Society
since 1914. The election of both as Honorary Members was unanimously carried.

ELECTION OF HONORARY AUDITORS

Council recommended the re-election of Messrs Price, Waterhouse & Co., who had
kindly agreed to continue this work on an honorary basis. Their election was duly
carried and the President expressed the Society’s gratitude for their services.

ANY OTHER BUSINESS

The position of Field Secretary was discussed, and it was decided to wait until such
time that Council thought it fit to revive the office.

Miss C. M. Rob voiced the feelings of all members of the Society when she thanked
Dr Dony, not only for all that he had done for the B.S.B.I. during twenty years in
office, culminating in his two years as President, but also for his great help to British
Botany and to Weeds, and the thanks were carried in loud acclamation.

There being no further business, the Meeting closed at 12.50 p.m.

I.K.F.
EvY.

NORTH-WEST REGION MEETING, 1969

A meeting of members resident in the North-west Region was held in the Library,
Merlewood Research Station, Grange-over-Sands, on Saturday, 3rd May, 1969
following the Annual General Meeting of the Society.

The Secretary of the Society opened the meeting and E. F. Greenwood was elected
to the Chair. The main business was to elect a Regional Representative to serve on the
Society’s Council in place of the retiring Representative, Dr C. D. K. Cook.

The only nomination had been A. Newton, proposed by E. F. Greenwood and
seconded by Miss V. Gordon. Mr Newton was declared elected.

After D. E. Allen and the Chairman had made a few comments on the purpose of
Regionalisation, the meeting was closed.

E. F. GREENWOOD.

REPORTS 83

COMMITTEE FOR THE STUDY OF
THE SCOTTISH FLORA

FOURTEENTH ANNUAL REPORT TO 31ST DECEMBER 1968

The Committee met twice, each time in Perth, and an Exhibition meeting was held
jointly with the Andersonian Naturalists of Glasgow on Saturday, 9th November, at
the Department of Botany, University of Glasgow, by kind permission of Professor
P. W. Brian.

Of the nine field meetings scheduled to take place during the 1968 season, one had
to be cancelled (River Mouse (Cranley Estate), Lanarkshire. Leader: Mr W. A. Scott)
because of lack of support. The others were to The Avon and Cander Valleys, Stone-
house, Lanarkshire, June 15th, led by Mr S. A. Birnage; Argyllshire, 22 and 23rd June,
led by Mr A. G. Kenneth; Upper Nithsdale, 29th and 30th June, led by Dr H. Milne-
Redhead; Lammermuir Deans, 29th and 30th June, led by Miss E. P. Beattie; Isle of
Tiree, 3rd—11th July, led by Miss U. K. Duncan; Isle of Mull, 13th—20th July, led by
Mr A. A. Slack; Gosford Bay, East Lothian, 20th July, led by Dr J. Milne; Crichton,
27th July, led by Miss E. P. Beattie and Cobbinshaw Reservoir, Midlothian, 3rd August,
led by Dr P. Myerscough.

The Exhibition meeting was attended by about 100 people of whom about 50 were
present for the evening session. A short meeting was held at 2 p.m. of B.S.B.I. members
normally resident in Scotland in order to elect the Regional Representative and three
other members to serve on the Committee for the next four years.

The following exhibits were shown:

B.S.B.1. Symphytum Survey.
Biological Records Centre’s Recording Schemes currently in operation.
Vice county Boundaries of Scotland.
Herbarium sheets of various plant groups including Potamogeton and Zostera.
Miss U. K. DUNCAN

Map and photographs of Inchcailloch, Loch Lomond National Nature Reserve.
E. T. IDLE
Elatine hydropiper L.—new to Scotland. Found near the mouth of the R. Endrick, on

the east side of Loch Lomond, on sand and mud exposed by the 1968 summer drought.
E. T. IDLE, J. MITCHELL & A. McG. STIRLING

28 herbarium sheets of Scottish plants. R. MACKECHNIE
Specimens illustrating the distribution of tetraploid (2n = 68) and hexaploid (2n = 102)
Campanula rotundifolia in the British Isles. H. A. MCALLISTER

Mull 1961- Map showing areas visited by the C.S.S.F. party and specimens of the
interesting plants found.
A. SLACK
Interesting plants found on the Isle of Mull C.S.S.F. Field Meeting 1961.
D. A. RATCLIFFE, B. W. RIBBONS & A. McG. STIRLING
The Filmy Ferns—Hymenophyllum tunbrigense and H. wilsonii. Fresh material was
exhibited from near Tarbert, Loch Fyne, where both species occur in some

abundance. A. McG. STIRLING
Herbarium specimens: (a) Pilosella officinarum, P. aurantiaca and P. x rubrum.
(b) Wool adventives and grain aliens. Miss M. MCCALLUM WEBSTER

Messrs E. B. Bangerter and J. F. M. Cannon gave a most interesting and stimulating
illustrated account of the immense amount of botanical work being done by the staff
of the British Museum and their many helpers, both professional and amateur, in
collecting data for the Museum’s Flora of the Isle of Mull.

After the most enjoyable Buffet Supper, slides were shown by a number of people of
the flora of Britain, the Faeroes, the Pyrenees, etc.

F*

84 REPORTS

A full programme of field meetings has been arranged for 1969, some of which are
in conjunction with other Societies. There will also be a few Junior meetings organised
by Miss Ailsa Burns, the B.S.B.I. Junior Meetings Secretary.

It is a pleasure to express thanks to the officers of the B.S.B.I. and B.S.E. and to all
those who have generously given of their free time to help the Committee to carry out

its activities.
E. P. BEATTIE.

EXHIBITION MEETING, 1968

An Exhibition Meeting was held at the Department of Botany, British Museum
(Natural History), London, S.W.7, on Saturday 23rd November 1968 from 12 noon
to 5.30 p.m. About 220 members and guests attended.

FUNGI OF NORTHAMPTONSHIRE WOODLANDS

Twenty Ferraniacolor slides were exhibited, being part of a series being produced of
Northamptonshire fungi. The following species, all of which are common, were
illustrated :

Aecidium compositarum (x5), Amanita phalloides, Auricularia auricula, Clavaria
cristata, Collybia distorta, C. fusipes, C. peronata, Empusa sp. (x 5), Hygrophoropsis
aurantiaca, Hypholoma fasciculare, Inocybe griseo-lilacina, Laccaria amethystina,
Lactarius quietus, Lycoperdon perlatum, Marasmius ramealis, Mucor sp. (x3),
Scleroderma aurantium, Sparassis crispa, Stereum rugosum, Trametes abietina.

A Praktina camera fitted with an 8in. lens on home-made bellows is used for all
normal work. This gives a much greater working distance, depth of field and better
perspective than the normal 2in. lens. All the slides were taken using electronic
flash. T. B. BARRATT.

Festuca altissima IN MERIONETH
The Azlas shows a single record for North Wales of Festuca altissima All., a 19th
century one of Thomas Ruddy’s from Llandderfel, Merioneth. Another from the same
period (Llanymawddwy, Merioneth, Miss May Roberts) was traced after the Atlas
was published. Both of these localities were lost.

Over the last few years, however, F. altissima has been found by P. M. Benoit,
Miss A. Burns, Miss V. Gordon and Mrs K. M. Stevens to be widespread in the
Dolgellau district of west Merioneth, in the less accessible parts of the glens with
basic (mostly dolerite) rocks. The following 10 stations in four 10 km squares are
currently known to exist in the county. These include the old station at Llanymawddwy,
which was rediscovered in 1968. Ruddy’s original locality has not yet been traced but is
also likely still to exist.

23/61. Panteinion glen, 1967, PMB & KMS. Bron Meirion waterfalls, 1966, PMB.
Arthog waterfalls, 1966, PMB. Abergwynant glen, 1957, VG; 1966, PMB. Ty’n y
Ceunant, 1967, PMB & KMS.

23/71. Ty Mawr, near Dolgellau, 1968, PMB. Torrent Walk, 1966, PMB & AB.

23/71 & 23/72. Nant Helygog, 1966, PMB & KMS.

23/82. Afon Eiddon, 1967, PMB & KMS. Llanymawddwy, 19th century, Miss May
Roberts; 1968, PMB & KMS.

(23/93. Llandderfel, 19th century, Thomas Ruddy.)

A specimen and distribution map were exhibited.
P. M. BENOIT.

REPORTS 85

CHROMOSOMES OF Spartina pectinata LINK IN IRELAND

The exhibit indicated the number and meiotic appearance of chromosomes in Irish
Spartina pectinata Link (Boyle 1968a, b).

Photographs of pollen mother cells at first meiotic metaphases and anaphases
showed 2n = 41. 20 bivalents and a univalent were present at metaphase. Anaphase
showed two groups of 20 chromosomes with the univalent lagging between them; in
one case the univalent was the same as at metaphase; in another the univalent appeared
to have divided into two parts which were very close to each other.

Chromosomes of Irish S. pectinata are similar to those described by Marchant
(1968) for S. pectinata recently introduced to Kew from Canada. Marchant interprets
the univalent as a B-chromosome.

REFERENCES

Boy Le, P. J. (1968a). Preliminary account of Spartina pectinata Link new to Ireland. Jr. Nat. J.,
16: 74-75.

BovLeE, P. J. (1968b). Spartina pectinata Link in Ireland. Proc. bot. Soc. Br. Isl., 7: 505.

MARCHANT, C. E. (1968). Evolution in Spartina (Gramineae), 3. J. Linn. Soc. (Bot.), 60:

411-417.

PATRICIA J. BOYLE

YELLOW-BERRIED Crataegus monogyna

Living and dried material was shown of Crataegus monogyna Jacq. bearing fruit
varying from yellow to scarlet from asolitary bush discovered during November 1960
by J. H. Chandler growing beneath conifers by the roadside at Essendine in Rutland.
A herbarium specimen with similarly coloured berries found by Miss I. M. Hayward
near Galashiels, Selkirk, in 1911 was also shown. A herbarium specimen collected by
J. P. M. Brenan of the true yellow-berried C. monogyna which was discovered by the
Cam Brook near Midford, N. Somerset, by Miss F. M. Barton during 1946 was
exhibited for comparison.

REFERENCES

GILBERT, J. L. (1966). Yellow-Berried Haw. Gdnrs’ Chron., 160: 12. (This paper gives several

further references.)
HADFIELD, M. (1966). Orange-fruited hawthorn. Gdurs’ Chron., 160: 16.
J. H. CHANDLER & J. L. GILBERT.

INTROGRESSIVE HYBRIDISATION BETWEEN BRITISH ANNUAL Senecio
SPECIES

The evidence for introgression from Senecio squalidus into S. vulgaris was given. This
is believed to result in variants of the latter species including var. hibernicus Syme.
Possible mechanisms for this introgression were discussed, and one of these was
illustrated, using the progeny from the triploid S. x Jondinensis (S. viscosus x
S. squalidus) as an example. The implications of the investigation are discussed in a

short Note (p. 47).
P. Crisp & B. M. G. JONES.

MORPHOLOGICAL DEVELOPMENT OF Spartina *X’

The exhibit comprised four herbarium sheets of fertile Spartina ‘x’ C. E. Hubbard
from Sandy Haven Pill, Pembrokeshire. These demonstrated the seedling stage, the
appearance of the first rhizomes, the production of the first inflorescence, and finally a
flowering and a seeding head, including husked seeds, from a mature plant.

86 REPORTS

Few fertile seeds per head have been found but seedlings occur in great abundance.
It is by seed that S. ‘x’ is dispersed so widely where it has become established; the
spread, slow in a newly colonised locality, becomes increasingly rapid as more and

more patches develop. 3
T. A. W. DAvIs

B.S.B.I. MEMBERSHIP

Five maps and charts were exhibited giving the following facts and figures:-
1. Rise and fall in membership, 1942 (343 members) to 1968 (1796+).
2. World map showing distribution of 164 Overseas members.

3. Proportion of men to women at 1st November 1968:

Men .. ne 063 G9-2,7,)
Women 7s 22" 53916007)
Subscribers .. 194 (10:8 %)

4. List of 79 present-day members who joined before 1939 when the Society was
known as the Botanical Exchange Club. Our most senior member is Mr G. C. Brown,
Colchester, who joined in 1911.

5. Regional representation:

% Of total members per 100,000

membership of population

South-east 701 members 38-9 4-30
South-west 240 A 13-4 4-92
Midlands 171 Bs 9-6 1:99
North-east 130 5 2 1:82
North-west 136 es 7:8 1-94
Scotland 120 on 6:7 2°38
Wales 94 Ne a2 3-55
Ireland 40 as 22, 0-95
Overseas 164 9-0

Mrs J. G. Dony.

Cardamine raphanifolia IN EAST GLOUCESTERSHIRE

Herbarium sheets of Cardamine raphanifolia Pourret (C. latifolia Vahl) were exhibited
from a site at Lower Slaughter in East Gloucestershire, reported by Mr R. C. L.
Howitt in June 1968. This species, which was not officially added to the British List
until about 1949, is a native of the mountains of southern Europe. In Great Britain
C. raphanifolia has been recorded from several localities in the Ambleside and Winder-
mere area, also from Dundee, Scalby and Goathland in north-east Yorkshire, and
Tonbridge in West Kent. It has been known from near Ambleside since 1932. At Lower
Slaughter it is well-established for half a mile along the banks of the River Eye, a
tributary of the River Windrush, in an area of downwash limestone gravels deposited
on lower lias clay at an altitude of 425 ft on an easterly dip-slope of the north Cotswold
escarpment. A few fruiting capsules were found in June 1968, but seeds were not then
fully developed.

Mrs R. DUDLEY-SMITH.

REPORTS 87

Lathyrus maritimus BIGEL. SUBSP. acutifolius (BAB.)PEDERSEN
IN EASTERN SCOTLAND

The specimens came from two large colonies on fixed dunes in Angus (Forfar).
Unfortunately the slightly fleshy character of the leaves was lost when dried. It should
be stressed that, although the plant was checked by an expert, it has not yet been seen
by Dr Pedersen and it would seem desirable that fresh material should be sent to
him. Miss U. K. DUNCAN.

Senecio x ostenfeldii DRUCE IN SCOTLAND

Two specimens from Angus, one from East Ross-shire and one from East Sutherland
were selected from a range of herbarium specimens to illustrate the variation in the
hybrid swarm which may occur where S. jacobaea and S. aquaticus are found together.
The variation in the size and number of the flower-heads and the shape of the leaves is
paralleled by the variation in the degree of hairiness of the achenes.

Miss U. K. DUNCAN.

Cardamine POLLEN
See Plant Notes (p. 45). R. P. Etuis & B. M. G. Jones.

Carex aquatilis IN THE CENTRAL AND EASTERN HIGHLANDS

Sheets of the small upland and tall lowland forms of Carex aquatilis were exhibited.
The upland form is only known to occur on the plateau around Clova and Caenlochan,
except for one isolated record from Ben More, Mid-Perth. Evidence was presented that
these forms represent parts of a continuous series, in which the size of several organs
decreases with increasing altitude.

Upland plants of C. aquatilis grow together with the closely related species C. nigra
and C. bigelowii in the bogs on the Clova plateau. An isometric scatter diagram has
been devised to assist the analysis of populations, which may include hybrids
between the three species. Its application to this Carex population was described.
The scatter diagram suggests that there may be hybridization between all three pairs
of species, particularly C. aquatilis and C. nigra, but little or no double hybridization.
The technique may be useful as a basis upon which to simplify the population into
species pairs, by eliminating all the plants showing characteristics of one species. Pairs
of species can then be treated with a more sensitive bipolar hybrid-index technique,
preferably one involving standardization of continuous variables. J. S. FAULKNER.

STEREOPHOTOGRAPHS OF BRITISH GENTIANACEAE AND OF
Minuartia recurva

J. H. FREMLIN.

SOME NEW BOTANICAL POSTAGE STAMPS
Mrs A. N. Gipsy.

Scabiosa columbaria SHOWING ‘‘BIRD’S NEST’” PROLIFERATION

A herbarium specimen was shown of Scabiosa columbaria L. exhibiting “‘bird’s nest’’
proliferation which was discovered by T. E. Patston and J. L. Gilbert at Barnack,
Lincolnshire, during 1957. Each plant first bore a single lateral stem carrying one
capitulum about 3:5 cm in diameter surrounded by leafy bracts longer than the
corolla. Later the stem branched apically into two similar branches each bearing

88 REPORTS

similar inflorescences. On the later-formed central or main stem of each plant a
collection of leaves formed a “‘bird’s nest’’, instead of a capitulum, from the base of
which there were up to 20 stems bearing smaller, normal capitula 1:5 cm in diameter.
As all the original plants were affected I suspected that the character had been inherited
and have since proved that this mutant always comes true to seed.

REFERENCES

GILBERT, J. L. (1958). Botanical Records, District 7, 1957. J. Northampt. nat. Hist. Soc., 33:
214-215.
GILBERT, J. L. (1967). Odd Scabious. Gdurs’ Chron., 161 : 15.
J. L. GILBERT.

SOME FLORAL ABERRATIONS IN Oxalis acetosella L. IN EPPING FOREST

The exhibit featured a series of annotated drawings of consecutive transverse sections
through an abnormal chasmogamous flower (floral formula KS C(5+1) A(5+1+5)
G(6)). A pictorial representation of the vascular skeleton was constructed and a
number of conclusions regarding the nature of the aberration were drawn.

1. Hexamery was based upon a structural pentamery.

2. Additional floral parts were arranged such as to produce a superimposed zygo-
morphy. This and other structural peculiarities require further study.

3. Preliminary studies of somatic chromosomes failed to demonstrate convincingly
that the aberration was correlated with an abnormal karyotype. The genetical basis
could not be inferred in the absence of population statistics, though this feature
characterised both chasmogamous and cleistogamous flowers in the whole local
population (clone ?). R. J. GILES.

A STUDY OF SOME WESTERN EURASIAN GLADIOLI

The Eurasian gladioli are a natural group of approximately a dozen species that are
centred on the Mediterranean Sea. They are separated from the majority of the genus
(a further 230 species) in Africa by the Sahara Desert. The western Eurasian gladioli
are four species which show considerable variation in many characters. They fall
naturally into two groups: 1) The cornfield weed Gladiolus italicus Mill. (G. segetum
Ker-Gawl.), the seeds of which are unwinged; 2) Three species with winged seeds,
G. byzantinus Mill., G. illyricus Koch and G. communis L.

The exhibit was devoted to the second group and distribution maps and colour
photographs of each taxon were shown. It included a hypothetical scheme for the
evolution of this group based on morphology, leaf chromatography and cytology.
It was suggested that the tetraploid G. illyricus, which shows a range of tepal shapes
from obovate to oblanceolate, may have given rise to three evolutionary lines: 1) to
hexaploid G. illyricus in Britain, with obovate tepals, 2) to tetraploid and octoploid
G. communis with rhomboidal tepals, and 3) to hexaploid and octoploid G. byzantinus.

A. P. HAMILTON.

A TREE SURVEY IN THE LONDON BOROUGH OF HARROW

The purpose of this exhibit was to bring to the attention of members of the Society the
survey being carried out in Harrow. The aim of this survey is to map the trees in and
around Harrow. This includes not only common roadside trees but some of the rarer
trees, such as Liriodendron and Taxodium, found in parks and gardens. The Council has
recently been planting such unusual trees as Ginkgo and Metasequoia. The distribu-
tions of certain trees was demonstrated on a large-scale map of Harrow using coloured
pins. Also shown was a collection of miniature trees in pots which included Viburnum
sp. and Taxus. HARROW COUNTY SCHOOL For Boys.

REPORTS 89
BRITISH SPECIES FROM SPITSBERGEN
B. M. G. JONES.

STILL MORE ABOUT MULL

The exhibit illustrated some aspects of the progress made during 1968 in the survey of
Mull and its adjacent small islands by the Department of Botany, British Museum
(Natural History).

The highlight of the season’s fieldwork was a series of visits to the Treshnish Isles,
using a helicopter placed at the disposal of the department through the courtesy of the
Royal Navy. Apart from a very few isolated records, the only previous information on
this group of islands was a B.S.B.I. Maps Scheme card for Lunga, the largest of the
islets. For the eleven islands from which lists are now available, a total of 204 species
of flowering-plants and ferns has been recorded. This may be compared with the total
of c. 850 species for the area as a whole. Some provisional attempts were made to
correlate the size and constitution of the floras of the individual islands with their
relative positions, size and ecological diversity. Maps, photographs of the islands and
a tabulation of all known records were exhibited.

The second section of the exhibit was concerned with the Pteridophytes of Mull.
A map showing some distribution patterns was exhibited together with specimens of
two subspecies of Asplenium trichomanes (subsp. trichomanes and subsp. quadrivalens
D. E. Meyer emend. Lovis) and sheets of the Dryopteris filix-mas aggregate as
represented on the island.

The third section was concerned with the species-pair Juncus bulbosus/J. kochii, one
of the critical problems that received special attention during 1968. J. kochii appears
to be very common and widespread on Mull, while J. bulbosus is apparently much
more restricted both in range and ecology. Subsequent examination of material in
the British Museum herbarium suggests that the latter species (as delimited in Clapham,
Tutin & Warburg, ed. 2) is apparently a rare plant with an arctic-alpine type of distri-
bution. The characters used to delimit these taxa are as listed by Benoit and Allen
(Proc. bot. Soc. Br. Isl., 7: 504 (1968)), except that in our experience the stamen number
for J. kochii is sometimes three and less value has been placed on the pigmentation of
the floral parts. As however there is a considerable divergence between the distribution
patterns as indicated above and that suggested by Benoit and Allen, it is clear that
much more study is necessary and the exhibit was offered in the hope that it will
stimulate further interest in these taxa. Living material of J. bulbosus would be
welcomed by Mr A. Eddy.

DEPARTMENT OF BOTANY, BRITISH MUSEUM (NATURAL HIsTory).

THE FLORA OF GUERNSEY

The exhibit comprised: (1) specimens of four of the notable discoveries in 1968,
(2) specimens of seven of the island’s specialities which should be seen on the expedition
in April 1969, and for which it was hoped to find fresh localities, (3) the current
check-list of Guernsey plants, (4) copies of articles on the flora of Guernsey in the
Gardeners’ Chronicle (March 29, April 5, April 12, (1968)), (5) a copy of an article on
x Asplenophyllitis microdon (Br. Fern Gaz., 10: 1-8 (1968), (6) a copy of The Natural
History of Guernsey by Mr N. Jee.

The four discoveries were:

(a) Exaculum pusillum. Seen in a second hollow in a new area, towards the east end
of L’Ancresse Common by Dr H. J. M. Bowen.

(b) Eryngium campestre, of which Dr Bowen found a very small colony on the
coast of L’Ancresse.

(c) Geranium cf. submolle Steud. Found by Dr Bowen at La Garenne. This species
is discussed in Plant Notes (p. 47).

90 REPORTS

(d) Geranium cf. robertianum L. A tall octoploid form of G. robertianum was
discovered by Mr P. Girard well naturalised in a waste area on the southern outskirts
of St. Peter Port.

The seven special plants were x Asplenophyllitis microdon (two more plants dis-
covered in 1968), Anogramma leptophylla, Ophioglossum lusitancum, Isoetes histrix,
Trifolium occidentale, Poa infirma and Milium scabrum.

D. MCCLINTOCK.

Senecio vernalis WALDST. & KIT. IN LEICESTERSHIRE (V.C.55)

This European species was found at two sites near Market Harborough in 1968. In
both cases a roadside verge had been newly seeded with imported grass, some of which
came from Holland and Denmark. At one site there was some evidence of hybridisation
with local S. vulgaris, but this requires a further study. Material of the species and of
the putative hybrid was exhibited, together with material from a similar roadside
locality in north Devon collected in 1961. K. G. MESSENGER.

Hypericum canadense L. IN THE BRITISH ISLES

This plant was first recorded for the British Isles by Prof D. A. Webb, who found it in
1954 on wet, acid ground by the side of Lough Mask, Co. Galway. It has now appeared
in a second locality, more than a hundred miles away from the first, in a wet sloping
meadow close to the car-park of the Eccles Hotel, Glengariff, Co. Cork. It was found
here, on 6th August, 1968, by Mr & Mrs K. L. Butcher.

Hypericum canadense, a native of eastern North America, has also been found in
the Netherlands and in the Vosges. Like Juncus tenuis and Sisyrinchium bermudiana, it
is most probably a recent arrival in Europe, actively extending its distribution, and not
a relict like Eriocaulon septangulare. R. D. MEIKLE.

THE NATIONAL WOODLAND SURVEY

The preservation of examples of the climax vegetation over much of Britain, namely
woodlands, constitutes an important part of our national conservation policy. The
acquisition and management of woodland nature reserves, therefore, features promin-
ently in the work of both the Nature Conservancy and County Trusts. Although
censuses have been made by the Forestry Commission and others, these do not give
sufficient information on which to base a woodland conservation policy.

To obtain this information the Nature Conservancy, in conjunction with the County
Naturalists’ Trusts, have organised a woodland survey, the purpose of which is to obtain
a record of the present day extent, location and characteristics of native-type wood-
lands. Details are noted on specially designed cards under three broad headings: site
description, trees and shrubs, and ground vegetation. This information is stored and
processed at the Biological Records Centre.

Amateur naturalists are encouraged to participate in the survey and Nature Con-
servancy staff will attend meetings to explain the techniques. Lists, cards and instruc-
tion sheets will be sent on request.

There are over 20,000 broad-leaved woodlands (over 10 acres) in Britain and in one
season records of 2,000 of these have been made.

NATURE CONSERVANCY, MONKS WooD EXPERIMENTAL STATION.

MONKS WOOD NATIONAL NATURE RESERVE

Twelve coloured slides of vascular plants from the Monks Wood collection were
exhibited. These show that whilst Monks Wood is mainly a woodland National
Nature Reserve, many other habitats are represented within the boundaries of the area.
There are extensive rides and a series of ponds, some open fields which have been cultiva-
ted in the recent past as well as the grassland which surrounds the Nature Conservancy’s

REPORTS 91

Experimental Station. This leads to a richness in vascular plant species. About 330 are
recorded from the Reserve including the following rare or local species:-

Platanthera chlorantha Carex pallescens
Colchicum autumnale Dipsacus pilosus
Centaurium pulchellum Rumex palustris
Peplis portula Neottia nidus-avis
Melampyrum cristatum Sorbus torminalis

Members were reminded that those who wish to visit the Wood must obtain permits
in advance. They should write to the Regional Officer of the Nature Conservancy,
60, Bracondale, Norwich, NOR 58B, and not to Monks Wood Experimental Station.

NATURE CONSERVANCY, MONKS Woop EXPERIMENTAL STATION.

THE B.S.B.1. Symphytum SURVEY

The Symphytum Survey began early in 1968 as one of the new Network Research
projects. By the end of October over 400 completed cards had been returned. The
exhibit was based on these and personal observations.

The initial results show that S. asperum Lepech. is extremely rare, whilst
S. x uplandicum Nyman is widespread and occurs throughout the British Isles. The
two forms of S. officinale L., the cream, var. ochroleucum DC., and the reddish-purple,
var. purpureum Pers., have different distribution patterns, the former mainly occurring
in eastern England whilst the latter occurs in the South and West. Introgression takes
place between the two, and mixed colour populations are extremely abundant in the
area of overlap, particularly in Berkshire, Hampshire and Dorset. Many of the
populations measured are intermediate between S. x uplandicum and the various
colour forms of S. officinale.

The scheme will continue in 1969 and 1970 and those who wish to take part were
invited to write for further record cards to the Biological Records Centre, Monks
Wood, Huntingdon. F. H. PERRING.

VASCULAR PLANTS NEW TO IRELAND

Two species which have been discovered for the first time in Ireland this year were

exhibited:

(i) Juncus compressus Jacq. Discovered by D. M. Synott by the south bank of the
River Boyne at Rosnaree, Co. Meath, on 5th August. The plant was seen again
during a Field Meeting of the Irish Regional Branch of the B.S.B.I. at Slane,
Co. Meath, on 17th August.

It seems very likely that this is a species native to Ireland which has previously
been overlooked. Its occurrence in eastern Ireland is what would be expected on
phytogeographical grounds as elsewhere in the British Isles it is mainly confined
to southern and eastern England.

A full account of the discovery appears in the Irish Naturalists’ Journal (16:
92-93 (1968)).

(ii) Carex flava L. This was discovered by F. H. Perring when collecting with D. A.
Webb and R. Goodwillie on 15th July 1968 in a calcareous marsh by the River
Corrib, near Menlough in North-East Galway.

The area is on limestone over which the river periodically floods, giving rise to
a rich fen vegetation in which the plants are often standing in water. Carex flava is
locally abundant reaching a height of 65 cm (2 ft). C. lepidocarpa Tausch also
occurs and it seems likely that hybridisation is taking place between the two.
Specimens which are intermediate in fruit size have been collected.

F. H. PERRING.

92 REPORTS

LOCAL FLORAS CURRENTLY AVAILABLE FOR SALE

A list of 42 Local Floras which can be bought new from various addresses was on
display as well as many examples of them from the Botanical Library at the Natural
History Museum. The list was published in Proc. bot. Soc. Br. Isl., 77: 615-617, (1969).

F. H. PERRING.

VARIATION IN THE Vicia sativa AGGREGATE

Work on this aggregate was reported by means of text, histograms, karyotype ideo-
grams and herbarium specimens.

Vicia sativa s.l. is well differentiated from its closest relatives in this country (V. lutea
and V. lathyroides) but recognition of taxa within it is extremely difficult, even though
the range of variation is very large. Previous attempts show wide divergences in the
ranges attributed to each taxon. This is hardly surprising because the variation of
most of the characters is such as to produce a unimodal curve, the peak usually
coming close to the artificial limits placed on the infraspecific taxa.

Preliminary results suggest that if one wishes to recognise infra-specific taxa the
whole aggregate can be divided into two (‘angustifolia’ and ‘sativa’) or three (‘bobartii’,
‘segetalis’ and ‘sativa’) subspecies or varieties. Further work is progressing on the
vegetative and reproductive morphology, the karyotype, and on breeding experiments.

E. PICKERING & C. A. STACE.

THE UPPER TEESDALE FLORA IN JUNE

Twenty colour slides, taken with a Praktica IV and a Meritar 2-9 lens, were selected
to show representative plants of various habitats.

Flowering plants of damp or marshy lowland pastures included Trollius europaeus,
Orchis mascula, Dactylorhiza incarnata, and Primula farinosa on drumlins.

Viola lutea and Coeloglossum viride were from drier, grazed pastures while Polygonum
viviparum, Pedicularis palustris and Mimulus guttatus were photographed by the sides
of becks.

Representative of the riverside flora were Bartsia alpina, Potentilla fruticosa, on
shingle, Antennaria dioica, on whin-sill pavement, and Geranium sylvaticum in
hayfields.

The upland vegetation was represented by Thalictrum alpinum, on gravel flushes, and
Dryas octopetala on sugar limestone. Miss H. M. PROCTOR.

““IS YOUR PLATE CAMERA REALLY NECESSARY ?”’

A number of whole-plate (64 x 84 in) and 15 x 12 in enlargements were exhibited, to
demonstrate that 35 mm negatives will, as a matter of routine, produce photographs
of plants and vegetation of acceptable quality for most normal purposes, including
publication. Up to whole-plate size, 35 mm negatives on slow and medium-speed
films are capable of giving prints virtually indistinguishable from those yielded by
negatives of any larger size. The large camera will always have an advantage in produc-
ing large exhibition prints of the highest quality, but the advantage is completely lost in
half-tone reproductions. On the other hand, 35 mm equipment has great advantages in
portability and ease of use, important in getting the photograph when conditions are
right, and in minimising damage to the habitat. M. C. F. PROCTOR.

SLIDES OF ALPINE PLANTS AT BOURG ST. PIERRE ALPINE FLORA
M. C. F. PROCTOR.

REPORTS 93

Taraxacum SECTION ERYTHROSPERMA IN THE BRITISH ISLES

Herbarium specimens, maps and comments were exhibited for each of the 26 species
in this section that are known to occur in the British Isles. These are as follows:-

T. acutum Richards in sched. T.argutum Dahlst. *T. austriacum v.S. (introduced)
*T. brachyglossum Dahlst. T. canulum Hagl. T. commixtum Hagl.

T. degelii Hagl. T. disseminatum Hagl. *T. dunense v.S.

T. fulviforme Dahlst. T. fulvuum Raunk. T. glauciniforme Dahlst.
*T. gotlandicum Dahlst. *T. hispanicum Lind. f. T. lacistophyllum Dahlst.

T. laetiforme Dahlst. T. laetum Dahlst. T. oxoniense Dahlst.

T. placidum Richards in sched. *T. proximiforme v.S. T. proximum Dahist.

T. pseudolacistophyllumv.S. *T. retziiv.S. T. rubicundum Dahlst.

T. silesiacum Dahlst. T. simile Raunk.

The asterisked species are new to the British Isles. T. acutum and T. placidum are new
species; T. acutum and T. argutum are thought to be endemic. The commonest species
are T. oxoniense, T. brachyglossum, T. rubicundum and T. lacistophyllum, probably in
this order. Several of the main phytogeographical elements in the British flora,
including the Lusitanian, are represented in this section. A. J. RICHARDS.

RECORDING IN PERTHSHIRE

The exhibit demonstrated the 5 km square system, currently in use by the Botanical
Section of the Perthshire Society of Natural Science under the author’s leadership, for
recording plants and bryophytes in the county. Species record sheets which include a
map (c. 280 squares) are filled in with habitat and distribution data derived from field
cards used on all excursions. Seven species were chosen to illustrate this, as well as for
their own intrinsic interest. They are all alien to Britain and have different spreading
potential but all occur well established in competition with native species in natural or
semi-natural habitats. Herbarium specimens were displayed in conjunction with the
record sheets of Geum macrophyllum, Tolmiea menziesii, Salvia glutinosa, Petasites
albus, P. japonicus, Aster longifolius and Juncus tenuis. A. W. ROBSON.

COLOURED DRAWINGS OF PLANTS OF THE WESTERN DESERT AND TUNISIA
Mrs B. H. S. RUSSELL.

TWO SPECIES NEW TO CAMBRIDGESHIRE

Cystopteris fragilis (L.) Bernh.

This fern is common in upland limestone areas in Britain but very rare over much of
lowland England where it is largely confined to artificial habitats. Like several other
British ferns, it shows a partiality for railway platforms, which often supply a very
special moist and calcareous microhabitat.

The specimen shown was from a small but vigorous population under the overhang
of a concrete platform on the abandoned Old North Road Station on the former
Cambridge—Bedford railway. It is curious that this platform was the sole Cambridge-
shire locality for Adiantum capillis-veneris, which grew there between 1920 and 1953.

Ceratophyllum submersum L.

This rare aquatic was found completely filling Horseshoe Pond in Madingley Park.
Two variants were found: those with spinulose projections on the fruit (var.
haynaldianum (Borbas) G. Beck) and those without (var. submersum). There was no
definite evidence that they occurred on the same individual.

P. D. SELL & S. M. WALTERS.

94 REPORTS

SOME ALIEN EUPHORBIAS IN THE BRITISH ISLES

The following species and hybrids were exhibited:

E. esula L. subsp. esula

E. esula L. subsp. tommasiniana (Bertol.) Nyman (E. virgata Waldst. & Kit., E. uralensis
Fischer ex Link) x subsp. esula(=E. x pseudovirgata(Schur)So6). This hybrid
has often been misidentified in the British Isles as EF. uralensis.

E. corallioides L. The ‘Slinfold Spurge’.

E. ceratocarpa Tenore

E. oblongata Griseb. A rare casual. A. R. SMITH.

A PROPOSED LIST OF BRITISH HYBRIDS

Sample pages were exhibited of a proposed work to list and provide outline information
for all British interspecific hybrids. The proposal stems from a symposium held in
September 1968 on ‘The Teaching of Experimental Taxonomy in Universities’. Its
aim would be to provide a single source of information on hybrids, and to focus
attention on gaps in our knowledge. The exhibit was illustrated byspecimens of Primula
veris x vulgaris and Juncus balticus x cf. inflexus. C. A. STACE.

THE DISTRIBUTION OF Juniperus communis IN SOUTHERN ENGLAND

The exhibit showed the results of a survey in progress on the distribution of juniper
in southern England. The survey included details about the topography of each site and
the juniper colony, particularly the age structure and numbers of bushes. A map gave
the present-day distribution of juniper colonies on a 1 km square basis with the
approximate numbers of bushes where known. This showed the restriction of juniper
to calcareous soils, and the low numbers in certain areas such as the Cotswolds and the
North Downs. Another map gave sites recorded in the literature since 1750, and a
third showed sites with regeneration. Although incomplete in parts, these three maps
illustrated the general decrease in sites, their concentration in particular areas, and the
poor future outlook for juniper. The exhibit indicated the need for active conservation
measures. LENA K. WARD.

SLIDES OF ORCHIDS FROM RAVENSTHORPE, WESTERN AUSTRALIA

All the orchids are terrestrial, the two principal genera represented in the district being
Pterostylis (Greenhoods) and Caladenia.

Greenhoods are predominantly green and grow in shade. They vary in height from
3 in to 18 in and have a gnome-like appearance due to the large ‘head’, long ‘ears’ and
upright growth. Pollination is by gnats.

Caladenia is a varied genus which includes The White Spiders (with beautiful fragile
flowers with long ‘antennae’), The Dragons (variously coloured and with ‘antennae’ of
varying length), and The Fairies (lacking ‘antennae’, with slender spikes or large
single flowers). The Pink Fairy is a lovely orchid of wet ground; in contrast The Blue
Fairy flowers best after fire has destroyed the Bush. The only yellow-flowered species is
the Cowslip. Mrs. L. E. WHITEHEAD.

REPORTS 95

PHOTOMICROGRAPHS OF A SECTION OF Orchis simia ROOT

A series of photomicrographs of a section of root of Orchis simia were exhibited, the
magnification being from 350x to 2200x. The large number of crystals (calcium
carbonate?) in the cells made the material unsuitable for electron microscopy. The
septate hyphae of the mycorhizal fungus were clearly seen within each cell; several
hyphae were branched. With the assistance of Mr P. E. Smith, attempts were made to
culture the fungus but without success so identification proved impossible.

H. M. WILKS.

From 4.30 p.m. the following members showed colour slides in the Lecture Hall.
Flora of the Camargue .. si Ws af Miss L. KENDRICK
Conservation—a Postscript and a Preiude : as sau J. E. LOUSLEY
B.S.B.I. Junior Field ee 1968 Bi ae oe C. T. PRIME
Alderney plants... ae: a ra an ve M. C. F. PROCTOR

Some Alpine flowers ah ne £2 ae Ee .. Mrs B. H. S. RUSSELL

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LYNCH’S HOTEL

A.A. R.LA.C,

Lisdoonvarna, Co. Clare, Ireland
Telephone: Lisdoonvarna 10

This Hotel is personally recommended by Elizabeth Nicholas as
an ideal base for exploring the Burren Country (Botanists Paradise).

Exclusive informative map of Botanical, Geological, Historical
and Archaeological interest. Groups welcomed.

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Large Private Car Park and Garage Free.

Special Terms: April, May, June
Brochure Mrs. M. Lynch

KEANE’S HOTEL, THE BURREN

Lisdoonvarna, Co. Clare, Ireland
Telephone: Lisdoonvarna 11

Two hundred and fifty square miles of limestone—a natural rock-
garden where you will find northern-and southern-type flora growing
side by side, such as: Dryas octopetala, Gentiana verna, Erinus
alpinus, Epipactis atrorubens, Potentilla fruticosa, etc.

This Hotel (4th generation) caters especially for naturalists and their
comforts. Central Heating. Fully Licensed. Fruit and Vegetables
from our own garden. Jersey Herd. Packed Lunches a Speciality.
Good Wine List. Elizabeth Nicholas recommended. Cordon Bleu.

We can supply you with:

Lectures. Slide Shows. Annotated Ordnance Survey Maps. Lists of
Wild Plants in season. Details of Birds and Butterflies. Use of
Reference Books. Competitions. Private Car park.
Reductions for April, May.
Brochure:

Mrs. Mary Angela Keane, Keane’s Hotel, Lisdoonvarna, Co. Clare

Irish Naturalists’ Journal

A MAGAZINE OF NATURAL HISTORY
Published every quarter by the I.N.J. Committee

Edited by Miss M. P. H. Kertland, M.Sc., with the assistance
of Sectional Editors

Annual Subscription 10/- post free. Single Parts, 3/6

All communications to be addressed to
THE EDITOR, DEPARTMENT OF BOTANY, QUEEN’S UNIVERSITY, BELFAST

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Level

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Watsonia
January 1970 Volume eight Part one

Contents
THE JOURNALS OF THE BOTANICAL SOCIETY OF THE BRITISH ISLES

Cranpr, JA, Fa AC ee See
teris assimilis S. Walker in Britain AS = Se as

Jones, B. M. G. & NEWTON, L. E. The Status of Puccinellia ee a
distans (Crép.) Jansen & Wachter in Great Britain aS oe

BALL, P. W. & BRown, K. G. A Biosystematic and Ecological Study
of Salicornia in the Dee Estuary .. ae = S oe

YEO, P. F. Euphrasia brevipila and E. borealis inthe BritishIsles .. |

SHORT NOTES ... aE ae 2 =e oe “a eh
Wess, D. A. The Irish Vice-Counties oS me Sica
PLANT REcoRDS. Compiled by E. C. WALLACE 8 ie
Book REVIEWS .. SS = ae we a = aor 4 SE
OBITUARIES fy oo ee
REPORTS
Report of the Council for 1968 a es ee Peeceree
Teesdale Defence Committee Report ee se tte
Annual General Meeting, May 3rd, 1969 .. ae ue ae -
North-West Region Meeting, 1969 .. me i ae oo =
Committee for the Study of the Scottish Flora .. ee

Exhibition Meeting, 1968 4,

Published and sold by the Botanical Society of the British Isles,
c/o Department of Botany, British Museum (Natural History),
London, S.W.7.

Watsonia, 8 (1). Price Two pounds

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is A member of the Tinlins Printing: Group

WATSONIA

Botanical Society of the British Isles

Patron: Her Majesty Queen Elizabeth the Queen Mother

Applications for membership should be addressed to the Hon. General Secretary:
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Officers for 1970-71

Elected at the Annual General Meeting, 9th May 1970
President, E. Milne-Redhead

Vice-Presidents, Prof. J. G. Hawkes, Prof. D. H. Valentine, J. E. Lousley, Dr F. H.
Perring

Honorary General Secretary, Dr I. K. Ferguson
Honorary Treasurer, J. C. Gardiner

Honorary Editors, E. F. Greenwood, Dr G. Halliday, Dr M. C. F. Proctor,
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Honorary Membership Secretary, Mrs J. G. Dony

All enquiries concerning sales of, or advertising space in, the Society’s publications
should be addressed to D. H. Kent, 75 Adelaide Road, West Ealing, London, W.13.

Watsonia, 8, 97-120 (1970). 97

Notes on the Hawkweeds (Hieracium sensu lato) of
western Scotland

A. G. KENNETH and A. McG. STIRLING

Stronachullin, Ardrishaig, Argyll; 17 Austen Road, Jordanhill, Glasgow W3

ABSTRACT

The distribution of species of Hieracium sensu stricto and the subspecies of Pilosella officinarum
C. H. & F. W. Schultz in west and north-west Scotland is indicated for those areas and taxa of
which the authors have personal knowledge. Corrections are included for the Distribution Maps
of certain species as shown in the Critical Supplement. Ecological information is given for the
species of Hieracium and their distribution is discussed in relation to the areas visited within
each vice-county. A key to the genus Pilosella and an Index of Place Names are given as
Appendices.

INTRODUCTION

Our knowledge of the distribution of the genus Hieracium in Scotland owes
much to the explorations of certain English botanists in the period between 1850
and 1918. Notable among these were J. Backhouse Jnr, E. F. and W. R. Linton,
Henry Groves, F. J. Hanbury, E. S. Marshall, W. A. Shoolbred and G. C.
Druce, all of whom collected in the central and north western Highlands.

After the first world war little attention was paid to this genus by British
field botanists, due mainly to a lack of taxonomists competent to undertake
determination of specimens. It was not until the commencement of the B.S.B.I.
Distribution Maps Scheme in 1954 that they were encouraged by the intensive
taxonomic study of the genus being undertaken by P. D. Sell and Dr C. West
to resume the study of hawkweeds in the field on anything approaching the
scale of fifty years earlier. Progress has been so great in recent years, both in
taxonomic and field work, that sufficient information became available to
make possible the publication in the Critical Supplement of distribution maps
of all the hawkweed species then known to occur in the British Isles (Perring
& Sell 1968).

During the years 1959-1968 the authors have made extensive field investiga-
tions into the hawkweed flora of western and northern Scotland, resulting in a
very large number of determinations by Sell and West. We have thought it
desirable to place the results of these researches on record in a form which
supplements the information given on the distribution maps in a number of ways.
We must emphasise that this account is in no sense a Hieracium Flora of western
Scotland and we have not included records contributed by the earlier botanists
referred to above. We have treated our records on a vice-county and district
basis and appropriate topographical, geological and habitat data have been
included. We have not included any records from the following vice-counties
within our area: v.c. 97 Westerness (with the exception of that part which is in
Argyll), v.c. 100 Clyde Isles and v.c. 110 Outer Hebrides. Many of the records
are additional to those shown on the relevant maps in the Critical Supplement,

i>
0CT9 1970
LIBRARIES

98 A. G. KENNETH AND A. McG. STIRLING

and the opportunity has also been taken to indicate certain alterations necessary
to the published maps. It will be apparent that while certain areas are well
recorded there are many other districts which are relatively underworked.
Knowledge of the Scottish Hieracia is by no means complete, and an appreciable
_ number of specimens remain undetermined, some of which may be undescribed
species.

In the area under consideration there is no lack of the sort of rocky habitats
favoured by most species of Hieracium. In such situations, particularly where
basic conditions occur, a good varied hawkweed flora may be expected. Steep,
rocky stream-sides and gullies usually afford particularly favourable conditions.
It is seldom possible to be specific on the ecological requirements of individual
species; where we have found it possible to draw conclusions on the basis of
personal observation we have included these. In our experience at least some
species do appear to be capable of colonising and persisting in a diversity of
habitats. Good examples of this are H. vulgatum, H. cravoniense and H.
maritimum (which occurs in a very wide range of habitats, including river shingle,
a wall top, heathery grassland, montane rock ledges and a sandy loch shore).

The principal basiphilous groups are: certain of the Section Oreadea, and
most species of the Sections Subalpina and Cerinthoidea. It would appear that
species of the Section Alpina, though generally regarded as tending to be
acidiphile, are not exclusively so, and that some at least may prefer neutral
or even slightly basic conditions. They are, however, rarely to be found in the
mountain gullies so much favoured by species of the Section Subalpina.

Altitude is a very important factor influencing the distribution of hawkweeds,
but anomalies are occasionally encountered. For example, on Meall a’ Bhuiridh
in Glen Coe species of the Section Alpina have been noted below the highest
recorded station for H. subumbellatiforme at 2,000 ft, while species of the same
section occur near Loch Sloy, Dunbartonshire at around 1,800 ft.

The nomenclature used in these notes is that of Sell and West (Perring & Sell
1968), which differs significantly from that of Dandy (1958), and in accordance
with current views the subgenus Pilosella is here recognised as of full generic
rank.

REGIONAL ACCOUNTS

VICE-COUNTY 72, DUMERIES

With the exception of the area of the Moffat hills, where in the past a number of
interesting and very local species have been recorded, comparatively little is
known of the hawkweed flora. Investigations of some areas in the west of the
county during the summer of 1968 suggest that further search of suitable
localities may yield interesting results.

The localities examined in 1968 included rocky outcrops and gullies by the
Crawick Water and in the Dalveen Pass, both in the Lowther Hills area, and
Glenwhargen Craig, a rocky hill in the valley of the Scar Water, a western
tributary of the Nith. H. caledonicum was found to be not uncommon in the
Lowther Hills localities, and a fair amount of H. duriceps was also noted. The
most interesting plant from the Dalveen Pass is H. pseudosarcophyllum, a
hawkweed previously known only from the vicinity of Moffat in the same
county. H. caesiomurorum was also noted. Glenwhargen Craig proved to have
H. leyi in considerable quantity, hundreds of plants of this fairly distinct

}

HAWKWEEDS OF WESTERN SCOTLAND 99

species being scattered over a large area of otherwise floristically rather poor
rock. H. Jeyi is a glaucous-leaved species of the Section Oreadea which had not

previously been recorded for Dumfriesshire. This was accompanied in a few

places by H. rubiginosum, a hawkweed of rather common occurrence in the
south-west of Scotland.

A rather puzzling plant with close affinities to H. subhirtum was found in
two places in the Mennock and Scar valleys.

VICE-COUNTY 73, KIRKCUDBRIGHT

One of us (A. McG. S.) was able, during the years 1959 to 1962, to examine
the hawkweeds of this county with some interesting results, the more rewarding
records being obtained in the lowland and coastal areas of the county. A large
proportion of the north-western hill country is composed of acid granite rocks
and is not conducive to the development of a good hawkweed flora, but the
Silurian rocks, which predominate in the north-east, are rather more productive
as basic conditions are not infrequent. Our limited examination of these hill
areas has yielded only three species: H. sparsifolium was found near High Bridge
of Ken, Dalry, and also near Lauriston, in the former locality being accompanied
by H. euprepes, while H. rubiginosum was found by the Poldores Burn in an
aberrant form, accompanied by a plant which awaits identification. Since
H. holosericeum has been reported from one of the higher hills of the Kells
range, it seems likely that if the area was more thoroughly searched a good
selection of species, perhaps including some of the Section Subalpina, would
prove to be present.

One of our earliest finds in the county is of considerable interest. A plant
obtained on rocks by the River Dee near Tongland Bridge, Kirkcudbright,
in 1959 was found to be H. vagense, a very distinct species previously known
only from limestone areas in Wales. In the same year a hawkweed found on
sea-cliffs at Douglas Hall, Colvend, was identified as H. subplanifolium, this
being the first Scottish record for the species. The Kirkcudbrightshire coast
from Colvend in the east to the county boundary in the west is for the most
part rocky, with massive cliffs occurring in many places. The rock is mainly
of Silurian age, its rather basic nature and sunny southern exposure producing
a varied and interesting flora including a number of hawkweed species. In
addition to H. subplanifolium already mentioned, H. subrude and H. orimeles
have been found on shore rocks near Kirkcudbright, while H. schmidtii and
H. cravoniense have been identified from shore rocks near Gatehouse-of-Fleet.

In the landward area of the county, H. diaphanoides occurs on the bank of
the River Urr near Corsock, and H. diaphanum on the railway bank near
Gatehouse-of-Fleet station. The latter species also occurs by the Grey Mare’s
Tail, Newton Stewart. Of the aphyllopodous species, H. umbellatum has been
recorded from two sites, one a railway bank near Kirkcudbright, the other an
area of waste ground near Dalbeattie. H. perpropinquum appears to be not
uncommon, having been detected in at least three widely separated localities.

VICE-COUNTY 74, WIGTOWNSHIRE

Practically nothing is known of the hawkweed flora of the county, but our
impression is that it is likely to be poor in comparison with that of its neighbours,
mainly due to a lack of exposed rocky habitats in the landward area. However,

100 A. G. KENNETH AND A. McG. STIRLING

the coastal cliffs, if examined at the right season, might prove to harbour a
few species, as they are similar in character to those of the Kirkcudbright coast.

VICE-COUNTY 75, AYRSHIRE

Like Dumfriesshire, Ayrshire, though possessing much hill country with rocky
habitats suitable for hawkweeds, has been little explored for these plants. One
of us (A. McG. S.) has examined rocky sites in the neighbourhood of Dalmel-
lington and Straiton with moderate results, and a few species have been noted
on coastal rocks in the south-west of the county. The Dalmellington district
has yielded H. ampliatum, H. duriceps and H. rubiginosum, the latter species also
occurring on Craig Hill, Straiton, where it appeared to be the only species
present, though H. duriceps is on riverside rocks nearby. H. lissolepium, a
species of the Section Tridentata rather rare in Scotland, was recorded at
Stinchar Bridge. Hill country with apparently suitable ground near Rowantree
Hill, Barr proved to be disappointing, the only species present being H. duriceps.

In the south-west of the county H. schmidtii grows on rocky banks at
Kennedy’s Pass near Girvan, and H. caledonicum occurs in an old quarry close
to the county boundary near Cairnryan. In the north near Largs H. subrude
grows on basalt rocks. The aphyllopodous species H. umbellatum and H.
perpropinquum have been noted in some quantity on sandy waste ground near
Kilwinning.

VICE-COUNTY 76, RENFREW

This county has not been the subject of particularly intensive investigation so
far as the hawkweed flora is concerned. Although a good deal of high ground
with rocky outcrops occurs within its boundaries, the rocks, which are mainly
basalt lavas, seem not to be so productive of Hieracium species as the same
formation just across the Clyde in Dunbartonshire; however, where limited
exposures of more basic rock occur, as in Shielhill Glen, Inverkip (Calciferous
Sandstone Series), good results may be expected. In the locality just mentioned
a good colony of H. dipteroides was found in 1967. Another find of some
interest was made in 1964 by Mr D. McClintock, who noted H. chloranthum
growing on a wall near Lochwinnoch—a most unusual habitat for a species
which normally favours rocky sites, and which grows in such habitats by the
River Calder nearby. H. /atobrigorum has been noted at Lochwinnoch, and also
near Neilston.

VICE-COUNTY 86, STIRLING

Suitable hawkweed ground is plentiful in the county, which in this respect is
very similar to Dunbartonshire. In the south, extensive exposures of basalt
crags occur on the Campsie and Fintry hills, while to the east of Loch Lomond
the higher hills, composed mainly of schists, calcareous in places, provide
good rocky habitats, both on the exposed crags at the higher levels and on the
sides of the numerous streams. However, our knowledge of the hawkweeds of
the highland area of Stirlingshire is very limited, and Ben Lomond, the county’s
highest summit, and other neighbouring hills should produce excellent results
with further investigation.

It is on the basalts of the Campsie and Fintry hills that the greatest display of
Hieracium species is to be found, and certain particularly favourable areas
have been rather carefully examined. The best localities have a warm, southern

a

HAWKWEEDS OF WESTERN SCOTLAND 101

exposure. H. caledonicum is common on the craggy basalt overlooking Lennox-
town, and other species occurring are H. rubiginosum, H. euprepes, H. duriceps,
H. subhirtum and the very rare H. britanniciforme, for which the only other
known Scottish station is in Kintyre. In the nearby Campsie Glen the Section
Tridentata is represented by a colony of H. stewartii, and H. lissolepium, which
belongs to the same section, has been recorded from the Gonachan Burn,
where H. rubiginosum and H. pictorum also occur. In both localities the rare
H. rhomboides has been noted. The best ground in the Fintry hills has proved
to be a range of basalt cliff known as the Double Craigs. Here the interesting
feature of the hawkweed flora is the occurrence, apparently to the virtual
exclusion of other species, of three species belonging to the Section Oreadea,
all of which are distinctly local in Scotland. These are H. Jasiophyllum, H.
jovimontis and H. dicella. The last-named species also occurs in Ballagan Glen
in the Campsies, where H. petrocharis, a member of the Section Subalpina,
also grows at the surprisingly low altitude of 700 ft. H. duriceps appears to be
generally distributed over the area just described.

VICE-COUNTY 97, (ARGYLL)

Our knowledge of the hawkweeds of this portion of the vice-county of Wester-
ness is not extensive. Of the few hills we have visited, perhaps the most rewarding
is Garbh Bheinn in Ardgour where the following varied assortment of species
has been recorded: H. holosericeum, H. alpinum, H. senescens, H. marshallii,
H. pseudanglicum, H. shoolbredii, H. sommerfeltii, H. chloranthum, H. argenteum,
H. yulgatum and H. sparsifolium. Of these, perhaps the most interesting are
H. alpinum, which is rather rare in the western Highlands, and H. sommerfeltii,
a local plant usually growing at high altitudes. On the low ground in Ardgour,
H. latobrigorum and H. subcrocatum of the Section Foliosa and H. perpropin-
quum of the Section Sabauda occur at Gearradh.

In Kingairloch we have looked over Creach Bheinn and Maol Odhar. These
two hills are contiguous and they produce H. marshallii, H. ampliatum, H.
shoolbredii, H. sommerfeltii, H. chloranthum, H. pictorum and H. vulgatum.
Meall nan Each has H. anglicum and H. shoolbredii.

The district of Morvern offers a good deal of interesting ground, much of the
country rock being basalt of Tertiary age supporting a good flora. On Beinn na
Beathrach H. chloranthum, H. senescens, H. ampliatum, H. pictorum and H.
strictiforme have been noted—a disappointing list, while Sithean na Raplaich
and the vicinity of Loch Dur na Marst have H. anglicum and H. subhirtum.
Sithean na Raplaich would probably produce better results with more careful
searching as it is a particularly promising area. Beinn na h-Uamha near the
head of Loch Teacuis has produced H. pseudanglicum, H. anglicum, and H.
duriceps, while on the low ground H. shoolbredii has been found near Killundine.

The district as a whole may be compared topographically with the neighbour-
ing Isle of Mull, but though the number of species of Hieracium so far found
there is similar to that of its neighbour, in their composition there is a marked
difference. For instance, no species of the Section Alpina has so far been detected
on Mull, and only one named species of the Section Subalpina has been noted
there, yet in Ben More the Mull hills attain as great an altitude as those of the
adjacent mainland. The eighteen species recorded probably represent only a
small proportion of the total.

102 A. G. KENNETH AND A. McG. STIRLING

VICE-COUNTY 98, ARGYLL (MAIN)

This large vice-county contains a great diversity of habitats suited to Hieracia,
and probably supports a larger number of species than does any other. Although
we have investigated several areas in considerable detail, these only represent
a small proportion of the potentially good hawkweed ground, and this particu-
larly applies to the higher hills. Nevertheless we have assembled a considerable
mass of data concerning the genus in Argyll, and we propose to deal with this at
some length.

It is convenient to consider the hawkweeds of Argyll under two main
headings: first, the species of the low hills and coastal rocks between Crinan
Canal and Connel Ferry, including the islands of Seil, Luing and Kerrera,
and bounded on the east by Loch Awe, and secondly the various mountain
areas which we have examined.

The lower hills and islands

Commencing in the south of this area, the first locality to be considered from
which hawkweeds have been determined in quantity is Creag nam Fitheach
near the head of Loch Craignish. This hill is almost entirely composed of
limestone and, together with the Bealach Mor, which should be considered as
essentially the same locality, it produces a most unusual assortment of Hieracia.
The impressive list from this hill consists of: H. petrocharis, H. dasythrix, H.
ampliatum, H. shoolbredii, H. sarcophylloides, H. dicella, H. jovimontis, H.
caledonicum, H. angustisquamum, H. chloranthum, H. duriceps, H. subhirtum,
H. vennicontium, H. euprepes, H. rubiginosum, H. vulgatum, H. cravoniense and
H. ebudicum. At the Bealach Mor the somewhat undistinguished-looking
rarities H. rhomboides and H. triviale occur. H. ebudicum is the prevailing
hawkweed of the limestone cliffs; previous to its discovery here and in v.c. 101
it had been known only from the Outer Hebrides. It appears to be capable of
attaining a greater size than that indicated in the description by Pugsley (1948).
Extensive exposures of basic andesite rocks occur in the area to the north of
Kilmelford, and these produce an interesting hawkweed flora. The sites particul-
arly examined are the Pass of Melfort, Creag an Sturra and Cruach an Nidd. In
the first of these localities the species noted are H. dasythrix, H. petrocharis,
H. dicella, H. chloranthum, H. sanguineum, H. pseudostenstroemii, H. rubiginosum,
H. subhirtum, H. pictorum, H. vulgatum, H. cravoniense and H. rhomboides.
H. uisticola and H. sanguineum occur on rock faces to the west of Kilmelford,
while Creag an Sturra in the same area has H. dicella and H. petrocharis.
Cruach an Nidd, in addition to some of the commoner plants mentioned above,
has H. uiginskyense, H. caledonicum, and H. strictiforme. These localities for
H. sanguineum represent its only Scottish station, the species being previously
known only from limestone areas in north-west England, south Wales and the
west of Ireland. H. jovimontis, which appears to be a rather scarce species in
Argyll, has been noted on cliffs by Loch Tralaig, to the east of Kilmelford.
The andesite offers similar rewarding hawkweed ground on the islands of
Seil and Luing and on the adjacent mainland, the sites in this case being close
to the sea. On Seil, the coastal rocks between Cuan Ferry and Dun Nucaig
produce H. hebridense, H. ampliatum, H. shoolbredii, H. dasythrix, H. argenteum,
H. sarcophylloides, H. caledonicum, H. vulgatum and H. caesiomurorum.
The area about Dun Mor, Ellenabeich, has H. hebridense, H. anglicum,

HAWKWEEDS OF WESTERN SCOTLAND 103

H. pseudanglicum, H. ampliatum, H. shoolbredii, H. dicella and H. caledonicum,
also the form of H. vulgatum with floccose leaves which is treated as a species
by some continental botanists under the name of H. coniops. Near Balvicar, on
Seil, H. subcrocatum has been found. This species has also been noted on the
mainland north of Seil Sound. On Luing only H. argenteum and H. vulgatum
have been noted, the latter occurring in the form referable to H. coniops as
well as in its typical form.

To the north of the area just mentioned there are extensive stretches of
rather basic rock about Gallanach, south of Oban. These produce H. anglicum,
H. ampliatum, H. shoolbredii, H. fratrum, H. sarcophylloides, H. argenteum,
H. caledonicum, H. uisticola, H. vulgatum and H. strictiforme. Of these, H.
fratrum is the most interesting species, its only other stations being in the Moffat
hills in Dumfriesshire and in the north-east Highlands. Nearer Oban, on the
mainland, there is some good ground about the Kerrera Ferry where the species
so far noted are: H. dasythrix, H. ampliatum, H. shoolbredii, H. langwellense,
H. hebridense, H. argenteum, H. chloranthum, H. sarcophylloides, H. dicella,
H. caledonicum, H. scoticum and H. euprepes. Although not extensive, these
cliffs have an interesting flora. The island of Kerrera lies opposite, and while
not particularly good for hawkweeds it has produced H. anglicum, H. lang-
wellense, H. caledonicum, H. chloranthum, H. subhirtum, H. vulgatum, H.
latobrigorum and H. strictiforme. Some of the plants referred to H. vulgatum
from Kerrera are decidedly odd. They may be similar to the floccose-leaved
H. coniops, but this taxon seems to be of doubtful status owing to the existence
of intermediate forms. In our experience it appears, unlike H. vulgatum sensu
stricto, to be limited to natural habitats.

To the north of Oban there is an area of coastal cliff at Ganavan which has
produced H. ampliatum, H. hebridense, H. rubiginosum and H. vulgatum. Of the
other coastal areas in the vice-county to the north of Loch Etive, these are
virtually terra incognita, but mention should be made of the limestone island
of Lismore at the mouth of Loch Linnhe, which has a considerable extent of
exposed rock and from which H. cymbifolium and H. hebridense have been
recorded. The former also grows on the same limestone formation on the
adjacent island of Bernera, where H. petrocharis is another species of some
interest. H. cymbifolium is otherwise known only from the Carboniferous
limestone of central and northern England and from Raasay. Another hawk-
weed from Bernera has yet to be determined.

The frequency of H. dicella over the whole of this low ground area is remark-
able; there are two further localities for it in the vicinity of Ford, at the south
end of Loch Awe. In this area, near Kirnan, H. cheriense formerly grew on a
bridge where it was probably an introduction.

The following species of the Sections Tridentata and Foliosa have been
noted in localities other than those already dealt with: H. uiginskyense (Glen
Gallain and Lag na Cille), H. strictiforme (Crinan Ferry, north side), H.
latobrigorum (frequent about the River Add and near Kilmartin), H. subcrocatum
(canal bank near Lochgilphead; Creag an Tairbh, Ford), H. drummondii
(Ford—Kilmartin road junction), H. maritimum (River Euchar, near Raera).

The higher hills

Many of the Argyllshire hill areas have been examined and a number have
proved particularly rewarding, especially for species of the Sections Alpina and

104 A. G. KENNETH AND A. McG. STIRLING

Subalpina. Ben Cruachan at the head of Loch Awe has produced H. lingulatum,
H. senescens, H. gracilifolium, H. sinuans, H. callistophyllum, H. dasythrix,
H. pseudanglicum, H. pseudanglicoides, H. nigrisquamum, H. centripetale, H.
anfractiforme, H. lintonianum, H. subhirtum, H. euprepes, H. rubiginosum,
H. dewarii, H. sparsifolium and H. strictiforme. The area searched is in the
vicinity of the Hydro-Electric Board’s new loch, and most of the plants recorded
grow in an extensive ravine running east from this. The total absence of species
of the Section Alpina in the area examined was rather surprising, but the
occurrence of no less than eleven of the Section Subalpina must make this
one of the best areas for this group. As usual they are to be found about stream
sides and on rock ledges of the better rock. The presence of H. lintonianum
here is noteworthy as its previously known localities are in the east central
Highlands.

Beinn Buidhe, north-east of Inverary has much good rock, and we found the
Brannie Burn particularly rewarding. Again no species of the Section Alpina
was noted, the species recorded being: H. lingulatum, H. senescens, H.
pseudanglicum, H. langwellense, H. uistense, H. chloranthum, H. subhirtum,
H. dipteroides, H. oxyodus and H. dewarii. Here H. dipteroides and H. oxyodus
are the most unusual species. The hills between Arrochar and Cairndow have
been searched, in some cases systematically, and we are treating them here as
a single group, though indicating on which hills the more interesting species
occur. The rock is schist, calcareous in places, which accounts for the com-
parative wealth of species, particularly on Beinn an t’Seilich and Stob an Eas,
where there are some very attractive basic gullies. The following species have
been detected on these hills: H. Jingulatum, H. senescens, H. petrocharis, H.
dasythrix, H. anglicum, H. ampliatum, H. shoolbredii, H. chloranthum, H.
pictorum, H. subhirtum, H. piligerum, H. caesiomurorum, H. vulgatum, H.
sparsifolium, H. dewarii and H. strictiforme. Of more particular interest are
H. holosericeum (Beinn Luibhean and Binnean an Fhidhleir), H. eximium
(Beinn Narnain), H. cuspidens, H. gracilifolium, H. vennicontium and H.
dipteroides (Stob an Eas), H. anfractiforme (Beinn an t’Seilich), H. glandulidens
(Beinn Ime), H. marshallii (Binnean an Fhidhleir), H. aggregatum (Ben
Chorranach), and H. nigrisquamum (Ben Donich).

The Glen Coe district has long been noted for a prolific hawkweed flora.
Many of the records of the earlier specialists are localised by the use of a place-
name which might apply to several hills over a wide area. In particular it has
been interesting to compare the species found during an investigation of Meall
a Bhuiridh and Sroin na Creise with the records cited in Pugsley’s Prodromus
(1948) for ‘Kingshouse’. Our list for this hill area is: Section Alpina—dH.
eximium, H. calenduliflorum, H. memorabile, H. tenuifrons and a distinct-
looking undescribed species which has also been collected by Dr West near
Glenfinnan (v.c. 97); Section Subalpina—H. lingulatum, H. dasythrix, H.
senescens, H. anfractiforme, H. callistophyllum, H. petrocharis, H. vennicontium
and an interesting apparently undescribed species from high on Sroin na Creise.
Other species in this area include H. shoolbredii, H. nitidum, H. dewarii and
H. subumbellatiforme. Despite the attentions of the earlier collectors, species
of the Section Alpina grow in fine profusion over a wide area. H. tenuifrons
occurs in two well-marked forms differing in shape of head and indumentum
of the phyllaries.

A day on Bidean nam Bian, the highest mountain in Argyll, produced H.

HAWKWEEDS OF WESTERN SCOTLAND 105

pseudanglicum, H. senescens, H. shoolbredii, H. subhirtum and an ‘unknown’
species of Section Subalpina. The only species of Section Alpina noted was
H. eximium, and the hill is clearly much less productive of this section than
some of its neighbours, notably Meall a Bhuiridh. In Glen Etive, H. shoolbredii
has been noted near Dalness, while H. sparsifolium occurs on a roadside bank.
H. marshallii grows at a moderate altitude at the entrance to Fionn Glen,
and it has also been noted on Buchaille Etive Mor. On Aonach Dubh nearby
the rare H. aggregatum has been recorded, and on rocks by the roadside near
the Meeting of the Waters H. reticulatum grows in fair quantity. Also in the
vicinity H. pseudanglicoides occurs on Stob nan Cabar.

All the hills of the Glen Coe district mentioned so far are composed of
rocks of volcanic origin, mainly andesites and rhyolites. In contrast to this, the
limestone hill Creag Bhan, near Ballachullish, may be cited as an example of a
very basic hill with a rich flora of hawkweeds as well as other flowering plants.
The species noted here include H. senescens, H. clovense, H. pseudanglicum,
H. sommerfeltii, H. pictorum, H. euprepes, H. piligerum and H. subhirtum.
With the exception of H. clovense, which is a rare species, and perhaps H.
sommerfeltii, these are all typical of the Argyll hills. The most remarkable fact
about their occurrence on Creag Bhan is their comparative abundance, provid-
ing ample proof of the fact that most Hieracia, though not strict calcicoles,
undoubtedly flourish best in basic conditions.

The hills about Bridge of Orchy were known to the earlier botanists as
good hawkweed ground, and Raven (Raven & Walters 1956) has described
their hawkweed flora. Our own investigations in recent years have included
Beinn a’ Chaisteal and Beinn Dothaidh to the east of the Tyndrum—Glen Coe
road, Coire Chailein on Beinn Bheag northwest of Tyndrum, and Beinn Udlaidh
in Glen Orchy. The first has produced H. gracilifolium, H. dasythrix, H.
langwellense and H. anglicum, but undoubtedly further search would produce
better results. Beinn Dothaidh has proved rather disappointing, only H. eximium
and H. ampliatum being noted. Coire Chailein has the following: H. cuspidens,
H. dasythrix, H. pseudanglicum, H. pseudanglicoides and H. rubiginosum. Of
these H. cuspidens is a rare plant which has also been recorded in the Glen
Kinglas area, and H. pseudanglicoides is another uncommon species of very
limited distribution. A fine north-facing corrie on Beinn Udlaidh has a
good selection of species including H. senescens, H. dasythrix, H. lingulatum,
H. pseudanglicum, H. nigrisquamum, H. chloranthum, H. anglicum, H. shoolbredii
and H. pictorum. On Beinn a’ Cleibh, just to the west of Beinn Laoigh, H.
senescens, H. gracilifolium, H. hastiforme, H. oxyodus, H. lintonianum and H.
subhirtum have been noted. Of these H. hastiforme is particularly noteworthy
as it has not been observed for many years. No doubt this hill would provide a
large number of species if systematically searched. Meall Garbh, a hill near
the head of Loch Creran, has produced H. tenuifrons, H. vennicontium and H.
callistophyllum.

Various species, mainly belonging to the aphyllopodous sections of the
genus, occur in more or less isolated localities in the county. H. cravoniense
grows near Dalvuie, Benderloch. It is widespread in the Crinan—Connel area
and usually stylose-flowered. Further south, in v.c. 101, it has fully developed
ligules. H. grandidens occurs as an introduction on walls at Tighnabruaich.
Of the Section Foliosa, H. strictiforme occurs occasionally on roadsides, as
near St Catherines, Loch Fyne, and between Inverary and Dalmally, H.

106 A. G. KENNETH AND A. McG. STIRLING

maritimum grows on stabilised shingle at the head of Loch Etive, probably
washed down from some higher site in the surrounding hills, and H. latobrigorum
occurs as the forma angustifolium Pugsl. at the north end of Loch Awe. The
status of this latter plant is unknown and the locality has now probably been
destroyed by road alterations. The typical plant is widespread.

Two alterations are now necessary to the maps published in the Critical
Supplement. The material on which the record for H. oxyodus in square 17/80
was based has now been redetermined as a form of H. subhirtum, a species
frequent in the area. The record of H. schmidtii from the same square is also
an error, the plant so named from Creag nam Fitheach having since been
referred to H. ebudicum.

To date we have detected 70 species in the vice-county, made up as follows:
Section Alpina 6, Subalpina 18, Cerinthoidea 6, Oreadea 9, Vulgata 20, Alpestria
1, Tridentata 2, Foliosa 7, Sabauda 1.

VICE-COUNTY 99, DUNBARTON

In Dunbartonshire two geological formations provide exposed rock with a
good variety of hawkweed species. These are (1) the basaltic lavas of Carboni-
ferous age which give rise to the prominent craggy escarpments of the Kil-
patrick Hills, and which are similar in character, though not in hawkweed
species, to those of the Stirlingshire hills, and (2) the schistose rocks which
occur over most of the county north of the Highland Boundary. Between these
two formations lies a smaller tract of Old Red Sandstone rocks, but although
there are good exposures of these in places we have not so far recorded any
Hieracia from them.

The Kilpatrick Hills area has several exposures of basalt which are con-
siderably more basic and consequently richer as to flora than neighbouring
outcrops. These sites have yielded H. caledonicum, H. duriceps, H. vulgatum,
H. chloranthum, H. rubiginosum, H. caesiomurorum and H. cravoniense, but
perhaps their most interesting and attractive hawkweed is H. saxorum, a hand-
some species with glaucous, blotched leaves and quite large flowers which adorns
the basalt crags above Glenarbuck and near Cochno Loch reservoir. This
appears to be its only locality in the west of Scotland, though recorded as not
uncommon in the northeast.

The schistose rocks of the area between Loch Lomond and Loch Long
produce a varied hawkweed flora, mainly in the numerous rocky gullies formed
by the mountain streams. Such sites are frequently quite basic, and among the
commonest species occurring are H. dasythrix, H. subhirtum, H. duriceps and
H. vennicontium, while on the bank of the Auchengaich Burn, Glen Fruin,
the very local H. breadalbanense was discovered in 1967. This last species was
previously known only from a few central Highland localities.

Dunbarton has only three hills of any appreciable altitude providing suitable
habitats for the more montane species, Ben Vorlich (3,092ft), Ben Vane
(3,004ft) and Beinn Dubh (2,509ft), all situated in the northern extremity of
the county. The ubiquitous H. lingulatum is common, and we have also recorded
the following other members of the Section Subalpina on one or more of these
three hills: H. senescens, H. dasythrix, H. gracilifolium, H. callistophyllum,
H. anfractiforme, H. nigrisquamum and H. pseudanglicum. Of these H. dasythrix
is exceedingly common, H. pseudanglicum frequent, and the others much less so.
Of the Section Cerinthoidea, H. anglicum is quite common, and H. ampliatum

HAWKWEEDS OF WESTERN SCOTLAND 107

much less so, while, surprisingly, H. shoolbredii has not yet been detected in the
county, though frequent in the west of Scotland generally. Three species of the
Section Alpina have so, far been found—a very creditable list for an area so
close to the southern limit of the Highlands. H. holosericeum occurs, apparently
in small quantity, on Ben Vorlich, and H. eximium and H. alpinum have been
found in the Ben Vane—Beinn Dubh area at the unusually low altitude of
1,800 ft. The Section Oreadea is very poorly represented on the Dunbartonshire
hills, the only species so far noted being H. chloranthum. The otherwise wide-
spread H. argenteum, which might reasonably have been expected, has not so
far been noted. A number of species of the Section Vulgata occur on the schist
hills. Of these H. pictorum and H. subhirtum are probably the most frequent
species, and in addition to H. breadalbanense already mentioned, H. duriceps,
H. piligerum and H. caesiomurorum occur.

Only two of the aphyllopodous species are known to occur in the county,
and these have been recorded from the low ground only. AH. latobrigorum
grows near the mouth of the Glenmallan Burn on Loch Long-side, while H.
perpropinquum is quite common on some of the islands in Loch Lomond and
on the adjacent shores, usually in rocky places and accompanied by H. vulgatum.
Another ‘leafy-stemmed’ species, H. dewarii, is found locally in the northern
part of the county, occurring typically in rocky gullies on the lower slopes of
the hills, as by the Stuckgowan Burn, Tarbet. It is interesting to note that although
H. dewarii was originally described from specimens obtained near Dollar,
Clackmannanshire, the earliest collection of this plant was probably from a
Dunbartonshire locality (Arrochar) by Prof. J. H. Balfour and a student party
in 1842. Prof. Balfour again collected this species in the county in 1847, near
Inverarnan.

This account of the Dunbartonshire hawkweeds can now be concluded with
the mention of two species occurring locally on old walls at Rhu, near Helens-
burgh. These are H. diaphanoides and the introduced species H. grandidens.

VICE-COUNTY 101, KINTYRE

This vice-county may conveniently be divided into two distinct topographical
areas: Knapdale, lying between the Crinan Canal in the north and West Loch
Tarbert in the south, and Kintyre proper comprising all the country situated to
the south of West Loch Tarbert. Knapdale consists largely of hilly ground
rising to 1,840 ft with numerous rocky exposures, whereas in Kintyre the best
hawkweed ground occurs on the sea cliffs.

While most of the rock in Knapdale is base-poor schist, there are areas of
better rock, and it is on such areas that the majority of our collections have
been made. No species of the Section Alpina has been noted, which is not
surprising in view of the moderate elevation of the hills. On this hill ground
we have examined several distinct areas, of which the best is Cruach Lusach
and the valley of the Lussa, near Achnamara. Here the following species have
been recorded: H. glandulidens, H. dasythrix, H. shoolbredii, H. iricum (stylose-
flowered), H. argenteum, H. caledonicum, H. orimeles, H. chloranthum, H.
uistense, H. duriceps, H. pictorum, H. piligerum, H. rivale, H. ebudicum, H.
subhirtum, H. rubiginosum, H. vulgatum, H. solum, H. reticulatum and H.
maritimum. This lengthy list includes an unusually large number of species
of rare occurrence in the vice-county, and indeed, in the country as a whole.
H. solum calls for special mention. First found in this area in 1962 it proved

108 A. G. KENNETH AND A. McG. STIRLING

to be an undescribed species of the Section Alpestria, its closest allies growing
in the Shetlands. It favours dry rock ledges and apparently does not require
basic conditions.

To the south of Cruach Lusach there are some lower hills composed of
rather poor rock where several interesting Hieracia occur. This range, Meall
Ruadh, stretches from west of Loch an Dughaill to Loch an Dobhrain, and
H. subrude, H. dissimile, H. duriceps, H. uistense, H. cravoniense, H. vulgatuim,
H. chloranthum and H. solum have been recorded there. This is the original
locality for H. solum, and H. dissimile has here its only known British station.
Although belonging to the Section Subalpina, H. dissimile has a superficial
resemblance to forms of H. vulgatum, but its heads are quite different. Stiil
further south our next principal area comprises Sliabh Gaoil, the highest hill
in Knapdale, and the Artilligan Ravine which drains from it to the northeast.
In this area have been observed H. dasythrix, H. hebridense, H. langwellense,
H. chloranthum, H. rivale, H. duriceps, H. piligerum, H. rubiginosum, H.
caesiomurorum, H. subhirtum, H. uistense, H. vulgatum, H. cravoniense, H.
latobrigorum and H. strictiforme. The only known stations for H. langwellense
and H. hebridense in the vice-county are here, and an interesting undetermined
species of the Section Subalpina has also been found.

Some hawkweeds have been noted in Knapdale from localities other than
those already mentioned. Cnoc na Seamraig, an isolated outcrop of limestone
at about 700 ft, produces H. subplanifolium in some quantity. Rather surpris-
ingly, no other species of interest is present. There are outcrops of epidiorite
north of Tayvallich, and these produce H. dicella and H. petrocharis, while
shore rocks near Stronefield have yielded H. vennicontium at a remarkably
low altitude. H. ebudicum grows on hill ground near Scotnish, having only
recently been detected on the mainland of Scotland. H. cravoniense, usually
stylose-flowered, has quite normal flowers in its Knapdale localities. Of the
aphyllopodous species, H. maritimum in addition to its Cruach Lusach station,
has been recorded from the Wishing Ledge, near Carse, and from the Isle of
Danna, while the rarer H. drummondii grows in some quantity in rocky wood-
land and on walls at Crinan. H. umbellatum occurs in rocky woodland near
Carse, not far from where its close ally H. maritimum is found at the Wishing
Ledge, and H. reticulatum occurs at Ormsary.

In Kintyre there is good hawkweed ground on the rocky coastline between
Machrihanish and Southend, the best localities here being the fairly massive
outcrop of a cornstone of the Old Red Sandstone known as ‘The Gauldrings’,
near Ballygroggan, and the extensive limestone cliffs on the coast west of
Largybaan. At Ballygroggan the outstanding species of interest is H.
britanniciforme, known from only one other Scottish locality. Other species
occurring here are H. anglicum, H. flocculosum, H. caledonicum and a plant
whose identity at present remains obscure. The hawkweed flora of the Largybaan
cliffs includes four species of the Section Cerinthoidea—H. anglicum, H. floccu-
losum, H. ampliatum and H. shoolbredii. In addition, H. subrude, H. rubiginosum,
H. dicella and H. sarcophylloides occur, the latter in its only Kintyre locality.
Previous to its discovery here it had been known only from the extreme north
of Scotland. Apart from these two outstanding areas, a good colony of H.
subrude is found on the rocks at Keil, Southend, and also on high ground near
the Mull lighthouse where it grows with H. argenteum, H. euprepes and H.
rubiginosum. Between Ballygroggan and Largybaan the cliffs have produced

HAWKWEEDS OF WESTERN SCOTLAND 109

H. rubiginosum, H. caledonicum and H. uisticola. The last-named is a rare
species, and this is its only known station in v.c. 101.

Considerable exposures of Old Red Sandstone occur in the southern half of
Kintyre. However these areas do not appear to be particularly productive for
hawkweeds, although on cliffs near Achinhoan Head south of Campbeltown
we have noted H. rubiginosum, H. vulgatum and H. euprepes, and on similar
rocks in the Bellochantuy—Westport area H. rubiginosum, H. caledonicum
and the apparently rare H. angustisquamum, which has only two other Scottish
localities.

Of the aphyllopodous species, H. drummondii grows near Muasdale and
in Barr Glen, H. perpropinquum near Ballochroy, and H. umbellatum at Peninver.

Since the publication of the Critical Supplement further material has been
collected which has led to a reassessment of the identity of certain plants. The
relevant maps should be amended as follows: delete H. scoticum in square 16/60.
The material on which this record was based is now considered to be a form of
H. rubiginosum; delete H. senescens in 16/60, the plant concerned being H.
subrude. H. senescens should also be deleted from square 16/78, as should
H. saxorum from the same square. Other deletions are: H. ampliatum from
16/60, H. caesiomurorum from 16/64, and H. subplanifolium from 16/78.

In all, 43 species have so far been recorded from Knapdale and Kintyre.

VICE-COUNTY 102, SOUTH EBUDES

The island of Islay and the Garvelloch Isles are the only parts of this vice-
county where we have examined hawkweeds, and our investigations have only
been of a limited nature.

There is an extensive exposure of dolomitic limestone on the northeast
coast of Islay between Rudha Bholsa and Rudha Mhail, but it has so far proved
to be disappointing. We have seen H. caledonicum in quantity, and H. scoticum
in a single locality. There is also a species of Section Tridentata which may be
H. gothicoides, but sufficiently mature plants have not yet been obtained. In
the south of Islay, on the Mull of Oa near Lower Killeyan, the following occur:
H. iricum, H. caledonicum, H. orimeles, and H. subcrocatum. H. vulgatum has
been observed in many Islay localities, including the small isle of Texa. Our
impression is that the hawkweed flora of Islay is surprisingly poor considering
the extent of suitable ground on the island.

The Garvelloch Isles botanically speaking belong to v.c. 102, and since
they are largely composed of limestone might be expected to have a good
hawkweed flora. However, in the course of a brief visit in August 1966 the
only species detected was H. shoolbredii.

VICE-COUNTY 103, MID EBUDES

Apart from H. shoolbredii recorded from Sorisdale, Isle of Coll, all our hawk-
weed records from this vice-county have been made in the Isle of Mull. Despite
some fairly extensive areas of basic rock, the hawkweed flora of Mull, so far
as we have been able to judge, is remarkably poor. No species of the Section
Alpina has been found, and only one named species of the Section Subalpina.

S-Airde Bheinn in the north of the island has extensive outcrops of a rather
poor igneous rock. H. sommerfeltii is the most interesting of its hawkweeds,
and H. shoolbredii, H. chloranthum, H. rubiginosum and H. subhirtum also occur.
The Ben More area, including Beinn Fhada and Choire Bheinn, produces

110 A. G. KENNETH AND A. McG. STIRLING

H. anfractiforme, H. shoolbredii, H. orimeles, H. chloranthum, H. caledonicum,
H. subhirtum, H. rubiginosum and H. maritimum. In addition there is an
‘unknown’ species of the Section Subalpina on Beinn Fhada which is closely
allied to H. marshallii. The rock is mostly poor, but this is a large area and
there must be a number of other species present. The most interesting of those
mentioned are H. anfractiforme from Beinn Fhada and H. maritimum from
Choirc Bheinn. Fine basic cliffs about Creag a’ Ghaill, south of Gribun, have
H. iricum, and this also occurs on shore rocks nearby. Allt na Teangaidh has
H. subhirtum and H. ampliatum; the latter also grows on Beinn na h-IJolaire.
On Creach Bheinn, a rather poor hill in the south of the island, we have noted
H. shoolbredii and H. chloranthum.

H. shoolbredii is fairly generally distributed, and some forms have more or
less glabrous leaves (as on s-Airde Bheinn) which lack the usual stellate hairs on
the undersurface. There are numerous localities throughout the island for
H. chloranthum, and H. vulgatum is, as usual, widely distributed. One of the
few rarities, H. sarcophylloides, was noted on basalt cliffs near the mouth of
Loch Spelve in the southeast of Mull where it has been observed in flower in
the last week in May, an early date for any hawkweed in the western highlands.

Of the aphyllopodous species, H. /atobrigorum grows at Aros Bridge, also at
Knock. H. strictiforme of the same section has been noted only on Iona.

The total number of species recorded by us in Mull to date is only 15, which
compares unfavourably with apparently very similar mainland areas.

VICE-COUNTY 104, NORTH EBUDES

Our observations in this vice-county have been limited to the Isle of Skye
and Raasay. In the course of a number of short visits to the Isle of Skye several
species have been noted, mainly from the lower slopes of the basalt hills or
from rocky ground and cliffs around the coast. Outcrops of limestone, both
Jurassic and Cambrian, occur in some areas, and these have also provided a
few records.

The basalt hills have yielded H. anglicum, H. chloranthum, H. shoolbredii
and H. orimeles, while H. rubiginosum has been found in basalt boulder scree
below sea-cliffs near Duntulm on the north coast. On the Jurassic strata H.
piligerum and H. pictorum have been recorded from the lower slopes of Blaven,
and near Elgol H. shoolbredii occurs. The areas of Cambrian (Durness) lime-
stone near Broadford and elsewhere have proved rather disappointing in the
number of species noted, though one comparatively rare species, H. rhomboides,
has been noted near Loch Cill Chriosd. Other species recorded from the
Cambrian are H. duriceps, H. vulgatum, H. langwellense and H. reticulatum.

Of the Section Subalpina the only species yet encountered has been H.
lingulatum, the most frequently occurring member of the section. This was
found on Blaven. In spite of the altitude reached by the Cuillins these hills
appear to be almost destitute of hawkweeds, the rock being gabbro which
supports a very poor phanerogamic flora. No species of the Section Alpina
has been found on the island.

During the course of a week’s visit to Raasay at the end of June 1969 a
number of hawkweeds were recorded. These were observed mainly on lime-
stone and other associated rocks of the Jurassic series which provide the richest
habitats on the island. The species observed were H. ampliatum, H. langwellense,
H. shoolbredii, H. cymbifolium, H. duriceps, H. pictorum, H. caesiomurorum,

HAWKWEEDS OF WESTERN SCOTLAND 111

H. pollinarioides, H. diaphanoides, H. cravoniense, H. rubiginosum, and AH.
vulgatum. The occurrence of H. cymbifolium represents a considerable north-
ward extension of range, while that of H. pollinarioides adds a third vice-county
to the distribution of this interesting north-west Scottish endemic.

The Isle of Scalpay, neighbour to Raasay and Skye, proved in the course of a
brief visit to be rather poor in Hieracium, only H. shoolbredii and H. vulgatum
being noted. This island is composed mainly of Torridonian Sandstone.

VICE-COUNTY 105, WEST ROSS

This is an extensive vice-county with much good hawkweed ground. Our
investigations have been mainly concentrated on some of the hills. In the north,
Cul Beag and Ben More Coigach have been systematically searched. Much of
the rock (Torridonian Sandstone) is poor, and hawkweeds are largely confined
to the better areas. However the lists for both hills are interesting. On Cul Beag
the following species occur: H. holosericeum (an unusual form with dark heads),
H. alpinum, H. lingulatum, H. gracilifolium, H. anglicum, H. senescens, H.
nitidum, H. sarcophylloides, H. caledonicum, H. subhirtum, H. piligerum and
H. pictorum. There is also at least one undescribed plant from this hill. On Ben
More Coigach and Beinn an Eoin there is a formidable assembly consisting of:
H. marginatum, H. globosiflorum, H. hanburyi, H. eximium, H. lingulatum, H.
centripetale, H. pseudanglicum, H. glandulidens, H. hebridense, H. ampliatum,
H. shoolbredii, H. argenteum, H. uistense, H. euprepes, H. rivale, H. subtenue,
H. duriceps, H. camptopetalum, H. pictorum, H. piligerum, H. rubiginosum,
H. caesiomurorum, H. caledonicum and H. subhirtum. This must be one of the
best localities in the north-west for hawkweeds, and, like Cul Beag, it too has
at least one distinct-looking plant not referable to any of the described species.

Opposite Ullapool on the Scoraig peninsula, Beinn Gobhlach produces
H. hanburyi, H. lingulatum, H. nitidum, H. subtenue, H. subhirtum, H. euprepes,
and H. dipteroides. This last is a rare plant and quite new to this area. Much
further south, in Glen Shiel, Sgurr an Lochain has H. globosiflorum with very
dark heads and styles, H. piligerum, H. subtenue, and the scarce H. oxyodus,
while a little further east from this hill, Coire an t’ Slugain yielded a good
selection including H. anfractiforme, H. sinuans, H. lingulatum, H. nigris-
quamum, H. euprepes, H. subtenue and H. vulgatum. Near Inverewe the hills
of the Coulin Forest have H. holosericeum, H. hanburyi, H. globosifolium and
H. lingulatum.

The extensive area of Cambrian limestone near Kishorn has been fairly
well worked, the following species occurring: H. lingulatum, H. ampliatum,
H. hebridense, H. shoolbredii, H. chloranthum, H. caledonicum, H. piligerum,
H. duriceps and H. vulgatum. This area has an extremely interesting general
flora (Slack & Stirling 1963), and it is therefore surprising that the hawkweeds
there include none of the very local and interesting species known from the
same geological formation in the Durness area of Sutherland. Also in the
Kishorn area the epidiorite rocks of An Sgurr and Bad a’ Chreamha have
yielded a rather similar list, although H. lingulatum, H. piligerum and H.
duriceps appear to be absent. On the rocks of Sgurr a’ Gharaidh, just above
the upper limit of the limestone, H. dasythrix was found, but this hill was not
closely examined, nor were the Applecross hills to the west, H. nitidum and a
few of the commoner species being all we have noted there.

The aphyllopodous species H. Jatobrigorum and H. perpropinquum occur on

112 A. G. KENNETH AND A. McG. STIRLING

rock outcrops by roadsides near Dornie. This is the most northerly station yet
recorded for the latter species.

Some of the West Ross plants call for special comment. The colony of
H. alpinum on Cul Beag appears to be the most northerly at present known,
while the record of H. eximium from Ben More Coigach represents a great
extension of range, being the first from north of the Great Glen. All populations
of H. globosiflorum seen in v.c. 105 have been dark-styled, and this is in fact
the only form we have seen in the western Highlands. H. camptopetalum, which
we have noted from Blarnaleyoch, near Ullapool, as well as from the Ben More
Coigach area already mentioned, appears to be a typical hawkweed of alpine
or montane streamsides, and may be more widespread than at present supposed.
The number of species noted by us in West Ross is 36.

VICE-COUNTY 108, WEST SUTHERLAND

This large and interesting vice-county deserves a much fuller investigation
than we have been able to make. The limestone areas of Inchnadamph and
Durness have been thoroughly covered by others, as has much of the northern
coast, and we have not spent much time in these areas. Our researches have
been concentrated on the hills which, apart from Ben Hope and Ben Loyal,
have been rather neglected.

On the hills of the Parphe, the district south of Cape Wrath, including
Creag Riabhach, An Grianan, Fashven, Cnoc na Glaic Tarsuinn and Farrmheall,
the following species occur: H. globosiflorum, H. glandulidens, H. senescens,
H. dasythrix, H. ampliatum, H. shoolbredii, H. caledonicum, H. argenteum,
H. praetermissum, H. pauculidens, H. euprepes, H. subtenue, H. nitidum, H.
rivale and H. rubiginosum. The north-facing cliffs of Creag Riabhach where,
by the way, H. ampliatum is a most unusual-looking plant, may well be worth
further investigation. This is a very wild area. The range of hills comprising
Cranstackie, Beinn Spionnaidh and Beinn Ceannabeine yields the following:
H. lingulatum, H. euprepes, H. anglicum, H. ampliatum, H. shoolbredii, H.
caledonicum and H. subrude.

Possibly the most promising ground for hawkweeds is Foinnaven, including
Cnoc a’ Mhadaidh. In addition to several plants of doubtful identity, the
following occur: H. marginatum, H. centripetale, H. glandulidens, H.
pseudanglicoides, H. lingulatum, H. senescens, H. shoolbredii, H. sarcophylloides,
H. argenteum, H. nitidum, H. subtenue, H. subhirtum, H. rubiginosum and H.
dovrense. This is the third Scottish station for H. dovrense, a Scandinavian
species otherwise known only from Ben Loyal and near Rhiconich in the same
vice-county. It belongs to the Section Alpestria. H. marginatum is otherwise
known only from Ben Loyal, Ben More Coigach and Ben Klibreck. A.
sarcophylloides in an unusual montane locality, and H. pseudanglicoides are
other noteworthy species. The occurrence of the latter on Cnoc a’ Mhadaidh
greatly extends its known range. There is an interesting open ravine near
Rhiconich which produces H. glandulidens, H. ampliatum, H. shoolbredii, H.
nitidum and the rare species H. oxyodus and H. dovrense.

A brief survey on Ben Stack produced only H. caledonicum, H. duriceps and
H. subtenue, though there was a species of Section Alpina, unfortunately not
in determinable condition. H. iricum is by Loch Stack, and HA. sparsifolium
by the River Laxford. The coastal areas about Oldshoremore have #H.
pollinarioides, H. iricum and H. nitidum, while the interesting Sandwood area

HAWKWEEDS OF WESTERN SCOTLAND 113

produces H. jovimontis, H. praetermissum, H. argenteum and H. rubiginosum.
This last-mentioned area requires examination earlier in the year than we have
so far found possible. North of the Sandwood area, in spite of apparently
suitable ground, the coast and streamsides seem devoid of hawkweeds until
Cape Wrath is reached. Then in the area Geodha na Seamraig to Kearvaig
we find H. shoolbredii, H. sarcophylloides, H. caledonicum and H. sparsifolium.
On the coast north of Loch Inshore, H. caledonicum and H. strictiforme occur,
while near the Cape Wrath ferry H. iricum, H. shoolbredii, H. praetermissum,
H. argenteum and H. latobrigorum have been recorded. Near Durness, at Smoo,
H. pollinarioides and H. caesiomurorum have been noted, and further east at
Traigh na h-Uamha H. praetermissum and H. variifolium are present along
with H. iricum and other species. H. praetermissum and H. pollinarioides are
confined to this northwest corner of Sutherland, and H. variifolium is practically
so, having one outlying station. H. pollinarium, which is also confined to West
Sutherland where it is very local on the north coast, has been noted near the
mouth of the River Naver at Bettyhill.

Several identifications have been made from other areas of the vice-county.
Quinag has H. globosiflorum (again the dark-styled form), H. glandulidens, H.
anglicum, H. argenteum, H. pictorum and H. subtenue, and Glas Bheinn in the
same area has H. holosericeum. On the limestone near Ledmore H. angustis-
guamum has been found; this is an interesting isolated locality for a species
whose only other Scottish occurrences are in Argyll and Kintyre. Ben Hope
has been visited, and that well-worked hill has given H. shoolbredii, H. nitidum,
H. praetermissum, H. variifolium, H. subtenue and H. camptopetalum. Further
to the east an interesting ravine near Kirtomy has produced H. nitidum, H.
scoticum, H. subrude and H. duriceps.

Some general comments on the distribution of species in the vice-county
may be of interest. It is curious that H. vulgatum is apparently absent from the
Rhiconich—Durness area. We have seen it by Loch Stack, and on the east side
of Loch Eriboll. Elsewhere in the Highlands it seems to be ubiquitous. H.
caledonicum is frequent in the areas we have examined, while H. iricum appears
to be rather more frequent than it is in other parts of the west Highlands. Its
preference seems to be for base-rich sites on the low ground. H. anglicum and
H. shoolbredii are predictably well distributed in the lower parts, and H.
argenteum is also frequent.

H. dovrense, to which some reference has already been made, appears to
favour ravine conditions at medium to high altitudes, and may well occur more
widely than the records suggest. We do not know it in its Ben Loyal station
where it seems to be a rare plant, but at Rhiconich and on Foinnaven it grows
in some quantity. These sites are much like those favoured by H. dewarii, but
in complete contrast to those of H. solum.

Up to the present, 37 species have been noted by us in the vice-county.

GENERAL OBSERVATIONS

Having dealt with the species on a vice-county basis, it is now appropriate to
make some observations on their distribution and ecology in relation to the
west of Scotland area as a whole.

The preference of species of the Section Alpina is generally for the more
exposed rocky situations, or even, as in the case of H. holosericeum, for exposed

B

114 A. G. KENNETH AND A. McG. STIRLING

stony summits; however, they sometimes occur in the higher gullies. The lowest
altitude at which we have observed species of this section in the west central
Highlands is on Ben Vane, Dunbartonshire, where H. alpinum and H. eximium
grow at about 1,800 ft. In the northwest Highlands however, H. globosiflorum
occurs on the Cape Wrath hills down to 1,300 ft.

Members of the Section Subalpina are generally found on the hills from
about 1,000 ft upwards, although in a number of localities certain species occur
at much lower altitudes. The species nearest to Section Alpina are unlikely
to display this tendency. Undoubtedly the most frequent species are H.
lingulatum, H. senescens and H. dasythrix, and these may reasonably be expected
on any hill with a good hawkweed flora, except perhaps in the extreme north,
where the species belonging to the Section Subalpina seem to be distinctly
scarcer. On most of the hills of the west central Highlands H. pseudanglicum
is rather frequent.

The species of the Section Cerinthoidea are not essentially plants of the hills.
While some appear to be indifferent to base status, e.g. H. shoolbredii and
H. langwellense, others such as H. anglicum, H. ampliatum and H. iricum are
distinctly basiphile. We have not observed plants of this section at a greater
altitude than 2,000 ft.

Three species of the Section Oreadea are common and widespread in the
west Highlands, namely H. caledonicum, H. argenteum and H. chloranthum,
though the latter is absent from the extreme north. The species of this section
vary in ecological preference, the three just mentioned not appearing to restrict
themselves to basic sites, while the group centred on H. dicella seem to be
found only on basic rock. As a whole the section is best represented at low
to moderate elevations, H. sommerfeltii reaching the greatest elevation of the
species we have seen.

The Section Vulgata contains a larger number of species than any other
section and is a confusing assemblage of plants with few obvious characteristics
common to all. The most frequent species are H. subhirtum, H. pictorum, H.
duriceps and H. vulgatum. The first two are typical of the west Highland hills,
and H. duriceps, though often seen in hill gullies, also occurs on exposed rock,
sometimes at low elevations. H. vulgatum is a most variable and ubiquitous
species almost throughout our area. It occurs in many habitats, both natural
and artificial, and reaches at least 2,000 ft. Hawkweeds growing on walls and
similar artificial sites practically always turn out to be this species, though as
already mentioned H. chloranthum has once been obtained from a wall. Plants
of the H. exotericum complex are not common in our area. They are introduc-
tions and probably spreading, and like H. vulgatum they often occur on walls
or disturbed ground. The only representative dealt with in our account is
H. grandidens.

Of the Section Tridentata, H. sparsifolium is widespread and frequent at
moderate elevations in our area, and H. uiginskyense is of sporadic occurrence,
though it has not been observed outside the Highland area. Of the remaining
species, two have been noted, but these were not within the Highland area.

H. latobrigorum is perhaps the most frequent of the Section Foliosa, and it is
not uncommon particularly on roadside habitats. H. strictiforme is almost as
frequent but is perhaps more inclined to occur in natural habitats and at a
greater elevation. H. subcrocatum and H. reticulatum are distinctly less common,
and H. subumbellatiforme is apparently a rare plant. Mention has already been

HAWKWEEDS OF WESTERN SCOTLAND 115

made of its occurrence at the unusual elevation of 2,000 ft on Meall a’ Bhuiridh,
Glen Coe.

The Section Umbellata is represented by H. umbellatum, which is sparingly
distributed through much of our area, usually at a low altitude both in natural
situations such as rocky woodland and in artificial habitats (railway embank-
ments, waste ground etc.). We have not observed it north of Strontian, Argyll.

The only species of the Section Sabauda we have seen in our area is H.
perpropinquum which has a scattered distribution on the lower ground as far
north as Dornie in West Ross.

It is hoped that the foregoing information will serve in some measure to
expand the information given in the Critical Supplement, and it may be found
helpful when consulting the latter to use it in conjunction with the vice-county
overlay provided with the At/as. In mentioning the Critical Supplement it is
necessary to draw attention to an unfortunate error concerning two of the
Hieracium maps. Due to a transposition of the titles, the map of H. nitidum
shows the distribution of H. jovimontis and vice versa.

THE GENUS PILOSELLA

In accordance with the view of Sell and West, first propounded by the brothers
Schultz in 1862, we are treating the subgenus Pil/osella as of full generic rank.
The taxon Hieracium pilosella L. as a result of this treatment must now be
known as Pilosella officinarum C. H. & F. W. Schultz, of which a number of
distinct forms have been given subspecific rank (Sell & West 1967), and we
give here some indication of the occurrence of these forms within our area.
However, this at best is only a rough outline of their distribution as insufficient
material has been determined from most of our area, though Argyll and Kintyre
are well represented. It would therefore be unwise to draw any conclusions as
to the relative frequency of the various subspecies.

Pilosella officinarum sensu lato usually occurs on neutral to basic grassland,
walls, etc. Unlike most species of Hieracium it is usually a soil plant rather
than rupestral.

Pilosella officinarum subsp. officinarum

v.c. 98 Gallanach, near Oban; Seil; Cuan Ferry; Dun Nucaig; Stronmilchan,
near Dalmally

v.c. 104 Oskaig, Raasay

Pilosella officinarum subsp. concinnata

v.c. 98 Isle of Kerrera; Kerrera Ferry; Gallanach; Luing; Seil; Ben Buidhe;
Isle of Bernera, Lismore

v.c. 101 Isle of Danna; Largybaan; Keil, Southend; Loch an Dobhrain and
Cnoc na Seamraig, Knapdale

v.c. 102 Garbh Eileach, Garvelloch Isles

v.c. 103 Aros Bridge and Creag a’ Ghaill, Mull

v.c. 105 Kishorn

v.c. 108 Bettyhill; Traigh na h-Uamha; Sheigra; Culkein of Drumbeg

Pilosella officinarum subsp. trichoscapa
v.c. 98 Bealach Mor; Creag a’ Chapuill; Luing; Seil; Cuan Ferry; Dun
Nucaig

116 A. G. KENNETH AND A. McG. STIRLING

v.c. 101 Isle of Danna; Lussa; Achnamara; Point of Knap
v.c. 102 Bridgend, Islay

v.c. 103 Allt na Teangaidh, Mull

v.c. 104 Near Loch Cill Chriosd, Skye

Pilosella officinarum subsp. tricholepia

v.c. 72 Scar Water

v.c. 86 Gonachan Burn, Campsie Hills

v.c. 97 Drumnadrochit

v.c. 98 Kerrera Ferry; Creag nam Fitheach; Creag an Sturra; Ford—
Kilmartin road junction

v.c. 101 Keills, Knapdale

v.c. 102 Killeyan, Mull of Oa, Islay

v.c. 104 Near Loch Cill Chriosd, Skye; in several localities on Raasay

v.c. 108 Culkein of Drumbeg; Traigh na h-Uamha

Pilosella officinarum subsp. nigrescens

v.c. 72 Dalveen Pass

v.c. 86 Campsie Hills above Lennoxtown

v.c. 98 Luing; Seil; Kerrera; near Kirnan; Creag an Sturra; Gallanach;
Kilmichael Glassary; Isle of Bernera, Lismore

v.c. 101 Isle of Danna

Pilosella officinarum—unnamed subsp.

v.c. 97 Letterfinlay

v.c. 98 Coire Chalein, Beinn Beag; Kilmartin; Kirnan; Tibertich; Creagan-
tairbh Mor

v.c. 101 Isle of Danna; Ballygroggan; Campbeltown; Keil, Southend; Largy-
baan coast

v.c. 108 Sheigra

The distinguishing characters of these subspecies and of the other species of
Pilosella occurring in the British Isles are given in Appendix I. It should be
pointed out that, while many specimens can be referred to one or other of
these taxa, intermediates are common and have been ignored for the purpose of
the present notes. P. officinarum subsp. concinnata, the most frequent subspecies
in our area, occurs in two distinct forms, one with short pale glandular hairs,
and the other with short darker glandular hairs. This second form should not
be confused with the unnamed subspecies with long, dark glandular hairs.

ACKNOWLEDGMENTS

Our grateful thanks are due to Mr P. D. Sell and Dr C. West for the encourage-
ment and help they have given us at all stages of the production of these notes,
and for scrutinising the final manuscript, also for devoting so much time and
effort to the determination of the many specimens which we submitted for
identification. Although this account has been based solely on the authors’
personal records, they are greatly indebted to the following who have so
generously made their own Scottish hawkweed records available: Miss U. K.
Duncan, Miss C. W. Muirhead, Miss M. McCallum Webster and Mr. W. Scott.
This additional information has rendered the distributional pattern of many
species much clearer.

HAWKWEEDS OF WESTERN SCOTLAND EJ)
REFERENCES

Danby, J. E. (1958). List of British Vascular Plants. London.

PERRING, F. H. & SELL, P. D., ed. (1968). Critical Supplement to the Atlas of the British Flora.
London.

Pucs ey, H. W. (1948). A Prodromus of the British Hieracia. J. Linn. Soc. (Bot.), 54.

RAVEN, J. E. & WALTERS, S. M. (1956). Mountain Flowers. London.

SELL, P. D. & West, C. (1967). Taxonomic and Nomenclatural Notes on the British Flora.
Watsonia, 6: 313-314.

Stack, A. A. & STIRLING, A. McG. (1963). The Cambrian Limestone Flora of Kishorn,
West Ross. Proc. bot. Soc. Br. Isl., 5: 1-12.

APPENDIX I

ARTIFICIAL KEY TO THE BRITISH SPECIES AND SUBSPECIES
OF PILOSELLA HILL

This key is included by kind permission of the compilers, P. D. Sell and Dr C. West’
Taxa known to occur in Scotland are asterisked.

1. Ligules brownish- or purplish-red or orange (P. aurantiaca) :
Ligules yellow, often with a reddish stripe on the back of the outer face .

WN

2. Stolons long and abundant; phyllaries 6-8 mm long é‘
*P. aqurantiaca subsp. TarEeOCrecen
Stolons shorter and fewer; phyllaries 8-10 mm long
*P. aurantiaca subsp. nanitings

3. Plants with a single capitulum ona scape .. ; as 53 Bins 4
Plants with stems bearing more than one capitulum. . oa Ae sac 11
4, Phyllaries with numerous glandular hairs, without simple hairs .. ae 5
Phyllaries with simple hairs, with or without glandular hairs a 2 6

5. Small plant; glandular hairs of phyllaries up to 0-5 mm long, equal or almost so,
yellowish or blackish . *P. officinarum subsp. concinnata
Larger plant; glandular hairs of phyllaries up to 1 mm long, unequal, blackish
*P. officinarum unnamed subsp.

6. Phyllaries with numerous simple hairs and numerous glandular hairs .. 7
Phyllaries with numerous simple hairs and few or no glandular hairs _.... 8

7. Phyllaries with pale, simple hairs and pale, glandular hairs
*P. officinarum subsp. Of ana
Phyllaries with longer, dark simple hairs and more robust, dark glandular hairs
*P. officinarum subsp. nigrescens

8. Stolons short, thick and turning up at the ends (P. peleteriana) .. ce 9
Stolons long, slender and prostrate .. Se es as a ahs 10

9. Scape 10-20 cm high; leaves lanceolate, oblanceolate or elliptic; capitula large

and broad Rs P. peleteriana subsp. peleteriana
Scape to 30 cm high; leaves narrower, capitula smaller and narrower

P. peleteriana subsp. porurnenan

10. Hairs of phyllaries dark me sh *P. officinarum subsp. trichoscapa
Hairs of phyllaries pale .. an oy *P. officinarum subsp. tricholepia

118

A. G. KENNETH AND A. McG. STIRLING

11. Phyllaries 10-12 mm ae (P. Aacelae) "e oe ses ae i 12
Phyllaries 6-9 mm long . fe ae an = As: ake 13
12. Hairs of stem 2-3 mm long; hairs of phyllariesto1-Smmlong ..

*P. flagellaris subsp. Wiesehane

Hairs of stem up to 7-5 mm long; hairs of phyllaries to2-S mmlong ..
*P. flagellaris subsp. bicanitala
13. Stem with up to 8 capitula (P. lactucella) ne ae i sz a 14
Stem with usually more than 10 capitula ies ae 3 Bs A. 15
14. Stem 5-12 (—20) cm high P. lactucella subsp. lactucella
Stem over 20 cm high P. lactucella subsp. helveola
15. Leaves deep yellowish green, oblanceolate, widest leaves of plant 15-30 mm wide
*P. caespitosa subsp. colliniformis
Leaves caesius green, very narrowly elliptic or oblong, widest leaves less than
(usually much less than) 15 mm wide (P. praealta) wh a ae 16
16. Leaves glabrous or nearly so; phyllaries 5-6 mm long P. praealta subsp. spraguei
Leaves with hairs, at least on the margins and midrib; phyllaries 6-8 mm long 17
17. Stolons short and upturned; peduncles and phyllaries with no or few simple
hairs” 22 P. praealta subsp. praealta
Stolons long and prostrate: peduncles and ‘phyllaries with numerous simple
haies, ae Me Bs om aie ag *P. praealta subsp. arvorum

APPENDIX II

INDEX OF PLACE NAMES MENTIONED IN THE TEXT WITH

NATIONAL GRID REFERENCES

v.c. 72, Dumfries

Crawick Water 26/79-13 Mennock 26/81-08

Dalveen Pass 26/90:07 Scar Water 25/79 -99

Glenwhargen Craig 26/76-03

v.c. 73, Kirkcudbright

Colvend 25/88:54 Lauriston 25/68 -64

Grey Mare’s Tail, Poldores Burn 25/61-96
Newton Stewart 25/49:72 Tongland Bridge 25/69-53

High Bridge of Ken 26/61 -90

Kells range 25/5-8

v.c. 75, Ayr

Craig Hill, Straiton 26/38-01

Dalmellington 25/50-03

v.c. 76, Renfrew

Lochwinnoch 26/35-59

Neilston 26/47-57

Shielhill Glen 26/23-72

v.c. 86, Stirling

Ballagan Glen 26/57-79

Campsie Glen 26/61-80

Double Craigs 26/63 -87

Gonachan Burn

26/63-85

HAWKWEEDS OF WESTERN SCOTLAND

v.c. 97 (Argyll)
Beinn na Beathrach
Beinn na h-Uamha
Creach Bheinn
Garbh Bheinn
Gearradh

v.c. 98, Argyll
Aonach Dubh
Balvicar

Bealach Mor
Beinn a’ Chaisteal
Beinn a’ Chleibh
Beinn Buidhe
Beinn Dothaidh
Beinn an Lochain
Beinn Luibhean
Ben Cruachan
Ben Donich

Ben Ime

Ben Narnain
Beinn an t-Seilich
Beinn Udlaidh
Bernera

Bidean nam Bian
Binnean an Fhidhleir
Buchaille Etive Mor
Coire Chailean
Creag Bhan
Creag a’ Chapuill

v.c. 99, Dunbarton
Ben Vane

Ben Vorlich

Beinn Dubh
Auchengaich Burn

v.c. 101, Kintyre
Artilligan Burn
Achinhoan Head
Bellochantuy
Carse

Cnoc na Seamraig
Cruach Lusach
Danna

The Gauldrings, Ballygroggan
Keil

Largybaan cliffs

v.c. 102, South Ebudes
Garvelloch Isles

Lower Killeyan

Rudha Bholsa

17/75-57
17/68-53
17/87-57
17/90-62
17/96-61

27/15-56
17/76-16
17/83-04
27/3436
27/24-25
27/20-18
27/32-40
27/21 -07

27/24-07

27/09-30
27/21-04
27/25-08
27/27-06
27/20-07
27/27-32
17/79-39
27/14-54
27/21-10
27/21-54
27/31-33
27/10-53
17/85-02

27/27-09
DUIZIND
HAYA
26/27-90

16/85-77
16/76-17
16/66-38
16/73-61
16/80-77
16/78-83
16/69-78
16/62:19
16/67-07
16/59-15

17/66-12
16/27-43
16/38-78

Killundine

Loch Dur na Marst
Maol Odhar

Meall nan Each
Sithean na Raplaich

Creag nam Fitheach
Creag an Sturra
Creagantairbh
Cruach an Nidd
Cuan Ferry
Dalvuie

Dun Nucaig
Ellenbeich
Gallanach
Ganavan

Glen Gallain
Kerrera Ferry
Lag na Cille
Loch Tralaig
Meall a’ Bhuiridh
Meeting of the Waters
Pass of Melfort
Raera

Sroin na Creise
Stob an Eas
Tibertich

Cochno Loch
Glenarbuck
Glenmallan
Kilpatrick Hills

Lussa (river)

Loch an Dobhrain
Meall Ruadh
Mull of Kintyre
Ormsary

Scotnish

Sliabh Gaoil
Stronefield
Wishing Ledge

Rudha Mhail
Texa Isle

119

17/58-49
17/66-52
17/88-57
17/88-56
17/64-52

17/83-04
17/82-14
17/18-01
17/85-14
17/75-14
17/92-36
17/75-15
17/74-17
17/82-25
17/85-32
17/85-18
17/83-28
17/85-25
17/87-16
17/24-50
27/1756
17/84-15
17/82-20
27/23-52
27/18-07
17/84-02

26/47-75
26/45-74
26/25-96
26/4-7

16/77-85
16/80-79
16/79-80
16/60-07
16/73-71
16/75-88
16/82-74
16/71-73
16/72-61

16/42-79
16/39-43

120 A. G. KENNETH AND A. McG. STIRLING

v.c. 103, Mid Ebudes
Allt na Teangaidh
s-Airde Bheinn

Aros Bridge

Beinn Fhada

Ben More

Choire Bheinn

v.c. 104, North Ebudes
Blaven

Broadford

Duntulm

v.c. 105, West Ross
An Sgurr

Bad a’ Chreamha
Beinn Gobhlach
Ben More Coigach
Blarnaleyoch

Core an t-Slugain

v.c. 108, West Sutherland
An Grianan

Beinn Ceannabeine
Beinn Spionnaidh

Ben Hope

Ben Loyal

Ben Stack

Cape Wrath ferry

Cnoc na Glaic Tarsuinn
Cnoc a’ Mhadaidh
Cranstackie

Creag Riabhach
Culkein of Drumbeg
Farrmheall

Foinaven

Geodha na Seamraig

17/45-32
17/47-54
17/55-44
17/54-35
17/52-33
17/48-32

18/53-21
18/6-2
18/41-74

18/85-38
18/85-36
28/05-94
29/10-04
28/14-90
28/05-08

29/26-62
29/42-64
29/36-57
29/47-50
29/58-48
29/27-42
29/37-66
29/29-65
29/32-52
29/35-55
29/28-64
29/11-34
29/30-59
29/31-50
29/28-73

Creach Bheinn
Creag a’ Ghaill
Knock

Loch Spelve
Sorisdale, Coll

Elgol
Loch Cill Chriosd

Coulin Forest

Cul Beag

Dornie

Kishorn

Sgurr an Lochain
Sgurr a’ Gharaidh

Glas Bheinn
Kearvaig
Kirtomy
Ledmore
Loch Eriboll
Loch Inshore
Oldshoremore
R. Laxford
Quinag
Parphe
Sandwood
Sheigra

Smoo

Traigh na h-Uamha

17/63-26
17/45-32
17/54-39
17/73-27
17/27-63

18/52-14
18/61-20

18/9-5

29/14-08
18/88-26
18/83-40
28/00-10
18/88-44

29/25-26
29/29-72
29/76-62
29/24-12
29/40-54
29/33-70
29/20-58
29/24-47
29/19-29
29/25-60
29/22-65
29/19-60
29/41-67
29/44-65

Watsonia, 8, 121-134 (1970). 121

A revision of some of the taxonomic characters of
Polypodium australe Fée

R. H. ROBERTS

51 Belmont Road, Bangor, Caerns.

ABSTRACT

In addition to the presence of paraphyses three other characters have been emphasized for
identifying Polypodium australe. These are the number of indurated annulus cells, the length
of the rhizome scales and frond shape. A reassessment of these characters from cytologically
determined plants shows that the variation in all of them is greater than has been supposed,
and the overlap with P. interjectum Shivas is such as to reduce their taxonomic value con-
siderably. It is also evident that most of the plants formerly thought to be morphologically
intermediate between P. australe and P. interjectum, and consequently regarded as hybrids,
fall within the ambit of P. australe. The shape of the rhizome scales as an accessory taxonomic
character is discussed, and support is given to the view that the large, morphologically distinct
paraphyses of P. australe are its best single distinguishing character.

INTRODUCTION

The existence of three distinct taxa in Europe within the aggregate species
Polypodium vulgare L. was first recognized by Rothmaler (1929). He ranked
them as subspecies, namely subsp. vulgare, subsp. prionodes Rothm. and subsp.
serratum* (Willd.) Christ. Cytological support for this view was provided some
eighteen years later by Manton (1947, 1950), who showed that three cytotypes
occurred within the European aggregate. It is well to recall, however, that the
form shown by Manton (1950) to be diploid had been recognised as a distinct
entity as long ago as 1810 by Willdenow, who had named it P. vulgare L. var.
serratum. On several occasions subsequently specific rank has been proposed
for it, for example by Futo (1905). As a result of cytogenetic and biometric
studies Shivas (1961) raised the three cytotypes to specific rank, the diploid
becoming P. australe Fée (= P. vulgare L. subsp. serratum (Willd.) Christ),
the tetraploid P. vulgare L. sensu stricto and the hexaploid P. interjectum Shivas
(= P. vulgare L. subsp. prionodes Rothm.). She also described their morphologi-
cal characters and most Flora descriptions published since have been based largely
on her work, e.g. Warburg (1962) and Valentine (1964).

Nevertheless, during the examination of a large number of specimens, the
writer encountered considerable difficulties. These were particularly apparent
in the case of the diploid, P. australe Fée, whose published descriptions seemed
to contain some serious errors, especially with regard to the number of indurated
annulus cells and frond shape (Roberts 1966). Two characters had been relied
upon to identify the specimens on which my observations were based. The first
was the presence of paraphyses among the sporangia; the second was the

* The correct name for this taxon at the subspecific level is actually subsp. serrulatum
Arcangeli.

122 R. H. ROBERTS

contracted appearance of the flattened annulus in P. australe compared with
P. interjectum Shivas. These annulus differences have been studied by Benoit
(1966), who gives excellent figures of them, and by Roberts (1966). However,
paraphyses were reported among the sporangia of P. interjectum in the Nether-
lands (Meinders-Groeneveld & Segal 1967), and in those of both P. interjectum
and P. vulgare in Germany (Lenski 1962, 1964). The presence of paraphyses
could thus no longer be regarded as an absolute diagnostic character of P.
australe, as I had assumed in 1966. The possibility arose that some of the
specimens on which my observations had been made might have been referable
to P. interjectum. It was clear that the solution could be found only by the
reappraisal of these characters in plants which had been determined cytologi-
cally.

MATERIALS AND METHODS

Several of the plants used in my previous study had been kept in cultivation.
Further collections of living plants were made subsequently, making a total of
twenty-eight plants from ten localities. Fixations of young sporangia from all
of them were made in 1:3 acetic-alcohol and examined as soon as possible
afterwards by squashing two or three sori in aceto-carmine. Suitable prepara-
tions were preserved for a few hours by ringing the cover-slip with paraffin wax.
A few were made permanent in Euparal. Spore mother cells showing well-spread
chromosomes were drawn under an oil-immersion objective and counts made
from the drawings.

Meiosis was found to be perfectly normal in all of these twenty-eight plants.
There was no evidence of lagging chromosomes or any other irregularities
found during meiosis in hybrids. Moreover, they all showed 37 bivalents, as

a 2<* ie
wy Yay
0” & off

FiGurE 1. Diagram of the photomicrograph of meiosis, Plate la.

TAXONOMIC CHARACTERS OF POLYPODIUM AUSTRALE FEE 123

can be seen in the photomicrograph of meiosis in one of them (Plate la and
Fig. 1).

Twenty plants each of P. interjectum and P. vulgare were also determined
cytologically and used for comparison with the diploid plants.

SPORANGIUM CHARACTERS

The number of annulus cells* in each of the diploid plants was counted in fairly
large, random samples of sporangia when these had dehisced later in the
season. The results are given in Table 1. They differ widely from those given by

TABLE 1. THE NUMBER OF INDURATED ANNULUS CELLS IN
TWENTY-EIGHT PLANTS OF P. AUSTRALE, ALL WITH A
CHROMOSOME NUMBER OF 2 = 37

No. of
Locality Range Mode sporangia
examined
Caernarvonshire, v.c. 49
Conway No. 1 3-13 6 300
Conway No. 2 4-14 7 330
Conway No. 3 3-9 5 450
Conway No. 4 3-10 6 370
Conway No. 5 3-8 5 220
Criccieth No. 1 3-10 6 200
Criccieth No. 2 5-14 9 360
Criccieth No. 3 5-15 9 250
Criccieth No. 4 4-12 — 8 200
Glanwydden No. 1 3-10 6 300
Glanwydden No. 2 3-11 7 450
Great Orme 4—10 6 300
Little Orme No. 1 2-8 5 500
Little Orme No. 2 4-12 8 540
Little Orme No. 3 4-16 9 530
Little Orme No. 4 3-13 8 610
Penrhynside No. 1 3-10 6 200
Penrhynside No. 2 3-13 il 450
Anglesey, v.c. 52
Lligwy Wood No. 1 4—13 7 770
Lligwy Wood No. 2 5-10 7 220
Lligwy Wood No. 3 3-10 6 250
Lligwy Wood No. 4 4-8 6 200
Pentraeth No. 1 3-11 6 350
Pentraeth No. 2 3-10 6 450
Pentraeth No. 3 4-14 9 250
Pentraeth No. 4 6-18 10 350
Denbighshire, v.c. 50
Llanddulas 5-14 9 260
North Somerset, v.c. 6
Cheddar 3-9 6 240

*To avoid tiresome repetition ‘indurated annulus cells’ are referred to in most of this
paper as ‘annulus cells’

124 R. H. ROBERTS

Futo (1905), Manton (1950), Shivas (1961) and Rothmaler & Schneider (1962),
who give the following values respectively: 6, (4-)5(-6), (3-)6(-8), and (4-)
6(-9). On the other hand they agree remarkably well with the counts of annulus
cells in paraphyses-bearing plants given by Martens (1950) and Fernandes (1968).

From these results it is clear that the range of this character in P. australe
completely overlaps the range in P. interjectum, as may be seen from the histo-
grams of the number of annulus cells in the three taxa (Fig. 2). The modal class
in many of the diploid plants is as high as that found in those of the hexaploid,
and in one instance higher. Indeed, in some hexaploid plants the average
number of cells in the annulus is as low as 7, as was also found by Shivas (1961).

It is noteworthy, too, that large differences in the number of annulus cells
are found between plants of P. australe in the same locality and clearly belonging
to the same breeding population. For instance, the four plants from the Little
Orme, Caernarvonshire, included in Table 1, were all found growing on the
limestone outcrop within 120 yards of each other. The distributions of the
number of annulus cells in three of the Little Orme plants are shown in Fig. 3.
That of the fourth plant from this locality is nearly identical with one of them
and is therefore omitted. A similar distribution of this character was found in
most of the plants included in Table 1; in only four of them was the skewness

OS)
ee
35 ate
yoo
| |
ae ial!
t ' |
30 if ot Lae: Qn
pia ol
' 1 5 SF Soe = 4n
t : | |
eae ee ee ee 6n
: :
SOS : | |
< : i.
© t 1 I |
za I 1
< : S25) E
« 20 ioe
© ! J | :
a 1 |
“
re) ' t |
!
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a ' ' l
! ' °
= ; |
= H :
10 : =
:
'
'
J

2 3.4.8.6 7 8 910 11 12 13 14 1S anemia
NUMBER OF ANNULUS CELLS

FIGURE 2. Histograms of the number of annulus cells in the three
species of Polypodium. (Based on 9,900 sporangia of P. australe
and 1,000 sporangia each of P. vulgare and P. interjectum).

PLATE 1. (a) Meiotic chromosomes in two spore-mother-cells of P. australe from Llanddulas,
Denbs., showing 37 bivalents. (b) Flattened annuli from the same plant as (a), with 6-12 indu-
rated cells (appearing dark). (c) Flattened annulus of P. australe from the Little Orme, Caerns.,
with 16 indurated cells. (d) A single annulus from the same sample as (b), with 12 indurated cells
and showing one of the characteristic paraphyses. (e) Flattened annuli of P. interjectum from
Anglesey, shown at the same magnification as (b), for comparison.

a
fy
ey
F
3

PLATE 2. Rhizome scales of (a) P. australe from the Little Orme, Caerns.; (b) P. interjectum
from Beaumaris, Anglesey. Both at the same magnification.

TAXONOMIC CHARACTERS OF POLYPODIUM AUSTRALE FEE 125
180

160

140

120

100

80

60

NUMBER OF SPORANGIA

Pe@eeee we wae a= =

40

20

ieee A 5G 7 68 O10. IT 12” 13 14 Sy 16
NUMBER OF ANNULUS CELLS

FIGURE 3. Histograms of the number of indurated cells of the
annulus in three of the plants of P. australe from the Little Orme,
Caerns. Numbers as in Table 1.

more markedly positive. The annulus cells in some of the Conway plants had
been counted when they were first gathered several years ago. When these counts
were repeated in 1968 they showed no significant differences from the previous
ones, although the length of the fronds had increased in cultivation from around
12 cm to 28 cm or more.

Other sporangium characters, however, have proved to be of considerable
taxonomic value. Lenski (1964) found a constant difference between the tetra-
ploid and the other two species in the number and size of the ‘basal cells.’
According to her, both the diploid and the hexaploid have usually from two
to four large cells separating the sporangium stalk from the annulus; the tetra-
ploid has none or one (rarely two), which, when present, is smaller. Counts
made from the material used in the present study (Fig. 4) agree well with those
of Lenski. P. vulgare has almost invariably only one small basal cell, which is
usually as wide as, or very little wider than the annulus. P. interjectum has
mostly 2 or 3 and P. australe 3 or 4, and in both of these the basal cells are
much wider than the annulus, as can be seen from the photomicrographs of
dehisced sporangia (Plate 1b and e). This character is thus absolutely diagnostic
for P. vulgare, but is only occasionally of use to distinguish between P. australe
and P. interjectum. The figures of sporangia given by Rothmaler & Schneider
(1962) are incorrect regarding this detail.

The difference between the diploid and hexaploid in the appearance of the

126 R. H. ROBERTS

NUMBER OF SPORANGIA

0) ee 2 ss ARS
NUMBER OF BASAL CELLS

FIGURE 4. Histograms of the number of basal cells of the
sporangium in the three species of Polypodium.

flattened annulus has already been mentioned. This can only be observed when
completely dehisced sporangia are subjected to moderate finger pressure
between two glass slides. Many of the annuli will then be flattened, i.e. spread
out to their full length. The indurated cells of P. australe are shorter (mean
from ten annuli 21—26um) and broader (mean of five annuli 81-100um), thus
giving the annulus a distinctive, contracted appearance compared with that of
P. interjectum. In the latter the indurated cells are longer (mean from ten annuli
28-35um) and the annulus narrower (mean of five annuli 76-86um) (Benoit
1966, Roberts 1966). This difference can be appreciated by a glance at the
photomicrographs of the flattened annuli (Plate 1b and e).

Lastly, the three species differ in the colour of the annulus, a character to
which Benoit (1966) first drew attention as the best non-microscopic character
for separating P. vulgare (‘red-brown’) from P. interjectum (‘pale golden brown’).
In P. australe the colour of the annulus, seen under the microscope, is usually
bright yellow and shows little variation. In P. interjectum, on the other hand, it
varies considerably, not only from one plant to another, but even among the
sporangia on the same plant. Usually a pale buff or golden brown (when it
hardly differs in colour from the rest of the sporangium), it varies to pale or
bright yellow, and occasionally to a deeper orange-yellow. This variability is
perhaps the most striking feature of annulus colour in the hexaploid and is in
itself often a useful diagnostic character. Thus, despite its variability in the
hexaploid, annulus colour can be of great value in separating the diploid from
the hexaploid. In P. vulgare, as Benoit (1966) has indicated, the annulus is a
dark reddish-brown, which contrasts so sharply with the rest of the sporangium
that it can be seen clearly as a dark line even with a x 10 lens.

TAXONOMIC CHARACTERS OF POL YPODIUM AUSTRALE FEE oy.

PARAPHYSES

Martens (1950) first drew attention to the value of the branched paraphyses
as a diagnostic character of P. australe. Shivas (1961) also found paraphyses
of the type illustrated by Martens in all the plants which she determined cyto-
logically as diploid. Lenski (1962, 1964), however, stated that paraphyses were
present in 35 out of 50 hexaploid plants and even in 10 out of 67 tetraploid
plants examined by her. Meinders-Groeneveld & Segal (1967) have also
reported paraphyses in the hexaploid in the Netherlands, but failed to find
them in the tetraploid.

In view of these conflicting reports, a careful search was made among the
sporangia of the cytologically attested plants of P. interjectum. These were
supplemented by a further eighty plants not determined cytologically, making
a total of one hundred. No paraphyses similar to those of P. australe were found
in any of them. In many instances (38 of the 100 plants) the minute, glandular
hairs which occur scattered on the lower surface of the frond in all the European
species of Polypodium, were found either among the sporangia or close to the
base of the sorus. In rare instances they were even found attached to the base
of the sporangium stalk. A search of the sori of 100 plants of P. vulgare gave
a similar result. However, these organs are quite distinct from the paraphyses
of P. australe. They are much smaller—usually only 3 or 4 cells long—with
their glandular, inflated, terminal cells mostly coloured a dark reddish-brown,
and as long as the other cells: from 45 to 80um. They are, in fact, identical
with the illustrations of the ‘paraphyses’ of P. interjectum given by Lenski
(1964). In the paraphyses of P. australe the terminal cells are invariably much
shorter than those at the base, which are mostly from 100um to 240um long.
In their illustrations of the paraphyses of P. interjectum, Meinders-Groeneveld
& Segal (1967) appear to show both kinds: a few (Fig. 5a-c) are somewhat
similar to those of P. australe; the others (Fig. 5d-i) are exactly like the small,
glandular hairs described above. It is not clear what criterion was used by these

NUMBER OF SAMPLES

TS) rn) ay peepee are ama ee
OFS Fal Om "20- 30- 40- "50-60 60— 70- 80— 90- 100- 110— 120- 130- 140-— eisoen 160- 170- 180-
SRO 82.9. SINEAD SI CO WSS SON IOD ENO) 1295139) VAI 159) 169) N79 189

LENGTH (IN UNITS OF 10~zm)

Ficure 5. Histograms, in 10um units, of the length of (a) the glandular
hairs of P. interjectum, and (b) the paraphyses of P. australe.

128 R. H. ROBERTS

FIGURE 6. Paraphyses of the hybrid P. australe x interjectum
(a—b); of P. australe (c—e); the glandular hairs (‘paraphyses’)
of P. interjectum (f—j). All to the same scale.

authors to identify the plants in which they found the larger, branched para-
physes. If they relied solely on the number of annulus cells, it would now
appear desirable that these plants should be examined cytologically.

The large difference in length between the glandular hairs of P. interjectum
and the paraphyses of P. australe is shown in the histograms (Fig. 5), and the
difference in their morphology in the figures (Fig. 6) drawn to the same scale.
The mean length of the paraphyses of P. australe (970 um) is nearly seven times
that of the glandular hairs of P. interjectum (140 »m). Because of their large
size and very distinctive morphology, together with the relative ease with which
they can be found, the paraphyses of P. australe remain its best single diagnostic
character, as was originally claimed by Martens (1950).

FROND SHAPE

The shape of the frond in P. australe is described as ovate to triangular-ovate,
with narrow, acute and usually dentate pinnae. Shivas (1961) states that the
longest pair of pinnae is the second or third from the base.

However, both the diploid and hexaploid are extremely variable in frond
shape. The ratio of length to breadth of the blade varies from 1-2 to 2-4 (mean
1-6) in the diploid plants, and from 1-5 to 3-2 (mean 2:1) in the hexaploid plants.
The ratio of length to breadth of the longest pinna on each frond was found to
vary from 3-0 to 5-9 in the diploid, and from 2:3 to 4:3 in the hexaploid. In
both these characters there is thus a large overlap between these two species.
The variation in the ratio of length to breadth of the blade seems to be partly
genetic and partly environmental. Poorly grown specimens generally have lower
values of this ratio in both species. Thus this character is of least help with the
plants most likely to be confused.

Pinna serrations in the diploid plants also vary widely, as Shivas (1961) has

TAXONOMIC CHARACTERS OF POLYPODIUM AUSTRALE FEE 129

already observed. They are entirely lacking in two of the twenty-eight plants in
Table 1, and so slight as to be scarcely noticeable in another three of them. The
well-marked serrations usually shown in illustrations of P. australe (e.g. Hyde &
Wade 1962) occur in only eight of these plants.

A frond character of the diploid which is often very striking in the field is
the tendency for the lowest pair or several pairs of pinnae to be inflexed (Manton
1950), often with the lowest pair crossing. But even this is entirely lacking in
some plants. This characteristic remains constant in cultivation, suggesting that
it is genetically controlled and not the result of environmental factors.

The silhouettes of fronds (Fig. 7), taken from six of the diploid plants, illus-
trate the range of variation in almost every detail of frond shape which occurs
in this species.

The number of bifurcations of the secondary veins has been stated to be

FiGurE 7. Silhouettes of fronds from six of the plants of P.
australe listed in Table 1.

130 R. H. ROBERTS

taxonomically useful, for example, by Rothmaler & Schneider (1962). But
the variation in this character in both the diploid and the hexaploid plants is so
great that it is evidently of very little use for separating these two species.

THE RHIZOME SCALES

The length and shape of the rhizome scales have been studied by Rothmaler |

& Schneider (1962). According to them the scales are 3-7mm long in P.
vulgare, 3—6mm long in P. interjectum, and 5-1lmm long in P. australe.
Valentine (1964) cites this as a key character with only slight modification:
36mm in both P. vulgare and P. interjectum, and 5—11mm in P. australe.

Measurements of rhizome scales were made from the twenty plants of P.
interjectum determined cytologically and from the plants of P. australe listed in
Table 1. All measurements were made under the microscope, the width being
taken from the widest part of the scale near its base. The results are shown in the
scatter diagram (Fig. 8), but for convenience the measurements of scales less
than 4-5 mm long have been omitted. The ranges of scale length found were:
3-5-11 mm for P. interjectum, 5-16 mm for P. australe. Only twenty-five scales
of P. vulgare were measured. In these the length was 3-6 mm, which agrees
with the values given by the other authors; but the ranges found for P. inter-
jectum and P. australe differ a good deal from their results, especially in the
case of the hexaploid. The overlap is so much greater than they have indicated
that the taxonomic value of scale length on its own is considerably reduced.

On the other hand the shape of the rhizome scales appears to provide a useful
accessory diagnostic character, although it is not as absolute as the illustration
given by Rothmaler & Schneider (1962) indicates. In P. australe the scales
are mostly linear-lanceolate. In P. interjectum they are often rather abruptly
narrowed just above the dilated and relatively broader base, as can be seen in
Plate 2. The difference between the two species in the ratio of the length to the
width of the scales is evident from Fig. 8.

Two other differences between the rhizome scales of the diploid and hexaploid
were noticed. Those of P. australe are thinner and have appreciably shorter
cells than those of P. interjectum. The means of the lengths of ten cells (measured
near the middle of the scale) vary from 1654m to 200um in P. australe, from
230um to 320um in P. interjectum. These results suggest that it may be possible
to separate the two species on this character alone, but further observations
are needed to verify this.

HYBRIDS

Wild hybrids of P. australe with both P. vulgare and P. interjectum have been
reported by Manton (1950, 1961) and Shivas (1961), and confirmed cyto-
logically. Perring & Sell (1967) state that these hybrids have also been recog-
nized on morphological grounds. On several occasions during the present study
plants have been collected which on frond characters were suspected to be
hybrids of P. australe with one of the other species. However, cytological
examination has shown them to be only P. australe. Three of them are among
those listed in Table 1.

So far only one plant has been found which appears to be a hybrid involving
P. australe. This is very conspicuous in the field because it forms a large, leafy

TAXONOMIC CHARACTERS OF POL YPODIUM AUSTRALE FEE 131

mass in a locality (Lligwy Wood, Anglesey) where both P. australe and P.
interjectum occur in abundance. Its robust growth, vigorous vegetative spread
and large fronds (many of them over 60 cm long) suggest hybrid vigour. In
frond shape it more closely resembles P. interjectum, but the blade is thinner
and its colour lighter than is usual in the hexaploid. Microscopic examination
showed its spores to be highly sterile, and the few (usually 3 or 4) fully-developed
spores present in some sporangia are uneven in shape and size. The mean
number of annulus cells is 7 with a total range of 4-9. Paraphyses as long as
those of P. australe are present (Fig. 6), but are not so branched nor as abundant
as in the diploid. This is contrary to the view expressed by Shivas (1961) that
paraphyses are absent from hybrids of P. australe.

Meiosis in this plant was examined on several occasions. It was found to be
highly irregular, but only approximate counts were obtained. These showed
about 32-36 bivalents and around 70 univalents. They agree fairly closely
with the counts of 37 bivalents and 74 univalents found during meiosis in the
tetraploid hybrid by Shivas (1961). Although further cytological examination
of this plant is desirable, the evidence certainly suggests that its parentage is
that inferred from its morphology.

DISCUSSION

The suggestion first made by Futo (1905) that P. australe (as P. vulgare var.
serratum Willd.) could be identified by the small number of annulus cells
(mostly 6) was reaffirmed by Manton (1950) and by Shivas (1961). Martens
(1950), however, found much higher numbers of annulus cells—from 4 to 20—
in plants which he identified as var. serratum by the large, branched paraphyses
among their sporangia; and similar values were obtained by Roberts (1966).
Three plants determined as diploid from root-tip counts by Lenski (1964) were
found to have correspondingly high numbers of annulus cells: from 4 to 17;
and a study of herbarium material by Fernandes (1968) has given equally high
counts.

Probably because both Manton (1950) and Shivas (1961) had made their
observations on cytologically determined material, most recent Flora writers
have accepted their evaluation of this character. For example, Hyde & Wade
(1962), Valentine (1964) and Warburg (1962) all cite a range of 4-7 annulus cells
for P. australe. Perring & Sell (1968) have also used this character for separating
the three European taxa. Furthermore, these authors state that specimens with an
‘intermediate’ number of annulus cells were also found to have a number of
aborted and empty spores. In view of the results presented here, it is clear that
most of the ‘intermediates’ referred to by Perring & Sell (1967, 1968) are not
hybrids involving P. australe, as these authors seem to have assumed. Some
degree of lowered spore fertility can often be observed in plants of all three
species of Polypodium, and is due to factors other than hybridity—probably
environmental conditions during the early stages of spore formation. However,
such plants usually have some of their sporangia with a full complement of
good spores. The symptoms of spore sterility in hybrids have been clearly
described by both Manton (1961) and Shivas (1961). Not only are the great
majority of the spores aborted, but the few full spores produced are highly
irregular in shape and size.

In view of the frequent occurrence of the pentaploid hybrid, P. vulgare x

132 R. H. ROBERTS

interjectum, the apparent rarity of hybrids involving P. australe is rather unex-
pected. Yet such appears to be the case. The diploid species reaches the northern
limit of its distribution in the British Isles, where it has been recorded as far
north as Westmorland (near Lowood, J. Sidebotham, July, 1846, MANCH),
Cringlebarrow in Lancashire (LANC) and on Ingleborough in Yorkshire
(Manton 1950). In Wales it occurs in small, isolated patches, most of which
grow on mortared walls, limestone, or other rocks with a high base content.
Only in rare instances has it been found on an apparently base-deficient sub-
stratum or as an epiphyte on trees. This is in agreement with Manton’s obser-
vations on its occurrence in limestone habitats in northern Europe. Its sporadic
distribution in Wales is in marked contrast to that of the other two species,
both of which are widely distributed and often abundant. For the survival of
such small populations of P. australe the formation of hybrids will, presumably,
be disadvantageous. They will tend to compete with the diploid parent in a
habitat which is already severely restricted. Under such conditions any genetic
barrier to hybridization would tend to be strengthened.

LENGTH (mm)

0:6. 08) 1:0 1-2, 14, 1-6, 1-8. 2:0. 2:2 24,265 2:8 )3'0) 3:2nese
WIDTH (mm)

Ficure 8. Scatter diagram of the length and width of the rhizome scales of
P. australe (solid dots) and of P. interjectum (open dots).

TAXONOMIC CHARACTERS OF POL YPODIUM AUSTRALE FEE 133

Perring & Sell (1967) have reduced the three cytotypes to the rank of sub-
species on the grounds that the presence of hybrids between them makes field
identification impossible. If ease of identification in the field is accepted as the
criterion of a good species, then this view is to a great extent justified, for the
large overlap between P. australe and P. interjectum makes it impossible in
many instances to separate them on frond characters. Nevertheless there are
several microscopic characters which enable P. australe to be identified with
certainty. From P. interjectum it can be separated by the large, distinctive
paraphyses, as well as by the colour and appearance of the flattened annulus,
and the overall difference in the shape of the rhizome scales. From P. vulgare
it can always be separated by the number of basal cells of the sporangium;
but confusion with this species is seldom likely in any case.

In spite of the much greater overlap between P. australe and P. interjectum
in some of the characters hitherto used to separate them, Shivas’s taxonomic
conclusions regarding the diploid are, in my opinion, not affected. The fact
remains that P. australe also differs from the other European taxa in chromo-
some number, in its ecological requirements and in its geographical range.
Moreover, Shivas (1961) has shown that it is not closely related to P. vulgare
sensu stricto. To place it as a subspecies of P. vulgare is to ignore the cytogenetic
evidence. While, therefore, there is no sound biological reason why the diploid
should be reduced to the rank of subspecies, this must remain, like the species
concept itself, a matter of opinion.

ACKNOWLEDGMENTS

Thanks are due to Dr W. S. Lacey, University College of North Wales, Bangor,
for reading the first draft of this note in manuscript and for some helpful
suggestions; to Mr W. Elfyn Hughes for information about some of the localities,
and to Mr W. Neville and Mr B. Gower for taking the photomicrographs used
in Plate 1.

REFERENCES

BENOIT, P. M. (1966). Some recent work in Wales on the Polypodium vulgare aggregate.
Br. Fern Gaz., 9: 277-282.

FARQUET, P. (1933). Materiaux pour servir a la connaissance du Polypodium vulgare et de ses
sous-espéces. Ber. schweiz. bot. Ges., 42: 484-506.

FERNANDES, R. B. (1968). O genero Polypodium L. em Portugal. Bolm. Soc. broteriana,
2a sér. 42: 35-158.

FuTo, M. (1905). Polypodium vulgare L. und Polypodium vulgare var. serratum Willd. Hedwigia,
44: 106-111.

Hype, H. A. & WADE, A. E. (1962). Welsh Ferns, 4th ed. Cardiff.

LENSKI, I. (1962). Nachweis von Paraphysen-tragenden Polypodien in Deutschland. Ber. dt.
bot. Ges., 75: 189-192.

LENSKI, I. (1964). Merkmalspriifungen an den europdischen Zytotypen von Polypodium
vulgare L. s.l. Flora, 154: 245-266.

MANTON, I. (1947). Polyploidy in Polypodium vulgare. Nature, Lond., 159: 136.

MANTON, I. (1950). Problems of Cytology and Evolution in the Pteridophyta. Cambridge.

MANTON, I. (1961). Evolution in the Pteridophyta, in WANSTALL, P. J.,ed. A Darwin Centenary.
London.

MARTENS, P. (1950). Les paraphyses de Polypodium vulgare et de la sous-espéce serratum.
Bull. Soc. r. Bot. Belg., 82: 225-262.

MEINDERS-GROENEVELD, J. & SEGAL, S. (1967). Pteridologische aantekeningen. 3. De eikvaren
(Polypodium vulgare L. sensu lato) in Nederland. Gorteria, 3: 183-199.

134 R. H. ROBERTS

PERRING, F. H. & SELL, P. D. (1967). Taxonomic and nomenclatural notes on the British flora.
Watsonia, 6: 292.

PERRING, F. H. & SELL, P. D., ed. (1968). Critical supplement to the Atlas of the British Flora.
London.

RosERTS, R. H. (1966). Some observations on Polypodium australe. Br. Fern Gaz., 9: 283-287.

ROTHMALER, W. (1929). Die Pteridophyten Thiiringens. Mitt. Thiir. Bot. Ver., 38: 92-118.

ROTHMALER, W. & SCHNEIDER, U. (1962). Die Gattung Polypodium in Europa. Die Kultur-
pflanze, Beih., 3: 234-248.

Suivas, M. G. (1961). Contributions to the cytology and taxonomy of species of Polypodium
in Europe and America, I: Cytology; II: Taxonomy. J. Linn. Soc. (Bot.), 58: 13-25, 27-38.

Suivas, M. G. (1962). The Polypodium vulgare complex. Br. Fern Gaz., 9: 65-70.

VALENTINE, D. H. (1964). Polypodiaceae, in TuTIN, T. G., et alia, ed. Flora Europaea, vol. 1,
p. 23. Cambridge.

WARBURG, E. F. (1962). Polypodiaceae, in CLAPHAM, A. R., TUTIN, T. G. & WARBURG, E. F.
Flora of the British Isles, 2nd ed., pp. 39-41. Cambridge.

WILLDENOW, C. L. (1810). Species Plantarum, 4th ed., vol. 5, part 1, p. 173. Berlin.

Watsonia, 8, 135-144 (1970). 135

Studies in Potentilla anserina L.
D. J. OCKENDON and S. M. WALTERS

Department of Botany, Imperial College, London, and Botany School, University
of Cambridge

ABSTRACT

The variation of Potentilla anserina L. in Britain was studied using material of known wild
origin. Both tetraploid and hexaploid cytodemes were found; the latter were much less common
but showed no geographical pattern and could not be distinguished with certainty on any
morphological character.

INTRODUCTION

Surprisingly little is known about the morphological, cytological and genetical
variation of a number of common British plants. In recent years a few such
species have been studied by the final-year class at the Botany School, Cam-
bridge University. These studies do not permit an exhaustive investigation of
the species concerned, but they often bring to light a number of interesting
facts which could form the basis of a more complete study. The work reported
here on Potentilla anserina L. was carried out during the summers of 1967 and
1968 by the final-year class and the authors. We are grateful to a number of
colleagues who helped to collect samples for cultivation.

The experimental taxonomic study of Potentilla anserina by Rousi (1965)
provided a very useful basis for our work, and should be consulted for further
information on the taxonomy, morphology and distribution of the species. Most
of the 51 samples which Rousi studied came from Europe and North America.
Three of these proved to be hexaploid (2n=42) and the rest tetraploid (2n=28).
The hexaploids had a low pollen fertility (about 20%) and failed to set seed,
while the tetraploids had a much higher pollen fertility (60-98 %) and produced
fertile seed. The only British plant studied by Rousi (from Nether Poppleton,
near York) was hexaploid and had irregular meiosis and low pollen fertility
(9-97):

Very few of the herbarium specimens of British P. anserina in the Cambridge
Herbarium (CGE) have fruit, and the scarcity of fruit in natural populations
has been observed by several botanists. This suggested to us that in Britain
the hexaploid might be commoner than the tetraploid, although the reverse
is true for the two areas (Finland and California) studied in detail by Rousi
(1965).

MATERIALS

The living plants used in this study are listed in Table 1. Most have been culti-
vated in the Cambridge University Botanic Garden; the remainder include two
large population samples from Fulbourn Fen and Wicken Fen, Cambridgeshire.

. J. OCKENDON AND S. M. WALTERS

136

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Botanical Society of the British Isles, and Thomas Nelson & Sons Ltd., taken from their

Atlas of the British Flora, and updated by the Biologica
Rousi (1965) considers a number of morphological characters

and reproductive, selecting particularly those characters wh
have used to distinguish taxa within the P

characters were stud

Ficure 1. Distribution of Potentilla anser
results follows.

138 D. J. OCKENDON AND S. M. WALTERS

1. LENGTH/BREADTH RATIO OF TERMINAL LEAFLET

Rousi found that this ratio ‘was lower in all three hexaploid plants than in
the related tetraploids from the same or neighbouring areas’. The figures for
the two populations sampled by us were:

Standard

No. of Standard Error of
plants Range Mean Deviation Mean
Fulbourn 73 1-4—3-5 2:02 0-41 0:048
Wicken S7/ 1-3-3-9 DoD 0:49 0:065

In each case a single leaf was measured. The range of values for plants in
cultivation (1-5-3-1, Table 2) is somewhat less than that found in a single
population sample. The hexaploids have values of 2-0 or below, which supports
Rousi’s finding, but, as many of the tetraploids also have values below 2:0,
this character alone cannot be used to distinguish the two ploidy levels.

2. NUMBER OF TEETH

The number of teeth on one side of the terminal leaflet, plus the terminal tooth,
was counted. The figures for the population samples were:—

Standard
No. of Standard Error of
plants Range Mean Deviation Mean
Fulbourn 36 9-16 12.97 1:30 0-22
Wicken 36 8-18 11-64 1-85 0-31

In each case a single leaf was used. Again the ranges of values for the population
samples are greater than the range of single values for the plants in cultivation
(Table 2), and there is no correlation between number of teeth and ploidy
level. However, the two lowest values (7 and 8) are on plants from native
localities in the western Scottish islands (plants nos. 25 and 24 respectively).

3. EPICALYX SEGMENTS
The following grading was adopted in order to measure the variation in the
toothing of the epicalyx segments:
Grade 0—all segments entire
Grade 1—some but not all segments toothed
Grade 2—all segments toothed or laciniate, but not exceeding the calyx
Grade 3—all segments laciniate, some or all exceeding the calyx

The results for the population samples were as follows :—

Total no.
of flowers Grade Mean Grade
0 1 Dp) 3
Fulbourn 10 0 1 9 0 1:90
Wicken 50 1 5 39 5 1-96

The plants in cultivation showed the same sort of range for this character as
the population samples, but the west Scottish and Lizard plants (25, 26, 28, 23)
showed low grades of 0 or 1, in contrast to the hexaploid from Yorkshire (19a)
and the tetraploid from Teesdale (16), both of which showed grade 3 (Table 2).

STUDIES IN POTENTILLA ANSERINA 139
TABLE 2. MORPHOLOGICAL CHARACTERS

In several cases, the sample cultivated consists of more than one original plant. Such
cases are indicated in the remaining tables by adding a, b, etc. to the reference number.

Terminal Leaflet

Plant Ref. re Epicalyx
No. Length/breadth No. of teeth Grade
ratio
1 271 — —
2 22 — —
6 2-4 oo —
7 1-5 — —
9a (6x) IZ — —
9b (6x) 1-6 — —
14 (6x) 2:0 = 1
15 2:1 13 2
16 2:0 12 8)
17 2°8 10 2
18 7 12 2
19a (6x) 1-6 12 3
19b (6x) 1-9 10 3
20 1-9 11 2
21a 2:0 12 —
21b 1-6 12 2
22a 2:4 11 0
22b 2°4 11 2
22C 3-1 — 2
22d UG 11 0
23 ZA 12 1
24 2-0 8 _
25 1-8 7 0
26 iS 10 1
27 7) _ —
28 I — 1

Further evidence of a difference in the epicalyx between coastal Scottish and
lowland English populations can be seen by comparing the following figures
scored in the field in August 1968 with those given above for the Wicken and
Fulbourn populations:

Total no.
of flowers Grade Mean Grade
0 1 2 3
Loch Linnhe,
Fort William 10 1 6 2 1 1-3
Lochaline,
Morvern, Argyll 10 0 8 2 0 2
North outskirts
of Oban (1) 10 4 6 0 0 0:6

North outskirts
of Oban (2) 10 0 6 4 0 1:4

140 D. J. OCKENDON AND S. M. WALTERS

A noteworthy feature of the coastal Scottish populations is that they approach
the arctic P. anserina subsp. egedii (Wormsk.) Hiit., which is characterised by
entire or shallowly toothed epicalyx segments and a low number of teeth on the
terminal leaflet. Plant no. 13 from northern Iceland is referable to this subspecies,
and plants from Shetland in the Cambridge Herbarium (CGE) are identifiable
as subsp. egedii using the diagnostic description provided by Rousi. Rousi has
found populations in Finland with a mixture of anserina and egedii characters,
a condition which is approached by some of the coastal Scottish populations.
The occurrence of such intermediate populations, and the ease of production
of artificial hybrids between subspp. ‘anserina’ and ‘egedii’ which Rousi found,
support his conclusion that the appropriate rank for these two taxa is that of
subspecies. If the variation between these two subspecies is more or less con-
tinuous, and has a geographical basis, then the two subspecies may be regarded
as the relatively extreme end-points of a topocline.

CHROMOSOME NUMBERS

P. anserina is not easy cytological material, and precise counts are difficult to
make. Approximate counts are usually sufficient for a determination of the
ploidy level, however, and as most of our counts are approximate the ploidy
level rather than the chromosome number is given in Table 1. All our counts
were made from squash preparations of root tips. Root tips of P. anserina can
easily be obtained by allowing runners to root in water. The root tips were
pre-treated with a-bromonaphthalene for 2 hours, fixed in Carnoy’s solution
and stained in alcoholic hydrochloric acid carmine (Snow 1963).

Four of the 33 samples which have been counted are hexaploids, the rest being
tetraploid (Table 1, Fig. 1). One of the hexaploid samples (19) is from the
same place as Rousi’s British hexaploid. The other three hexaploids are from
Great Cumbrae, Bute (9), Lleyn Peninsula, N. Wales (14) and Burwell, Cam-
bridgeshire (30). There is thus no clear pattern in the distribution of the two
ploidy levels in Britain. This agrees with Rousi’s finding that his 3 hexaploids
were not confined to any one area.

In the relatively few instances where more than one plant per sample was
counted, there was no evidence for mixed populations of tetraploids and
hexaploids. |

POLLEN FERTILITY

Pollen fertility was estimated by counting 100-200 pollen grains stained with
cotton blue in lactophenol. In contrast to Rousi’s results we have found that
the pollen fertility of a single plant varies at different times of the year (Table 3),
and that the fertility of plants in a population differs at any one time (Table 4).
Because of variation in pollen fertility during the flowering season, it is impor-
tant to note the date when the pollen fertility is determined. The 3 counts made
for plant 25 (Table 3) suggest that the pollen fertility falls off towards the end
of the flowering season. Nevertheless, counts made late in the season may still
show a high fertility (e.g. 97% for plant no. 24 on 23/7/68).

The range of pollen fertility in single populations is unusually large (Table 4)
and may result from individual plants losing their pollen fertility at different
rates. The difference in average pollen fertility between the Fulbourn Fen and

STUDIES IN POTENTILLA ANSERINA 141
TABLE 3. POLLEN FERTILITY (% OF ‘GOOD’ GRAINS)

Plant Ref.

No. Date Fertility Date Fertility Date Fertility
la July 67 45 27/5/68 92
1b July 67 86 27/5/68 99
2a July 67 80 27/5/68 ST
2b July 67 86

2c July 67 35

3 July 67 64 27/5/68 97
4 July 67 90 27/5/68 92
5 July 67 27/5/68 48
6a July 67 76 27/5/68 75
6b July 67 62

7 July 67 62

8 July 67 30 27/5/68 35
9a July 67 28 27/5/68 14
9c July 67 11

10a July 67 45 27/5/68 58
10b July 67 47 27/5/68 50
10c July 67 47 27/5/68 35
11 July 67 58 July 67 87 27/5/68 93
12 July 67 86

13 July 67 32 27/5/68 79
14 6/7/68 11

15 6/7/68 98

16 6/7/68 95

17 28/6/68 39

18 27/5/68 26

19a 26/7/68 2

19b 15/8/68 10

20 28/6/68 80

21a 27/5/68 86

21b 28/6/68 98

22a 6/7/68 98

22b 28/6/68 97

22c 6/7/68 85

22d 28/6/68 98

22e 9/7/68 97

23 6/7/68 97

24 23/7/68 97

25 6/7/68 97 19/7/68 83 26/7/68 68

26 6/7/68 97

28 9/7/68 97 19/7/68 90

TABLE 4. VARIATION IN POLLEN FERTILITY IN POPULATIONS SAMPLED

No. plants Range of Average

Population Date sampled fertility fertility
Ditton Park Wood, Cambs. 31/7/67 12 37-94% 19%
Fulbourn Fen, Cambs. 25/7/68 35 20-97% 53%

Wicken Fen, Cambs. 25/7/68 30 41-99% 82%

142 D. J. OCKENDON AND S. M. WALTERS

Wicken Fen populations is also striking, and indicates that there is variation
between populations in this character, as well as within populations.

This variation in pollen fertility in P. anserina tends to obscure the clear-cut
difference between the pollen fertility of tetraploids and hexaploids found by
Rousi (1965). The pollen fertility of our hexaploids is 2-28 %, while the lowest
value recorded for the tetraploids is 26% (plant no. 18). Plants with pollen
fertilities of 25-35% could be tetraploid or hexaploid and this is not a safe
way of determining the ploidy level unless several pollen fertility counts are
made at different times of the year. The majority of the tetraploids have a
maximum pollen fertility above 80%, but some plants (e.g. 8 and 10) have a
consistently low pollen fertility, indicating a marked degree of male sterility.
Rousi (1965) records similar male sterility in one of the populations which he
studied.

FRUIT PRODUCTION

Almost all the tetraploid plants in cultivation which have flowered have also
produced fruit (Table 1). Of the hexaploids, both plants from Nether Poppleton
(19a and b) have not produced fruit, but all three plants from Great Cumbrae
(9a—c) and the single plant from the Lleyn Peninsula (14) have fruited. Fruit
production by these hexaploids is unexpected and needs to be further investi-
gated. Rousi mentioned the possibility that this might occur, although he did
not publish any evidence for it.

Rousi selfed a number of plants and found that they were all self-incompatible.
We have found the same results with the 11 plants (10 tetraploids, 1 hexaploid)
which we have tested. Each of these plants has fruited in open cultivation, and a
cross between plants 15 and 16, made at the same time as the self-pollinations,
has produced 10 achenes. It is reasonable to conclude that the failure of fruit
production following self-pollination results from self-incompatibility rather
than sterility.

MODES OF REPRODUCTION

P. anserina is capable of vigorous vegetative spread, and the difficulty of finding
fruiting material in the field raises the question of the relative importance of
sexual and asexual modes of reproduction under natural conditions. In both
the Wicken and Fulbourn Fen populations fruiting heads were found after a
careful search, but could easily have been overlooked. At Wicken, P. anserina
was found growing on a track, in tall grass near the track, and at the margin
of the track, but it was only at the margin that any quantity of fruit was found.
Most of the plants grown in pots in Cambridge fruited well during the summer
of 1968, but those grown in beds fruited poorly in 1968, although they were
vegetatively very vigorous that year and had fruited the previous year. The
plants in pots also flowered much more freely than those in the beds. These
facts suggest the possibility that sexual reproduction is much reduced in plants
which are growing very vigorously (and reproducing asexually) because of poor
flower production. P. anserina does not appear to have a very definite flowering

STUDIES IN POTENTILLA ANSERINA 143

period in Cambridge, and flowers may be produced sporadically from May to
September. Flowering in the wild is also often rather poor, less than about
10% of the plants being in flower at any one time, and those that are flowering
having only one or two flowers open at once.

In addition to poor flowering, low fertility of the flowers which are produced
might also reduce fruit production. It is of interest that fruits were easier to
find in the Wicken population than in the Fulbourn population, and that the latter
had a much lower average pollen fertility. However, plants with consistently
low pollen fertilities (such as the three plants from St Catherine’s, Loch Fyne,
Argyll (10a-c)) have produced good fruit, and there is no evidence that female
sterility is associated with male sterility.

The achenes produced by the plants in cultivation are fertile, and up to 80%
germination has been obtained using the method suggested by Rousi (1965).
Evidence for the successful sexual reproduction of P. anserina in the field is
scanty, but seedlings were found in a damp meadow at Soham, Cambridgeshire
in July 1968. The seedlings occurred in a small compact group, suggesting that
they had grown from the achenes of a single fruiting head.

The precise rate of vegetative spread of P. anserina is not known, but runners
up to 1m in length may be produced during one growing season. Vegetative
reproduction is undoubtedly very important in producing the dense, and some-
times quite extensive, patches of P. anserina which are commonly seen in the
field. Although long-distance dispersal of vegetative parts may occur, it is likely
that sexual reproduction is more important than asexual for the establishment
of new populations.

DISCUSSION

The three morphological characters investigated all vary somewhat on different
parts of the same plant and may vary widely within a single population. This
severely limits their use as simple taxonomic characters and suggests that
statistical methods should be applied if these characters are to be studied in
more detail. It is clearly important to have sizeable population samples with
which to compare the data collected from individual plants from different
localities. There are no clear discontinuities in the morphological variation of
P. anserina in Britain, and the high variability of single populations tends to
obscure the geographical pattern of the variation.

The low and variable pollen fertility of P. anserina is unusual for a species
which reproduces sexually and is not agamospermous. Some other species of
Potentilla (e.g. P. tabernaemontani) are pseudogamous (Smith 1963), but there
is no evidence of this in P. anserina. The low pollen fertility could be interpreted
as indicating an increasing emphasis on vegetative reproduction as opposed
to sexual reproduction. It is possible that mutations leading to reduced male
fertility also give increased vegetative vigour and hence are not selected against
as strongly as they would be in species which relied on sexual reproduction
alone.

The distribution of the hexaploids in Britain does not show any obvious
pattern (cf Fig. 1), and this agrees with Rousi’s suggestion that hexaploids have
had a polytopic origin from the tetraploids. Their reproduction is almost entirely
by vegetative propagation and their high sterility probably prevents them from
spreading far from their source of origin.

144 D. J. OCKENDON AND S. M. WALTERS
CONCLUSION

Both hexaploid and tetraploid plants of P. anserina occur in Britain, the latter
being much the commoner. The two cytodemes cannot be distinguished with
absolute certainty by any character other than the chromosome number.
However, plants exhibiting all or most of the following characteristics are
likely to be hexaploid: low pollen fertility, poor achene production, relatively
broad terminal leaflets, high tooth number, and laciniate epicalyx segments.
Rousi’s single sample of P. anserina from Britain was atypical in being hexa-
ploid and could give a misleading impression of the species in this country. This
illustrates the danger of making generalisations based on insufficient evidence.
Much remains to be discovered about the variation of P. anserina in Britain,
and more information about its ecology and mode of reproduction in the field
is highly desirable. In particular, coastal populations in natural habitats in the
north and west could be much more effectively sampled to see to what extent
a topocline between subsp. anserina and subsp. egedii exists in Britain.

REFERENCES

Rous, A. (1965). Biosystematic studies on the species aggregate Potentilla anserina L. Ann.
Bot. Fenn., 2: 47-112.

SMITH, G. L. (1963). Studies in Potentilla L. I. Embryological investigations into the mechanism
of agamospermy in British P. tabernaemontani Aschers. New Phytol., 62: 264-282.
SNow, R. (1963). Alcoholic hydrochloric acid carmine as a stain for chromosomes in squash

preparations. Stain Technol., 38: 9-13.

Watsonia, 8, 145-153 (1970). 145

Plant records from Mull and the adjacent small islands. 2
J. F. M. CANNON and E. B. BANGERTER

Department of Botany, British Museum (Natural History)

ABSTRACT

References are made to additional early publications on the flora of the area. Herbarium and
field work subsequent to the earlier paper (Cannon & Bangerter 1968) is summarised. Some
species formerly known in the area only from Iona are recorded from localities on the mainland
of Mull, and significant extensions of range of some alpine species are noted. A systematic
list of the more interesting new records is given, including thirty-six new vice-county records.

INTRODUCTION

This paper reports the new and interesting records of vascular plants that have
resulted from fieldwork during 1968 undertaken by our Department and our
collaborators in the Mull Survey Project, details of which may be found in our
previous paper (Cannon & Bangerter 1968). Further research into the literature
and specimens preserved in this and other herbaria has also provided additional
significant information.

We have already given a brief account of the relevant botanical literature
from the time of John Ray to the present day. Among additional early publica-
tions the following have been found to contain records from our area. J.
Walker (1808) gives a list of plants found on Iona in his Essays on Natural
History and Rural Economy; this list antedates at least one published vice-
county record by over a hundred years. W. J. Hooker (1821) gives Mull localities
for a number of species in his Flora Scotica. W. Keddie (1850), Lecturer on
Natural Sciences in the Free Church College, Glasgow, in an appendix entitled
‘Botany of Iona and Staffa’ to his Staffa and Iona Described and Illustrated. . .
gives an extensive list including some cryptogams; unfortunately, he only rarely
indicates whether the species recorded were from Iona, Staffa or both islands.
The volumes of the third edition of English Botany edited by J. T. I. Boswell Syme
contain a number of records from Mull, some of which are based on his own
visit to the island in 1848 and these are mostly substantiated by specimens in his
herbarium, which has recently been incorporated into the British Museum
herbarium. We have been able to add a few records for Sorbus and Mentha to
those indicated in the recently published Critical Supplement to the Atlas of the
British Flora (Perring & Sell 1968).

In the course of an exhaustive search during a visit to the herbaria at the
Royal Botanic Garden, Edinburgh, and the Department of Botany, University
of Glasgow, we found a number of Mull specimens, some of which were of
great interest. In his Glasgow Catalogue, Ewing (1892, 1899) indicates the exist-
ence of specimens to support some of his records for v.c. 103. We had hoped
to trace many of these in Ewing’s herbarium at Glasgow, but we were only

D

146 J. F. M. CANNON AND E. B. BANGERTER

able to find a few. We are grateful to the authorities of these institutions for
their hospitality and generous co-operation. We should particularly like to thank
Mr Allan Stirling for generous help in searching the Glasgow herbarium.

Through the courtesy of Dr A. J. Richards, who organised a search of the
herbarium at the Department of Botany, University of Oxford, we were able
to examine nine specimens collected by Mr John Chapple during the 1939
B.E.C. Excursion to Mull, reported on by Wilmott (1942). The whereabouts
of the other voucher specimens from this excursion still remains a mystery as
Mr Chapple, who led the party, can only suggest in correspondence with us
that they should be at Oxford.

During 1968, the following fieldwork additional to that of our own parties
was undertaken. An excursion organised by the Committee for the Study of
the Scottish Flora and led by Mr A. A. Slack visited the island for one week
in July. Dr C. A. Stace of the Department of Botany, University of Manchester,
led a party of students during August. Mr Michael Mullin worked on the
island for six weeks under the vacation student scheme sponsored by this
Museum. Miss Ursula K. Duncan continued her very close co-operation with
the survey. Our list also contains records made in 1967 by Mrs Joan Duncan
and in 1968 by Mr Hugh McAllister of the University of Glasgow, and by
Dr R. Watling of the Royal Botanic Garden, Edinburgh, who is a major con-
tributor to the mycological work of the project.

Our parties, which were recording mainly vascular plants, were in the field
for two periods, each of a fortnight, during one of which visits were made to
the botanically almost unknown Treshnish Isles, lying between Mull and the
islands of Coll and Tiree. Through the courtesy of the Royal Navy, a helicopter
was placed at our disposal, greatly facilitating access to these small islands,
which are set in seas notorious for their changeable and unpredictable character.

From the species listed in Cannon and Bangerter (1968) as apparently
restricted to Iona, the following can now be deleted as they have been found
elsewhere in the area: Allium vineale, Brassica napus, Bromus sterilis, Carduus
acanthoides, Centranthus ruber, Fumaria bastardii, Papaver dubium, Raphanus
raphanistrum, Sedum telephium.

During the 1968 season, work in the field has revealed some significant
extensions to the distributions of some of the montane species in the Mull
flora. Visits by our own staff to the mountains on the north-east side of Glen
Forsa (Dun da Ghaoithe, Mainnir nam Fiadh and Beinn Bheag), by Dr C. A.
Stace and party (Sgurr Dearg, etc.), by the C.S.S.F. party (Dun da Ghaoithe)
and by Mr M. Mullin (Beinn Chreagach Mhor and Beinn Chreagach Bheag),
have demonstrated the presence on this eastern side of the island of the following
species, hitherto recorded only from the west and central parts of the main
massif, with some extensions to the mountains around Loch Buie: Alchemilla
alpina, Cardaminopsis petraea, Carex bigelowii, Cherleria sedoides, Epilobium
alsinifolium, Juncus triglumis, Luzula spicata, Lycopodium alpinum, L. clavatum,
Oxyria digyna, Salix herbacea, Saussurea alpina, Saxifraga hypnoides, S.
stellaris and Thalictrum alpinum. A visit to Cnoc an da Chinn above Achleck
by a museum party resulted in the first records of plants in this montane group
from the northern half of the island. In this locality three species, Lycopodium
alpinum, Salix herbacea and Thalictrum alpinum, were found during the course
of a short and rather hurried exploration. It is clear that there is still plenty to
be learned about the distribution of mountain plants in Mull.

PLANT RECORDS FROM MULL 147
SYSTEMATIC. LIST

In the following list, which follows the order of Dandy (1958), the British
Museum (Natural History) Department of Botany Mull Survey is referred to
as BMMS. Specimens in the British Museum Herbarium are indicated by
(BM) or by BMMS and number. Specimens in the herbarium of the Royal
Botanic Garden, Edinburgh are indicated by (E) and those in the herbarium
of the Department of Botany, University of Glasgow by (GL). Localities unless
otherwise stated are on the mainland of Mull. Further details (e.g. National
grid references) are available in most instances in our files.

The following members of the B.M. Survey are referred to in the list:
Lorna F. Bowden (now Mrs L. F. Ferguson), J. F. M. Cannon, A. Eddy,
I. K. Ferguson, E. W. Groves, P. W. James, A. C. Jermy, A. Melderis, Theresa
E. Parker, N. K. B. Robson, Patricia A. Sims and A. R. Vickery.

Thanks are due to the Biological Records Centre and to Mr D. H. Kent
for help in the checking of potential new vice-county records.

1/3. LYCOPODIUM ANNOTINUM L. We are indebted to Dr Dennis Ratcliffe for
clarification of the record from the Laggan Deer Forest. The reference to Diphasium
alpinum (Lycopodium alpinum) in the account of the 1961 B.S.B.I. Field Meeting,
to which we drew attention in our previous paper, has now been explained as an error
for Lycopodium annotinum. The record for the latter species published in Proc. bot.
Soc. Br. Isl. (1962) 4: 419 is correct.

15/6. ASPLENIUM VIRIDE Huds. Noted in Appendix II of the Atlas as a probable
error for v.c. 103. A specimen in the Edinburgh herbarium provides a firm record:
Iona, nunnery, 1906, R. C. DAVIE (E). New vice-county record.

45/1. CLEMATIS VITALBA L. In a hedgerow at Achnadrish, near Dervaig, 1967,
BMMS 333 (JERMY). New vice-county record; no doubt originally as an introduction.

58/1. PAPAVER RHOEAS L. Accepted by the AZlas for v.c. 103 but not plotted.
North central Iona, in potato field, 1968, MULLIN.

58/2. PAPAVER DUBIUM L. Previously known from Iona but now recorded from
the mainland of Mull, at three piaces in or around Tobermory, 1967, BMMS
(MELDERIS).

58/6. PAPAVER SOMNIFERUM L. Salen, waste ground between village and pier, 1966,
U. K. DUNCAN;; north of Fishnish, on rubbish dump, 1967, A. G. KENNETH
(BM). New vice-county record.

64/f. DICENTRA FORMOSA (Haw.) Walp. Established in a wood at Kilfinichen,
1968, BMMS 2287 (U. K. DUNCAN & JAMES). New vice-county record.

74/3. RAPHANUS SATIVUS L. Rubbish tip 150 yards west of Carsaig Bay, 1967,
BMMS 343 (BOWDEN). New vice-county record.

85/1. TEESDALIA NUDICAULIS (L.) R.Br. Previously known only from Coire nan
Each on the south side of Ben More, recorded by Wilmott (1942). Now recorded from
the lower end of Glen Forsa (GR.17/6040) 1968, MULLIN.

98/3. BARBAREA INTERMEDIA Bor. Three records from rubbish dumps around
Tobermory, 1968, BMMS 2307 (U. K. DUNCAN & JAMES); D. KINGSTON s.z.

148 J. F. M. CANNON AND E. B. BANGERTER

(BM) on C.S.S.F. excursion; BMMS 3144 d. K. FERGUSON). New vice-county
record.

100/4. ARABIS HIRSUTA (L.) Scop. Recorded without locality by Ross (1877) and
during the Maps Scheme from Iona. Now recorded from the mainland of Mull:
Gribun, 1961, B. W. RIBBONS; Cnoc Mor near Uisken, 1968, MULLIN.

[113/11. VioLA LUTEA Huds. W. Keddie (1850) included this species in his list of
plants from Staffa and Iona; G. Ross noted it in his 1877 and 1878 lists, localising it as
Calgary Bay in the latter. Wilmott (1942) says ‘would be N.C.R., but the f. curtisii
(Forst.) Drabble may have been intended’. Gerrans (1960) records this species from
the grounds of Glen Forsa House, but the specimen, Gerrans 606 (BM), has been
re-identified as V. tricolor. We have been able to examine a specimen collected by
J. A. C. Chapple during the 1939 B.E.C. excursion which has been confirmed by
R. D. Meikle as V. tricolor subsp. curtisii. It seems very probable that this specimen
was the ultimate origin of Wilmott’s information, although only known to him
through ‘one or other of the lists contributed’. In the light of these observations and
bearing in mind the number of new records of V. tricolor, it seems unlikely that V. lutea
occurs in our area. ]

115/4. HYPERICUM CALYCINUM. L. Iona, Sandeels Bay, 1906, R. C. DAVIE (&),
conf. Robson. New vice-county record. There is also a Distribution Maps Scheme
card made by Mrs Harris in 1955 in which this species is recorded from Calve and
Lovage islands in Tobermory Bay.

123/1. SILENE VULGARIS (Moench) Garcke Listed by Wilmott (1942) without
locality and ‘would be N.C.R.’. We have located a specimen: Iona, Shelly Bay, 1906,
R. C. DAVIE (E).

125/1. AGROSTEMMA GITHAGO L. In our previous paper we commented on the loss
of certain agricultural weeds, including this species which was then known only from
the 1877 list by G. Ross. We now know from a specimen: Iona, west, coast north of
village, 1906, R. C. DAVIE (E), that corncockle occurred in the area at least into the
early part of the twentieth century.

136/3. SAGINA MARITIMA Don Shown in the Aflas as a pre-1930 record in the Tober-
mory area, presumably based on Ross (1878). During 1968 this species was discovered
by members of the C.S.S.F. party, by the BMMS and by others in the west of our
area, from Mingary in the extreme north to Jona in the south and including three of
the Treshnish Isles.

137/1. MINUARTIA VERNA (L.) Hiern The occurrence of this species is now con-
firmed, although not as yet in the Iona locality (see Cannon & Bangerter 1968). Ben
More, mountain ledge, 1500 ft, 1956, J. V. SUTHERLAND (E). We are indebted
to G. Halliday for confirmation of this determination and for pointing out that it
represents an interesting extension of range to the west of Scotland. New vice-county
record.

179/1. RHAMNUS CATHARTICUS L. Rubh a’ Ghuirmein near Torosay Castle, 1968,
one tree only, BMMS 3437 (GROVES). Presumably an introduction. New vice-county
record.

192/17. TRIFOLIUM HYBRIDUM. L. Additional to the first vice-county record from
Ulva, we can now give the following from the mainland of Mull: Glac Mhor, Mishnish,
1967, BMMS 2095 (MELDERIS); Torloisk, 1968, C.S.S.F. Party; Tobermory, by
ruined Whisky Distillery, 1968, BMMS 3278 (JAMES).

PLANT RECORDS FROM MULL 149

206/1. VICIA HIRSUTA (L.) Gray Not plotted in the Atlas but recorded by G. Ross
(1878) from Sorne in Mishnish. Its presence in another locality has now been estab-
lished: Iona, Baile Mor, 1967, BMMS 2145 (MELDERIS).

207/6. LATHYRUS SYLVESTRIS L. Included in Appendix II of the Atlas but not
plotted from lack of precise data. There is a specimen in the Boswell Syme herbarium:
under cliff on the bottom of Glen Liebetal between Loch Spelve and Loch Buy (Buie),
1848, SY ME (BM). This specimen is presumably the basis for the record with imprecise
locality in English Botany where Syme remarks ‘it is certainly wild’ in this area.

211/3. RuBUS ARCTICUSL. The Rev. Dr J. Walker is reported by J. E. Smith (1806)
to have collected this species in 1782 ‘in rocky mountainous parts of the isle of Mull’.
Syme (1864), under ‘excluded species’, records ‘Ben More in the isle of Mull... but
recent search has failed to discover it.” Both Druce (1920) and Harley (1956) refer
to these records. Harley mentions two specimens in the British Museum herbarium,
neither of which is from Mull. A third specimen has recently come to light in the
Hortus Siccus Britannicus Fasc. 19 (1802) no. 6 of James Dickson (BM), of which a
duplicate bearing a label ex Herb. Mus. Brit. (presumably cut from another copy of the
Hortus Siccus) is in the Edinburgh herbarium. The Dickson specimens have original
printed labels which read ‘Isle of Mull’. R. M. Harley has kindly confirmed the
identity of the specimen at the B.M. The relegation by Druce (1920) of this species to
‘the dubious plants of Britain’ was perhaps rather premature and Polunin (1953)
included it in his list of arctic species likely to be found in Britain. Its status in Britain
must await further evidence, but clearly a special watch should be kept for this species
in Mull, especially as Koenigia islandica, another arctic species, was found in Mull, for
only the second time in Britain, as recently as 1956.

224/1. ACAENA INERMIS Hook.f. (A. microphylla var. inermis (Hook.f.) Kirk)
This New Zealand species appears to be well established at the track side of the south
drive of Glengorm Castle and has persisted there for at least ten years: 1959, C. W.
MUIRHEAD s.n. (FE); 1968, BMMS 2554 (ROBSON). New vice-county record.

227. COTONEASTER At least four species of introduced Cotoneaster are now
known to occur in Mull. C. microphyllus Wallr. ex Lindl. was recorded by Gerrans (1960)
from the walls of the derelict Aros Castle and the grounds of Glen Forsa House. We
can now add: Wall at entrance lodge to Aros House, 1966, U. K. DUNCAN; 1968,
BMMS 3275 dd. K. FERGUSON & GROVES); near Tostarie, 1968, BMMS
(ROBSON & VICKERY). The three following species are here recorded as new to the
vice-county. C. simonsii Bak.: Track by beach at Loch Buie, 1966, BMMS (CANNON
& GROVES 96); Dervaig, near Dun Auladh, 1967, BMMS 1051 JERMY); these
confirm the tentative record in Wilmott (1942). C. frigidus Wallr. ex Lindl.: Woods
near Kilpatrick House, Torosay peninsula, 1967, BMMS 1012 (JAMES). C. bullata
Bois.: Track to Aros Castle, 1968, BMMS 3086 (ROBSON).

232/4(1). SORBUS INTERMEDIA (Ehrh.) Pers. Shown in the Critical Supplement for
Iona. Known to us from the mainland of Mull at Tavool House, by National Trust
track, 1967, BMMS 1010 (JAMES); Pennyghael Woods, 1968, BMMS 2289 (U. K.
DUNCAN & JAMES).

232/5(1). SORBUS ARIA (L.) Crantz | Knock, by track south of bridge, doubtless
as an introduction, 1968, BMMS 3238 (JAMES). New vice-county record.

241/1. TOLMIEA MENZIEsII (Pursh) Torr. & Gray Naturalised in a damp ditch near
estate lodge cottage in Aros House grounds, 1966, BMMS (CANNON & GROVES
4); 1967, BMMS 315 (JAMES & BOWDEN). New vice-county record.

150 J. F. Me CANNON AND E. B. BANGERTER

301/1. AETHUSA CYNAPIUM L. In Appendix II of the Atlas. Salen, weed in garden
of new house, 1968, MULLIN. A further record for western Scotland, where this
species is very sparsely distributed.

311/2. HERACLEUM MANTEGAZZIANUM Somm. & Levier sensu lato. By stream a
hundred yards south of Torosay Castle, about eight plants, 1968, BMMS 3267
(JAMES). Kindly confirmed by R. K. Brummitt. New vice-county record.

320. POLYGONUM The following introduced species occur on Mull. P.
baldschuanicum Regel occurs at Penmore Mill, Mornish, 1968, BMMS 3435 (R.
WATLING). P. cuspidatum Sieb. & Zucc. occurs in several localities throughout the
island. P. sachalinense F.Schmidt Dervaig, ditchnear garden, 1967,A.G. KENNETH;
1968, BMMS 3165 (JAMES). P. polystachyum Wallr. ex Meisn. Tobermory, 1967,
A. G. KENNETH; BMMS 2081 (MELDERIS); Aros House woods, by lake, 1967,
BMMS 321 (JAMES & BOWDEN). P. campanulatum Hook.f. Near Calgary, 1965,
A. G. KENNETH; near Dun Auladh, on Dervaig to Calgary road, 1967, BMMS
1050 (JERMY); and 1968, BMMS 3246 (JAMES). With the exception of P. cuspidatum,
these appear to be new vice-county records.

358/4. VACCINIUM OxYCOCCUS L. Accepted by the Azlas for v.c. 103 but not plot-
ted. Loch na Criadhach Moire, 1968, BMMS 2545 (EDDY).

360/1. ORTHILIA SECUNDA (L.) House Allt Mor Coire nan Eunachair, on vertical
rocks by stream, 1968, STACE (BM). New vice-county record; previously known from
the adjacent part of Morvern across the Sound of Mull.

370/3. LYSIMACHIA VULGARIS L. Accepted by the AZlas for v.c. 103 but not plotted.
Lightfoot (1777) in his Flora Scotica remarks ‘I observed it by the sides of lochs in
the island of Mull’. G. Ross included it in his 1877 list but not in the 1878 list. We
have only recently encountered it in the course of our work near Lochnameal, in dry
areas of wet marshy pasture, 1968, BMMS 2481 (CANNON, JERMY & PARKER).

370/5. LyYSIMACHIA PUNCTATA L. One mile south of Salen, 1967, A. G.
KENNETH; 1968, BMMS 3425 (L. F. FERGUSON); Kilfinichen, grassy area outside
walled garden, 1968, BMMS 3215 (JAMES); ditch between Torosay Castle and the
sea, 1968, BMMS 3265 (JAMES). New vice-county record.

402/1. MERTENSIA MARITIMA (L.) Gray In addition to the now well-known locality
at Ensay Burn, we have been able to verify two other older records and have located
two new stations on the Treshnish Isles. Only one plant (with a single leaf) was found
to exist at the Port Haun station. The colony at the head of Loch na Keal discovered
by R. Kidston in 1889 (GL) was also found to be far from flourishing, consisting
only of two plants in poor condition. On the Treshnish Isles this species was found on
Lunga by Ferreira during the Maps Scheme era and by Templeman in 1939. A large
colony was found on Sgeir a Chaisteil, 1968, BMMS 2802 (PARKER) and a smaller
one on Fladda, 1968, BMMS (EDDY). Pennant (1772), recording this species from
Iona, stated ‘the beautiful sea bugloss makes the shores gay with its glaucous leaves
and purple flowers’. In recent times it was recorded for Iona during the Maps Scheme
but has not been observed there during the period of our own survey.

409/1. LycruM BARBARUM L. (L. halimifolium Mill.) Iona, main street of Baile Mor,
1968, BMMS 3394 (JAMES). Confirmed by W. T. Stearn. New vice-county record.

415/L. LYCOPERSICON ESCULENTUM Mill. Growing by sewage outlet south of Iona
Town (Baile Mor), 1968, BMMS 3378 (JAMES). New vice-county record.

PLANT RECORDS FROM MULL 151

425/2 x c. MIMULUS GUTTATUS DC. x CUPREUS Dombrain Gribun, south of
Clachandu, 1967, A. G. KENNETH (BM). We are grateful to R. H. Roberts for
confirming this identification. This record came too late for inclusion in his paper
(Roberts 1968) and forms the first record for Mull, this hybrid only previously being
known in v.c. 103 from Coll.

425/3. MIMULUS MOSCHATUS Dougl. ex Lindl. Listed by Wilmott (1942) from
Kellan, but with the comment ‘must await confirmation from the collection’. We
can now give a firm record from another locality: Garbh Choire, north of Gruline
House, 1968, BMMS 2341 (I. K. FERGUSON).

432/2. PEDICULARIS SYLVATICA L. subsp. HIBERNICA D. A. Webb We have two
specimens of this subspecies from Mull, distinguished by ‘calyx and pedicels which
are uniformly clothed with rather long white curled hairs’ (Webb 1956). Abhain a
Ghlinne, Brolass, in dense Molinia, 1968, BMMS 2928 (JERMY); Loch Squabain,
wet Molinia slopes, 1968, BMMS 3077 (JERMY). A. G. Kenneth also reports in a
letter (August 1967) that he has collected this subspecies on Beinn Fhada, on low
ground near Allt Coire nan Gabhar. These appear to be the first records for v.c. 103.

445/4 x 3. MENTHA X VERTICILLATA L. Accepted by the Critical Supplement for
v.c. 103 but not plotted. Allt na Searmoin, 1966, U. K. DUNCAN; between Gruline
bridge and Loch Ba, 1966, U. K. DUNCAN;; Allt Ardnacross, river bank, 1968,
BMMS 3123 (JAMES). Specimen confirmed by R. M. Harley. Both parent species
M. arvensis and M. aquatica are well distributed in Mull.

445/5. MENTHA SPICATA L. Salen, waste ground between road and shore, 1966,
U. K. DUNCAN. This locality provides a second record to that indicated in the
Atlas. We have a specimen, confirmed by R. M. Harley, Kilfinichen Bay, roadside,
1968, BMMS 3220 (JAMES) which is probably from the same area as the A/las
locality.

445/6 x 7. MENTHA X NILIACA Juss. ex Jacq. Forms of this hybrid, the correct name
for which according to R. M. Harley is probably M. xvillosa Huds., have been found
in Mull, but we are leaving the publication of new vice-county records until Harley
has published the results of his current work.

475/1. CAMPANULA LATIFOLIA L. Wilmott (1942) gives two localities which are
presumably the basis for the Atlas dots. We can provide two additional localities:
Torosay Castle, 1967, BMMS (BOWDEN); Pennyghael Woods, 1968, BMMS 3209
(JAMES).

485/10. GALIUM ULIGINOSUM L. Recorded without locality by Ross (1877) and
otherwise known previously only from Coll and Tiree in v.c. 103. Carsaig Bay, 1967,
J. DUNCAN;; Creag nan Croman, Sgulan Breac and the east end of Loch Peallach,
1967, BMMS (MELDERIS); Lochnameal, 1968, BMMS 2477 (CANNON, JERMY
& PARKER). The four last localities are all in the northern part of the island within
three miles of Tobermory.

487/3. SAMBUCUS RACEMOSA L. Carsaig Bay, 1967, U. K. DUNCAN (BM), also
observed in the same year by J. DUNCAN. New vice-county record.

506/2 x 1. SENECIO X OSTENFELDII Druce (S. aquaticus Hill x S. jacobaea L.) Grass
Point, behind house at foot of pier, 1968, BMMS 3259 (I. K. FERGUSON). Both
parent species were observed in the neighbourhood and indeed are common through-
out Mull. The hybrid is likely to be encountered elsewhere in the area. New vice-
county record.

152 J. F. M. CANNON AND E. B. BANGERTER

509/2. PETASITES ALBUS (L.) Gaertn. Torosay Castle, streamside by bridge on main
road, 1966, U. K. DUNCAN. This species, which is commonly recorded in eastern
Scotland, has only very rarely been observed in the west. New vice-county record.

509/4. PETASITES FRAGRANS (Vill.) C. Presl Near Torloisk House, 1968, BMMS
2278 (U. K. DUNCAN & JAMES). New vice-county record.

512/1. INULA HELENIUM L. This species, which is now known to occur in seven
localities in our area, was published in Trans. Proc. bot. Soc. Edinb. (1913) 26: xxv as
a new vice-county record, and was repeated, with the addition of the locality, one
mile east of Bunessan, in Rep. botl. Soc. Exch. Club Br. Isl. (1914) 3: 388. In fact, it
was recorded from ‘near Aros’ in Hooker’s Flora Scotica (1821) and even earlier
from Iona by J. Walker (1808) in his Essays.

539/4. CARDUUS ACANTHOIDES L. Previously known from Iona but now recorded
from three stations around Tobermory, 1967, BMMS (MELDERIS).

557/3. CICERBITA MACROPHYLLA (Willd.) Wallr. Kilfinichen, abundant as weed in
walled garden and on roadside, 1968, BMMS 3218 (JAMES); Tobermory, garden
escape, 1968, STACE. New vice-county record.

560/H. HELIANTHUS ANNUUS L. An Crosan. Plants 4—6 ft high, along the drift zone
in a bay close to the abandoned village at Shiaba, 1968, MULLIN. New vice-county
record.

576/2. ZOSTERA ANGUSTIFOLIA (Hornem.) Reichb. While Z. marina has been
known from several stations around Mull, dating back to the lists of G. Ross, this
species has only recently been found: North-east side of Loch Don, near old pier, 1966,
U. K. DUNCAN (BM); Lochdonhead, 1967, KENNETH & STIRLING (BM).
Both these have been confirmed by T. G. Tutin. New vice-county record.

577/7. POTAMOGETON ALPINUS Balb. Recorded by Wilmott (1942) from Tobermory
reservoir. Two specimens, determined by J. E. Dandy, provide additional localities:
Lochnameal, 1968, BMMS 2490 (CANNON, JERMY & PARKER); Lochan north-
west of Cnoc an da Chinn, 1968, BMMS 3305 (1. K. FERGUSON).

577/15. POTAMOGETON BERCHTOLDII Fieb. Previously known from Iona, but now
recorded for the first time from Mull itself: Outlet of Loch na Cuilce, 1968, KENNETH
(BM). Determined by J. E. Dandy.

579/2. RUPPIA MARITIMA L. This species, which has previously been recorded in
the vice-county from Coll and Iona, has now been located on Mull and the Treshnish
Isles: Near Garmony, 1965, A. G. KENNETH; Lochdonhead, 1967, KENNETH
& STIRLING (BM); 1968, BMMS 3270 (I. K. FERGUSON) and in 1968 in three of
the Treshnish Isles, Sgeir an Eirionaich, BMMS 2593 (CANNON & JERMY); Bac
Beag, BMMS 2617 (ROBSON & SIMS); and Fladda, BMMS 2823 (EDDY). An
immature specimen, collected at Lochdonhead, may be R. cirrhosa but confirmation
of this record must await the collection of mature fruiting material.

607/5. ALLIUM VINEALE L. Included in our previous paper as known only from
Iona. Located during the last season in four additional localities: the island of Erraid,
seaside cliff, 1968, McALLISTER; Tober a Mhoire, 1967, BMMS (MELDERIS);
Criadhach Mhor, 1967, BMMS (MELDERIS); between Kilninian and Tostarie,
1968, BMMS 2748 (ROBSON).

649/1. ARUM MACULATUM L. Torloisk House, woods and adjacent hedgerows at
roadside, 1968, BMMS 2277 (U. K. DUNCAN & JAMES). Presumably an introduc-
tion, this plant is the form without spots on the leaves. New vice-county record.

PLANT RECORDS FROM MULL 153

663/17. CAREX VESICARIA L. In our previous paper, we showed that the record for
this species was based on erroneous evidence. During the past season, the first firm
locality was established: Reudle, 1968, BMMS 2743 (EDDY). Confirmed by A. C.
Jermy. New vice-county record.

676/9. POA COMPRESSA L. Recorded on a Distribution Maps Scheme card from
Calve & Lovage islands (Tobermory Bay) by Mrs S. Harris in 1955 but not taken
up in the Atlas. Recorded in 1967 from Tobermory itself by A. MELDERIS. Doubt-
fully native in our area. New vice-county record.

683/12. BROMUS THOMINII Hardouin. _In addition to the locality discovered by
A. G. Kenneth in the Gribun shown in the Critical Supplement, we have two further
records: Tobermory village and Erray House near Tobermory, 1967, BMMS
(MELDERIS).

707/5. PHLEUM ARENARIUM L. Iona, on shell sand banks at the northern end of the
island, 1968, MULLIN. New vice-county record. The only record shown in the
Atlas for the west of Scotland is in grid square 27/04.

REFERENCES

CANNON, J. F. M. & BANGERTER, E. B. (1968). Plant records from Mull and the adjacent
small islands. Proc. bot. Soc. Br. Isl., 7: 365-372.

Danpy, J. E. (1958). List of British Vascular Plants. London.

Druce, G. C. (1920). The extinct and dubious plants of Britain. Rep. botl Soc. Exch. Club
Br. Isl., 5: 731-799.

EwIna, P. (1892). Glasgow Catalogue of Native and Established Plants, being a contribution to
the Topographical Botany of the western and central counties of Scotland. Glasgow.

EwIna, P. (1899). Glasgow Catalogue ..., 2nd ed. Glasgow.

GERRANS, M. B. (1960). Notes on the flora of the isle of Mull. Proc. bot. Soc. Br. Isl., 3: 369-374.

Har ey, R. M. (1956). Rubus arcticus L. in Britain. Watsonia, 3: 237-238.

Hooker, W. J. (1821). Flora Scotica. London.

KeppiE, W. (1850). Staffa and Iona described and illustrated . . . Glasgow, Edinburgh and
London.

LIGHTFOOT, J. (1777). Flora Scotica. London.

PENNANT, T. (1774). A Tour in Scotland and Voyage to the Hebrides 1772. Chester and London.

PERRING, F. H. & SELL, P. D., ed. (1968). Critical Supplement to the Atlas of the British Flora.
London.

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

POLUNIN, N. (1953). Arctic plants not yet found in Britain. Watsonia, 3: 34-35.

RATCLIFFE, D. (1963). B.S.B.I. Field Meeting in Mull, 1961. Proc. bot. Soc. Br. Isl., 5: 94.

ROBERTS, R. H. (1968). The hybrids of Mimulus cupreus. Watsonia, 6: 371-376.

Ross, G. (1877). Isle of Mull, Mid Ebudes. Rep. Bot. Loc. Rec. Club for 1876, 188-192.

Ross, G. (1878). On the flora of Mull. Trans. Proc. bot. Soc. Edinb., 13: 234-242.

SMITH, J. E. (1790-1814). English Botany. London.

SyME, J. T. I. Boswell (1863-1886). English Botany, 3rd ed. London.

TEMPLEMAN, A. (1942). Iona and Lunga, in Report on Excursions arranged in 1939. Rep.
botl Soc. Exch. Club Br. Isl., 12: 249-250.

WALKER, J. (1808). Essays on Natural History and Rural Economy. London and Edinburgh.

Wess, D. A. (1956). A new subspecies of Pedicularis sylvatica L. Watsonia, 3: 239-241.

WILMoTT, A. J. (1942). V.C. 103. The Island of Mull, in Report on Excursions arranged in
1939. Rep. botl Soc. Exch. Club Br. Isl., 12: 236-249.

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Watsonia, 8, 155-162 (1970). 155

Exotic Amelanchier species naturalised in Europe and their
occurrence in Great Britain

F.-G. SCHROEDER

Systematisch-Geobotanisches Institut, G6ttingen, Untere Karspiile 2,
West Germany

ABSTRACT

In Europe, three Amelanchier species of American origin are naturalised to a considerable
extent: A. spicata (Lam.) K. Koch, A. lamarckii Schroeder and A. confusa Hylander. Only
one of them, A. l/amarckii, has been found hitherto in the British Isles. Its distribution is
restricted mostly to southeastern England, especially to those parts where acid soils prevail.
A detailed key and a preliminary list of localities are given.

INTRODUCTION

For more than 80 years, several exotic cultivated species of the genus
Amelanchier (Rosaceae) have been naturalising themselves more and more in
various parts of western, central and northern Europe. The taxonomy and
distribution of these neophytes, however, has remained mostly doubtful and
confused. In the last few years I have tried to clear up this interesting problem.
The results are presented in my paper ‘Amelanchier-Arten als Neophyten in
Europa’ now in the press. As this publication is written in German, I am giving
here a short summary in English with a key to the more important species and
a list of the localities known to me of the only species that occurs in England.

TAXONOMY

In Flora Europaea, Franco (1968) gives an account of the genus Amelanchier
which unfortunately contains several mistakes. Besides the indigenous A. ovalis
Medic., he mentions two species as naturalised: A. grandiflora Rehder and
A. spicata (Lam.) K. Koch, the latter being considered conspecific with A.
humilis Wieg. In reality, however, there are three well-separated species which
are naturalised. Moreover, my studies on nomenclature have led to some
conclusions which are rather different from the opinions expressed in Flora
Europaea. In order to prevent further confusion in the difficult taxonomy of
the genus, I think it necessary to give the following corrections:

1. The statement in Flora Europaea that A. spicata (Lam.) K. Koch is con-
specific with the American species A. humilis Wieg. is based on two views
advocated only by Jones (1946). The first is that A. stolonifera Wieg. and A.
humilis should be united, an opinion refuted by most other authoritative
American taxonomists. The other is his conviction that A. spicata and A.
stolonifera are synonyms, which is definitely wrong (Fernald 1946). To sum up,
in my opinion A. spicata is not conspecific with A. stolonifera, much less with
A. humilis. Neither of the two latter taxa has so far been found naturalised in
Europe.

156 F.-G. SCHROEDER

2. As I have pointed out briefly in Taxon (Schroeder 1968) and discussed
more fully in the paper mentioned above, the name 4A. grandiflora Rehder
cannot be applied to any of the species naturalised in Europe. The original
description (Rehder 1920) agrees in several respects (but not in all) with the
species I have named A. lamarckii (see below). A herbarium sheet stated in
Rehder’s handwriting to be the ‘type’ (Herbarium of the Arnold Arboretum)
is, however, quite different both from his description and from A. lamarckii.
It seems to belong to the true A. canadensis (L.) Medic. The marked difference
between the description and the type, which may be due to a confusion of
herbarium material, makes it impossible to decide now what Rehder really meant
by his new species. Therefore the name A. grandiflora is to be considered a
nomen confusum.

3. The taxon which is most widely cultivated in western and central Europe
and which is extensively naturalised in the Netherlands and north-west Germany
as well as in parts of southern England, has hitherto usually been erroneously
called A. canadensis (L.) Medic. In recent years it has often been wrongly
transferred to A. laevis Wieg., A. intermedia Spach, A. confusa Hyl. or A.
grandiflora Rehder. It has, however, already been described very adequately
by Lamarck (1783) as Crataegus racemosa. As the specific epithet racemosa is,
for nomenclatural reasons, not applicable in the genus Amelanchier, I have
replaced it (Schroeder 1968) by the nomen novum, A. lamarckii.

4. In Sweden, another Amelanchier taxon has been cultivated under the
name A. canadensis. This too is also naturalised to a considerable extent. It
has been described as a new species, A. confusa, by Hylander (1966). A. confusa
is very different from A. /amarckii and may by no means be treated as a synonym
of the latter (or included under A. grandiflora as in Flora Europaea). It is, in
fact, a third naturalised species which must be considered as a permanent
member of the European flora.

To sum up, the four important Amelanchier species native or naturalised in
Europe are:

A. ovalis Medicus, Gesch. Bot., p. 79 (1793)

A. spicata (Lam.) K. Koch, Dendrologie, 1: 182 (1869)
A. confusa Hylander, Lustgarden, 45/46: 273-274 (1966)
A. lamarckii Schroeder, Taxon, 17: 633 (1968)

These can be identified using the following key:

1. Styles very short, about 1 mm long, not exceeding the hypanthium. Petals
often woolly outside. Leaves 2-5-5 cm long, mostly rounded at the apex,
coarsely crenate with about 3-5 rounded teeth per cm. Upright or spreading
shrub upto 3 mi se ; 3 = A. ovalis

Styles 2-5-4 mm long, clearly deceias He ‘pate connate at least at
the base, often to 4 or even 4 of their length. Petals glabrous or only ciliate
at the apex. Leaves finely or coarsely serrate with acuminate teeth. Taller
shrubs up to 6 m or more es by ce ie ae a Z

2. Inflorescences erect, dense, the lowest pedicels not much longer than the rest.
Petals 6-10 mm long, obovate, about twice as long as wide, ciliate at apex.
Top of ovary densely woolly. Calyx on mature fruits erect or spreading.
Young leaves at flowering time light green or somewhat olive green, with

EXOTIC AMELANCHIER SPECIES IN GREAT BRITAIN M7

dense floccose yellowish tomentum beneath. Mature leaves 3-6 cm long,
broadly elliptical or obovate to nearly orbicular, rounded to cordate at base,
rounded or shortly acuminate at apex, mucronate, finely serrate with about
5-7 teeth per cm. Fastigiate shrub up to 8 m, without conspicuous autumn
colouring .. i ee : ‘ Be ae Ag A. spicata

Inflorescences ascending or fate lax, the lowest pedicels often much longer
than the rest. Petals 10-15 mm long, sloamesclete. at least three times as
long as wide, glabrous. Young leaves at flowering time glabrous or only
silky-hairy beneath as re iy - a A i: 3

3. Top of ovary woolly. Calyx on mature fruits strongly recurved. Young
leaves at flowering time mostly flat, light green, often with light red patches
between the veins, glabrous on both sides or with a few residual areas of thin
white silky tomentum beneath. Mature leaves 3-5-5 cm long, ovate, broadly
cuneate to subcordate at base, acuminate from the middle, mucronate, irreg-
ularly and mostly coarsely serrate with about 2-6 teeth per cm. Fastigiate
shrub up to 6 m, without conspicuous autumncolouring .. A. confusa

Top of ovary glabrous. Calyx on mature fruits erect or spreading, not re-
curved. Young leaves at flowering time about half open, intensely purplish or
copper-coloured, white silky-hairy beneath. Mature leaves 4-5-8-5 cm long,
obovate to oblong or elliptical, rounded or subcordate at base, shortly
acuminate in the upper third, mucronate, finely and regularly serrate with
about 5-7 teeth per cm. Tall shrub or small tree up to 10 m, with divergent
branches and very conspicuous yellow, orange or crimson autumn
colouring Si, ae Ms ‘Pe at. b2 se: A. lamarckii

The three naturalised taxa are undoubtedly of North American origin, but
none of them is known as a distinct species in North America. They must be
considered as microspecies, which have been overlooked or taken for hybrids
in their native country because of their intermediary positions between several
other well-known and widely distributed species. The suggestion that they might
be hybrids that have arisen in European gardens (neogenic endemics) cannot
be taken seriously. In Europe, the three species have been found naturalised in
the following countries (abbreviations as in Flora Europaea):

A. spicata: Au Be Cz Da Fe Ge Ho No Rs(B) Rs(C) Su
A. confusa: Su
A. lamarckii: Be Br Da Ge Ho Su

Besides these three widespread neophytes, some other exotic taxa occur as
ornamentals in gardens and parks. A. laevis Wieg. and A. asiatica (Sieb. &
Zucc.) Endl., in particular, are now increasingly cultivated, as is to a lesser
extent A. stolonifera Wieg. Therefore these species may be expected to escape
from gardens in the future. In the 19th century A. alnifolia Nutt. and A.
lamarckii var. scharnkeana (Aschers. & Graebn.) Schroeder (Schroeder 1968)
were planted in several large parks and gardens. Both taxa, which seem to be
cultivated no longer, have been found growing wild in two and three localities
respectively. In contrast, the true A. canadensis (L.) Medic. exists only as a
rarity in some botanic gardens, and A. arborea (Michx.f.) Fern. seems to occur
more rarely still. A key to most of the exotic species (though containing the
confusing incorrect names A. spicata and A. stolonifera) may be found in
Kriissmann (1960).

158 F.-G. SCHROEDER

Ficure 1. Amelanchier lamarckii, typical leaves.

EXOTIC AMELANCHIER SPECIES IN GREAT BRITAIN 159

PRELIMINARY REPORT ON THE DISTRIBUTION IN ENGLAND

In British floristic literature, several records of naturalised Amelanchiers may
be found. As in other European countries, there is no unanimity concerning
the taxonomic position of the naturalised specimens. The modern Floras (e.g.
Clapham, Tutin & Warburg 1962) mostly quote A. Jaevis, but other species
such as A. canadensis, A. confusa, A. intermedia or A. alnifolia are also mentioned
as naturalised. The descriptions given in the Floras are often combinations of
field notes and extracts from American literature, and therefore it is rather
difficult to decide which species are really present as neophytes in Great
Britain.

In the course of my investigations, I had the opportunity to see the material
of several important British herbaria, and in 1968 I visited the wild Amelanchier
population at Hurtwood in Surrey. All the specimens I have seen—dried and
living ones—belong clearly to Amelanchier lamarckii, the species that is so
frequent in the Netherlands and north-west Germany. Therefore it may be
supposed, for the present, that A. /amarckii is the only species naturalised in
Great Britain. Certainly it is the most widely distributed one, even if individuals
of other species should be detected in the future.

In order to facilitate identification, I have, in addition to the description in
the key, illustrated typical leaves of A. lamarckii (Fig. 1). A drawing which
depicts the habit of flowering and fruiting twigs very well may be found under
the name of A. /aevis in Clapham, Tutin & Warburg (1960), while the drawings
by Hadfield (1959, 1969) are not so characteristic, though they give some details
more accurately. A translation of my synoptical table showing the differences
between A. lamarckii, A. laevis, A. arborea and A. canadensis has been published
by McClintock (1969). For further details I refer to my publications mentioned
above.

The only records of naturalised Amelanchiers in the British Isles which I
could get refer to England, and most of them are from the south-eastern part
of the country. Even in this district the distribution is not uniform. Obviously
there are distribution centres in areas with predominating acid soils: the Eocene
beds of the New Forest and the Bagshot-Camberley region, and the areas of
Cretaceous sandstones. This reflects the ecological requirements of A. /amarckii,
which, both in Britain and on the Continent, is restricted to acid, mostly sandy
soils of rather variable water content. In Hurtwood, Surrey, A. lamarckii is a
member of forest and hedge communities belonging to the vegetation type called
Querco-Betuletum by continental botanists, and the descriptions indicate
similar vegetation also in several other localities. This corresponds well with
the situation in the Netherlands and in north-west Germany, where the Querco-
Betuletum and the shrub communities replacing it are the typical habitat of
A. lamarckii.

The following preliminary list of records that I have so far been able to obtain
together with the distribution map (Fig. 2) do not claim to be complete. I
intend only to draw attention to the problem. The list has been compiled from
the data I found in herbaria and literature, and, for the most part, from many
private records of several botanists, which I obtained with the aid of Mr D.
McClintock and Dr H. J. M. Bowen. It is a pleasure to me to acknowledge
the friendly assistance I obtained from these two gentlemen and likewise from
the keepers of the herbaria mentioned below.

160

F.-G. SCHROEDER

LIST OF RECORDS

ABBREVIATIONS AND SIGNS USED IN THE LIST

Herbaria:

BM___ London, British Museum (Natural History)
K Kew, Royal Botanic Gardens

LTR Leicester University, Department of Botany
OXF Oxford University, Botany School

Literature (for titles see under References):

B Bowen 1968

DR _Druce 1926

DU Dunn 1893

KL Kent & Lousley 1953
S Salmon 1931

WD Wolley—Dod 1937

Other sources:

—

V.C.

V.C.

V.C.

BL Bowen by letter

MC McClintock by letter

R Data from an unpublished Flora of Kent by Dr F. Rose, communicated by
McClintock

ny: Data from an unpublished record list by Mrs Yule, Godshill, Hants.,
communicated by McClintock

= Specimens seen by the author and identified as Amelanchier lamarckii

9, DorsET: Upton near Poole, a few trees in a hedge, 1956 (BL); Osmington
near Weymouth, 1956 (MC); Broadstone near Poole, on the main road A349
(MC).

11, SouUTH HAMPSHIRE (all records from Y, unless indicated otherwise): Ashley
Heath area; St Ives, heathy roadsides, well established, on the borders of the
New Forest (BL); Iford area near Christchurch; Nea Croft area near Christ-
church; Holmsley, 1933 (OXF, as A. /aevis)!; Wootton; Sway, Set Thorne;
Bolderwood; Hollands Wood, Ramnor; Clay Hill; Fletchwood; Ashurst,
Langley Wood; Marchwood; Stanswood and Hillhead; Rownhams; Chil-
worth; Ampfield near Romsey (MC); north of Chandlers Ford; each side
of Fareham—Winchester road near Fair Oak.

12, NorTH HaAmpsHIRE: Liss Forest, Woolmer Forest (Y); Passfield, Headley
Down (Y); Kingsley, Fleet, 1950 (K, as A. lJaevis)!; Fleet area, Aldershot,
between Blackwater and Hartley-Wintney, about a mile south of where the
main road goes over Hartford Bridge Flats, two specimens growing on a wild
bit of heath but with the roots on the edge of a boggy ditch, 1923 (OXF,
as A. canadensis or A. laevis)!; north of Mortimer to county border (Y.)

. 13, West Sussex: Midhurst, growing in a wood, where there seems to be a lot

of it, probably either planted or bird sown, 1923 (BM, OXF, as A. florida)!;
Blackdown, above Valewood House amongst planted pines, 1924 (OXF, as
A. canadensis)!; Storrington, in a wild spot, 1926 (OXF, as Crataegus sp.)!;
Slinfold near Horsham (WD).

. 14, East Sussex: Mayfield (WD); forests on sandstone south of the county

border near Tunbridge Wells, 1968 (communicated by D. W. Shimwell).

. 16, West KENT: Tunbridge Wells Common, 1937 (R); Kenward Green, Pembury,

near Tunbridge Wells, 1962 (R); Cranbrook, Angley Wood, 1956 (R); Parsons
Marsh, Brasted (near Scords Wood), a strong colony, 1960 (R); Hosey
Common, 1953 (KL); Crockhamhill Common, 1939 (R).

EXOTIC AMELANCHIER SPECIES IN GREAT BRITAIN 161

v.c. 17, SURREY: Oxshott Heath (KL); Bagshot Heath, naturalised at Lightwater,
1958 (K, as A. arborea)!; Windlesham, self-sown in thickets by the road on
acid soil, 1969 (BL)!; Deepcut Camp, before 1921 (BM, as A. canadensis)!;
Copses about Witley and Thursley (DU); Hurtwood: from this forest area
on Lower Greensand naturalised Amelanchiers have been known since as long
ago as 1893 (DU). Clapham, Tutin & Warburg (1962) record ‘A. Jaevis’ as
‘common over several square miles in the Hurtwood’. My visit in 1968 showed,
however, that the distribution is not so extensive. The densest population of
A. lamarckii that I found was in the parts lying west of the road from Peaslake
to Ewhurst, especially in a strip extending from the margin of Peaslake about
1 mile to the south-southwest, where many large shrubs and numerous
seedlings are present. On a herbarium label written by Dr Warburg (OXF)
this locality is referred to as “Bentley Copse near Shere’, and the forest
structure is quite adequately described there as “Oak coppice with pine,
rowan, whitebeam, ling and bilberry’, to which may be added holly, birch
and alder-buckthorn. In other parts of the Hurtwood area I saw scattered
specimens on Holmbury Hill and about 4 mile east of Leith Hill. Further
records exist from Park Wood near Albury (S), Farley Green (S) and Westcott
Heath (LTR)!.

( ee \ oP

XN
4
¢
p)

\

PSs ) |
rN GNSS Gm
¢ Nv

id, 4 )

\
Bp)

-—-

FIGuRE 2. Distribution of Amelanchier lamarckii in southern England.

GREATER LONDON. V.C. 16, WEST KENT: Hayes Common, 1911 (KL), 1918 (OXF, as
A. canadensis)!; Bromley, Fishers Wood, 1961 (R).v.c. 17, SURREY: Richmond,
Ham Common, 1943 (KL), 1953 (BM, as A. Jaevis)!. v.c. 18, SouTH ESSEx:
Walthamstow, Cook’s Folly Wood, 1899 (BM, as A. canadensis)!. v.c. 21,
MIDDLESEX: North End, Hampstead, 1916, 1949 (KL); Kenwood, 1950 (KL);
East Finchley Wood, 1887, 1912 (KL).

V.C. 22, BERKSHIRE: ‘Amelanchier, when naturalised in Berks, grows in small colonies
which spread quite fast if not destroyed by fire or the axe. Such colonies are
confined to acid, fairly dry soils similar to those in N. Surrey, the New Forest

162 F.-G. SCHROEDER

and also on the N. German plain. Associated woody plants include Betula
pendula, B. pubescens, Frangula alnus, Sorbus aucuparia, Calluna vulgaris and
Vaccinium myrtillus, together with the well naturalised alien Rhododendron
ponticum’ (BL); Newbury, Inholmes (B); Reading, Ufton park, acid wood
margin, 1961 (B, BL); Whiteknights Park, 1969 (BL)!; Ashley Hill near
Henley, acid woodland (B, BL); Crowthorne, Heath Lake and Wellington
College, woods on acid, sandy soil (B, BL); Sandhurst, woodland on very
acid soil (B, BL); Ascot, quite naturalised in woods, 1915 (OXF, BM, as
A. canadensis)!.

v.c. 24, BUCKINGHAMSHIRE: In a wood near Lane End, 1915 (DR; OXF, as A.
canadensis) !.

v.c. 69, WESTMORLAND and NorTH LANCASHIRE (not shown in Fig. 2): Broughton-
in-Furness, on peat mosses, 1962 (K, as A. Jaevis)!; Islands of Windermere,
1891 (BM, as A. canadensis)!.

REFERENCES

BowENn, H. J. M. (1968). The Flora of Berkshire. Oxford.

CLAPHAM, A. R., TUTIN, T. G. & WARBURG, E. F. (1960). Flora of the British Isles. Illustrations,
vol. 2, p. 21, fig. 631. Cambridge.

CLAPHAM, A. R., TUTIN, T. G. & WARBURG, E. F. (1962). Flora of the British Isles, 2nd ed.,
p. 422. Cambridge.

Druce, G. C. (1926). Flora of Buckinghamshire. Arbroath.

Dunvn, S. T. (1893). Flora of Southwest Surrey. London.

FERNALD, M. L. (1946). Amelanchier spicata not an American species. Rhodora, 48: 125-129.

FRANCO, J. Do A. (1968). Amelanchier, in TUTIN, T. G., et alia, ed. Flora Europaea, vol. 2,
pp. 71-72. Cambridge.

HADFIELD, M. (1959). British Trees. London.

HADFIELD, M. (1969). Amelanchier lamarckii. Gdnrs’ Chron., 166(1): 13-14.

HyLANDER, N. (1966). Nagra anmarkningar om de i Sverige odlade och férvildade Amelanchier-
arterna, sarskilt A. confusa Hyl. Lustgarden, 45/46: 266-277.

Jones, G. N. (1946). American species of Amelanchier. Illin. Biol. Monogr., 20 (2).

KENT, D. H. & Loustey, J. E. (1953). A hand list of the plants of the London area. London
Naturalist, 32, Suppl.: 114.

KRUSSMANN, G. (1960). Handbuch der Laubgehdlze, vol. 1. Berlin.

LAMARCK, J. B. DE (1783). Encyclopédie méthodique. Botanique, vol. 1, p. 84. Paris.

McCuintock, D. (1969). A new Amelanchier. Gdnrs’ Chron., 165 (18): 11-12; 165 (20): 5.

Reuper, A. (1920). New species, varieties and combinations from the Herbarium and the
collections of the Arnold Arboretum. J. Arnold Arbor., 2: 42-62.

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

SCHROEDER, F.-G. (1968). Zur Nomenklatur in der Gattung Amelanchier (Rosaceae). Taxon,
17: 633-634.

SCHROEDER, F.-G. (in press). Amelanchier-Arten als Neophyten in Europa. Abh. Naturw.
Ver. Bremen, 37 (3).

WoLLeEyY-Dop, A. H. (1937). Flora of Sussex. Hastings.

Watsonia, 8, 163-164 (1970). 163

Short Notes

362/1. MONOTROPA HyYPoPITys L.—New to Fife (v.c. 86).

A colony of Monotropa hypopitys sensu lato was found in Tentsmuir Forest, Fife
(GR.37/50.24) on 24th June, 1967. It has been under observation since that time and
produced some 190 stems in the summer of 1968.

The nearest record to this one is in Cleish parish, Kinross, attributed to A. Robertson
(Young 1936), but it is quite unsubstantiated and may be an error since it is not in
Robertson’s list published by Balfour (1894), the only source of Robertson records
referred to by Young. The nearest authentic records, based on specimens in the
Perth Museum are Perth (McRae, 1904) and Killin (McNab, 1857). The most northern
record in Scotland as far as I can ascertain is Cawdor Woods, Nairnshire.

The Tentsmuir plants are growing under Pinus sylvestris, 35—40 feet high, planted
about 40 years ago over old sand dunes. The sand is now covered with a deep litter
consisting mainly of decaying pine needles. The ground flora in and around the
colony is dominated by mosses, chiefly Pleurozium schreberi, Plagiothecium undulatum,
Hypnum cupressiforme, and Rhytidiadelphus triquetrus. Other associates are Dicranum
scoparium, Peltigera canina, Goodyera repens, Festuca ovina, F. rubra, Carex arenaria
and occasional plants of Veronica officinalis and Senecio jacobaea.

The plants appear to be intermediate between Monotropa hypopitys L. and M.
hypophegea Wallr. as described by Clapham, Tutin and Warburg (1962). The question
of intermediates is discussed in the Critical Supplement to the Atlas of the British Flora
in the light of work done by D. J. Wicker. A taxonomic investigation is now being
made at the St Andrews University Department of Botany and it is hoped soon to
obtain a chromosome count. This should enable us to determine the status of the
Fife plants.

Dried and spirit material from the Tentsmuir colony has been deposited in the
herbarium of the Department of Botany, University of St Andrews.

REFERENCES

BALFour, I. B. (1894). An old list of ‘Stations of Rarer Plants ascertained to grow around
Inverkeithing and north of the Forth. By A. Robertson’. Trans. Proc. bot. Soc. Edinb.,
20: 84-90.

CLAPHAM, A. R., TUTIN, T. G. & WARBURG, E. F. (1962). Flora of the British Isles, 2nd ed.
Cambridge.

PERRING, F. H. & SELL, P. D., ed. (1968). Critical Supplement to the Atlas of the British Flora.
London.

Youna, W. (1936). A List of the Flowering Plants and Ferns recorded from Fife and Kinross
(v.c. 85). Trans. Proc. bot. Soc. Edinb., 32: 98.

A. ANGUS

716/g. SPARTINA GLABRA Muhl.—In the British Isles.

Spartina glabra is a member of the S. alterniflora complex and has been treated by a
number of American authors as S. alterniflora var. glabra. A study of the population
at Southampton Water by Marchant (1967) and Marchant & Goodman (1969) has
convinced these authors that the taxon deserves specific rank.

Marchant & Goodman (1969) give the following detailed description—‘Spartina
glabra is a large, graceful grass 100-120 cm tall, with a drooping habit rather than
the erect rigidity of the other British Spartina species. The matted roots are penetrated
by long, steeply descending and ascending rhizomes with long internodes. The leaves

E*

164 SHORT NOTES

are longer and laxer than in S. alterniflora, but are otherwise similar in the two species.
The inflorescences are 20-35 cm long, with eight to twelve spikes. The spikes (5-15
cm long) are spaced out in a raceme and distantly inserted along the rachis. The
spikelets are 12-14 mm long, by 1-14 mm wide, each covering about half the length
of the spikelet above. The lower glume is almost subulate and ciliate to the tip, 6 mm
long, without a membranous border, one to three nerved and glabrous to the naked
eye. The upper glume is very narrow, ciliate to the tip, and with cilia extending beyond
as a crest: this glume is 12-14 mm long by 1-14 mm wide, without a membranous
border, three to five nerved and, though glabrous to the naked eye, distinctly pilose
under the binocular microscope. There are three stamens, with anthers 5-54 mm long,
exserted at maturity and dehiscing to produce good pollen. The fruit is a caryopsis
8—10 mm long and viable.’

Details of the British distribution given by Marchant & Goodman are as follows:
v.c. 9, Dorset: plants from America were planted in Poole Harbour by Prof. F. W.
Oliver in 1924. They apparently failed to survive for very long. Tillers from the
Southampton water population were planted in the Nature Conservancy’s ‘Spartina
garden’ at Keysworth, Poole Harbour in 1963. The plants survive and are spreading.
v.c. 11, S. Hants: tillers introduced from America were planted at Eling, Southampton
Water by Prof. S. Mangham in 1924. The only surviving stand of this original introduc-
tion is a single clump about 10 m in diameter. The clump is centred on the original
planting site and is surrounded by S. x townsendii and Scirpus maritimus into which
it is slowly spreading. v.c. 27-28, Norfolk: material from America was planted at
Blakeney Point by Prof. F. W. Oliver in 1924. The plants died in 1925. v.c. 106, E. Ross:
introduced at Udale Bay, Cromarty Firth in 1947 or 1948.

Hitchcock (1951) considers that the taxon occurs throughout the southern native
range of the S. alterniflora complex. This complex extends down the eastern seaboard
of North America from Quebec and Newfoundland to Florida and Texas.

The synonymy of S. glabra is as follows:—

Spartina glabra Muhl., Desc. Gram.: 58 (1817).
Dactylis maritima Walt., Fl. Carol.: 77 (1788), non Curtis (1787).
Spartina stricta (Ait.) Roth var. glabra (Muhl.) Gray, Man., 2nd ed.: 552 (1856).
Spartina alternifiora Lois. var. glabra (Muhl.) Fern., Rhodora, 18: 178 (1916).
Spartina maritima (Curt.) Fern. subsp. glabra (Muhl.) St Yves, var. glabra St
Yves, Candollea, 5: 48 (1932).

REFERENCES

Hitcucock, A. S. (1951). Manual of the Grasses of the United States. Misc. Publ. U.S. Dept.
Agric., 200: 511-512.
MARCHANT, C. J. (1967). Evolution in Spartina (Gramineae). 1. The history and morphology
of the genus in Britain. J. Linn. Soc. (Bot.), 60: 1-24.
MARCHANT, C. J. & GOODMAN, P. J. (1969). Spartina we Muhl., in Biological Flora of the
British Isles. J. Ecol., 57: 295-297.
D. H. KENT

Watsonia, 8, 165-167 (1970). 165

Plant Records

Records for publication should be submitted to the appropriate vice-county Recorder and
not direct to the Editors.

Records are arranged in the order given in the List of British Vascular Plants (L.B.V.P.) by
J. E. Dandy (1958). Herbarium abbreviations are those used in British Herbaria by D. H. Kent
(1957).

The following signs are used :—

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

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

T 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.

9/1. CRYPTOGRAMMA CRISPA (L.) R.Br. ex Hook. 45, Pemb.: Glogue, Clydey.
On quarry waste. GR 22/221.327, T. A. W. Davis (68/1185), 1968, NMW. Second
vice-county record. (1969, Nature Wales, 11: 140).

25/1. POLYPODIUM AUSTRALE Fée *60, W. Lancs.: north end of Cringlebarrow
Wood, Yealand Redmayne. On north-facing, shady limestone outcrop. GR 34/49.75,
G. Halliday (6/69), 1969, LANC. Conf. J. A. Crabbe.

46/16. RANUNCULUS HEDERACEUS L. 104, N. Ebudes: Talisker, Isle of Skye. In
ruts of cart track near the farm. GR 18/32.30, G. Halliday (124/68), 1968, LANC.
This is the second record for the island.

102/4. RoRIPPA ISLANDICA (Oeder) Borbas *45, Pemb.: Castlemartin Corse.
On seasonally flooded spoil from a fen ditch. GR 11/90.99, T. A. W. Davis (68/1165),
1968, NMW. (1969, Nature Wales, 11: 140).

185/2. GENISTA ANGLICA L. 50, Denb.: bog near Llyn Creiniog, Llansannan.
GR 23/926.652, B. Seddon, 1962, field record. First post-1930 record.

192/19. TRIFOLIUM FRAGIFERUM L. *H15, SE. Galway: south side of Kilcolgan
river. In saline marsh below the bridge. GR 93/48, D. A. Webb, 1969, TCD.

~254/13. EPILOBIUM NERTERIOIDES Cunn. 45, Pemb.: Penrallt-y-gardde, Nevern.
GR _ 22/134.410, Miss Arundel, 1968, field record. (1969, Nature Wales, 11: 140).
Second vice-county record.

7425/1 x 425/2. MimuULUS GUTTATUS DC. x M. LUTEUS L. *H16, W. Galway:
southwest corner of Narrow Lake of Lough Mask near Drishaghaun. In marshy
ground by the lake shore. GR 94/03, G. C. Argent & G. Halliday, 1965, TCD. Det.
D. A. Webb. This specimen was originally published as M. luteus (1967, Proc. bot.
Soc. Br. Isl., 6: 396).

7425/2. MIMULUS LUTEUS L. [*H16, W. Galway: southwest corner of Narrow
Lake of Lough Mask near Drishaghaun. The specimen on which this record was
based has been redetermined as M. guttatus x M. luteus (see above).]

166 PLANT RECORDS

432/2. PEDICULARIS SYLVATICA L. subsp. HIBERNICA D. A. Webb *104, N.
Ebudes: Talisker, Isle of Skye. Common in moorland north of the farm growing
mixed with subsp. sylvatica. GR 18/32.30, G. Halliday (93/68), 1968, LANC, TCD.

4526/1. ANTHEMIS TINCTORIA L. *9, Dorset: Coldharbour, Sherborne. Top of
ditch in field. GR 31/646.172, A. C. Leslie, 1968, field record.

$58/1/40. HIERACIUM LANGWELLENSE F. J. Hanb. H16, W. Galway: one mile
northwest of Owenduff Bridge, southwest side of Lough Fee. Growing on rocks above
a waterfall. GR 84/83, G. Halliday (441/69), 1969, CGE, LANC. Det. P. D. Sell.
Second vice-county record.

558/1/154. HIERACIUM MACULATUM Sm. *9, Dorset: Lenthay district of
Sherborne. On a wall near the school playing fields. GR 31/633.161, A. C. Leslie,
1969, herb. R. D’O. Good. Conf. C. E. A. Andrews.

577/7. POTAMOGETON ALPINUS Balb. 44, Carm.: Taliaris Pool. GR 23/634.283,
B. Seddon, 1966, BM, NMW. Det. J. E. Dandy. Second vice-county record. This
specimen was originally determined as P. praelongus (1967, Proc. bot. Soc. Br. Isl, 6:
397).

577/8. POTAMOGETON PRAELONGUS WuIf. [*44, Carm.: Taliaris Pool. The
specimen on which this record was based has been redetermined as P. alpinus (see
above). There is no record of P. praelongus in Carmarthenshire.]

577/13. POTAMOGETON PUSILLUS L. *H15, SE. Galway: turlough, Race Course,
Gort. GR 93/47, D. A. Webb, 1969, TCD. Det. J. E. Dandy. *H16, W. Galway:
mouth of Eglinton Canal, Galway. GR 93/39, D. A. Webb, 1965, TCD. Det. J. E.
Dandy. Larger lake, Omey Island, northwest of Clifden. GR 84/63, D. A. Webb, 1968
TCD. Det. J. E. Dandy. Second vice-county record. *H17, N.E. Galway: River
Corrib, left bank, Galway. GR 93/39, F. H. Perring and D. A. Webb, 1968, TCD.
Det. J. E. Dandy.

577/16. POTAMOGETON TRICHOIDES Cham. & Schlecht. 52, Anglesey: Llyn
Coron. GR 23/378.699; Llyn Padrig, GR 23/364.727, B. Seddon, 1962, NMW.
Det. J. E. Dandy. These are respectively the second and third vice-county records. The
first record was in 1813.

577/19. POTAMOGETON CRISPUS L. 45, Pemb.: abundant in Stackpole Lake.
GR 11/97.95, T. A. W. Davis (68/1145), 1968, BM, NMW. Det. J. E. Dandy. (1969,
Nature Wales, 11: 140). This is the second vice-county record and the first since 1887.

577/21. POTAMOGETON PECTINATUS L. 45, Pemb.: near Ludchurch. GR 22/11,
D. A. Cadbury, 1966, BM. Det. J. E. Dandy. (1969, Nature Wales, 11: 140). This is
the second vice-county record. Abundant in Stackpole Lake. GR 11/97.95, T. A. W.
Davis (68/1143, 1144), 1968, BM, NMW. Det. J. E. Dandy. (1969, Nature Wales,
11: 140) This provides confirmation of Babington’s first Pembrokeshire record from
Stackpole Lake which does not appear to be substantiated by an herbarium specimen.

580/1. ZANNICHELLIA PALUSTRIS L. *H15, SE. Galway: south side of Kilcolgan
river. Growing in a small saline pool below the bridge. GR 93/48, G. Halliday (449/69),
1969, LANC, TCD.

589/3. POLYGONATUM MULTIFLORUM (L.) All. +45, Pemb.: Clarbeston Road.
On a roadside hedgebank. GR 22/011.207, O. Vaughan, 1968, NMW. Det. G. Ellis.
(1969, Nature Wales, 11: 140). This is the second vice-county record.

PLANT RECORDS 167

7605/2. JUNCUS TENUIS Willd. [*H27, W. Mayo: by the road linking Lough
Mask and Lough Nafooey (1968, Proc. bot. Soc. Br. Isl.,’77: 198). Although politically
in Co. Mayo, this site is in H16, West Galway. Juncus tenuis has not yet been recorded

from West Mayo.]

629/1. NEOTTIA NIDUS-AVIS (L.) Rich. *104, N. Ebudes: In a birchwood just
above the old track from Fearns to Rudha na’ Leac and Hallaig, Raasay. GR 18/53,
S. Wegmiiller, 1969, herb. Mrs C. W. Murray.

663/33. CAREX LASIOCARPA Ehrh. *50, Denb.: Llyn Creiniog, Llansannan.
GR 23/927.652, B. Seddon, 1962, NMW.

684/2. BRACHYPODIUM PINNATUM (L.) Beauv. *45, Pemb.: Broad Haven,
Bosherton.. A large patch on sandy peat overlying carboniferous limestone. GR
11/97.94, T. A. W. Davis (68/1179), 1968, BM, NMW. Det. G. Ellis. (1969, Nature

Wales, 11: 140).

168 Watsonia, 8, 168-180 (1970).

Book Reviews

Perspectives in Phytochemistry. Edited by J. B. Harborne and T. Swain. Pp. 235.
Academic Press, London. 1969. Price £3. 3s. Od.

The contents of this volume will be of much greater interest to plant biochemists
than to botanists, but they make it clear that recent advances in the chemical analysis
of plant substances are likely to uncover a whole new series of taxonomic characters.
At present, these characters appear to be more useful in demonstrating relationships
or lack of relationships between families than in distinguishing individual species.
On the basis of chemical evidence, Callitrichaceae, Hippuridaceae and Plantaginaceae
may all be closely related to the Scrophulariaceae, while there may be grounds for
dividing Cyperaceae into two families, based on the Scirpoideae and Caricoideae.

The book consists of ten chapters by experts on various aspects of plant biochemistry,
physiology and chemotaxonomy. Botanists will find food for thought in the general
reviews of Erdtman and Hegnauer, and in the provocative article on Monocotyledonous
plants by Bate-Smith. There is a chapter on the metabolism of cinnamic acid in fungi,
but no review of the recent developments in chemotaxonomy of lichens. An interesting
paper by Galston, based on detailed studies of etiolated pea seedlings, suggests that
flavonoids may have important physiological functions and are not simply waste
products of metabolism.

This book is well produced and not very expensive by modern standards. I did not
notice many misprints, but was surprised to learn that Hymenophyllum is in the
Pittosporaceae (p. 124). It should be consulted by all professional taxonomists who
are interested in phylogeny.

H. J. M. BOWEN

An Introduction to Plant Taxonomy. C. Jeffrey. Pp. vi +128. J. & A. Churchill Ltd.,
London. 1968. Price £1. 4s. Od.

In this book the author has set out to provide a true beginner’s introduction to the
classification of plants, demanding of his readers an absolute minimum of pre-
knowledge of technical terms and procedures. The first two chapters explain the
practical necessity for classification in very general terms and outline the concept of
relationships at varying levels of affinity. In chapters 3 and 4, we are introduced to
the hierarchy of units used in classification and a brief review is given of biological
factors such as reproductive isolation and hybridisation in relation to speciation.
In chapter 5, the theoretical problems involved in the formal processes of naming
plants are explained, while in chapter 6 the practical problems experienced in using
keys and other aids to identification are discussed. The book concludes with chapters
on the principal current systems for classification at higher levels and an attempt to
place the taxonomist in his context in society.

As an explanation to the student and the interested layman of the kind of work
that takes place in a major institute of taxonomic research, such as the herbarium
at Kew where the author is himself employed, this book is certainly a major success.
The clear picture it presents of modern work in the more traditional fields of systematics
should be of real value to the student, and it can be specially recommended to the
amateur botanist and the gardeners who frequently wonder why ‘they’ keep changing
the names of familiar plants.

On the debit side, we find that while subjects like keys and the rules of nomenclature
are excellently considered, little or no mention is made of the ‘modern’ tools of
taxonomy such as statistics, genetics, cytology and phytochemistry. This is a great

BOOK REVIEWS 169

pity, as I fear it may cause lecturers to underestimate the excellent qualities of this
book, when considering texts to commend to their students. In this respect, one is
reminded of the author’s recent review of a publication covering a rather similar field.
On that occasion Mr Jeffrey criticised the other author’s excessive preoccupation with
the more exotic manifestations of current work. In his own book, however, he has
failed to range far enough into the frontier regions of modern research.

There are a number of quite pleasing, though sometimes rather naive line drawings.
The artist seems more at home among the plants than the animals; thus the supposedly
human baby in plate 17 seems to have a head of hair and facial ‘stubble’ worthy of
a grown man! Several of the photographic illustrations are very poorly reproduced
and some of them merit serious criticism on other counts. Thus while plate 1 does
indeed show an excellently prepared and documented herbarium specimen, plate 2,
which claims to demonstrate ‘the housing of specimens in a modern herbarium’
actually illustrates Kew cabinets of 1932 vintage. Surely, even in a subject with its
roots as firmly in the past as systematic botany, 1932 can hardly be regarded as
modern. It would not have been difficult to obtain a photograph of cabinets embodying
the advantages that have become available through modern manufacturing techniques.
Plates 4 to 6 illustrate three very dismal specimens of members of the Cucurbitaceae.
While this choice, together with the special problems they illustrate, doubtless stems
from the author’s interest in and consequent expert knowledge of this family, I am
sure that it should have been possible to illustrate the topics so well discussed in the
text with some more attractive and better collected specimens. These pictures are
scarcely likely to improve the image of taxonomic botany, which I take to have been
one of the author’s intentions in writing this book.

While these criticisms have been quite severely stated, it is hoped that they will not
mask a genuine admiration for this very worthwhile introduction to a subject at present
poorly equipped with elementary texts.

J. F. M. CANNON

Plant Anatomy: Experiment and Interpretation. Part I. Cells and Tissues. Elizabeth G.
Cutter. Pp. vii + 168 with 83 figures. Edward Arnold, London. 1969. Price £2. 8s. Od.

Another book has been published on Plant Anatomy. One’s first reaction is to ask if
it is really necessary. This question is soon answered in the affirmative by a rapid
glance through Elizabeth Cutter’s book Plant Anatomy: Experiment and Interpretation.
Any lingering doubts are completely dispelled on a closer reading. The author
has set out to link results of experimental work with classical, descriptive anatomy,
in as far as such experimental data are available. This has proved a most instructive
exercise, and the book should encourage others to carry out research into some of
the many problems which remain to be answered. Indeed, it is refreshing to have a
text book on plant anatomy which does not give the impression that everything has
been done already (probably in Germany in the late 19th century!).

Whether used as an introductory text by the advanced student or interested layman
or read by comparative anatomists, this book is sure to prove of value and interest.

The presentation of the text is clear and concise, and the illustrations are ample.
By an obvious slip, Robinia is described as having a non-storied cambium (fig. 11.1).
Apart from this, very few mistakes of any consequence have been noted.

This is the first volume of two, and deals with cells and tissues. I look forward to
seeing volume 2, on organs.

D. F. CUTLER

Field Natural History—A Guide to Ecology. Alfred Leutscher. Pp. 248 + 83 text
figures. G. Bell & Sons Ltd., London. 1969. Price £3. Os. Od.

The number of ecology books that have come from the publishing houses in recent
years has increased. The majority of these, however, has been college and university

170 BOOK REVIEWS

textbooks, usually of a high standard but often too advanced for the amateur
naturalist—certainly so if he be only at the beginning of his interest in this discipline.
Those written especially for his use have been fewer in number and most have fallen
far short of providing him with the wide, yet reasonably deep, background to the
subject that he needs.

Alfred Leutscher now provides a book with the right requirements, the text of which
bears his unmistakable stamp of conciseness and clarity that has become polished
and perfected with his years as Chief Guide Lecturer to members of the public visiting
the British Museum (Natural History).

The author has arranged his text-matter in three parts. In the first, which is headed
‘The Organism and its Environment’, he describes under four sub-headings the form
and function of both animals and plants, how they have evolved, the nature of soils
and rocks, and the ecological factors of their environment. In Part 2 (which to my
mind has been extremely well written) all the major habitats found within the British
Isles are described. Lastly, Part 3 is entitled ‘Field Work’ in which, for most of the
projects suggested, Mr Leutscher has drawn upon his broad experience of survey work
in Epping Forest, near his home.

Nowadays it is unusual for a book of this kind to be published without at least
some coloured photographs. In fact there is a tendency for authors to lean somewhat
heavily on such illustrations, often to bolster up what is in fact the inadequacy of their
own text. This is not so with Mr Leutscher, for throughout his book the 80-odd figures
drawn by himself (some, albeit, better executed than others) succeed well in com-
plementing what he has written.

Some useful appendices follow the main text, covering such diverse topics as instruc-
tions for making plaster casts of animal footprints or cleaning a skeleton, a list of
Epping Forest fungi, guidance on making an herbarium, packing and sending animals
by post, and measuring the height of a tree!

Eight pages giving a bibliography (listing works mostly still in print) and a useful
address list of societies round off what is an excellent book for the botanist, zoologist
or general naturalist seeking ecological guidance and well worth the sixty shillings
asked.

E. W. GROVES

Synékologische Studie der Waldgesellschaften auf Amphibolitgestein. Vegetace CSSR
A3. M. Husova. Pp. 188. Czechoslovak Academy of Sciences, Prague. 1968. Price
Kés 36,00.

This is one of an extensive series of volumes on the vegetation of Czechoslovakia. It
deals with a vegetation type that is little represented in Britain—semi-natural wood-
land on base-rich, non-calcareous rock. An account is given of one particular hilly
area in south-west Bohemia, long known for its species-richness—the Branzovsky
hvozd. The major bedrock is a slowly-weathering amphibolite (a metamorphic rock
with hornblende and plagioclase as major constituents); it has a high content of
calcium and magnesium. There are several dominant tree species, of which Tilia
platyphyllos is perhaps the most interesting. Many of the woods are developed on
screes with ranker soils; flatter areas have eutrophic braunerde. Comparisons are
made with woodland on nearby siliceous rocks with oligotrophic braunerde. The
account contains many physical and chemical analyses of soils, extensive phyto-
sociological accounts of the major associations, some data on the activities of soil
micro-organisms and comments on the role of molluscs in making calcareous soil-
horizons over base-rich but non-calcareous rocks. There are discussions of the classi-
fication of the associations in the Ztirich-Montpellier system and of the effects of
amphibolite on the composition and growth of the vegetation. The top soil on
amphibolite has a pH of 4.7-7.0 (mostly 5.0-6.0); the amounts of exchangeable cations

BOOK REVIEWS 171

per unit dry weight vary enormously with the humus content. Calcifuges such as
Calluna, Vaccinium and Deschampsia flexuosa are almost completely absent. Many
calcicoles are present, e.g. Daphne mezereum, Inula conyza, Polygonatum odoratum,
Rhamnus catharticus and Viola hirta. The author goes some way to explaining the
absences and occurrences of these species but much remains to be established. She
incidentally gives us ideas on what our forests on base-rich rocks might have been
like before they were destroyed and the Tilia spp. in particular lost (cf. Pigott, C. D.
(1969). The status of Tilia cordata and T. platyphyllos on the Derbyshire limestone.
J. Ecol., 57: 491-504). Altogether there is much in this attractively produced book to
interest the British botanist.

P. J. GRUBB

Flora of Derbyshire. Edited by A. R. Clapham. Pp. 382 with a coloured frontispiece,
8 plates and 7 pages of maps. County Borough of Derby Museum and Art Gallery,
1969. Price £2. 5s. Od.

This Flora, which was started by a committee of 13 in 1949, is a welcome contribution
to our knowledge of this floristically particularly interesting county. The previous
Flora (Linton 1903) was of a standard far beyond many of its date, and so, as is noted
in the Introduction, ‘. . . it is perhaps not difficult to understand why his Flora of
Derbyshire has been neither revised nor replaced in all the sixty-five years since it
appeared.’ Linton’s Flora had, however, become sadly outdated, as is indicated by the
addition of 135 species and hybrids (excluding Rubus and Hieracium) in the new Flora.
Of these only 29 have reasonable claim to be regarded as ‘native’, but Aceras anthro-
pophorum, Anagallis minima, Aphanes microcarpa, Galeopsis bifida, Hymenophyllum
wilsonii and Vaccinium xintermedium are particularly gratifying additions. On the
debit side, however, 55 species (including 26 which may have been native) have not
been refound in recent years, and others (e.g. Huperzia selago, Ranunculus arvensis)
appear to have declined in numbers.

Polemonium caeruleum provides the coloured frontispiece (photograph by Mr
R. H. Hall), and precedes six introductory chapters (Introduction, Geology, Geomor-
phology, Climate, Vegetation, and Geographical Distribution) which give a useful
body of background information to the Flora. The Introduction itself summarises
botanical studies in the county since 1903 and includes a biographical account of
Dr E. Drabble. A more complete list of contributors would have been welcomed here
as an acknowledgement particularly to the many amateurs who have sent lists of
species into the committee over many years.

The chapter on Geology is excellent as a geological account but much of the data
is hardly pertinent to flowering plants: not many species have distributions correlated
with particular fossil facies of the Coal Measures! It is also perhaps unfortunate that
relatively little attention is given to the superficial deposits which are so much more
important in this context. A map of the ‘drift’ as well as the solid geology would have
been valuable. The list of references is rather incomplete and the omission of Edwards
& Trotter (1954), Ford & Mason (1967) and Bradley & Ford (1968) are particularly
unfortunate for anyone wishing to learn more about the geology of the county. The
section on Geomorphology is better but again suffers from a tendency to irrelevancy.
The inclusion of a map of the river systems is a particularly welcome feature. The
Flora was very fortunate in securing the services of Professor A. Garnett for the
chapter on Climate, and her thoroughness makes this a useful reference work for
biologists and meteorologists alike. One does, however, wonder whether the citing of
month by month figures of air pollution for only one year was necessary. Such figures
are So variable that mean values calculated over the last 3-4 years might have been of
more significance. The observation that ‘. . . some farmers record the unduly rapid
corrosion of fences and farming implements, even in the heart of the High Peak...’
(p. 31) should not pass underemphasised, and it is interesting that the distribution of

172 BOOK REVIEWS

lichens in the county (see Hawksworth 1969) correlates with the view of high air
pollution in non-urban as well as urban areas.

The chapter on Vegetation, by Dr T. T. Elkington, bears testimony to the tradition
of plant ecologists at Sheffield University, and will be particularly valuable for future
generations as it includes 84 detailed Habitat Studies. The use of some approximately
quantitative scale rather than the categories ‘abundant’, ‘frequent’, ‘occasional’
and ‘rare’ might have been an improvement here and one is struck by the very few
records of bryophytes and lichens in the lists, although these appear to be very
important constituents of the vegetation on some mor and rendzina soils. It is also
regrettable that there is no easily readable summary of the vegetation types and that
the amateur must still look to Hall (1962), pp. 69-95 for an introduction to the county
of this type. The phytosociological classification of the vegetation of the limestone
dales by Shimwell (1968a, 1969) might also have been referred to, at least in passing.
Dr Elkington also prepared the chapter on Geographical Distribution, which is particu-
larly good. Maps of representative species in the county are printed adjacent to ones
of the same species from the Atlas of the British Flora. Although only 17 species are
treated in this way, they were evidently very carefully selected and admirably suited
to this type of presentation which renders the chapter much more informative than the
pages of small intra-county distribution maps which have swelled several recent
county Floras.

Having, on page 96, reached the end of the introductory chapters one cannot help
feeling that at least some of this space might have been better employed: no dis-
cussion of the effects growth, and extent of urbanisation and industrialisation is given,
and some indication of built-up areas on one of the seven maps of the county included
would have been helpful to those unfamiliar with the county. It is also rather surprising
that no chemical analyses or classification of soil-types is given, although most of the
data needed for this has been published by Bridges (1966); that there is no data on
agriculture and forestry; no information on the extent of woodland; and no biblio-
graphy. The only papers cited are those specifically referred to in the introductory
chapters and a list of Drabble’s scattered works. Data included in the Biological Flora
of the British Isles, and other papers (e.g. Anderson 1965; Mills 1969; Shimwell
1968b) would have been a useful continuation of the annotated bibliography given by
Linton (1903).

The Flora itself is based on the 10 km square divisions of the National Grid, and
localities are given only for species which are not regarded as ‘common’. One locality
in each 10 km square is given for ‘less common’ species, and full lists are given for
‘rare’ ones. Beneath the data on each species ‘Linton: followed by a list of localities
appears. A comparison with Linton (1903) shows that these are localities given by
Linton in which the species has not been refound, and that they are not a complete
list of Linton’s sites as may at first appear. Linton did, after all, know more than one
locality for Pinus sylvestris! 1 km or 10 km square grid references are given after every
locality name, which is perhaps rather unnecessary for sites which only fall in 1 km
square. An appendix with localities and six figure references might have been an
improvement. The citing of four figure references together with the punctuation system
adopted leads to some ambiguities: e.g. ‘Fenny Bentley to Bradbourne 1950, 1850,
Black Rocks 2955’ (p. 345)—if one does not appreciate that dates are generally
omitted!

For the bulk of the records there is no indication when between 1903 and 1969 they
were made. Thus Hypericum elodes is stated to be ‘perhaps extinct’ (p. 141) although
an undated ‘recent’ record is given. It would also have been helpful to know who
determined the specimens, where the herbarium sheets supporting them are, what
herbaria have been searched, and what records are based only on published sources.
It seems that if the master set of record cards housed in Derby Borough Museum and
Art Gallery is lost or damaged, much valuable information particularly interesting to
the specialist might be completely lost. These deficiencies render this section scien-

BOOK REVIEWS 173

tifically of a lesser standard than that set in the recent Floras of Berkshire (Bowen,
1968) and Norfolk (Petch & Swann 1968).

The nomenclature is stated to follow that of Flora Europaea yet, for example, the
genus Lycopodium is retained and the genera Huperzia, Diphasium and Lepidotis are
not mentioned.

A summary of numbers of species reported from each 10 km square would have
been valuable, and perhaps have emphasised underworked areas. One cannot avoid the
impression that the coverage of the county for critical taxa, especially in the lowland
areas, is not as complete as it might have been. This is indicated by the long lists of
localities of Rubus species given by Linton in which they have not been refound; and
the ease with which one can add new 10 km square records for species (e.g. Geranium
dissectum SK 358518, 1965, Hawksworth; Potentilla anglica, SK 358512, 1964-1966,
Hawksworth; P.xsuberecta, SK 358512, 1966, Hawksworth LTR). In the limestone
areas also, some substrates have been underworked, such as peat pockets (e.g. Blechnum
spicant, Eriophorum angustifolium, Molinia coerulea at SK 180650, 1968, Shimwell),
gritstone inliers (e.g. Stanton Moor) and chalybeate springs (e.g. Lotus uliginosus,
Viola palustris at SK 216635, 1968, Shimwell). Dr D. W. Shimwell also reports that
Gagea lutea still occurs at Meadow Place Wood and in the Bradford Valley where it
was reported by Linton (1903), although no post-1903 records are given from these
sites in the new Flora.

The introduced species included appear to have been rather arbitrarily selected, and
one wonders why, when Abies alba is included, Larix decidua and Picea sitchensis are
not.

A good feature is the 16 splendid monochrome photographs which were mainly
provided by Mr R. H. Hall, one of the foremost amateur botanists to have worked in
the county.

Printing and proof reading errors are inevitable in a book of this length, and are no
more numerous than to be expected. Letters of the wrong fount are the most frequent
error of this type, except for the omission of distributional data from Linton (1903)
for some species. We are not told whether Arabis caucasica, Draba muralis or Ilex
aquilifolium, for example, were found by Linton or not, and it is therefore often
necessary to use both Floras in conjunction to obtain a picture of the history of a
particular species in the county.

The binding and covers are good, and this Flora is an essential addition to the shelves
of any botanist interested in the county. It is perhaps unfortunate that more fieldwork
was not done, that herbaria were not searched, more botanists consulted, and that
some indication of the dates of records was not given, but, if all these points had been
adequately attended to (and the introductory chapters amended), then this badly
needed Flora might not have appeared for several more years. At least we now have
a fairly comprehensive account of the flora of this county which will provide a basis
for the future studies of both amateur and professional alike for many years to come.

REFERENCES

ANDERSON, D. J. (1965). Studies on structure in plant communities. I. An analysis of limestone
grassland in Monk’s Dale, Derbyshire. J. Ecol., 53: 97-107.

Bowen, H. J. M. (1968). The Flora of Berkshire. Oxford.

BRADLEY, P. C. S. & Forp, T. D., ed. (1968). The Geology of the East Midlands. Leicester.

BripGEs, E. M. (1966). The Soils and Land Use of the District North of Derby. Harpenden.

EDWARDS, W. & TROTTER, F. M. (1954). British Regional Geology: The Pennines and Adjacent
Regions, 3rd ed. H.M.S.O., London.

Forp, T. D. & Mason, M. H. (1967). Bibliography of the Peak District of Derbyshire (to
1965). Mercian Geol., 2: 133-244.

HALL, R. H. (1962). The Major Plant Associations, in EDWARDS, K. C.,ed. The Peak District,
pp. 69-95. London.

HAwkswortTH, D. L. (1969). The lichen flora of Derbyshire. Lichenologist, 4: 105-193.

174 BOOK REVIEWS

LINTON, W. R. (1903). Flora of Derbyshire. London.

MILs, J. N. (1969). A new species of Hieracium in Derbyshire. Watsonia, 7: 40-42.

PetTcH, C. P. & SWANN, E. L. (1968). Flora of Norfolk. Norwich.

SHIMWELL, D. W. (1968a). The Phytosociology of Calcareous Grasslands in the British Isles.
Ph.D. thesis. University of Durham.

SHIMWELL, D. W. (1968b). Notes on the distribution of Thlaspi alpestre L. in Derbyshire.
Proc. bot. Soc. Br. Isl., 7: 373-376.

SHIMWELL, D. W. (1969). The Vegetation of the Derbyshire Dales. Nature Conservancy,
Shrewsbury (cyclostyled).

D. L. HAWKSWORTH

Plant Breeding. W. J. C. Lawrence. Pp. 56. Studies in Biology no. 12. Edward Arnold
(Publishers) Ltd., London. Price 14s. Od. (boards), 8s. 6d. (paper).

This study of Plant Breeding is one of a series of publications sponsored by the Biologi-
cal Education Committee with an educational purpose based on a biological theme
and dealing with a rapidly developing discipline of utmost importance to man’s
survival.

The text is set out in a concise and logical manner beginning with the earliest and
unconscious influences of primitive man in his development of cultivated plants from
weedy species and culminating in the enormous impacts of present-day manipulation
of genetics and selection in plant breeding and crop improvement. A final plea is
made by the author for the future of plant breeding and he emphasises the need for
broadening the genetic basis for breeding work and maintaining a much more adequate
gene pool for commercial food crops than has been done in the past. The dangers of
narrow selection methods and lack of variability are strongly emphasised and their
consequences for the future world food problem are pointed out.

Chapters on Breeding Aims, Breeding Methods and Breeding Principles are clearly
and concisely sectionalised and material under each sub-heading is written in an
interesting and informative manner, with due regard shown for the agricultural
economic applications which are so much a part of plant breeding today. A nice
balance is maintained between principles and practice as indeed is desirable in a subject
where, as the author intimates, broad principles have been neglected in the past.

Although the book deals with a somewhat specialised and applied aspect of genetics,
all the principles of cytogenetics and hybridisation are involved and this is clearly
an introductory treatment of plant breeding for the more advanced student and the
teacher who are already conversant with the elements of such disciplines. Indeed
there is no definitive list of technical terms used, although many are defined in the text.

Illustrations are for the most part kept simple and often self-explanatory and hence
are effective in emphasising points in the text. Happily, the author has avoided complex
numerical data with which the student might be overwhelmed and confounded.
Obviously the aim throughout the book has been to maintain simplicity as far as
possible and he has for the most part been successful in making this material easily
assimilable by the non-specialist.

This is an adequately illustrated and generally acceptable little book of reasonable
value at the price of 14/-, which should not be too hard on the pocket of most students

of biology. ae
. J. MARCHANT

Developmental Plant Anatomy. Alan Gemmell. Pp. iv + 60.

Plant Symbiosis. George D. Scott. Pp. iv + 58.

Fungal Parasitism. Brian J. Deverall. Pp. iv + 57.

All Edward Arnold, London. 1969. Price 14s. (boards), 8s. 6d. (paper).

These latest volumes in the Institute of Biology’s series ‘Studies in Biology’ continue
to uphold the reputation that the series has earned for dealing with biological topics

BOOK REVIEWS is

in a concise and contemporary way for use in schools and elementary university
courses. Each of them gives, in relatively few pages, a sound introduction to the subject
and brings the reader up to date with modern views and discoveries, in each case

citing recent literature.
N. K. B. ROBSON

The History of British Vegetation. Winifred Pennington (Mrs T. G. Tutin). Pp. vili+
152, with 30 plates + 32 text figures. The English Universities Press Ltd., London.
1969. Price £1. 8s. Od.

Dr Pennington is to be congratulated upon her History of British Vegetation, in which
a great deal of ground is covered in what she calls a small book. It has grown out of a
course given to students in the University of Leicester and is offered ‘as an introduc-
tion to those two great classics of British botany, Professor Harry Godwin’s History
of the British Flora, and the late Sir Arthur Tansley’s The British Islands and their
Vegetation’. If the book must be called an introduction, it is an impressive and com-
prehensive one, dealing with a subject of great biogeographical interest and significance.

‘All history must begin somewhere’, says Dr Pennington and in chapter 1 she
carries the reader back into the mists of antiquity some 60,000,000 years ago, when,
in early Tertiary times, the vegetation of south Britain was not unlike that of the
Indo-Malayan region of today. By the end of the Tertiary period, however, in the
Upper Pliocene, the picture had altered profoundly from that shown in Plate 1, a
reconstruction of the landscape in London Clay times. While no plant fossils of
Pliocene age have been found in Britain, there is evidence from Continental sources
in support of the view that the flora of Britain during this period was rather similar
to that of the present day. At the opening of the Quaternary there were comparatively
few genera not now represented in the flora of Europe. But further changes were on
the way.

Our modern flora is essentially a small outlying portion of the European flora,
but its history has been remarkedly varied throughout the Pleistocene and indeed
throughout the whole of the Quaternary. Several chapters of Dr Pennington’s book
are devoted to an account of the changing nature of flora and changing pattern of
vegetation during the Pleistocene period, including not only stratigraphical detail
but also an examination of the factors involved in these changes. Glacial and inter-
glacial episodes are covered in Chapters 2, 3 and 4, and it becomes apparent that the
last Full-glacial period is of particular interest from the historical point of view.
The so-called ‘Arctic Beds’ of the Lea Valley and Barnwell Station in Hertfordshire,
which belong to this period, have furnished plant remains suggesting a vegetation
rather resembling grass-sedge tundra, devoid of trees apart from dwarf willows and
arctic birch, and including a significant number of other arctic-alpine species. Of these
two fossil floras, that of the Barnwell Station Beds is the more recent and is dated
17,550 B.c.

Then followed the Late-glacial period which is equally interesting and suggestive
in its floristic history. To the deposits and sediments of this and later periods the
modern technique of pollen analysis has been applied, while the radiocarbon method
of dating has provided reliable information about the age of the deposits. The period
is divided into the three pollen Zones I, II and III, Zone II being the relatively short
and mild Allergd interstadial. While the zones show noticeable differences in flora
and vegetation, it is clearly established that, in general, the flora of Late-glacial times
was a relatively rich one, including not only arctic-alpine plants such as Dryas octo-
petala, Salix herbacea, Empetrum nigrum and Betula nana, but also tree birches, at
least during the Allergd phase, together with many other species now found in open
habitats and commonly referred to as weeds or ruderal plants.

This Late-glacial flora was largely disrupted and many of its members restricted

176 BOOK REVIEWS

in range by the pressure of events during the following Post-glacial period, the transi-
tion to which is recorded between Zones III and IV around 8000 B.c. The change
was marked by a pronounced amelioration of the climate and the rise in temperature,
along with increasing stabilisation of soils, permitted the growth and relatively rapid
spread of trees and shrubs, birch being one of the pioneers, followed by pine, hazel,
elm, oak, ash, alder and lime. The climatic optimum was reached about 3000 B.c.
when mixed deciduous forest occupied much of the country, ascending to at least
2,500 ft, although over a large part of Scotland mixed forest was probably replaced
by birch and pine. Whatever the maximum extent of forest in the British Isles when
it reached its ‘climatic climax’, little of it was to survive. A deterioration of the climate
contributed to the decline of forest, but other complex factors were involved, edaphic
and biotic, and not least of all was man’s onslaught on the natural resources of the
country.

In Chapters 5—9 Dr Pennington traces these Post-glacial changes in considerable
detail and stresses the anthropogenic factor. She makes use of Blytt and Sernander’s sub-
divisions of the period into Pre-Boreal, Boreal, Atlantic, Sub-Boreal and Sub-Atlantic.
These divisions are not only related to pollen zones, but correlations with archaeo-
logical periods are clearly indicated. Interest will be aroused by the author’s reference
to the Boreal-Atlantic transition, at about 5500 B.c. as one of the most significant
horizons in the history of British vegetation. ‘Not only does it mark what appears
to have been one of the most pronounced of the Post-glacial climatic shifts, but it
marks the last occasion on which most plants could disperse into Britain by natural
processes. The flora at this time represents the true ‘native flora’, and subsequent
additions to the flora fall into two categories—a very small group which have a natural
dispersal mechanism capable of leaping sea barriers, and a rather large group of
plants which, on all available evidence, appear to have been introduced by man,
deliberately or accidentally.’ Generations of botanists have repeatedly discussed such
questions and I am reminded of the verdict of one of them. In an essay on the origin
of the flora of Perthshire written in 1891, Buchanan White had this to say: ‘If indi-
genous means, as it really does, aboriginal or autochthonous, we have no indigenous
plants.’ In the light of present knowledge this may now be regarded as an extreme
view, although the complete extermination of our flora during the last ice age was
postulated by several earlier writers.

The influence of man on flora and vegetation in Britain can be traced as far back
as early Neolithic times, about 3000 B.c., when he commenced the process of forest
destruction by axe or fire or both. The impact of different human cultures has continued
with varying intensity to the present day, and while Dr Pennington’s book is not
immediately concerned with the composition of our modern flora or with the status
of its members, it contains frequent references bearing upon these matters, a large
number of plants being referred to specifically.

Existing vegetation is dealt with briefly in Chapters 10 and 11, the former being
devoted to plant communities of dry land—forests, grasslands, heaths and coastal
habitats, while the latter is concerned with wetlands, such as fen, moor and bog.
Different examples are cited from different parts of the country and the approach is
again historical. Pine and birch have been in Britain for more than 9,000 years, and
in marked contrast to this beech is a relatively recent immigrant, its pollen having
been found only in Zone VIII, from about 500 B.c. onwards. The growth and extension
of blanket peat can be traced, at least on the Pennines, to the beginning of the Atlantic
period, when Britain became separated from the Continental mainland about 5500 B.c.
and when the climate probably became more markedly oceanic.

A good deal of British vegetation can be described as oceanic and is well represented
in Ireland and elsewhere. References to Irish vegetation and Irish plants are numerous,
including the problematic ‘Lusitanian’ element, and although the history of the
group is uncertain it is suggested that it may have migrated into south-west Ireland
in early Post-glacial times along oceanic coastal fringes now submerged. I doubt if

BOOK REVIEWS 177

all its members shared the same history, but evidence from fossil remains of the plants
is still incomplete.

The final chapter of Dr Pennington’s book is devoted mainly to a few plants which
are essentially ‘Highland’ and their present distribution is shown in maps reproduced
from Atlas of the British Flora. They are mostly arctic-alpine, and in view of their
history within the British area, they may well be called glacial relic species. It would
have added greatly to the interest of the maps had they shown also the known fossil
records of these same species from glacial or other deposits. In addition to the maps
and other text figures, the book is well illustrated by numerous photographs, although
a few of these are a little lacking in definition. References to literature take the form
of a list of books and published papers for ‘Further Reading’ arranged in sections,
but a few references in the text do not appear in the list. At least I could not find
them and their omission may be intentional.

It may be only a coincidence that this book has appeared when European Conserva-
tion Year was about to begin. All who claim the need for conservation will find much to
contemplate and the opponents of conservation should take heed of the message
which the book has to tell. To British botanists with an interest in history, be they
taxonomists or ecologists or both, the book will make an immediate appeal. It is well
produced and at 28 shillings is not expensive.

J. R. MATTHEWS

Waldgesellschaften des mitteleuropdischen Gebirgsraumes nordlich der Alpen. F. K.
Hartman and G. Jahn. Pp. 636; 37 tables + 2 maps in separate cover. Gustav Fischer,
Stuttgart. 1967. Price DM. 148.

This work summarises a very large amount of information on the floristic composition
of the forests of the mountainous regions of central Europe north of the Alps. It
consists essentially of two series of tables with detailed explanations, a set of tables
presenting nearly 1,300 original floristic analyses, principally by Prof. Hartman, from
a wide variety of forest types, and a set of constancy tables summarising the available
floristic information (from the authors’ own data and the published work of many
other phytosociologists) on central European forests. The area covered extends from
the Jura, Vosges and Ardennes to Lower Austria and the Sudeten region, and from
the hills of Lower Saxony to the Alps. A large proportion of the data are from western
Germany.

The methods of collection and presentation of the data follow the usual Braun-
Blanquet lines. The authors have been at pains to relate their vegetation units to
altitudinal zonation and, in general, each table embraces a group of physiognomically
and ecologically related associations, often differentiated geographically. Indeed,
the broad geographical comparison and integration of the data is one of the most
valuable features of this book. Detailed soil analyses (pH, exchangeable Ca, Mg, K,
citric acid-soluble phosphate, loss on ignition, mechanical analysis, etc., often from
several horizons in the profile) are given from representative sites from most of the
original tables. The text volume concludes with a 39 page bibliography, an index of
plant communities and a list of Polish and Czech equivalents of place-names.

This volume is to be followed by two further volumes, which will deal with ecological
questions and applications to forestry and land-use.

M. C. F. PROCTOR

The Evolution and Classification of Flowering Plants. Arthur Cronquist. Pp. xii + 396.
Nelson, London. 1968. Price £2. 5s. Od.

The appearance of another system of Angiosperm classification in the wake of those
recently published by Hutchinson, Takhtajan, Thorne (in outline only as yet) and
Others at once indicates that an agreed classification of the Angiosperms is still a

178 BOOK REVIEWS

distant goal and emphasises the peculiar fascination that the inter-relationships of
this group of plants have held for systematic botanists. The first reaction to a new
system is inevitably ‘How does it stand in relation to the well-known ones?’, to
which the reply in this case must be that it is not revolutionary, but builds a new
superstructure on an old foundation. Dr Cronquist has based his system on that of
Bessey which, in turn, was intended to express the Bentham & Hooker classification
in evolutionary terms. He has attempted ‘to develop a general system of Angiosperms
compatible with presently available information, and to put that system in a proper
theoretical frame of reference’. The available information includes some that has not
previously been incorporated into a general system, such as the sequence of stamen
development, the distribution of floral nectaries and the number of ovule integuments,
and the whole is treated from a frankly evolutionary standpoint. Among recently
published systems, the present one most closely resembles that of Takhtajan; and
this similarity ‘reflects a community of interest and outlook, bolstered by a mutual
exchange of views over the past ten years’. The Takhtajan and Cronquist systems are
not identical, however: Cronquist admits to being less of a splitter than Takhtajan.

The author prefaces the account of his system by three chapters which may be
regarded as a sort of ‘credo’, as they contain the principles and arguments that he
has employed in working out his system. The first one, “Taxonomic Principles’,
indicates that he believes in the existence of a Natural System, based on multiple
correlation and reflecting evolutionary relationships—a belief that is not fashionable
in some quarters today. Among several interesting concepts that are discussed, one
may mention ‘The monophyletic requirement’ and ‘Parallelism’ as particularly
noteworthy. Cronquist admits that a completely monophyletic system is probably
unattainable, so that taxonomy ‘can provide only a somewhat muddy reflection of
evolution’. He regards evolutionary parallelism as a help to taxonomic work rather
than a hindrance (the usual view), because it tends to indicate relationship, and from
this principle develops a telling argument in favour of mutation pressure (orthogenesis)
rather than natural selection as the cause of the evolution of major group characters
in the Angiosperms.

In the second chapter, ‘The Origin of the Angiosperms’, Cronquist dismisses the
Pinicae and Gneticae as possible Angiosperm ancestors, leaving only the Cycadicae
as contenders for that title. Within the Cycadicae only the pteridosperms (Lyginop-
teridales) appear to be suitable for further consideration. Having selected the pterido-
sperms by a process of elimination, he however baulks at ‘going the whole hog’ by
fixing specifically on the Glossopteridae, as some authors have done recently, but
merely refers to the idea as ‘interesting’.

The third chapter is devoted to a comprehensive review of “The Evolution of
Characters’, dealing first with the concept of primitive characters, then with the
characters which, in the author’s opinion, primitive angiosperms must have possessed,
and finally tackling the vexed question of how to interpret evolutionary trends. In
contrast to some modern workers, Cronquist does not regard such interpretation as
a hopeless task. Unlike those who stress the dangers of either constructing imaginary
trends or misinterpreting real ones by ‘starting at the wrong place’, he regards the
recognition of evolutionary trends as a legitimate and helpful taxonomic exercise,
but points out that many trends seem to be the result of ‘genetic predisposition’
rather than natural selection.

Not all the interpretations of primitive characters and the directions suggested for
trends will meet with unanimous approval, but all are cogently argued and worthy
of consideration. However, while it is understandable that Cronquist should be luke-
warm to a rival theory of Angiosperm evolution such as the Gonophyll Theory, it is
hardly sufficient to dismiss it in a sentence as something that may possibly result in
modified concepts in the future. The facts on which that theory is based merit serious
consideration now. The rest of the book consists of an outline of the system, with keys
to the subclasses, orders and families that are meant to be conceptual aids rather than

BOOK REVIEWS 179

means of identification, as well as discussions of relationships and explanations of
details of the system that make fascinating reading. Useful bibliographies are provided
for each order.

Cronquist makes much of the direction of stamen maturation in the flower, using
this character to distinguish his Subclass Rosidae (centripetal stamens) from the
Subclasses Caryophyllidae and Dilleniidae (centrifugal stamens). Leins (1964), however,
has shown that the direction of maturation depends on the side of the androecial
primordium on which the stamens arise. If they arise on the outer side, development
is centrifugal, if on the inside, then it is centripetal. This observation suggests that the
direction of maturation is not such a fundamental character as it would appear,
because a trend towards perigyny or epigyny (as in the Rosales and Myrtales) could
result in a change from centrifugal to centripetal development. Thus, if the sequence
of stamen development is the crucial fact in allocating a family to one or other subclass
(as it appears to be in separating the Myrtaceae (Rosidae) from the Lecithydaceae
(Dilleniidae)), then perhaps too much weight has been placed on this one character.

In several places Cronquist refers to grouping of stamens and the occurrence of
stamen trunks as constituting an intermediate stage in the reduction from numerous
to few stamens, refuting both the Telome Theory (in which the stamen groups are
regarded as primitive) and the Dédoublement Theory (in which they are regarded as
multiplications of single stamens). He is certainly wrong to interpret the trends in the
androecium of Hypericum in this way, and would find it difficult to give a satisfactory
explanation of androecial trends in some other taxa, e.g. the Malvales.

Despite the occurrence of several such statements and assumptions which appear
to be unwarranted, this is a most attractive book and should be read by all who are
interested in Angiosperm taxonomy, in particular those who wish to discover how
modern developments in morphology, cytology, embryology, etc. can be accom-
modated in the system. Apart from an annoyingly large number of misprints and a
few omissions (e.g. the footnote on p. 8 and definitions of ‘introrse’ and ‘extrorse’
from the glossary), it is well produced and, at 55/-, not expensive by modern standards.

REFERENCE
Les, P. (1964). Das zentripetale und zentrifugale Androecium. Ber. dt. bot. Ges., 77: 22-26.
N. K. B. ROBSON

Plant Variation and Evolution. David Briggs and Max Walters. Pp. 256 + 22 colour
plates + 61 figures. World University Library. Weidenfeld & Nicolson, London. 1969.
Price £1. 15s. cased; 18s. paper.

This is a most attractive book, elegantly written and beautifully produced.

It starts with the history of the study of plant variation and taxonomy, followed
by an account of the beginnings of quantitative descriptive methods and elementary
biometry. Mendelian genetics is outlined, again from a historical point of view, and
is brought up to date with an account of modern ideas about population genetics
and natural selection.

The various breeding systems found in plants are discussed, and this is followed
by a chapter on definitions of species, and one on infraspecific variation and its adaptive
significance. The distinction between gradual and abrupt speciation is emphasised,
and finally the complications due to polyploidy and hybridisation are discussed.

There are two continuing themes throughout the book. One is the historical back-
ground of the topic under discussion, and the other is its significance in the study of
evolution. Most of the text is concerned with clear exposition rather than with con-
troversy, but where there is any doubt about the proper interpretation of any particular

180 BOOK REVIEWS

piece of data this is always pointed out, and the impression given is one of fair and
balanced discussion.

I applaud the authors’ courage in pre-empting the term ‘species’ for taxonomic use,
in the interests of clarity. Geneticists in the habit of maintaining that the breeding
unit is the only true species are asked to make do with ‘hologamodeme’. One can
only hope that this clear and impartial account of the relation between the two will
help to keep the temperature down. !

One of the striking features of the book is the consistent use of the Deme terminology
of Gilmour, Gregor and Heslop-Harrison. This attempt is successful because it
concentrates on the more useful and clearly defined terms such as gamodeme, topo-
deme and cytodeme. The avoidance of this terminology by the majority of authors
may well be due to the difficulty of using such terms as genoecodeme with precision,
and in particular with the impossibility of incorporating continuous variability into
the system.

The only criticism which could perhaps be made about the authors’ use of these
terms concerns gamodeme, in the sense of the mendelian population of the geneticist.
It is used consistently in this sense, and there is even a section on “The delimitation
of gamodemes’, but the extreme difficulty of defining the extent of gene flow which
occurs in the wild is not faced, particularly in the case of semi-continuous distributions
throughout which genes presumably move slowly but continuously over considerable
distances. There must be very many situations where to refer to gamodemes at all
is pure optimism, and to use the term routinely is perhaps to imply a simpler situation
than really exists.

A wide range of pertinent examples is quoted in support of the ideas put forward.
Sometimes the quotation is very brief, occasionally it is in the form of a diagram
alone, and one or two authors may feel that their work has been condensed out of
all recognition. On the other hand full references are given, and I could find no obvious
case where extreme brevity had led to positive misrepresentation. This may well be
the best way to stimulate the intelligent student to read further. This is an excellent
students’ book. It could be recommended even to advanced students of botany as a
safeguard against gaps in their understanding.

D. A. WILKINS

Watsonia, 8, 181-182 (1970). 181

Obituary

JOHN WILLIAM HESLOP-HARRISON
(1881-1967)

John William Heslop-Harrison died on 23rd January 1967 at his home in Birtley,
Co. Durham, at the age of 86 and by his death the Society lost one of its most famous
members. He was a field naturalist par excellence and a pioneer in biological research.

Harrison was born in Birtley in 1881 and before he was 11 he was showing signs
of a keen interest in natural history. This tendency was carefully encouraged by his
family—his mother was a sister of the Rev. J. E. Hull, M.A., D.Sc., who had a con-
siderable international reputation for his work on spiders. By the time he won a County
Council Scholarship, which enabled him to enter Rutherford College, Newcastle, in
1893, he was spending more and more time in the gardens and greenhouses at the
Birtley House Estate, intrigued by the procedures he saw and eagerly learning all
he could about living things. Other events guided him towards a life dedicated to
biology and it was while he was at College that he met Charles Robson and J. R.
Johnson, distinguished local naturalists, who stimulated his emerging interests in
field studies. His enthusiasm and flair for biological investigation developed rapidly
and by the time he graduated in 1903 the pattern of his life had been determined and
the foundations of his future career laid.

Harrison entered the Durham College of Science (the Newcastle Division of Durham
University) in 1900 and three years later he graduated with a ‘double first’—
Education and Chemistry, with Medal. For the greater part of his life he served his
College and University in Newcastle; he was D.S.I.R. Fellow (1917-1923), Lecturer
(1920), Reader (1926) and Professor (1927-46). He was awarded the degrees of M.Sc.
(1916) and D.Sc. (1917); he was elected Fellow of the Royal Society of Edinburgh
(1921) and Fellow of the Royal Society (1928). On his retirement he continued to serve
his University as Senior Research Fellow (1946-9) and as Secretary of the Durham
University Schools Examination Board (1940-50); he was appointed Emeritus
Professor, first of the University of Durham and later of the University of Newcastle
upon Tyne.

In 1905 he joined the staff of Middlesbrough High School for Boys where he taught
with characteristic energy and success for twelve years. Equally characteristically,
and despite a heavy teaching load, he began research work into the geographical
distribution and genetics of moths of the Bistoninae. Also during this period he made
an ecological study of the Lower Tees marshes. This pioneer work, as well as being a
good example of Harrison’s best writing, has become a historical document, for Lower
Teesside is now completely changed into an urban and industrial site. Back in New-
castle from 1917 onwards he made outstanding contributions to biological science in
two fields. In entomology he carried out notable investigations into the induction,
incidence and inheritance of industrial melanism in Lepidoptera. His results con-
tributed to both Darwinian and Lamarkian views and he was a pioneer in the teaching
of experimental evolution in British universities. Harrison’s chief botanical work
during this period dealt with the following important topics: the hybridity and cyto-
genetics of roses; variation and sex determination in willows as modified by parasitic
mites; and the first demonstration of sex chromosomes in plants. Besides Rosa and
Salix he was interested in other critical plant genera and he initiated cytotaxonomic
research on several including Orchis, Rubus and Viola. In this way he and his school
pioneered one area of experimental taxonomy in Britain and gained considerable
reputation in so doing.

FE

182 OBITUARY

In 1935, as a result of joining an expedition from University College, Dundee,
led by his friend Professor A. D. Peacock the previous year, he initiated a detailed
ecological and biogeographical study of the islands of the Inner and Outer Hebrides.
These studies carried out with parties of students and colleagues continued for some
twenty years and ran parallel with similar work in the northern counties of England.
Much of Harrison’s work is contained in over two hundred scientific papers and these
are listed and appraised in Biographical Memoirs of Fellows of the Royal Society,
vol. 14, November 1968. In addition he made many thousands of biological records,
principally in The Vasculum but also in most suitable journals.

During his long life he was renowned as a gifted field naturalist; his memory of
facts was prodigious and his knowledge of flowering plants and insects was of a most
unusual depth. To be with him on one of the outings of the Northern Naturalists’
Union, which he helped to found in 1924 and of which he held office in one capacity
or another from then onwards, was a thrilling experience. Leading the party he just
took for granted. Wherever he went the place seemed to produce rare or interesting
plants because he missed nothing and always did his homework; both novice and
professional marvelled at his versatility and recognised the touch of genius. He was a
fine teacher who possessed superb self confidence and the ability of passing on to
others his love of natural history.

He married Christian Watson Henderson, who predeceased him, as did his elder
son, George, who was a first-rate entomologist. His daughter, Mrs H. H. Clark is
Lecturer in Plant Science at Newcastle University, and his younger son, Professor
John Heslop-Harrison F.R.S., now occupies a Chair of Botany at Wisconsin
University, U.S.A.

J. A. RICHARDSON

Watsonia, 8, 183-194 (1970). 183

Reports
FIELD MEETINGS 1969

ENGLAND

GUERNSEY. 11TH—20TH APRIL

A visit by any Botanical Society to Guernsey was very long overdue, none having
ever before taken place to this rich and underworked island. Since Babington’s time
130 years ago streams of visitors have come to see the rarities and there have been few
resident botanists with time and skill to make continuing observations since Marquand’s
Flora was published in 1901. As a result, the run-of-the-mill plants have been almost
wholely neglected, and areas away from those known to be good have been unsearched.

The extent to which this was so was demonstrated when, in nine days in April,
no less than 18 new taxa of reasonable status were added to the island list plus 13
dandelions and about 25 Narcissi. A further 10 were recorded which need re-seeing
for assessment of wildness or name—who will undertake Erophila? There was also
a hairy form of Vicia lathyroides, which Dr Coombe, who was in Guernsey at the
time, has taken for study. As to wildness, how should one evaluate two undoubted
and well grown seedlings of Olearia forsteri in an old quarry? The Narcissi cultivars
were named by local growers and samples exhibited, so that gatherings could be
matched. They made a remarkable living museum of plants cultivated commercially up
to 70 years ago, which have naturalised themselves gayly and often plentifully in fields
and hedges without, apparently, having in any way hybridised or been modified.

This achievement was helped by a force of over 60 participants at one time or
another, with about 40 out on most days. Between them, these visited at least once
each of the 79 squares on the island map and amassed about 10,000 records. That
there were not more was due partly to a commendable caution in noting only what
recorders were sure of, and partly to the extremely late season. As an example, only
a fortnight later in 1958 Orchis laxiflora was in good flower, whereas in 1969 nobody
even spotted a rosette.

Nevertheless, much useful basic work was done, especially among the more ordinary
plants. Of the additions, the best was, at last, a small patch of Poa bulbosa on the
western side of L’Ancresse Common. Mr A. Copping knew it in Suffolk, but deserves
credit for recognising its barely sprouting bulbils. Since no-one subsequently revisited
it, it is uncertain if it is of the viviparous form, but it looked likely to be so.

More remarkable perhaps was a larger patch, in Cobo, of a white-flowered Romulea,
larger than the well-known R. columnae (of which an albino example was seen on
L’Ancresse) and skilfully named by Mr R. D. Meikle as R. rosea Eckl. He sent the
specimens to Dr de Vos at Stellenbosch who confirmed the identification. A separate
note on this interesting plant will be published later. Clearly it had been in this locality
for a considerable time, there being corms and seedlings at all stages of vigorous
growth. A New Zealand fern, Microsorium diversifolium, is grown in a few Guernsey
gardens, but a good colony found on a wall was well away from any known in cultiva-
tion and happily naturalised. The source of Cyrtomium falcatum on the other hand
was Clearly in a bed above a retaining wall, behind the Royal Hotel.

Lady Anne Brewis and her uncle, the Hon. L. Palmer spotted Prunus cerasifera in
three or four hedges in the south, and Sedum stoloniferum, which is rarely to be seen
anywhere, in a wild area east of the reservoir. Montia sibirica, Tellima grandiflora and
Trachystemon orientale were found as weeds in shrubberies etc., Anemone nemorosa

184 REPORTS

‘Vestal’ was in hedges near Town, and several good plants of Clematis balearica on
one or two walls. There were three or four tall old trees of Salix daphnoides in a field
at Les Maingys—but is this species ever other than planted? Salix basfordiana, what-
ever the category of that taxon should be, was sorted from S. fragilis following the
lead of Mr and Mrs Howitt (who considered S. aurita x cinerea to be frequent). The
bamboos Arundinaria anceps and Sasa veitchii are better, both of them spreading
vigorously by rhizome, the former into quite a miniature forest. Ficus carica was
recorded three times as apparent seedlings on walls. This is a regular wall plant further
south and these occurrences need closer investigation. A small colony of Myosotis
sylvatica was an unexpected find in arable fields east of the Airport. Veronica sublobata
was looked out for, and plants matching it seen in two places.
All these taxa are new to Guernsey. Many more were seen in new places, including
a sixth plant of x Asplenophyllitis microdon in a fifth locality, or rediscovered after a
long interval, like Luzula forsteri—all this as well, of course, as the standard seasonal
rarities. This experience suggests that there are plenty of species remaining undetected
which might readily be refound. But the intention has to remain to finish field work on
the island for the Flora in 1970, and with it a last chance of easy and wanted records.
The excursion began with a joint meeting with the Société Guernesiaise and a
packed hall of 200, when Mr McClintock commented on a superb set of colour
transparencies of Guernsey plants taken by the island’s colour photographic club,
Spectrum. The Club has in hand the business of photographing every wild plant in
Guernsey, which would make a unique record. The States of Guernsey also entertained
the participants to a fine Vin d’Honneur, a generous and much appreciated welcome,
while Elizabeth College gave us the hospitality of its laboratory, and car park. Such
kindnesses, and all the work that was put in generally, gave cause for much gratitude.
D. MCCLINTOCK

ROANHEAD AND ROUDSEA, NORTH LANCASHIRE. 4TH MAY

On the Sunday following the Society’s Annual General Meeting, members and guests
drove west from Grange-over-Sands to the Roanhead sand-dunes on the east side
of the Duddon Estuary. It was a dull, overcast day and the exceptional lateness of
the spring meant that very few of the many specialities of these dunes were evident.
Equisetum variegatum was plentiful in the rather dry dune slacks. Viola tricolor and
the dwarf sand-dune form, subsp. dunensis, of Valerianella locusta, were in flower,
the latter being much admired. The rich dandelion flora included T. faeroense, T.
brachyglossum and T. euryphyllum. Pyrola rotundifolia subsp. maritima was frequent
and a few stems of Corallorhiza were found.

After lunch the party moved on to Roudsea Wood on the east side of the Leven
estuary where we were fortunate in being taken around the Reserve by the Nature
Conservancy staff. Specialities which we saw included Carex digitata, a few vegetative
clumps of Carex flava and Andromeda polifolia in full-flower on the Holker mosses.

G. HALLIDAY

MIDHURST, SUSSEX. 24TH-26TH MAY

This meeting was held jointly with the Sussex Naturalists’ Trust for the collection of
records for the Sussex Flora survey. The beautiful and interesting area around Mid-
hurst with its diverse habitats ranging from chalk downland to acid heath was chosen
because it was remarkably underworked.

Twenty-six members of the B.S.B.I. and/or the S.N.T. attended the meeting and a
hard-core of experienced recorders enabled nine or ten parties to record each day.
Twenty-seven two-kilometre ‘tetrads’ were visited, resulting in 4485 records.

The heavy rain of the first morning proved to be of short duration and the parties
enjoyed excellent weather for the rest of the meeting.

Through the kindness of the Headmaster, the dining-room of the Lower School,
Midhurst Grammar School, was available in the evenings, where the abundance of

REPORTS 185

tables and the excellent lighting greatly facilitated the examination of specimens and
the completion of cards. Use of the kitchen enabled welcome refreshments to be
supplied. Several local Trust members, who were unable to join the parties in the field,
attended the evening sessions; it is hoped that some of these will have been encouraged
to continue recording in this area.

The records made during the meeting form a valuable addition to knowledge of
the distribution of the flora and the Sussex Flora Committee is grateful for the help
given by the B.S.B.I.

P. C. HALL

FLATFORD MILL, ESSEX. 28TH JUNE

About 40 members met at the Field Centre for an excursion devoted primarily to
grasses and ably led by Dr C. E. Hubbard; 55 species were seen.

The meadows on the north bank of the River Stour contained anthocyanin-less
variants of Cocksfoot and Yorkshire Fog, and on the edge of the tidal reaches possible
hybrids between Agropyron pungens and A. repens were noted. Alopecurus bulbosus
was seen scattered on the river bank and brackish marshes on the south side of the
river. Crevices in the sluice of the sea-wall supported Puccinellia fasciculata with both
lax and compact inflorescences.

In Stour Wood a distinctive woodland variant of Anthoxanthum odoratum, with
lax inflorescences, was observed and an interesting mixed population of Holcus
mollis and H. lanatus was seen, though no certain interspecific hybrids were recognised.

Finally, at Wrabness Dr Hubbard and Dr Marchant recounted the interesting story
of Spartina xtownsendii. Although the Spartina plants were not in flower, it seemed
likely that both the sterile diploid S. x townsendii and its fertile polyploid derivative
were present.

B. M. G. JONES

EAST NORFOLK. 26TH—27TH JULY

Thirty-two members joined this joint excursion with the Norfolk and Norwich
Naturalists’ Society. Saturday morning was spent on the sand dunes at Winterton
where Corynephorus canescens and Ammocalamagrostis baltica were frequent on the
drier areas. In the dune slacks there was a surprisingly calcifuge flora, Erica tetralix,
Nardus stricta and Sieglingia decumbens occurring widely. Much Calamagrostis canes-
cens was seen but only an occasional patch of C. epigejos, raising a doubt as to the parent-
age of Ammocalamagrostis. In one area of the slacks Pyrola rotundifolia subsp. rotundi-
folia was found, not the previously recorded dune form, subsp. maritima, found on the
west coast.

Horsey maritime fen, visited in the afternoon, proved very dry. However, Osmunda
regalis was seen, looking out of place among the sand hills, and more Ammocala-
magrostis was found, this time growing in the wetter Calamagrostis habitat.

On Sunday morning at Horsey the party was shown Dryopteris cristata growing
with Thelypteris palustris and Dryopteris carthusiana, and large stands of Sonchus
palustris and Peucedanum palustre were seen, complete with caterpillars of the Swallow-
tail butterfly.

The final visit of the weekend was to Hickling where Naias marina was fished from
the dikes and a large area of flowering Utricularia vulgaris admired.

G. Tuck

BELVOIR AND CHARNWOOD FOREST, LEICESTERSHIRE. lsT-3RD AUGUST

A total of 22 members and guests attended this meeting. On the Friday evening all
assembled at the meeting headquarters in Loughborough for welcome and briefing.
Mr I. M. Evans gave an account, illustrated by colour slides, of Leicestershire and
its flora, and explained how and why the present Leicestershire Flora Committee
had come into being. The Rev. A. L. Primavesi outlined the methods being used for

186 REPORTS

recording, and explained the procedure for recording in the field. Recording areas were
then assigned and record cards distributed.

On the Saturday morning members dispersed to unrecorded tetrads in the northeast
of the County. This was a beautiful summer day. The whole party reassembled for
picnic lunch at Eastwell, by courtesy of Mr J. M. Stanley. After a further spell of
recording all returned to Loughborough where the whole party had dinner. After
dinner each group reported on their experiences; all had succeeded in recording well
over 100 species in their tetrads, some recording over 200.

On the Sunday morning, in pouring rain, we were led by Mrs P. A. Candlish round
the Ulverscroft Nature Reserve, by courtesy of the Leicestershire Trust for Nature
Conservation. Here we saw regions of unspoilt woodland, marsh and meadow, and
were introduced to many of the Charnwood plants which are rare or absent in other
parts of Leicestershire. In spite of the rain, all found this visit interesting and enjoyable.

After picnic lunch, taken in the cars because of the rain, all went to The Brand, by
courtesy of the Lord Lieutenant of the County, Colonel Sir Robert Martin, who
welcomed us on arrival. By this time the weather had improved, and we explored
these beautiful grounds in sunshine, led by Mr P. H. Gamble. In the formal parts of
the grounds it was significant that the party seemed to be more interested in the garden
weeds than the cultivated plants, apart from some fine clumps of Osmunda regalis
thoroughly naturalised by the streams. Most of The Brand, however, is wild rocky
woodland, with long-abandoned slate quarries. Some of the less active members of
the party were daunted by the feats of mountaineering expected of them, but all
found interest in a fascinating assembly of plant species. It was curious to find plants
such as Origanum and Carlina growing within a few feet of Calluna and Teucrium
scorodonia, and apparently on the same soil. We also saw Umbilicus rupestris in what
is believed to be its most easterly native station.

We were fortunate in having Dr Dony present at this meeting, with his wide
experience of work on Local Floras.

A. L. PRIMAVESI

CONFERENCE EXCURSION. 21ST SEPTEMBER

About 45 members attended this outing, which in the morning visited the Forestry
Commission Lands at Alice Holt, Bentley, Hants, where Mr M. Brown showed the
party some of the types of planting and management carried out by the Commission
and discussed the history of the forests. An area of waste ground provided some
interesting ruderals and aliens including Galinsoga parviflora, G. ciliata, Antirrhinum
orontium and Onopordum acanthium.

After lunch the party visited Thursley Common Nature Reserve where the warden,
Mr R. M. Fry, explained some of the difficulties of management and the recovery
of the vegetation since the war when the site was used as a tank training area. Some of
the interesting features included bogland from which 14 species of Sphagnum have
been recorded and a large area dominated by Narthecium ossifragum. Lycopodium
inundatum was found in abundance. Other interesting species seen included Utricularia
minor, Rhynchospora fusca, R. alba and Hypericum elodes.

I. K. FERGUSON

ALIEN HUNT ABOUT LONDON. 4TH-S5TH OCTOBER

On October 4th thirty members of the B.S.B.I. and L.N.H.S. with the aid of a hired
coach explored four refuse-tips in west Middlesex, and a very interesting selection of
unusual aliens was seen.

Shepperton tip introduced us to a wide variety of typical bird-seed aliens including
Camelina sativa, Cannabis sativa, Carthamus tinctorius, Centaurea diluta, Chenopodium
probstii and Setaria lutescens (= S. glauca). Cucumis melo, Cucurbita pepo, Kochia
scoparia, Physalis peruviana and Sisymbrium loeselli were also found, but bulldozers
had unfortunately destroyed three grasses present the previous weekend: Digitaria

REPORTS 187

sanguinalis, Panicum capillare (both determined by C. E. Hubbard) and Sorghum
halepense, the latter being unusually frequent this year. The fine Amaranthus plant that
we saw was later determined by J. P. M. Brenan as one of the various manifestations
of the hybrid A. hybridus x retroflexus.

West Drayton tip provided two unusual grasses: Agrostis semiverticillata and
Polypogon monspeliensis. Glycine max, Coriandrum sativum, Polygonum pensylvanicum
and Trachyspermum ammi were new to many of us, but the rare alien Amaranthus
standleyanus, was, alas, beneath tons of earth.

Harefield tip gave us two identification problems: Digitaria ciliaris (= D. adscen-
dens; determined by C. E. Hubbard) and Polygonum cf senegalense. The pretty
knotgrass, Polygonum arenarium (= P. pulchellum; determined at Kew as P. patulum)
was also admired, but Londoners in our party scarcely glanced at the familiar Bromus
carinatus. Citrullus lanatus (= C. vulgaris) was seen, but not in flower.

With time running out, we had a brief look at Newyears Green tip where we found
Abutilon theophrasti, Ambrosia artemisiifolia (very young) and Salvia reflexa. The
Chenopodiums included C. opulifolium and C. ficifolium.

Our final stop was in the City, opposite the Tower of London, where we searched
out the flower-bed weeds: Sisymbrium irio in plenty and a colony of Portulaca oleracea
were of particular interest. Both Galinsoga ciliata and G. parviflora were also present,
but we had already seen these on tips earlier in the day.

On the Sunday the hunt continued, but in Surrey. The party, reduced to nineteen,
explored Guildford tip where we collected Amaranthus blitoides, Psoralea americana
and Silphium perfoliatum.

After lunch we visited the nearby Worplesdon tip to see Amaranthus caudatus and
Chenopodium glaucum. A quick stop at a sandy nursery beside Smart’s Heath, Mayford,
confirmed the continued presence of plentiful Digitaria ischaemum.

A tip at Send was almost barren, apart from the surprising presence of one plant of
Xanthium strumarium. Depauperate Apera spica-venti was also present.

Our final visit was to May’s Green, near Effingham, where the garden escape
Chrysanthemum uliginosum (= C. serotinum) was in full flower.

E. J. CLEMENT & T. B. RYVES

WALES

LONG MOUNTAIN, MONTGOMERYSHIRE. 15TH JUNE

Most of the day was spent exploring the steep wooded banks of the Rowley Brook
which divides Shropshire from Montgomeryshire. The banks are of base-rich Silurian
shale and harbour some very interesting species of which, perhaps, Astrantia major
is the most notable. Its habitat alongside the road had suffered disturbance but the
plant had not been destroyed and several plants were seen though not yet in flower.
Paris quadrifolia was fairly plentiful amongst Mercurialis perennis and a large patch
up to 9 ft across and hitherto unrecorded was found in another part of the wood.
Two stands of Lathraea squamaria were seen, both parasitic on Corylus; Chryso-
splenium alternifolium was frequent along the streamside, but Cardamine impatiens,
previously found here, could not be located. The day ended with a visit to the magni-
ficent stand of Sequoia sempervirens in Leighton Park.

Mrs D. M. PAISH

TOWYN BURROWS, GLAMORGAN. 5TH JULY

The party of fifteen members and guests followed a route through the Forestry
Commission rides to the Observation towers and thence through fixed dunes and dune
slacks to the estuary point.

In the rides, despite a regrettable increase in coarse grasses and scrub, good stands

188 REPORTS

of Ophioglossum vulgatum were seen on the ridges with Anagallis tenella in the ditches.
Eleocharis quinquefilora was seen in a boggy cross ride while the open dunes produced
Viola tricolor subsp. curtisii and Lathyrus sylvestris, the latter 6 ft high and just coming
into flower. In the slacks Dactylorhiza fuchsii, D. incarnata (including subsp. coccinea),
D. praetermissa and D. purpurella were plentiful. In addition there were occasional
plants of Gymnadenia conopsea and Epipactis palustris was commencing flowering.
By the upper Observation tower there was plenty of Anacamptis pyramidalis in the
hollows with Centaurium littorale on the hummocks. Liparis loeselii was having a
successful season and Sisyrhinchium bermudiana appears to be increasing.

Two members found time to look at Linaria supina on the nearby railway halt
where it was found to be regenerating freely despite British Railways’ efforts with a
herbicide.

Mrs H. R. H. VAUGHAN

PEMBROKESHIRE. 16TH—17TH AUGUST

The meeting, a joint one with the West Wales Naturalists’ Trust, was attended by
eighteen members. The theme was moorland and bog vegetation. The first day was
spent on Brynberian Moor, a gently undulating plain, dissected by small streams
with associated bogs, beneath the northern flank of Preseli. The soil is a shallow layer
of peat overlying boulder clay and, though acid, it is not uniformly base-deficient.
Among the commoner bog plants seen were Hypericum elodes, Pinguicula vulgaris,
Wahlenbergia hederacea and Eleocharis multicaulis. Locally abundant here but far
from common in the county were Drosera intermedia, Myrica gale and Rhynchospora
alba and beside the streams Thelypteris limbosperma. We also searched for and found
Pinguicula lusitanica, Hammarbya paludosa and Lycopodium inundatum, the two latter
in new sites as well as where previously known. Galium uliginosum was a new record
for the 10 km square and the third for the county.

On August 17th we met at Ambleston Common which demonstrates that ungrazed
wet moorland may become dominated by Molinia caerulea. Patches of Achillea
ptarmica and Serratula tinctoria broke the monotony and several clumps of the hybrid
Juncus conglomeratus x effusus were examined. A stream in Wallis Moor had Rorippa
x sterilis, Myriophyllum alternifiorum and the aquatic form of Juncus bulbosus sensu lato
with long trailing red stems. Tre-Rhos Common is part of an extensive moorland
and is richer botanically than the other two areas, but it is at its best earlier in the
season when orchids and sedges can be appreciated. Hypericum undulatum and
Cicendia filiformis were new records here and Drosera intermedia was seen, but search
for Pinguicula lusitanica where it was seen last year failed. The soil of these moors is
much as on Brynberian where habitats for tiny plants are kept open by grazing of
sheep and ponies.

T. A. W. DAvIs

CAMDDWR VALLEY, CARDIGANSHIRE. 30TH AUGUST

On this excursion four members visited the upper Camddwr valley now doomed
to disappear under the Brianne Reservoir. They were led by Dr J. P. Savidge who has
done considerable research in this area. The steep valley sides are mountain pasture
(grazed by sheep and ponies) and largely dominated by Molinia but intersected by
slightly more base-rich flushes.

Typical plants of the streams and flushes are Pinguicula vulgaris, Carex demissa,
Drosera rotundifolia, Thelypteris limbosperma and, more locally, Carex hostiana,
Thelypteris phegopteris and Rhyncospora alba. Here, very locally, in suitable open,
wet and slightly sheltered habitats Dr Savidge has found Selaginella selaginoides, not
recorded before south of Merioneth.

Mrs H. R. H. VAUGHAN

REPORTS 189

CORS GOCH, ANGLESEY. 28TH SEPTEMBER

Following the Annual General Meeting of the Welsh Region the previous day, Mr
R. H. Roberts led 12 members to Cors Goch and nearby areas. Here we found a perfect
illustration of the previous day’s talks: habitats merging into each other and the
association of species with usually specialised requirements. The many interesting
plants seen included Gentiana pneumonanthe, Gentianella amarella, Parnassia palustris,
Viola lactea and the hybrid Equisetum x litorale.

Mrs H. R. H. VAUGHAN

SCOTLAND

PEASE DEAN, BERWICKSHIRE. 8TH JUNE

Pease Dean was once a magnificent Sessile Oakwood but the upper two-thirds were
felled and the area replanted with conifers—mainly Larix decidua—some decades
ago. The lowest third was left and it is still magnificent. Betula pubescens, Corylus
avellana, Ulmus glabra, Alnus glutinosa and Sorbus aucuparia also occur. It has quite
a good woodland flora including Geranium sylvaticum, Ajuga reptans, Lysimachia
nemorum and Corydalis claviculata. Equisetum telmateia and Marrubium vulgare occur
at the mouth of the glen. In the adjacent arm, Tower Dean, there is a fine colony of
Orchis mascula.

In the afternoon Dowlaw Dean was explored. The upper part is wooded and the
trees have been felled in the lower (seaward) half. The almost perpendicular sides of
this dean are covered in June with extensive patches of Endymion non-scriptus. Orchis
mascula was also seen and there were many plants of Sedum rosea. 12 people attended.

E. P. BEATTIE

BYRNE HILL. AYRSHIRE. 15TH JUNE

The main object of this meeting, which was attended by 19 members and friends,
was to attempt to refind Minuartia verna, which was first discovered on Byne Hill in
1963. In this we were ultimately successful when late in the afternoon Miss E. Conacher
discovered a few plants in full flower scattered over an area of about 1 m2. A specimen
has been deposited in the herbarium of the Botany Department, Glasgow University.
Most of the day was spent examining the flora of the serpentine grassland and
boulders, and attempting to fill in some obvious gaps in the record card for the square.
Among the more interesting plants seen were Antennaria dioica, Carex dioica, Crypto-
gramma crispa, Helianthemum chamaecistus, Helictotrichon pratense and Juniperus
communis. Altogether more than twenty new species were recorded for the square

(GR.25/19).
H. A. MCALLISTER

THE SHINGLE BEDS OF THE TUMMEL AND TAY, PERTHSHIRE. 21ST—22ND JUNE

Two beautiful days were spent systematically botanising over the rich shingle beaches
found between Meikleour on the Tay in the south and Aldclune on the Garry in the
north. Only four members (including the leader) attended. Members of the Perthshire
Society of Natural Science were present on both days. Lists for the 5 km square
records were made at each of six sites visited. These were (Saturday) The Inch near
Braecock, a site near Meikleour and another near Kercock, all on the Tay below
Dunkeld; (Sunday) near Moulinearn and Ballinluig Island, both on the Tummel, and
a site near Aldclune on the Garry.

The Tay shingles were gay with Lupinus nootkatensis in a remarkably wide range
of colour forms. Not many new records were made but the following are of interest.
Alchemilla vestita has not been recorded for shingle beds before and suggests that it
may have been overlooked. Tolmiea menziesii appeared at almost every site on the

190 REPORTS

Tay and Tummel and clearly demonstrates its recent spread. Poterium sanguisorba
at Aldclune probably represents the most northerly station for this species. Subsp.
majus of Thalictrum minus occurred at two sites. The danger to these interesting sites
was demonstrated at Meikleour where gravel-bed working with modern machinery
destroys the natural colonisation patterns, and even when recolonisation takes place
the unnatural redistribution of shingle into heaps will prevent for a long time the
reappearance of these patterns based upon river movements. On all the sites visited
grazing by domestic animals does not occur, but on sites where it exists it reduces
considerably their ecological interest.

A. W. ROBSON

ISLE OF RAASAY, INVERNESS-SHIRE. 28TH JUNE—-STH JULY

The purpose of this meeting was to check records appearing in Heslop-Harrison’s
Flora of Raasay (1937) for inclusion in the proposed Flora of Skye. In the 30 years
since, many botanists, single and in university groups, have visited Raasay, but little
of their work has appeared in print or is in the Monks Wood files.

Our party of eight usually split up to cover more ground, and local transport was
arranged, but not for the first day. On the Sunday we covered the area northwest of
Inverarish village, as far as Holoman. Arenaria balearica was still on its wall, for about
100 yards alongside the road just beyond Raasay House. The short turf at Oskaig
Point had Coeloglossum viride, Gymnadenia conopsea, and Botrychium, but the coast
beyond to Holoman looked wet and bleak and the party divided, one group to
confirm the acidity of the ground as far as Holoman Island, while the rest returned
south, past a good patch of saltmarsh, with Carex extensa, Aster tripolium, Puccinellia,
and Blysmus.

The following day the party divided near Brae, one group going on to Brochel while
the others followed a nameless burn eastwards to Druim an Aonaich, and proved that
the sharp bend indicated a gorge. This gorge was one of the best ‘finds’ of the week,
as the list recorded there included Pyrola secunda (a rarity we had hoped to confirm),
Viburnum opulus, not previously seen on Raasay, and Dryas, Draba incana, Epipactis
atrorubens, Thalictrum alpinum, Polystichum lonchitis, and Gymnocarpium dryopteris.
Meanwhile, the northern group had noticed a pool beside the Brochel road with a
large patch of Cladium mariscus, and Utricularia minor and U. intermedia, both in
flower. Below the Castle, Inula helenium still grew, not yet full-size or in flower. Arnish
and Torran turned out to contain more woodland than anywhere apart from Raasay
House—mainly birch and hazel, with Hymenophyllum wilsonii on the mossy boulders.
With the assistance of Mr Malcolm Macleod we located Lythrum salicaria—difficult
to find when only 4 in. high!

On Tuesday, in low cloud and drizzle, we started from Fearns, and followed the
old track to Rudha na’Leac and Hallaig. Our Swiss botanist collected Neottia from a
birchwood just above the road, and wondered why everyone else got so excited—it
is not so rare in Switzerland! The cliffs higher up were covered in Dryas, Saxifraga
spp. and Silene spp., with Polygonum viviparum, but they had to be abandoned in
thickening mist, as this area is deeply fissured and dangerous. At Rudha na’Leac,
Sorbus rupicola hadn’t grown much in 30 years, and several smaller specimens were
seen on the cliffs behind. We failed to find Asplenium septentrionale near Hallaig
waterfall, but did better on boulder scree below Loch a’Chadhacharnaich, where
Orobanche alba, Draba incana, Arabis hirsuta and Listera ovata were among those
recorded. The loch was disappointing, apart from Potamogeton filiformis, and one
group climbed higher to follow the cliffs (more Dryas etc.), while the others took the
moorland path below.

The next day was fine enough for crossing to Scalpay. Landing at Camus na Geadaig,
we followed the eastern arm of the burn but it was not very exciting apart from Listera
cordata in quantity and our only record during the week for Carex laevigata. At
Loch an Leoid, Carex limosa and C. pauciflora were found on the moor. Three parties

REPORTS 191

then explored in different directions. At Loch a’Mhuilinn Nuphar lutea (not N. pumila)
and Typha angustifolia (not T. latifolia) were still present although both looked planted.
Thalictrum alpinum was seen at Sithean Glac an Ime, while in the gorge of the Allt
Liath below the waterfall Osmunda, Viburnum opulus, Ilex and Prunus padus were
found. Further east, a large patch of Lycopodium annotinum was found in scrub,
and a second patch was seen by the group returning down the western arm of the
Allt Camus na Geadaig.

On Thursday we again started from Brae. The ‘limestone’ party followed the burn
west of the road and found themselves in what must be Heslop-Harrison’s ‘Leek
gorge’. This was so interesting that they never reached the coast as planned, the
highlights in the long list recorded being Pyrola minor and P. secunda (both in flower),
discovered by coming face to face with them among heather and bracken, on climbing
out of the gorge.

The ‘“‘Duncaan”’ party reached the top with little to record but more Carex limosa
and C. pauciflora around the lochans on the way. Lycopodium alpinum and Salix herbacea
appeared on the last slope below the flat summit, but a careful search on ground to
the northwest failed to produce any Tofieldia. Thalictrum alpinum occurred beside
Loch Meall Daimh, and all down the burn running out of Loch na Meilich to about
700 ft.

For our last day the majority vote was for Fearns again. There we separated; one
group went to Point of Eyre and back by the woodland above the shore, which was
mainly birch and hazel, with Alnus, Fraxinus, Quercus petraea, Salix atrocinerea, and
Populus tremula. Another looked at woodland between North Fearns and the shore,
which had scattered plants of Agrimonia, Geum urbanum, and Rumex sanguineus along
the edge, with quantities of Stellaria holostea. The last group went to test a memory
of finding Hymenophyllum tunbrigense in fissures above Fearns on a visit 20 years
before, and returned in triumph with a specimen—a new Raasay record!

Of the records made during this meeting, seven appear to be new to Raasay, the
most interesting being Neottia nidus-avis, which is also new to v.c. 104. There were
30 new 10 km square additions, while 36 earlier records, which do not appear in the
Atlas, many of them ferns, were confirmed. We failed to find Galium boreale, Saxifraga
oppositifolia, Trollius and Rosa pimpinellifolia on Raasay, all of which are plentiful on
Skye, but the latter was seen on Scalpay.

Mrs C. W. MurRAY

BRAES OF BALQUHIDDER, PERTHSHIRE. 12TH JULY

This was a joint meeting with the Andersonian Naturalists of Glasgow and the
Botanical Section of the Perthshire Society of Natural Science. A party of 13, con-
sisting almost entirely of members of these two societies, assembled at Balquhidder
village before proceeding to the west end of Loch Doine where the cars were left.
Glen Carnaig was then ascended towards Stob Coire an Lochain with the intention
of examining some exposures of good rock reported to be in that area. However, in
the face of rapidly deteriorating weather with poor visibility this plan was abandoned
at about 2,500 ft and the descent was made by the same route. Only the commoner
mountain plants were seen, perhaps the most noteworthy being Salix phylicifolia, a
few bushes of which were noted on the banks of the Allt Carnaig.
Species lists were compiled for two 5 kilometre squares to assist the P.S.N.S. record-
ing scheme for flowering plants.
A. Mc.G. STIRLING

SHETLAND. 1ST—8TH AUGUST

Seven people attended this meeting which was the second to be held in Shetland (the
first was in 1960) and, as then, the object was to assist the leader and Mr Palmer in
the compilation of their new Flora of Shetland. The weather was perfect, a fresh

192 REPORTS

breeze and blue skies every day and not a drop of rain during the entire week. Our
headquarters was at Lerwick, and transport was by hired minibus.

On our first day we looked for Rhinanthus serotinus subsp. apterus in an old station,
Ringasta, near the southern tip of Mainland, but without success. The rest of the day
was occupied in exploring the large stretches of sandy pasture behind the Bay of
Quendale. This produced Carex maritima and Gentianella amarella subsp. septen-
trionalis but there was no sign of Sea Holly on the shore where it was last seen well
over 80 years ago. Catabrosa aquatica and much naturalised Spearmint grew by the
burn from the Loch of Hillwell, while the very rare (for Shetland) Silene alba subsp.
alba was seen in small quantity bordering a sandy oatfield south of the loch. Back
in Lerwick the sharp-eyed Miss Webster added two new names to the local list by
finding Sisymbrium officinale on the Rubbish Dump and a great quantity of naturalised
Lotus pedunculatus in a field near the Loch of Clickimin.

The highest land in Shetland, Ronas Hill, 1,486 ft, in the north of Mainland was the
target on the second day. On the way a stop was made at Mavis Grind to see some of
the hawkweeds on the high rocks of Cliva Hill. The endemic H. hethlandiae was in
full flower and a mile north another endemic, H. praethulense, grew in a cleft of granite
rock. Also noted were H. orimeles and H. sparsifolium, the latter with purple-blotched
leaves, its usual Shetland form.

A stop at Heylor enabled us to admire the miles of red granite cliffs and crags on the
north side of Ronas Voe. This is one of the most scenic spots in the whole county and
home of half the Shetland hawkweeds. The rounded, stone-strewn and extremely
exposed summit of the hill provides the only real alpine flora in Shetland. Here
Loiseleuria procumbens, Alchemilla alpina, Vaccinium uliginosum, Juncus trifidus,
Luzula spicata, Salix herbacea and Diphasium alpinum were seen. Saussurea alpina,
usually represented by a few flowerless rosettes, stubbornly refused to put in an appear-
ance.

On the Sunday the party was back at the south end again, this time searching for
Eleocharis acicularis, which Druce dredged from the Loch of Spiggie in 1924. Mr
Dupree and the leader obtained a boat and spent some time dredging the shallow
sandy north end of the loch. Little was brought up, apart from the masses of stonewort
and Potamogeton filiformis. It was then decided to examine the entire margin of the
loch. Before long Miss Webster found a few minute flowering plants of Eleocharis
on damp sand near Littleness, on the west side. There is only one other known Shetland
station for this plant, the nearby Loch of Brow, also one of Druce’s records. The two
boatmen somewhat redeemed themselves by finding Ste/laria graminea on the edge of
a cornfield above the east side of the Loch of Spiggie, a new station for this scarce
introduction. A single small bush of spreading-leaved Juniper was found by Miss
Webster on a heathery roadside bank above the south end of the loch; the normal
Shetland plant is the prostrate appressed-leaved subsp. nana. At Easter Quarff we
admired the fine colony of Mertensia maritima on the beach and the beautiful coppery-
orange hybrid Mimulus cupreus x guttatus by a nearby stream.

A long run to the extreme west of Shetland’s main island, to a delightful place
called Sandness, started the fourth day of the meeting. We stopped en route to see the
endemic Hieracium pugsleyi on the steep grassy banks of the Burn of Weisdale. At
Sandness the party divided, two members worked the immediate vicinity while the
rest walked round the rock-bound coastline eastwards to West Burrafirth. The rocky
meadows and heathy pastures about Sandness were dotted with Hieracium attenuati-
folium, H. vinicaule (these two endemic), H. caledonicum and H. scoticum. The beautiful
Parnassia palustris grew by a tiny loch and Asplenium trichomanes, a Shetland rarity,
hid its fronds in the crevice of a rock. Native shrubs of any kind are scarce in the
islands but on rocks above the Bay of Brenwell the party saw stunted specimens of
Populus tremula, Sorbus aucuparia and Lonicera periclymenum, with Rosa canina
on the sea-cliffs nearby. Two more endemic hawkweeds were seen: H. dilectum and
H. zetlandicum. Hymenophyllum wilsonii was abundant among Sphagnum on the

REPORTS 193

damp east-facing hill-slopes. A loch near Burraview was full of the scarce Phragmites
australis. The last plant of an exciting day was the recently-discovered taxon Pilosella
flagellaris subsp. bicapitata in dry rocky heathy pasture at West Burrafirth.

On the Tuesday the party travelled north by bus and ferry to the sands of Burra
Firth, in the north of Unst. We looked for Polygonum oxyspermum subsp. raii collected
here, its only Shetland station, in 1868 but it soon became apparent that it had suffered
the same fate as the narrow-leaved form of Sea Pea which used to grow here. The
hawkweeds H. australius, H. gratum (both endemic) and H. sparsifolium were seen
at the north end of the Loch of Cliff. Near Baltasound we examined the unique
serpentine habitat and its interesting plants Arenaria norvegica subsp. norvegica and
the plant of Shetland Cerastium arcticum subsp. edmondstonii. It was disturbing to
see that a large area where the Cerastium used to grow had been re-seeded.

On the following day, as also by a strange coincidence in 1960, Lerwick was prepar-
ing for a Royal visit. Unlike the 1960 party we decided that plants definitely came first
and, leaving the flags and the crowds behind, we made our way to a remote spot
on the east side of Mainland. In glorious sunshine half the party worked the Lunna
Ness peninsula, the remainder walked the coastline from Lunning to Vidlin. At
Lunning Head Lonicera periclymenum was noticed and at Lunning itself the large
glossy-leaved Senecio smithii. On rocks west of this tiny hamlet Hieracium gothicoides
was making a fine show. In the Lunna Ness area the best find was Gymnocarpium
dryopteris among rocks south of Stour Hevda.

The last but one day of the meeting found the party in the Walls area, on the west
side. The leader visited a dozen small islands in the Walls harbour area but nothing
exciting was found. Puccinellia capillaris grew in many places and a fine show of
Ligusticum scoticum was seen on a small grassy island near the head of the voe at
Walls. The rest of the party examined the Lochs of Bardister and Kirkigarth, finding
Potamogeton rutilus, Subularia aquatica, Ranunculus trichophyllus subsp. trichophyllus,
Eleogiton fluitans and Zannichellia palustris.

The party spent the last day at the south end with excellent results. Potamogeton
Xzizii, new to Shetland, was found in the Loch of Hillwell by Mr Palmer. One of its
parents, P. gramineus, is common in Shetland but the other, P. /ucens, has no reliable
record. This loch also supports Myriophyllum spicatum, which is known from nowhere
else in Shetland. So ended a memorable week both as far as plants and weather was
concerned.

W. SCOTT

HANDA ISLAND AND DURNESS, SUTHERLAND. 2ND-3RD AUGUST

Five people attended this weekend meeting in West Sutherland (v.c. 108).

A crossing of Handa Sound and a day botanising on the island were blessed with
fair weather. A landing was made on a sandy shore in the southeast of Handa and
the first habitats examined were the complex of sand-dunes, cliffs and stacks in this
corner of the island. A thorough search was made for the hybrid grass Ammocalama-
grostis baltica in its only Scottish locality, where it was first seen in 1937, but with no
success. A striking feature of the vegetation of these cliffs is the vigorous growth of
some species. This is particularly noticeable with Succisa pratensis, which bears foliage
of great luxuriance when compared with plants in moorland sites. Botrychium lunaria
(a new species for the Handa list) was found in short turf on the way from the beach
to one of the freshwater lochans in the centre of the island. This particular lochan
supported a fine display of Nymphaea alba in full bloom. Isoetes lacustris was seen
later in the day in another lochan. A rather rushed return for the boat to the mainland
gave little opportunity to look in detail at the machair and dunes of the south coast
but we were fortunate in seeing Catabrosa aquatica growing in some wet sand.

Instead of visiting the cliffs near Durness on the following day, it was felt that an
excursion to a less known area would be more rewarding. Consequently, the party
went to Sandwood Bay. Areas of sandy pasture, dunes, small burns and cliffs were

194 REPORTS

examined; the cliffs were particularly rich in species. A wet area behind the dunes,
near Sandwood Loch, supported a fine stand of Juncus balticus, while areas of cal-
careous sand at the northern end of the bay were covered with extensive carpets of
Dryas octopetala. It was particularly interesting to find this species acting as a stabilising
plant of a sandy substrate.

Some members of the party visited the coast near Sheigra during the afternoon;
cushions of Silene acaulis and a small colony of Scilla verna were seen here. Finally,
a flying visit to Durness produced plenty of Primula scotica, with a few plants still in
flower, as well as fruiting specimens of Dryas octopetala and Epipactis atrorubens.

B. S. BROOKES

SKINFLATS, STIRLINGSHIRE. 31ST AUGUST

At Skinflats there is no beach, only mudflats and saltmarsh. We spent the morning
investigating the saltmarsh and some inland ponds. Aster tripolium is the dominant
plant. Its associates were Plantago maritima, Triglochin maritima, Spergularia marina,
Festuca rubra, Glaux maritima, Atriplex hastata, Suaeda maritima and Salicornia in
the mudflats. Inland a little, in another saltmarsh, Aster tripolium was again the dominant
species but here Puccinellia distans and Ranunculus sceleratus occurred in some abund-
ance. Several fine stands of Calamagrostis epigejos were seen in the area.

The afternoon was spent exploring the extensive area of wasteground at Grange-
mouth Docks. At the mouth of the River Carron there is a small strip of saltmarsh
and here Aster tripolium, Triglochin maritima, Plantago maritima, Juncus gerardii and
Cochlearia officinalis were seen; thus if the Skinflats saltmarshes are drained Stirling-
shire will still have a few saltmarsh plants. In one lagoon which had been drained there
were rafts of Salicornia dolichostachya and in another a grove of Atriplex littoralis.
Perhaps the best introductions seen were Chenopodium rubrum, Pastinaca sativa,
Saponaria officinalis, Senecio squalidus (1 plant) and a magnificent patch of Coronilla
varia. There were also some magnificent stands of Calamagrostis epigejos. 12 people
attended.

E. P. BEATTIE

ISLE OF TIREE, 1968—A CORRECTION

In the report for the field meeting, Isle of Tiree. 3rd-10th July (1969, Proc. bot. Soc.
Br. Isl., 7: 636-637), a portion of the penultimate paragraph was unfortunately
omitted. The paragraph should read as follows with an asterisk denoting a species
believed to be new to v.c. 103 and a dagger one believed to be new to Tiree.

On July 8th we decided to return to the base-rich Balephetrish area (17/04), where
a number of shallow lochs (Loch Riaghain, Cnu Lochanan, etc.) gave us Hammarbya
paludosat, Juncus balticus, Potamogeton coloratus, P. x nitens and P. x sparganifolius*.
Crossing into 07/94 we added to our card Potamogeton perfoliatus and P. pectinatus
in Loch a’ Phuill, also Lysimachia nummularifolium, Potamogeton filiformis, P.
x nitens and P. rutilus* at Loch an Eilein. Our last day (July 9th) was given up to the
Reef, an area of machair partly occupied by an airfield. Anacamptis pyramidalis and
Draba incana were found in very small quantity. Disappointed by the machair, we
returned to Loch a’ Phuill (07/94), where the Potamogeton enthusiast refound Heslop-
Harrison’s Potamogeton x suecicus, while the rest of the party got Gnaphalium
uliginosumt and Platanthera bifoliat.
EDITOR

A DARWIN CENTENARY

Edited by P. J. Wanstall

The purpose of this volume is to make available to a wider public the papers
read at the very successful Conference arranged by the Botanical Society of the
British Isles in 1959. The contributors include Lady Barlow, G. D. H. Bell,
Prof T. W. Bécher, Prof W. H. Camp, Prof N. Hylander, Prof I. H. Manton,
Prof T. G. Tutin and Prof D. H. Valentine.

Demy 8vo., 140 pages, with 7 half-tone plates and 11 maps and figures in the text.
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LOCAL FLORAS

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The purpose of this volume is to make available to a wider public the papers
read at the Conference held by the Botanical Society of the British Isles in 1961.
The contributors include D. E. Allen, Dr H. J. M. Bowen, Dr J. G. Dony,
J. S. L. Gilmour, Prof J. G. Hawkes, D. H. Kent, J. E. Lousley, D. McClintock,
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THE CONSERVATION OF THE
BRITISH FLORA

Edited by E. Milne-Redhead

The Report of an important and most successful Conference held by the
Botanical Society of the British Isles at Durham in April 1963. This report
includes most of the papers read at the Conference, together with a verbatim
account of the discussions that followed.

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July 1970 Volume eight Part two

Contents

KENNETH, A. G. & STIRLING, A. McG. Notes on te Hawks ree
(Hieracium sensu lato) of western Scotland ee

Roserts, R. H. A revision of some of the taxonomic charact rs
Polypodium australe Fee Sees ea ee :

the adjacent small islands. 2. one = So ee

SCHROEDER, F.-G. Exotic Amelanchier species naturalised in 1 EA
and their occurrence in Great Britain... Se

SHort NOUS = (20) US ee
PLANT: RECORUS (600. ee

Book REVIEWS ... ae ue eee ne rat

OBITUARY Se

REPORTS

Field Meetings 1969 oe ee oe

BE Ee

Published and sold by the Botanical Society of the British Isles,
c/o Department of Botany, British Museum (Natural History),
London, S.W.i7.

Watsonia, 8 (2). Price Two pounds

A WILLMER BROTHERS LIMITED, BIRKENHEAD
UA member of the Tinling Printing Group

ICa

f the Botan

ings oO

d
ish Isles

rit

see

wees

ida

Dane Se

t

a

Beri

Botanical Society of the British Isles

Patron: Her Majesty Queen Elizabeth the Queen Mother

Applications for membership should be addressed to the Hon. General Secretary:
c/o Department of Botany, British Museum (Natural History), Cromwell Road, London,
S.W.7, from whom copies of the Society’s Prospectus may be obtained or from the Hon.
Treasurer at 12/14 Wigmore Street, London, W.1.

Officers for 1970-71

Elected at the Annual General Meeting, 9th May 1970
President, E. Milne-Redhead

Vice-Presidents, Prof. J. G. Hawkes, Prof. D. H. Valentine, J. E. Lousley, Dr F. H.
Perring

Honorary General Secretary, Dr I. K. Ferguson
Honorary Treasurer, J. C. Gardiner

Honorary Editors, E. F. Greenwood, Dr G. Halliday, Dr M. C. F. Proctor,
Dr N.K.B. Robson

Honorary Meetings Secretary, Mrs M. Briggs

Honorary Membership Secretary, Mrs J. G. Dony

Alli enquiries concerning sales of, or advertising space in, the Society’s publications
should be addressed to D. H. Kent, 75 Adelaide Road, West Ealing, London, W.13.

Watsonia, 8, 195-203 (1971). 195

Presidential Address 1970

BOTANICAL CONSERVATION IN BRITAIN,
PAST, PRESENT AND FUTURE

E. MILNE-REDHEAD

Nineteen-seventy is European Conservation Year and it seems to me fitting
that my Address today should have conservation as its theme.

The B.S.B.I. has a long history. It evolved from the Botanical Society and
Exchange Club of the British Isles (conveniently abbreviated to B.E.C.) at a
Special General Meeting in October 1947. The B.E.C. somewhat similarly
evolved from the Botanical Exchange Club of the Thirsk Natural History
Society, a body which, in spite of its name, had a national coverage. The
B.E.C. came into existence in 1858 and, at the age of eight, lost its parent body,
the Thirsk Natural History Society, and started on its independent career, moving
its headquarters to London, and for 2 years only adding London to its name.
Twelve years later it openly stated it represented the British Isles, and in 1910
added the word ‘Society’ to its title.

It is interesting to recall that, when the Club started, J. G. Baker, later to
become Keeper of the Herbarium & Library here at Kew, was the B.E.C.’s
“Curator of Flowering Plants and Ferns’, and H. C. Watson, so well known for
his Vice-county system and whose herbarium is at Kew, was an active member.

Conservation was an unknown word to botanists of those early days. In
the first report published in 1859 there is a list of desiderata from which her-
barium specimens ‘varying in number from 10 to 50, according to degree of
rarity would be acceptable’. This list includes Ranunculus ophioglossifolius,
Draba aizoides, Linnaea borealis, Phyteuma spicatum, Spiranthes aestivalis,
Cephalanthera rubra, Epipogium aphyllum, Cypripedium calceolus and Polygona-
tum verticillatum, all I think being uncritical plants the collection of which served
no particular scientific purpose. It was of course the age of the private herbarium.
Travelling was difficult and amateur botanists, unable to get easily to consult
the national or university herbaria, built up their own reference collections by
making use of the exchange facilities offered by the Club. Unfortunately, as is
not uncommon with collectors, the undesirable factor of rivalry and one-up-man-
ship crept in. Certain botanists felt they had to possess a rarity which their
rival had not got. So the demand for rare plants grew and the numbers of some
of them undoubtedly suffered considerably from over collecting during this
period. In the Kew Herbarium alone there are 29 specimens of Spiranthes
aestivalis from British localities (excluding the Channel Islands) and Polygonatum
verticillatum is represented by no fewer than 12 British specimens. These figures
are likely to be considerably larger in the Herbarium of the Natural History
Museum, which is on the whole richer in British plants.

Even in these early days the recording of plants was considered important
and over the years the B.E.C’s emphasis has changed very gradually from collect-
ing and exchanging to recording and studying. When the name B.S.B.I. was
adopted in 1947 the word ‘exchange’ was deliberately dropped from the title,

, AWHSON/ 2
J MN

APRS U1971

LiIBRARIE?

196 E. MILNE-REDHEAD

but the exchanging activities continued on a very restricted scale largely dealing
with critical groups, until 1954, when the Society accepted the Council’s sugges-
tion that the Exchange Section be temporarily suspended. So it has remained
until this day, and it seems unlikely that it will ever be revived. At that time the
Distribution Maps Scheme was at its height, and records for the Atlas were pour-
ing in from Members, so the fading out of the exchange facilities were noticed
by very few.

The Society’s first official active steps in Conservation were taken in 1948
when a Special Committee consisting of the Officers (Mr J. S. L. Gilmour,
Miss M. S. Campbell, Mr J. E. Lousley, Dr E. F. Warburg, Dr J. G. Dony)
together with Mr A. J. Wilmott was appointed to deal with all matters connected
with threats to the British Flora. The following year Mr A. H. G. Alston
replaced Mr Wilmott. This Threats Committee had been preceded by the
Society being represented at certain important Public Enquiries, namely by
Mr N. D. Simpson on Purbeck and Mr G. W. Temperley on Ross Links
whilst Mr J. D. Grose had prevented possible damage to Cirsium tuberosum in
Wiltshire by his tactful approaches to the government departments concerned.

This Threats Committee had plenty of work to do, and in 1950 it changed its
name to the Conservation Committee, Mr Lousley becoming its first Secretary.

In March 1949, the Nature Conservancy was set up, following advice given to
the government by the Wild Life Special Committee in 1947. It was, and I quote,
‘to provide scientific advice on the conservation and control of the natural
flora and fauna of Great Britain; to establish, maintain and manage nature
reserves in Great Britain, including the maintenance of physical features of
scientific interest and to organise and develop the scientific services related there-
tox

One of the first urgent tasks confronting the newly formed Nature Conser-
vancy, apart from establishing National Nature Reserves, was a duty placed
on them by Section 23 of the National Parks and Access to the Countryside Act,
1949, to notify to the local planning authority concerned any area of land, not
for the time being managed as a Nature Reserve, which was considered to be of
special interest by reason of its flora, fauna or geological or physiographical
features. These sites were to be known as Sites of Special Scientific Interest
(S.S.S.1:s)

The Nature Conservancy had, as its Director-General, Captain Cyril Diver,
hitherto professionally Clerk of the House of Commons, but well known to
naturalists for his pioneer survey work on the ecology of Studland Heath in
Dorset. It occupied offices in Victoria Street, Westminster, whilst Scotland was
managed from an office established in Edinburgh. It was in these somewhat
dismal London surroundings that the first Liaison Meeting with the B.S.B.I.’s
Conservation Committee took place in November 1950. These Meetings, which
were held at six-monthly intervals, soon proved to be of great value to all
concerned. The Conservancy was able to gain direct help in botanical matters
from B.S.B.I. Members, and the Society’s representatives at the Liaison Meetings
were able to call the Conservancy’s attention at a high level to threats to impor-
tant habitats, thus supporting what local representatives were trying to achieve
at county or regional level.

The staff of the Nature Conservancy was relatively small in these early years,
and much time was taken in organisation, administrative matters and office
work. As a result the Conservancy was only too pleased to call upon the help

PRESIDENTIAL ADDRESS 1970 197

of B.S.B.I. Members who volunteered to give advice on suitable areas to be
set up as botanically important S.S.S.I.’s. This co-operation worked extremely
well as I know from personal experience, for I spent several days with a member
of the Conservancy’s staff in areas of Gloucestershire I know well, indicating
sites which I thought to be worthy of protection. It was gratifying later on to
learn that most of the sites had been notified under Section 23 of the National
Parks and Access to the Countryside Act, 1949.

At this time the nature conservation trust movement had not got under way,
and only three counties had established Naturalists’ Trusts, namely Norfolk
(1926), Yorkshire (1946) and Lincolnshire (1948), so the B.S.B.I. formed a most
useful function in active conservation of important habitats. No one will
suggest that the original list of S.S.S.I.’s was without omissions but I feel sure
that the omissions would have been much more numerous and much more
serious had the great store of habitat information contained in the membership
of our Society not been made available at this crucial period in the Conservancy’s
history.

The Conservation Committee has used its influence from time to time to
impress on the Conservancy the importance of a number of botanically valuable
sites. The 1956 threat to Upper Teesdale above Cauldron Snout was strongly
objected to, and alternative sites near Dine Holm Scar and Cronkley Farm were,
in consequence, to be surveyed. In the same year the danger of quarrying to the
limestone cliffs and associated turf of Berry Head, South Devon, was stressed,
and ultimately the Society was represented at a Public Enquiry in 1959 by
Dr M. C. F. Proctor, which resulted in the rejection of the application for further
quarrying. The Committee pressed (unsuccessfully) for the establishment of a
nature reserve on part of the area of Dungeness remaining after the construc-
tion of the power station, and objected (successfully) to a caravan site being set
up at Mochras dunes in Merioneth. In 1960 the Committee sent in a memoran-
dum to the Ministry of Agriculture Research Study Group on the use (or should
I say misuse?) of toxic chemicals in agriculture.

In spite of the 1956/7 threat to Upper Teesdale having been removed the
Committee remained worried about the lack of permanent protection to Upper
Teesdale and at a liaison meeting with the Nature Conservancy in 1962 again
urged them to set up a National Nature Reserve in this highly important
botanical site. In May 1963 the Nature Conservancy declared a N.N.R. covering
6500 acres on the Yorkshire side of the Tees in Upper Teesdale, but the Durham
side including Widdybank Fell and Cow Green was still left without protection,
a factor which adversely affected the B.S.B.I.’s case to prevent the Cow Green
reservoir being built. But the unsuccessful fight put up by the Teesdale Defence
Committee, set up in February 1965, resulting in three debates in Parliament,
the raising of nearly £25,000 by public appeal and the final donation of £100,000
by Imperial Chemical Industries for research in Upper Teesdale, is a story
which must be told elsewhere.

With the advent of a nation-wide coverage by nature conservation trusts,
the need for the Committee’s liaison meetings with the Conservancy had
declined, and in 1968 a Conservation Liaison Committee was set up by the
S.P.N.R. on which the B.S.B.I. Conservation Committee was represented by
three of its members, who would raise matters of national botanical importance
with the Conservancy, as had been done at the former B.S.B.I. liaison meetings.
It was understood, however, that the B.S.B.I. retained the right to approach the

198 E. MILNE-REDHEAD

Director-General of the Conservancy or his Deputy at any time should a suffi-
ciently important and urgent botanical matter arise.

So I feel it is safe to say that the B.S.B.I. is very conservation minded and is
playing an important part in the now very active conservation movement. I am
happy to represent the Society on the Council for Nature and I am one of the
three representatives of that Council who sit on the Committee for Environmen-
tal Conservation, conveniently abbreviated CoEnCo, on which the bodies
interested in nature conservation, amenity and recreation in the countryside get
together to understand each others points of view and to advise the Government
on matters of policy relating to the countryside. So even on this high-level
committee of wide coverage the importance of botany — of trees, of vegetation
and of the habitats of individual plant species—is not allowed to be overlooked.

To mark European Conservation Year the B.S.B.I. has produced for its
members a “Code of Conduct’ which I hope will have a beneficial effect on their
attitude to their activities in the countryside. It has been well accepted and I
hope it may be widely used by botany and biology teachers and leaders of field
meetings outside the Society. I think we can safely say that the attitude ‘there
was only one so of course I took it’ is a thing of the past.

And now I will turn to what other bodies have done over the years to assist
with plant conservation.

Before the B.S.B.I. (or rather the B.E.C. itself) took a leading part in conserva-
tion, other bodies had started working on this vital matter. In 1924 the British
Correlating Committee for the Protection of Nature was founded with Herbert
Smith as Secretary; a year later* Sir Maurice Abbott-Anderson founded Flora’s
League; the S.P.N.R. proposed a Society for the Protection and Preservation of
Wild Flowers, and sent out a leaflet to all County Councils pressing for bye-law
enactment. In 1927 the Council for the Preservation of Rural England (now
with the word Preservation happily replaced by Protection) took the lead so far
as plant protection was concerned.

It studied the educational aspect of the problem and concerned itself with
the bye-law position resulting in the approval of a new bye-law by the Home
Office later that year, and again in 1933. In 1930 the Society for the Protection
of Wild Flowers and Plants was formed and carried out propaganda in schools.
In 1931 the C.P.R.E. formed the Wild Plant Conservation Board (with Herbert
Smith as Chairman and Herbert Griffin as Secretary) as an integral part of its
organization ‘for the purpose of advising on matters of policy and counselling
in reference to activities connected with the conservation of wild plants’. Its
object was, and I quote, ‘to focus the best and most instructed opinion, scientific
and otherwise, on the problem, and thereby to provide well-informed and
authoritative information on the whole subject’. Unfortunately it appears to
have achieved very little. It had a private bill drafted which never gained support.

The list of bodies represented upon the W.P.C.B. is, however, a formidable
one and the reasons why the Board seems to have had so little impact can only
be explained by the general feeling of apathy reigning during the thirties. I
think it is of interest to record the 28 bodies concerned, almost a miniature
Council for Nature! They are in alphabetical order:

* 1925 is the date given by Mr J. E. Lousley (Rep. botl Soc. Exch. Club Br. Isl., 12: 18
(1939)), whilst Sir Maurice Abbott-Anderson (Rep. botl Soc. Exch. Club Br. Isl., 9: 209
(1930)) states that he founded Flora’s League ‘last June’. Maybe Sir Maurice’s paper had
been awaiting publication for some time!

PRESIDENTIAL ADDRESS 1970

Alpine Garden Society

Association for the Preservation of
Rural Scotland

Botanical Society & Exchange Club of
the British Isles

British Association for the Advance-
ment of Science

British Ecological Society

British Empire Naturalists’
Association

Commons, Open Spaces & Footpaths
Preservation Society

Council for the Preservation of Rural
Wales

Flora’s League

Green Cross Society

Lincoln Naturalists’ Union

Linnean Society of London

Men of the Trees

199

National Trust

Northern Naturalists’ Union

Roads Beautifying Association

Royal Botanic Gardens, Kew

Royal Horticultural Society

School Nature Study Union

Selborne Society

Society for the Promotion of Nature
Reserves

Society for the Protection of Wild
Flowers & Plants

South-Eastern Union of Scientific
Societies

South-Western Naturalists’ Union

Trustees of the British Museum
(Natural History)

Watson Botanical Exchange Club

Wild Flower Society

Yorkshire Naturalists’ Union

The Board produced in 1931 (with a second edition in 1937) an excellent little
booklet explaining the urgent need (even then) for some more active plant
protection due to the increasing population and a growing industrialization
destroying wildlife around our towns, the loss of damp habitats due to drainage
and the general lowering of the permanent water-table to mention a few reasons
all of which apply even more strongly today. During this period agricultural
methods were relatively stable, herbicides were unknown and there was no
ploughing up of pastures for re-seeding with rye-grass and clover. After discuss-
ing the role that teachers could play in achieving the objectives of the Board the
leaflet states “The preservation of our wild plants may be undertaken in three
different ways: by legislation, by education, and by the establishment of nature
reserves and sanctuaries’. The leaflet stresses the need for botanical material
for use in schools to be cultivated for that purpose and not gathered from the
wild, an admirable method which had by then been adopted by the London
Education Authority but which other local authorities have been slow to follow.
The British Wild Plant Nurseries, 6 The Strand, Derby, is quoted as an establish-
ment which provided seeds free of charge under the auspices of the W.P.C.B.
I doubt if these Nurseries survived the second world war.

The W.P.C.B. also compiled (with the help of Mr H. W. Pugsley) schedules of
plants needing protection county by county, continued to press for bye-law
adoption and even gave evidence at Royal Commissions. It is sad that all this
effort had so little effect on the practical conservation of plants. I seem to
remember that on the Council of the B.E.C. in the thirties the Club’s representa-
tive on the W.P.C.B. again and again had nothing to report, but it vigorously
protested about the uprooting of Lioydia serotina on Snowdon in 1935! Most
of its activities came to a halt in 1939, but the Board continued in name until
1950.

The British Ecological Society, which has never been particularly active
in conservation matters, published a Report in 1943 entitled Nature Conserva-

200 E. MILNE-REDHEAD

tion and Nature Reserves, a valuable document in which the considered opinions
of ecologists on the principals of conservation are expressed.

Forerunners in the establishment of nature reserves were the Selborne
Society and the Society for the Promotion of Nature Reserves.* At one time it
was thought that all that was needed to preserve a rare plant was to put a fence
round it to keep people and grazing animals out. Two outstanding examples of
this were reserves set up by the S.P.N.R., one at Dancer’s End, Hertfordshire
(1942) to preserve Gentianella germanica and the other at Badgeworth, East
Gloucestershire (1933) to preserve Ranunculus ophioglossifolius. Neither reserve
originally had a management plan, and both nearly failed in doing what they
set out to do. Active management has now succeeded in safeguarding the
populations of Gentianella and the Ranunculus. The S.P.N.R. was also respon-
sible for setting up the Mickfield Meadow reserve in East Suffolk where a fine
colony of the Snake’s Head Fritillary (Fritillaria meleagris) has been preserved.
Had this not been a reserve in 1939 it is highly probable that it might have
succumbed to the plough during the war years, a fate of many other similar

ritillary meadows. Here again lack of grazing resulted in scrub development
which at one time was threatening the population of fritillaries but conservation
management has now restored the meadow to the former satisfactory condition.
But the reserve at Perivale on the outskirts of Ealing was established by the
Selborne Society in memory of Gilbert White as early as 1904. This was not a
reserve to preserve a particular uncommon species, but an area known as
Perivale Wood, mainly an oakwood with a carpet of bluebells and an adjacent
meadow, preserved largely as a sanctuary for birds. Now it is an island of country
surrounded by factories and houses, an area where many common and beautiful
wild flowers can be seen growing naturally and one which has more recently
proved of great educational value.

Whilst talking of management of nature reserves, one must mention the
Council for Nature’s Conservation Corps, set up in 1959, which has employed
young volunteers from many walks of life to carry out active conservation
tasks throughout the country. It has made a tremendous contribution to the
cause, and recently it has separated from the Council for Nature, being managed
now by an independent charitable trust, The British Trust for Conservation
Volunteers. But many of the smaller management tasks are now handled
locally by the County Trusts many of which have their own Conservation Corps.

The National Trust and the Royal Society for the Protection of Birds have
both contributed considerably to the protection of the flora in this country,
although it has not been the main consideration directing their policies.

But by far the greatest impact on plant conservation has undoubtedly been
due to the county nature conservation trusts which now cover the whole of
England and Wales, whilst the Scottish Wildlife Trust acts similarly throughout
Scotland. The number of reserves managed for their botanical interest by
Trusts is today very considerable, and is, I am glad to say, steadily growing.

In the conservation of a rare or local plant one is often in doubt whether to
treat the location of the site with absolute secrecy or to let the landowner know
that he has a rare plant on his land. The fine locality for the Lizard Orchid
(Himantoglossum hircinum) in Bedfordshire was destroyed by the farmer

* Incidentally Dr G. Claridge Druce, whose devoted stewardship kept the B.E.C. going during
the first thirty years of this century, left half his estate, nearly £20,000, to the S.P.N.R.

PRESIDENTIAL ADDRESS 1970 201

clearing scrub unaware of the presence of the orchid, whilst a few years later
another farmer in Oxfordshire deliberately ploughed up a useless corner of a
field knowing it to contain a population of the Monkey Orchis (Orchis simia)!
So what is one to do?! All such cases should be judged on their merits and
generalizations should be avoided. I usually prefer to take the landowner into
my confidence and hope that he will behave responsibly whilst at the same time
one can discourage visits to the plant as much as possible by botanists and
particularly the less responsible botanical photographers who are apt to do
considerable damage to the site by exposing the subject to better view.

Possibly the most famous example of failure adequately to conserve a rare
plant is the case of Cypripedium calceolus in Yorkshire. The location of the
famous site was a great secret, yet some dozens of botanists knew of it. An un-
fortunate article in the national press some few years ago gave clues to the
botanical detectives resulting in the discovery of the site by people who otherwise
would not have found it. So as the years went by, Cypripedium in this famous
locality got fewer and fewer, yet it was still too secret for steps towards its
conservation to be considered by the Yorkshire Naturalists’ Trust, the very body
which should have been in the forefront of its protection. This somewhat
farcical situation came to a head last year when, after alarming rumours and
correspondence with a number of botanists concerned, the B.S.B.I. Conservation
Committee urged the Nature Conservancy to call a site meeting of all the interes-
ted bodies. This was a most successful manouvre and, following a visit to the
locality and an excellent lunch in the local pub, an amicable meeting was held
at which a policy was agreed. I will not go into the details now, but I would
like to say that recent developments suggest that the policy adopted stands a
chance of proving successful.

Conservation of rare plants is one matter, conservation of common yet
attractive plants is another. I believe the overall population of such plants
as primroses (Primula vulgaris) and cowslips (P. veris) in England is today a mere
fraction of what it was fifty years ago. Forestry and agricultural practices have
so changed the face of England that thousands of acres of favourable habitats
have gone. These two species have suffered greatly. The cowslip fields have been
ploughed up and turned into arable or else resown as leys with grasses and
clover. The replanting of deciduous woodland with conifers has enormously
reduced the habitat for primroses. Both species grew in hedgebanks and roadside
verges. The widespread destruction of hedges and the spraying of roadside
verges has reduced the populations in these habitats. The railway banks were
one of the last refuges for these delightful plants, but now that railways have
been so drastically reduced, many banks have lost their statutory protection,
fences have gone and the plants, if not overgrown by scrub, have to survive
attacks from the public. When the railways were being built last century these
and other plants migrated on to the banks with comparative ease from the
nearby fields and hedgerows. Now with the construction of motorways the
equally suitable banks cannot be colonized so easily, as these plants have gone
from many of the fields, whilst hedges are few and far between. It is, in my
opinion, important to get these, and other suitable plants established on
motorway banks and it is up to the B.S.B.I. members to urge their County
Trust to do this artificially. Machinery for doing this has been agreed on
between S.P.N.R. and the Ministry of Transport, and some introductions have
been made, but a lot more could be done in that direction.

202 E. MILNE-REDHEAD

One of the greatest threats to primroses is the mass picking and digging
up by the town dwellers who visit the countryside in spring in ever increasing
numbers. One has to go a long way from Kew now to see a good show, the one
exception being the bank of the electrified Southern Region railway at Surbiton
where the public cannot get at the plants! Education is surely the cure for this
trouble, but the enlightened press will have to treat the subject more seriously
than hitherto if the wayside flowers are to be left for others to enjoy.

In 1963 the theme for the Society’s biennial Conference was ‘The Conservation
of the British Flora’, and a very useful series of papers was read and discussed at
Durham, culminating in a resolution which I put to the Conference, namely
‘That this Conference resolves that a working party be set up, composed of the
B.S.B.I., the S.P.N.R. Naturalists’ Trusts’ Committee and the Council for Nature,
to consider certain urgent educational and legal problems concerning the con-
servation of the British flora’. This was seconded by Dr Margaret Bradshaw and
carried unanimously.

As a result of this the Wild Plant Protection Working Party (W.P.P.W.P.) was
set up and a lot of hard work was put into the collection of plant conservation
information from this and other countries. Mrs Gigi Crompton acted as secretary
and driving force was supplied by Dr Max Walters and Mr Hector Wilks.
Eventually with the help of Mr R. S. W. Pollard, a draft Wild Plant Protection
Bill was produced. W.P.P.W.P. prepared an interim report in 1965, which was
presented as evidence to the Countryside in 1970 Study Group No. 5 dealing
with legislation. The Study Group, on which I sat, agreed whole-heartedly
with W.P.P.W.P.’s conclusions that there should be legislation to protect wild
plants on the basis of three categories: national rarities, species forbidden to be
offered for sale, species protected in special areas. The Countryside in 1970
Conference on 12th November 1965 accepted the report of the Study Group
and recommended that it should be sent to the Ministers concerned. However
such legislation was omitted from the White Paper dealing with the amendments
to the National Parks and Access to the Countryside Act, so it was accordingly
decided to go ahead independently to present a private member’s Bill to Parlia-
ment.

In November 1967 it was learned that Mr Peter Mills, M.P., had drawn 24th
place in the parliamentary ballot for private members’ bills and was willing to
introduce the Wild Plant Protection Bill. To cut a long story short the Bill was
duly presented, was treated frivolously by the B.B.C. and the Press and
failed to get a second reading. Today the Bill has been redrafted, with welcome
financial assistance from the World Wildlife Fund, taking into consideration
certain clauses in the recent Theft Act and W.P.P.W.P. will be meeting shortly
to consider the next move. Perhaps in European Conservation Year the need to
have such a measure in the statute books may be appreciated more than it
was two years ago. Members could help the cause very considerably by explain-
ing the aims to their Members of Parliament and by emphasising the need for
legislation to protect plants by writing letters and articles to their local press.

And what to the future! I see no reason why any more native plants should
become extinct. It is probably safe to say that we are now more conscious of
the need for conservation than we have ever been. And we are gaining new
weapons to help us in the fight. Here at Kew we have set up seed stores under
controlled conditions and it is hoped to bank seed of rare and threatened species,
so as to keep them alive for we hope fifteen to twenty years with the possibility

PRESIDENTIAL ADDRESS 1970 203

_ of re-introducing and establishing the species into suitable wild localities or at
least growing it from time to time in botanic gardens or for scientific study. I
think the time is coming when we must set about making suitable habitats — to
replace those that have been destroyed. I feel that we have the technical know-how
and that small-scale attempts should be made in botanic gardens such as Kew.
The failure of Sir Edward Salisbury to establish artificially an area of chalk
grassland at Kew should not deter further attempts at that kind of exercise.
It is the kind of experimental work I would like to see being undertaken at Kew
and if properly explained to the public could, I feel sure, form a very interesting
feature of the Gardens.

There is an undoubted need for statutory protection for both our rarest
and some of our common plants; there is an even greater need to educate the
public, making the most of the wonderful media at our disposal. We must
somehow make it possible for the public to see and appreciate some of our
botanical treasures rather than keep them hidden. The R.S.P.B. has set us
excellent examples in their Osprey and Snowy Owl publicity. Why should we
not have people queuing on a bank holiday to pay their money to see a fine
display of monkey orchid or spring gentian? These are the thoughts I would
like to leave with you today.

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Watsonia, 8, 205-235 (1971). 205

Taxonomy of the Linum perenne group in Europe

D. J. OCKENDON*
Department of Botany, Imperial College, London

ABSTRACT

A new taxonomic treatment of the Linum perenne group in Europe is presented, based ona
detailed biosystematic study. L. alpinum Jacq., L. extraaxillare Kit. and L. montanum Schleicher
ex DC. are treated as subspecies of L. perenne L., and L. collinum Guss. as a subspecies of L.
austriacum L. Previous confusion between L. perenne subsp. anglicum (Miller) Ockendon and
L. leonii Schultz has been resolved. The morphological variation within each subspecies is
described and interpreted in terms of its distribution. Most of the morphological characters
which have been used in the past have proved to be taxonomically unreliable because they vary
almost continuously within the group and show considerable phenotypic plasticity. Chromo-
some number, heterostyly and pollen characters are taxonomically more reliable than the
quantitative morphological characters. The pattern of variation within the L. perenne group is
discussed in relation to its rarity, disjunct distribution, ecology and breeding system.

INTRODUCTION

The taxonomic difficulty of the Linum perenne group was mentioned in 1848 by
Planchon and has been recognised by many taxonomists since then. A two-fold
approach was made to the problem. The morphological characters which had
been used in the past were examined in detail to see which were taxonomically
reliable, and an attempt made to build up a general picture of the morphological
variation in the group. Further taxonomic characters were sought, including
those derived from a study of the cytology, pollen morphology, breeding system
and artificial hybridization of the group. This work is published in detail else-
where (Ockendon 1968a), but the results of it will be used here in-so-far as they
affect the classification of the group. The taxonomic account given here is
essentially the same as that in Flora Europaea, vol. 2 (Ockendon & Walters 1968).
The opportunity is taken here to give the evidence on which this account is
based and to discuss the taxonomic decisions taken.

The L. perenne group occurs in Asia, Europe and North America. All the
North American plants can be referred to the single homostylous species L.
lewisii Pursh. Most of the Asian members of the group have been dealt with by
Juzepezuk (1949) and Davis (1967). This paper deals only with the European
members of the group. A large number of populations have been studied in the
field and considerable quantities of material were cultivated at Cambridge
University Botanic Garden. All the known extant British populations were
visited, plus about 70 per cent of those in France and Germany, and others in
Austria, Czechoslovakia, Italy, Jugoslavia and Spain. Material from the
following herbaria has been examined: British Museum (BM), Bruxelles (BR),
Cambridge (CGE), Edinburgh (E), Florence (FI), Geneva (G), Kew (K), Man-
chester (MANCH), Paris (P) and Vienna (W). The coverage of material from
most of western and central Europe is fairly complete, but that for southern and
eastern Europe is less so.

* Present address: National Vegetable Research Station, Wellesbourne, Warwickshire.

206 D. J. OCKENDON

TAXONOMY

DESCRIPTION OF THE L. PERENNE GROUP

Glabrous perennials, sometimes caespitose and woody at base; stems up to
80 cm, erect, ascending, decumbent or procumbent. Leaves 1-3 (-5) mm wide,
linear or linear-lanceolate, entire, 1- to 3-veined. Sepals 3-5-6 mm, unequal,
with three main veins; the outer sepals narrower than the inner and with an
acute or acuminate apex, the inner with a rounded or mucronate apex and
entire, scarious margins. Petals blue, 3—4 times as long as the sepals, obovate or
obovate-lanceolate, tapering gradually to a yellow, pubescent claw. Stamens 5,
alternating with 5 rudimentary staminodes. Stigmas capitate. Capsules 3-5-7
(-8) mm, subglobose.

The L. perenne group is sometimes confused with L. bienne Miller. The latter
has linear or clavate stigmas and subequal sepals with a single prominent mid-
vein, in contrast to the capitate stigmas and unequal sepals with three main
veins of the L. perenne group.

GENERAL FEATURES OF THE L. PERENNE GROUP

The following general features of the L. perenne group should be borne in
mind when considering the taxonomic treatment which follows.

!

FicureE 1. Distribution of the Linum perenne group in Europe. A L. perenne subsp. alpinum;
m@ L. perenne subsp. anglicum; x L. perenne subsp. montanum; A L. perenne subsp. perenne;
© L. perenne subsp. extraaxillare; O L. austriacum subsp. austriacum; @ L. austriacum subsp.
collinum; (\ L. leonii; + L. punctatum.

LINUM PERENNE GROUP IN EUROPE 207

FIGURE 2. Distribution of binomials associated with limited areas. 1. L. bavaricum Schultz;
2. L. darmstadinum Alefeld; 3. L. julicum Hayek; 4. L. limanense Lamotte; 5. L. loreyi Jordan;
6. L. petryi Beyer; 7. L. provinciale Jordan; 8. .L. punctatum Presl; 9. L. pycnophyllum Boiss.;
10. L. ruscinonense Timbal-Lagrave; 11. L. saxicola Jordan; 12. L. tommasinii(Reichb.) Nyman;
13. L. trinervium Freyn.

1. The L. perenne group is a natural group of very closely related species which
can be easily separated from all other species of Linum by the combination of
characters given above. Species within the group tend to be recognised on the
basis of single characters which do not correlate well with other characters.

2. The characters showing most variation within the group are habit, stem
length, leafiness of stem, leaf width, inflorescence size, sepal size and shape,
petal size and colour, position of fruiting pedicels and capsule size. The
majority of these characters are quantitative and show almost continuous vari-
ation. Apparent discontinuities in the variation (which are found) are liable to
disappear when more material is examined.

3. The distribution of the group in Europe is markedly disjunct, especially
in lowland areas (Fig. 1). Fieldwork in western Europe has shown that the
discontinuities in the distribution are not an artifact of uneven collecting but
are real, and that considerable extinction has occurred in the last hundred years.

4. Most of the subspecies in the group are local or rare and many of the popu-
lations are limited in extent (Appendix). Repeated collections have been made,
over long periods of time, from single topodemes such as those at Puy Crouel,
Clermont-Ferrand and at Mt. Spaccato near Trieste. Certain binomials have long
been associated with plants coming from a very limited geographical area, e.g. L.
limanense Lamotte — area around Clermont-Ferrand; L. tommasinii (Reichb.)
Nyman — an area near Trieste. So constant is the association between some bino-
mials and particular geographical areas that it is possible to map the areas in which
the binomials have generally been applied (Fig. 2). Members of the L. perenne

208 D. J. OCKENDON

group have commonly been identified on the basis of their locality rather than
by their morphological characters. The strong geographical emphasis in the
taxonomy of the group is reflected in names such as L. anglicum Miller, L. bavari-
cum Schultz and L. provinciale Jordan.

PREVIOUS ACCOUNTS OF THE L. PERENNE GROUP

In the second edition of Species Plantarum (1762) Linnaeus recognised three
species in the L. perenne group, namely L. alpinum Jacq., L. austriacum L. and
’ L. perenne L. These three species are still widely recognised.

Reichenbach (1837) treated the L. perenne group as the genus Adenolinum.
Planchon (1848) and Juzepczuk (1949) recognised Adenolinum as a section of
Linum. There is very little to be said for treating Adenolinum as a genus and it is
probably better to treat it as a sub-section rather than as a section of Linum.

Planchon (1848) considered the L. perenne group on a world-wide basis and
treated it as a single species with 9 varieties.

Ascherson & Graebner’s (1914) full account of most of the European mem-
bers of the L. perenne group is apparently based on the work of H. Holzer, who
determined quantities of herbarium material but did not publish his work. Their
account may be summarised as follows:

L. alpinum Jacq. L. perenne L. L. austriacum L.

subsp. A eualpinum subsp. A perenne subsp. A austriacum
B montanum B_ extraaxillare B_ squamulosum
C anglicum C petryi C collinum

They included L. leonii with subsp. anglicum, while L. loreyi was included
with subsp. collinum. Their detailed account of the infraspecific taxa gives a
good impression of the morphological variability and taxonomic confusion of
the group. Hegi’s (1924) account is largely based on that of Ascherson &
Graebner.

Alefeld (1867) described 4 new species belonging to the L. perenne group, but
his treatment of the group as a whole is not very systematic. The importance of
his work is that he was the first to appreciate the value of heterostyly as a taxo-
nomic character, although he was apparently unaware of its biological signifi-
cance. He recognised dimorphous and trimorphous species, the latter including
‘mid-styled’ in addition to the long- and short-styled forms. However, the “‘mid-
styled’ forms in the L. perenne group are self-fertile and not comparable with the
mid-styled, self-sterile forms of tristylous species of Oxalis and Lythrum (Fig. 3).
Thus in the L. perenne group the basic distinction is between homostylous and
heterostylous species.

The importance of Alefeld’s work was recognised by Juzepczuk (1949), whose
treatment of the section Adenolinum in Russia can be summarised as follows:

Group 1. Homomorpha — Homostylous
Series 1. Pallescentes — Pedicels erect
Series 2. Mesostyla — Pedicels deflexed

Group 2. Heteromorpha — Heterostylous
Sub-group 1. Trimorpha
Sub-group 2. Dimorpha
Series 1. Perennia — Pedicels erect
Series 2. Austriaca — Pedicels deflexed

LINUM PERENNE GROUP IN EUROPE 209

a b
Oh (0

: )
|
Mid

FIGuRE 3. Tristyly in (a) Oxalis compared with distyly in (b) Linum.

|

Long Short Long Short Homostyled Linum

The sub-group Trimorpha contains a single, poorly known species from the
Caucasus, L. subbiflorum Juz. The claim that this species contains long-, short-
and mid-styled forms needs to be substantiated. If it is confirmed, it will be the
only known example of the occurrence of both heterostyled and homostyled
forms within a single species of Linum.

Juzepezuk recognises a total of 18 species, of which 6 are homostylous. No
infraspecific taxa are given, but it is admitted that the species recognised can
be grouped into series of forms which replace one another geographically.
Juzepcezuk’s account is valuable because it emphasises the taxonomic importance
of heterostyly and pedicel position in the L. perenne group.

TYPIFICATION

The search for type specimens has not been exhaustive but most of those still
in existence have been examined. The holotypes of the species described by
Alefeld, Beyer and Schultz appear to have been lost, but a lectotype is available
for L. leonii Schultz. The lack of some type specimens is not a serious handicap,
because the precise type locality can be ascertained in almost every case and
further material from the type locality is usually available. The chief exception
to this is L. laeve Scop., for which the type locality (‘in montanis Carnioliae
calidoris’) is not very precise and the type specimen is not available. Neither the
original description nor the illustration of L. /aeve are sufficiently detailed to
ascertain the taxon to which this name was originally applied. Since then the
name L. laeve has been used for several taxa, both within and outside the L.
perenne group, and to avoid further confusion should now be rejected.

For species described before 1800, the specimens selected as type specimens
are perhaps better considered as representative specimens because of the diff-
culty of verifying with certainty the specimens (if any) from which the original
descriptions were made. L. perenne L. is typified by a specimen from the Lin-
naean Herbarium (396/8) labelled ‘2 H.U. perenne’ by Linnaeus. This specimen
was most probably in Linnaeus’ possession at the time he wrote the first edition
of Species Plantarum, as indicated by the number ‘2’ written on the sheet,
corresponding to the number this species has in Species Plantarum.

Neither of the specimens of L. austriacum in the Linnaean Herbarium (396/22
and 23) were numbered by Linnaeus and both were probably acquired by him
subsequent to the writing of Species Plantarum. However, there is a specimen

210 D. J. OCKENDON

of L. austriacum in Burser’s Herbarium (Juel 1936) bearing the inscription ‘In
Austria inferiori’, corresponding to “Habitat in Austria inferiore’ given for L.
austriacum in Species Plantarum. As Linnaeus is known to have used Burser’s
Herbarium when writing Species Plantarum (Stearn 1957), the Burser specimen
has been selected as the type of L. austriacum.

KEY AND DESCRIPTIONS

1. Homostylous, with anthers and stigmas at about the same height Ae
4. L. leonii
1. Heterostylous, with anthers and stigmas at different heights
2. Leaves very crowded,6mmlongorless_... se 3. L. punctatum
2. Leaves moderately crowded, the longest at least 10 mm long
3. Pedicels deflexed or patent hy At es 2. L. austriacum
3. Pedicels erect, + straight.. ae A os .. 1. L. perenne

1. L. PERENNE L., Sp. Pl., p. 277 (1753). Type in Linnaean Herbarium, sheet
number 396/8 (LINN).

Stems 10-80 cm, decumbent, ascending or erect. Inflorescence few- to many-
flowered. Inner sepals acute or obtuse. Pedicels erect. Capsules 5-8 mm. Hetero-
stylous. Central and Eastern Europe, extending locally westwards to Britain
and the Pyrenees.

1. Middle cauline leaves usually more than 2-5 mm wide subsp. extraaxillare (e)
1. Middle cauline leaves usually less than 2-5 mm wide
2. Inner sepals very obtuse, longer than the outer (lowlands)
3. Stems erect, or ascending just at the base; pollen grains with
furrows ee Ne subsp. perenne (a)
3. Stems decumbent or ascending below: pollen grains with pores
. subsp. anglicum __(b)
Di, Inner sepals acute or obtuse, equalling the outer (mountains)
4. Pollen grains withfurrows .. wi .. Subsp. alpinum _ (c)
4. Pollen grains with pores =i ae .. subsp. montanum _(d)

la. L. PERENNE subsp. PERENNE

L. bavaricum F. W. Schultz, Flora (Regensb.), 21: 643 (1838). Type not

located.

L. darmstadinum Alefeld, Bot. Zeit., 25: 252 (1867). Type probably lost.

Stems (20-—) 30-80 cm, ascending or erect. Middle cauline leaves 1-2-5 mm wide.

Inflorescence 5- to 25-flowered. Inner sepals 4-5—5:5 mm long, obtuse or very
obtuse, exceeding the outer sepals by 0-5-1 mm. Petals 12-20 mm long, pale
blue or mauvish blue. Capsules 5—7 mm long. Pollen grains with three furrows.
2n=18. Light sandy or gravelly soils, often near large rivers. Central Europe,
extending westwards to west Germany.

1b. L. PERENNE subsp. ANGLICUM (Miller) Ockendon, Reprium nov. Spec. Regni
veg., 74: 20 (1967a).
L. anglicum Miller, Gard. Dict., 8th: no. 5 (1768). Type: “Linum perenne
majus caeruleum capitulo majore’ Chelsea Physick Garden No. 1976 (BM).

Stems (15—) 20-70 cm, decumbent, ascending or erect from an ascending base.
Middle cauline leaves 1-2:5 (-3:5) mm wide. Inflorescence 5- to 20-flowered.

LINUM PERENNE GROUP IN EUROPE 211

Inner sepals 4:5—6:5 mm long, obtuse or very obtuse, exceeding the outer sepals
by 0-5-1 mm. Petals 14-20 mm long, pale blue or mauvish blue. Capsules
5-5-7:5 mm. Pollen grains with six pores. 2n=36. Usually in chalk or limestone
grassland but occasionally on light sandy soils. Endemic to Britain.

lc. L. PERENNE subsp. ALPINUM (Jacq.) Ockendon, Reprium nov. Spec. Regni
veg., 74: 20 (1967a).
L. alpinum Jacq., Enum. Stirp. Vindob., 54: 229 (1762). Type: Austria,
‘Schneeberg’, Jacquin (BM).
L. julicum Hayek, Abh. Zool.-Bot. Ges. Wien, 4(2): 109 (1907). Type:
Austria, ‘Steiermark: Sannthaler Alpen, im Felsschutt am Fuss der
Merzlagora nachst der Okreselhiitte, 1400 m’, 31/8/1904, A. Hayek (GB).

Stems 5-30 (-40) cm, ascending or decumbent, often numerous and slender.
Middle cauline leaves 1-3 (-5) mm wide. Inflorescence (1—) 3- to 12-flowered.
Inner sepals 4-5-6 mm long, acute or obtuse, equaling the outer sepals. Petals
10-22 mm long, pale blue or bright blue. Pedicels usually erect but occasionally
flexuous or patent. Capsules 5-7 mm long. Pollen grains with three furrows.
2n= 18. Alpine grassland and stabilised scree slopes, usually above 1500 m but
occasionally descending to 800 m. Pyrenees, Alps, Apennines, Pirin Mountains.

1d. L. PERENNE subsp. MONTANUM (Schleicher ex DC.) Ockendon, Reprium
noy. Spec. Regni veg., 74: 20 (1967a).
L. montanum Schleicher ex DC., Prodr. Syst. Nat., vol. 1, p. 427 (1824).
Type in Herbarium De Candolle (G-DC).

Stems 15-40 cm, ascending or erect from an ascending base. Middle cauline
leaves 1-2-5 mm wide. Inflorescence 3- to 10-flowered. Inner sepals 5-6-5 mm
long, acute or obtuse, equalling the outer sepals. Petals 15-20 mm long, pale
blue. Capsules 5-7 mm long. Pollen grains with six pores. 2n=36. Basic grass-
land, usually above 1500 m. Confined to the Jura and northern Alps.

le. L. PERENNE subsp. EXTRAAXILLARE (Kit.) Nyman, Consp. Fl. Europ., Suppl.,
vol. 2(1): 71 (1889).
L. extraaxillare Kit., Linnaea, 32: 573 (1864). Type in Prague (PR).

Stems 25-50 cm, erect from an ascending base. Middle cauline leaves 2-4 mm
wide. Inflorescence 5- to 15-flowered. Inner sepals 5-6-5 mm long, acute, equal-
ling the outer sepals. Petals 14-18 mm long, pale blue. Capsules 6-5-8 mm long.
Pollen grains with three furrows. 2n=18. Basic alpine grassland, usually above
1500 m. Carpathians and mountains of the Balkan Peninsula.

2. L. AUSTRIACUM L., Sp. Pl., p. 278 (1753). Type in Burser’s Herbarium,
number XII. 44: ‘Linum Syl. angustifolium, foliis rarioribus Bauh. Linum
syl. IJ. Tab. In Austria inferiori’ (UPS).

Stems (6—) 10-60 cm, erect or ascending. Middle cauline leaves 0:5-2 mm
wide. Inflorescence many-flowered. Inner sepals 3-5-6 mm long, acute. Petals
10-18 mm long, deep blue. Pedicels deflexed or patent. Capsules 3:5—7:5 mm.
Pollen grains with three furrows. Heterostylous. Central and eastern Europe.

Capsules 3-5-5 mm ee ee Se .. subsp. austriacum (a)
Capsules 5-7-5 mm ay or a. di subsp. collinum (b)

DID D. J. OCKENDON

2a. L. AUSTRIACUM subsp. AUSTRIACUM
L. trinervium Freyn, Ost. bot. Z., 26: 228 (1876). Type: Romania, ‘Sieben-
biirgen, comitat Torda. Wiesen im Viragosthal, 350 m’, 14/6/1871, J.
Freyn (BRNM).

Stems (20-) 30-60 cm. Inflorescence 10- to 40-flowered. Capsules 3-5-5 mm
long, usually rounded at the apex. 2n=18. Dry rocky limestone soils or light
sandy soils. Central Europe, extending to West Germany and Switzerland where
it may have been introduced.

2b. L. AUSTRIACUM subsp. COLLINUM Nyman, Consp. FI. Europ., p. 125 (1878).

L. collinum Guss., Fl. Sic. Syn., vol. 2, p. 808 (1845), nom. provis. Type:

‘Linum collinum Guss. — perenne pr. fi. sic. non Lin. pedunculis fructiferis
prope reflexi. Sicilia.” (BM).

Adenolinum tommasinii Reichb., Icon. Fl. Germ., vol. 6, p. 66 (1844).
Type: ‘Praires pierreuses du Mt. Spaccato dans les montagnes calcaires
prés de Trieste,’ May-June, 1860, M. de Tommasini (K).

L. tommasinii (Reichb.) Nyman, Syll., p. 218 (1854).

L. saxicola Jordan, Cat. Jard. Dijon, p. 28 (1848). Type: France, ‘de Gap’,
1851, A. Jordan (K).

L. provinciale Jordan, Obs. Pl. Crit., vol. 7, p. 17 (1849). Type: France,
‘Avignon, bords de la Durance’, 1837, A. Jordan (LY).

L. ruscinonense Timbal-Lagrave, Bull. Soc. Bot. Fr., 7: 509 (1860). Type:
France, ‘Font de Comps, Pyr. Or.’, May, 1852. P. M. E. Timbal-Lagrave (P).

L. limanense Lamotte, Prod. Fl. Centr. Fr., vol. 1, p. 157 (1877). Type:
France, ‘Puy de Crouel’, VM. Lamotte (P).

Stems 6-40 cm. Inflorescence 3- to 20-flowered. Capsules 5-7-5 mm long,
usually acuminate at the apex. 2n=18. Dry rocky limestone soils, ascending to
1100 m in the Alps and 1300 m in the Pyrenees. Spain, south and central France,
Italy, Jugoslavia, Greece and Sicily.

3.. L. PUNCTATUM C. Presl in J. & C. Presl, Del. Prag., p: 58 (1822)5 liype an
Prague (PR).
L. pycnophyllum Boiss. & Heldr. in Boiss., Diagn., Ser. 2, vol. 3(1), p. 97.
Type: Greece, ‘Rariforme in lapidosis reg. alpinae, m. Kyllenes. 6500 feet,
8/7/1848. P. E. Boissier (G).

Stems up to 15 cm, very crowded and densely leafy throughout. Leaves
3-5 (-7) mm long, narrowly ovate. Flowers solitary or in terminal groups of
two or three. Inner sepals 5-6 mm long, obtuse. Pedicels erect or flexuous.
Petals 8-12 mm long, pale blue. Capsules 5-5-7 mm long. Pollen grains with
three furrows. Heterostylous. Restricted to a few mountains in Sicily (Madonie,
Nebrodie 1400-1800 m), Greece (Mt Smolika 2000-2500 m) and Turkey.

4. L. LEONI F. W. Schultz, Flora (Regensb.), 21: 644 (1838). Type: France,
“Pelouses séches et pierreuses, champs de Sanfoin sur les hauteurs a calcaire
oolithique aux environs de Metz, prés d’Ancy, de Chatel et d’Onville’,
15/7/1840, Léo 432 (K).

L. loreyi Jordan, Cat. Jard. Dijon, p. 27 (1848). Type not located.
L. petryi Beyer, Verh. Bot. Ver. Brand., 40: 91 (1898). Type probably lost.

Stems 5-30 (—50) cm, erect when young, later often decumbent or reclinate.

LINUM PERENNE GROUP IN EUROPE 213

Middle cauline leaves 0-5-2 mm wide. Inflorescence 1- to 6(—12)-flowered. Inner
sepals 3-5-6 mm long, acute. Petals 8-14 mm long, deep blue. Pedicels erecto-
patent, flexuous or patent. Capsules 5—7 (-8) mm long. Pollen grains with three
furrows. Homostylous. 2n=18. Dry rocky limestone soils, occasionally on chalk.
Endemic to France and Germany.

The foregoing taxonomic account includes all the European members of the
L. perenne group except those which are confined to Russia. These are omitted
because I have not been able to grow these plants and know them only in the
form of limited herbarium material.

DISCUSSION OF INDIVIDUAL TAXA

la. L. perenne subsp. perenne. This subspecies has often been confused with the
commoner L. austriacum subsp. austriacum and its precise distribution in eastern
Europe is not known. It is rarer than has formerly been supposed and many of
the earlier records need to be checked. It occurs in south Germany, Austria,
Hungary, Romania and Russia, but is probably absent from East Germany,
Czechoslovakia and the Balkan Peninsula. In Germany and Austria subspp.
perenne and austriacum are readily distinguishable both by their morphology
and ecology, but in Eastern Europe the distinction is not very clear. There are
plants from Romania and Russia which have the erect pedicels of subsp. perenne
but the small capsules of subsp. austriacum. Some of these plants have been
referred to L. euxinum Juz. and L. marschallianum Juz. by Juzepczuk (1949),
but their true affinities are obscure.

L. perenne subsp. perenne has become extinct in many of its localities during the
last hundred years and now has a very disjunct distribution. Some of the topo-
demes are morphologically distinct, as in south Germany where there are three
morphodemes, of which two have been given binomials (L. darmstadinum Alefeld
and L. bavaricum Schultz). The Darmstadt topodeme has stems 30-70 cm long
and very pale blue flowers. Topodemes along the Danube from Regensburg to
Plattling consist of small plants with stems up to 30 cm long and deep blue
flowers. The topodeme near Munich is intermediate, with a stem length of 25-
50 cm and medium-blue flowers. It is of interest that most of the Central Euro-
pean topodemes of subsp. perenne are associated with the Danube or its tribu-
taries (apart from the Darmstadt topodeme which is near the Rhine). This
suggests that these topodemes may have been derived from L. perenne subsp.
alpinum by seeds being carried down from the mountains. If topodemes of
subsp. perenne were derived independently in this way, this would help to ex-
plain the morphological differences between them.

1b. JL. perenne subsp. anglicum. Confined to eastern and northern England with
one topodeme in southwest Scotland. The latter is unique in being no more
than 30 m from the shore line. L. perenne subsp. anglicum is less disjunct and less
variable than subsp. perenne. Morphological separation of these two subspecies
is difficult, but they can readily be distinguished by the fact that subsp. anglicum
is tetraploid and has pollen grains with six pores, while subsp. perenne is diploid
and has pollen grains with three furrows. A fuller account of subsp. anglicum
is given in Ockendon (1968b).

lc. L. perenne subsp. alpinum. Locally common in the western and south-
eastern Alps, less common in the eastern and southern Alps and absent from

214 D. J. OCKENDON

Ficure 4. Distribution of the Linum perenne group in the Alps. Solid figures — herbarium
records; open figures — literature records. O@ L. perenne subsp. alpinum; AA L. perenne
subsp. montanum; _\@ L. austriacum subsp. collinum.

The contour line indicates the main land masses above 900m.

most of the north and central Alps. Scattered in the eastern and central Pyrenees
and rare in the Apennines. This is the most variable subspecies of L. perenne,
comprising a series of forms which replace one another geographically. Some
of these forms may deserve taxonomic recognition but it is virtually impossible
to key them out satisfactorily.

The distribution of L. perenne subsp. alpinum in the Alps is far from con-
tinuous (Fig. 4) and four main areas of its occurrence can be recognised. The
largest of these is in the western Alps where it is scattered from Lake Geneva
to the Alpes Maritimes. In the southern Alps it occurs in the mountains of
Trentino and Bergamo, in the southeastern Alps mainly in the Julian, Kara-
wanken and Steiner Alps, and in the eastern Alps mostly in the Styrian Alps.
Three of these four topodemes have been given binomials. L. alpinum Jacq.,
originally described from the Schneeberg in the eastern Alps (Jacquin 1762), has
short, decumbent or ascending stems up to 15 cm long, pale blue petals and erect
or erecto-patent pedicels. L. julicum Hayek, originally described from the Sann-
taler Alps (Hayek 1907), has ascending or erect stems up to 30 cm long, deep
blue petals and flexuous or deflexed pedicels. The topodeme from the western
Alps, which has often been called L. montanum Schleicher, is rather heterogen-
eous, but generally has ascending stems up to 30 cm long, pale or medium blue
petals and erect or erecto-patent pedicels. The topodeme from the southern Alps
is similar to that of the western Alps except that it has deeper blue petals and
patent pedicels.

Plants of L. perenne subsp. alpinum from the Pyrenees differ from those from
the Alps in having longer stems and broader leaves. The stem length of Pyrenean

LINUM PERENNE GROUP IN EUROPE 215

plants may reach 45 cm and is seldom less than 20 cm. Some of the Pyrenean
topodemes show an unusually large amount of intrapopulation variation in leaf
width and pedicel position and some plants are almost impossible to key out
properly. The leaf width may be as much as 5 mm and such plants will key out
as L. perenne subsp. extraaxillare. The pedicels may be flexous or patent and
such plants are liable to key out to L. austriacum subsp. collinum. Confusion
between L. perenne subsp. alpinum and L. austriacum subsp. collinum in the
Pyrenees could be serious were it not for striking differences in their ecology.
L. perenne subsp. alpinum occurs in closed mesic habitats, usually above 1500 m,
with species such as Anthoxanthum odoratum, Briza media, Helictotrichon
pubescens, Lotus corniculatus, Plantago lanceolata and Primula veris. L. austriacum
subsp. collinum occurs in open, much more xeric habitats, often on rocky
slopes with species such as Helictotrichon bromoides (Gouan) C. E. Hubbard,
Koeleria vallesiana (Honck.) Bertol., Stipa pennata L., Teucrium polium L. and
Thymus vulgaris L.

Topodemes of L. perenne subsp. alpinum from the northern Apennines are
similar to those from the western Alps, but topodeme F50* from the central
Apennines near L’Aquila shows several differences, notably the lack of sterile
axillary shoots normally characteristic of this subspecies, and the short, appressed,
glaucous leaves. This topodeme seems to vary in the direction of L. austriacum
subsp. collinum, both in its morphology and ecology. There is considerable
variation in subsp. alpinum in the Apennines and more samples are needed to
see if this variation has a north-south pattern.

The Bulgarian topodeme of subsp. alpinum, which is confined to the Pirin
Mountains and Slavianka, resembles the topodeme from the eastern Alps
in having erect stems up to 20 cm long, narrow leaves, medium blue flowers and
erect pedicels.

Topodeme F42 from the central Pyrenees is particularly unusual. It has
broader leaves than the other Pyrenean topodemes and is tetraploid (2n= 36),
while the three other Pyrenean topodemes investigated are diploid (Ockendon
1968a). Most of the pollen grains of F42 have three furrows, as do the diploid
members of the L. perenne group, but a few of them have six furrows, resembling
the pollen of the tetraploids. This is the sole exception to the rule that in the
L. perenne group diploids have pollen with three furrows and tetraploids have
pollen with six pores (Ockendon 1968a). Topodeme F42 is unusual in yet another
respect, namely that it produces extremely few fertile seeds when open-pollinated
or when artificially cross-pollinated. Meiosis was examined in two plants from
this topodeme. Of 53 pollen mother cells, 42 per cent had one or more univalents
and 35 per cent had one or more trivalents. Univalents and trivalents are norm-
ally absent or very rare in the meiosis of the tetraploid L. perenne subsp. anglicum.
Thus the poor seed fertility of topodeme F42 may be accounted for by its meiotic
irregularities. The evidence suggests that this topodeme has become tetraploid
rather recently and has not been in existence as a tetraploid long enough for
selection to have increased the fertility to the normal level.

Id. L. perenne subsp. montanum. Restricted to the Jura and one locality in the
south-eastern corner of Bavaria on the Austrian border. This subspecies is
morphologically very similar to subsp. alpinum from the western Alps and is

*See Appendix for details.

216 D. J. OCKENDON

distinguished from it by its chromosome number and pollen morphology.
Grenier (1865) thought that plants from the Jura differed from those from the
western Alps but he was unable to describe any precise difference. The pollen
morphology is more reliable than any other morphological character for dis-
tinguishing subsp. montanum from subsp. alpinum.

It should be noted that the name L. perenne subsp. montanum is being used
here in a somewhat different sense from that in which the name L. montanum
has been used in the past. The name L. montanum has been applied to plants
from the Jura, the whole of the western Alps and the Pyrenees, while L. perenne
subsp. montanum, as used here, applies only to the tetraploids in this area, the
diploids being retained in subsp. alpinum. Neither the original descriptions of
L. montanum (Candolle 1824) nor the type specimens in the Geneva Herbarium
give any clear indication of the particular specimens to which the name should
be applied, and as the description fits the tetraploids it was decided to call them
L. perenne subsp. montanum.

It is a little surprising that there are no substantiated records of subsp. mon-
tanum between the single topodeme in Bavaria and the topodemes in the Jura,
some 500 km away. The L. perenne group as a whole is largely absent from this
area, apart from a record of L. perenne from Liechtenstein (Handel-Mazzetti
1911).

I have recently collected this plant from near Triesen in Liechtenstein and
it has proved to be diploid rather than tetraploid. Hence it represents an interest-
ing outlying locality for L. perenne subsp. alpinum and not another record for
subsp. montanum.

le. L. perenne subsp. extraaxillare. Scattered in the Carpathians and the moun-
tains of Jugoslavia. L. extraaxillare was originally described from the Tatra
and distinguished from L. alpinum by its broader leaves and longer stems. In
Romania and Jugoslavia both broad- and narrow-leaved morphodemes occur,
with the former predominating. Some of the Romanian plants have much
shorter stems (10-20 cm) than the Czech plants (25-50 cm). Although leaf width
separates subsp. alpinum and extraaxillare, it is not always reliable. The existence
of broad-leaved forms of subsp. alpinum in the Pyrenees and narrow-leaved
forms of subsp. extraaxillare in Romania could be used as an argument for
treating these two subspecies as one. However, there are many subspecies which
are incompletely separated by their morphology, and one can make a clear
geographical separation between subsp. a/pinum in the Alps and Pyrenees and
subsp. extraaxillare in the Carpathians and mountains of Jugoslavia.

2a. L. austriacum subsp. austriacum. The commonest subspecies in central
and eastern Europe. It has been questioned (Hegi 1924) whether it is native in its
westernmost localities, which are in Germany and Switzerland. Known introduc-
tions are recorded for some localities, e.g. Friesener Warte, near Bamberg,
Bavaria (Gauckler in Jit.) and Mon, Denmark (Bécher 1946), but there are
several other localities for which no such records exist. Topodemes such as F18
at Eichstatt, Bavaria are certainly fully naturalised if not actually native.
Capsule size, which is used to distinguish the two subspecies of L. austriacum,
is not totally satisfactory, because some topodemes of subsp. collinum produce
capsules as small as those of subsp. austriacum. Rather than applying a single
taxonomic character rigidly, which would give an arbitrary separation between
the two subspecies, a more satisfactory distinction can be made on the basis of

LINUM PERENNE GROUP IN EUROPE IANA)

their geographical distributions. Subsp. co//inum occurs in southern Europe south
of the Alps while subsp. austriacum occurs in central and eastern Europe, north
and east of the Alps. Nowhere do the two subspecies meet.

Subsp. austriacum in central Europe is much less variable than subsp. co/linum,
but in eastern Europe variants occur which have paler flowers, narrower leaves
and more erect pedicels than is the case in central Europe. A broad-leaved vari-
ant of subsp. austriacum from Romania was described as L. trinervium by Freyn
(1876).

2b. L. austriacum subsp. collinum. A morphologically heterogeneous and
highly disjunct subspecies extending from northern Spain through southern
France to Italy, Sicily, Jugoslavia and Greece. Most of the topodemes of subsp.
collinum are very limited in extent (Table 1) and can be defined more easily by
their distribution than by their morphology. The characters that vary most in
subsp. collinum are stem length and capsule size (Table 1), but there is also varia-
tion in sepal shape, as shown by the topodeme from Trieste, which has long nar-
row sepals. The pattern and extent of variation in L. austriacum subsp. collinum is
similar to that in L. perenne subsp. alpinum but the greater rarity of subsp.
collinum has made the morphological differences between the topodemes more
conspicuous.

The name L. austriacum subsp. collinum Nyman calls for some comment.
L. collinum has been applied to plants in southern Europe which are similar to,
but distinguishable from, L. austriacum subsp. austriacum. It has not been applied
exclusively to a limited area, as have the binomials listed in Table 1, and hence
is a suitable name for the subspecies as a whole. Nyman (1878) applied the
name to plants from Spain, Sicily and Greece. Plants from France were treated
by him as L. provinciale Jordan and are here included within subsp. collinum.

L. collinum Guss. is only a provisional name because Gussone (1845) was
uncertain of the status of the Sicilian plants for which he suggested the name.
Nevertheless, when Nyman (1878) published the combination L. austriacum
subsp. collinum with precise reference to Gussone, he was both definitely accept-
ing the taxon and validating the name by reference to Gussone’s definition.
Hence the name L. austriacum subsp. collinum Nyman is validly published.

Some mystery surrounds the plant named L. ruscinonense from Font-de-
Comps, Pyrénées Orientales by Timbal-Lagrave (1860). This plant has never

TABLE 1. TOPODEMES OF L. AUSTRIACUM SUBSP. COLLINUM

Extant Stem Capsule
Topodeme local Binomial length size
populations cm mm
North Spain 30-50 5:5-7:0
Cerdagne, Pyrénées 4 15-40 5-5-7:0
Prades, Pyrénées 1 L. ruscinonense 25-40 7:0-8-0
Puy-de-Déme 3 L. limanense 15-40 4-5-6:0
Hautes Alpes 1 L. saxicola 30-40 6:5-8:0
Vauclause 0 L. provinciale 40-60 6:5-7:0
Capo di Noli, Liguria 1 40-50 5:0-6:0
Trieste 4 L. tommasinii 10-25 4-5-5°5

North Greece 20-30 5:0-5:5

218 D. J. OCKENDON

been reported with certainty from any other site and I do not know precisely
where Font-de-Comps is, although I do know that it is in the vicinity of Prades.
L. ruscinonense, as described by Timbal-Lagrave, has short decumbent stems 5-8
cm long, bearing 1-4 flowers each. I visited Prades in 1966 and found only one
population (F35) of the L. perenne group in the area. This was at about 1000 m.
just above Villefranche-de-Confluent, at a place which could have been Font-de-
Comps. However, F35 has ascending stems 25-40 cm long, with 10-20 flowers
per stem, and is treated here as subsp. collinum. This probably is the population
known by Timbal-Lagrave, but if so, why were the plants so much smaller in
1860 than 1966? Stem length in the L. perenne group is phenotypically highly
plastic, and grazing may have been an important factor contributing to the very
short stems produced in 1860. The habitat of topodeme F35 is very similar to that
of other topodemes of subsp. collinum in the Pyrenees and quite different from
that of L. perenne subsp. alpinum. Hence it appears that L. ruscinonense is simply a
dwarf form of L. austriacum subsp. collinum and not a form of L. perenne subsp.
alpinum as Timbal-Lagrave supposed.

Another dwarf member of the L. perenne group which is referable to L.
austriacum subsp. collinum occurs at 600 m on Monte Calvo, Gargano, south
Italy. It has stems 10-15 cm long with 2-4 flowers per stem. Herbarium speci-
mens resemble the type specimens of L. ruscinonense. It was treated by Nyman
(1878) as L. punctatum, and has some similarities to L. tommasinii from Trieste.
It is geographically very isolated and needs further study.

The concept of L. austriacum subsp. collinum used here is an inclusive one. The
topodemes which have been tested, though morphologically diverse, show
fairly high interfertilities (Ockendon 1968a). Some of the morphodemes have
been adequately characterised, but others are very poorly known and, until they
are better known, any subdivision of subsp. collinum is liable to be arbitrary.

3. L. punctatum. A geographically isolated member of the L. perenne group
confined to a few mountains in Greece, Sicily and Turkey. Formerly, the Greek
and Turkish plants have been distinguished as L. pycnophyllum Boiss. et Heldr.,
but they are morphologically very similar to the Sicilian plants. Davis (1967) has
treated the Turkish material as L. pycnophyllum subsp. kurdicum.

4. L. leonii. Endemic to western Europe, chiefly in east and central France and
south and central Germany. Extending west to Charente, south to Aveyron, east
to Thiringen and north to Niedersachsen (Fig. 5). At one time L. leonii was
widespread in parts of eastern France, but as a result of extinction in many places
it is now rather rare. It was almost unknown in Germany until recently but in
the past ten years it has been reported from Bavaria (Gauckler 1964), Hessen
(Nieschalk & Nieschalk 1963), Siidniedersachsen (Lewejohann 1967) and
Thiiringen (Rauschert 1967). Rauschert’s paper gives a full discussion of all the
recent German records.

Past treatments of L. leonii have been very confused (Beyer 1898). It was
treated as conspecific with L. anglicum by Ascherson & Graebner (1914) and
Hegi (1924) and as a variety of L. alpinum by several French botanists. One
reason for this confusion was the failure to recognise that L. /eonii is homostylous
whereas all the other European members of the L. perenne group are hetero-
stylous. Small plants of L. /eonii are highly characteristic and unlikely to be con-
fused with anything except dwarf forms of L. austriacum subsp. collinum. They
commonly have a single stem 5—10 cm long with one or two flowers and the cotyl-

LINUM PERENNE GROUP IN EUROPE 219

Ficur_E 5. Distribution of L. Jeonii. B Records confirmed since 1955.
L) Old records not recently confirmed.

edons still present at the time of flowering. Large plants of L. leonii with stems
20-40 cm are sometimes found in populations consisting mainly of small plants.
It was probably a robust plant of this sort which misled Beyer (1898) into describ-
ing L. petryi as a new species. L. Jeonii is a short-lived perennial which usually
flowers in the first year, but the other European members of the L. perenne
group are longer-lived and usually do not flower until the second or third year.

L. leonii is much less variable than L. austriacum subsp. collinum but does
show variation in stem length. The topodemes of L. leonii from Céte d’Or and
Charente consist mostly of robust plants with erect stems 20-50 cm long. This
morphodeme corresponds to L. loreyi Jordan. As stem length is very variable
within single populations of L. leonii, the robust morphodeme does not warrant
treatment as a separate taxon. The two morphodemes of L. leonii are completely
interfertile (Ockendon 1968a).

DISCUSSION OF THE TAXONOMIC TREATMENT

The taxonomic treatment given is an attempt to summarise the major features
of the variation in the L. perenne group, but it is not intended as a detailed
account of the morphological variation, this being given separately for each sub-
species. The more detailed a taxonomic treatment of the group is, the more
arbitrary is it likely to be. For this reason it has not been found desirable to use
any taxonomic category below that of subspecies.

220 D. J. OCKENDON

Some justification may be given for the decision to retain L. perenne and L.
austriacum as species while treating L. alpinum as a subspecies of L. perenne.
Although the morphological separation between L. perenne and L. austriacum
is not completely clear in eastern Europe, there is no problem in central Europe.
Furthermore, artificial hybrids between these two species are produced only
with some difficulty and have reduced fertilities (Ockendon 1968a). The morpho-
logical separation between L. alpinum and L. perenne is incomplete because of
intermediates such as L. perenne 351/65 from Bavaria and L. perenne subsp.
extraaxillare from the Tatra. The cross L. perenne x alpinum is highly fertile
(Ockendon 1968a). L. perenne and L. alpinum may be thought of as ecological
races and are conveniently treated as subspecies. The subspecific category has
also been employed in the L. perenne group for widespread geographical races
(L. austriacum subspp. austriacum and collinum) and for widespread cytological
races (L. perenne subspp. anglicum and montanum) but has not been used for
morphodemes of restricted distribution, which are not given formal taxonomic
recognition.

MORPHOLOGICAL VARIATION

Some of the morphological variation to be found within the different subspecies
has already been described, but for a general consideration of the total variation
of the L. perenne group it is useful to consider the variation character by charac-
ter rather than subspecies by subspecies. For taxonomic purposes it is important
to distinguish between those characters which can be modified by the environ-
ment to a large extent and those which are only slightly susceptible to environ-
mental modification. An estimate of the phenotypic plasticity of the characters
studied has been made by comparing the phenotypes produced in the field with
those obtained by cultivation under various conditions. Phenotypically plastic
characters are liable to be untrustworthy taxonomically, but may have to be
used if more stable characters are lacking.

The morphological variation has been studied in terms of population samples
rather than single herbarium specimens, because of the importance of assessing
the amount of variation within populations and comparing this with the variation
between populations. The local population is a particularly important unit in
the L. perenne group because it is easy to define and the total number of local
populations in western Europe is sufficiently small for a considerable proportion
of them to be studied in detail. The sources of the material used here are given in
Appendix 1.

HABIT AND STEM LENGTH

Both habit and stem length vary strikingly and have been used as taxonomic
characters. Habit is difficult to describe accurately because of its complexity and
because the different types of habit grade imperceptibly into one another. The
descriptive terms used here are illustrated in Fig. 6. A further difficulty is that
the habit of a plant may appear to change as it becomes older. Thus, early in the
season the stems of L. perenne subsp. anglicum growing in open beds spread
radially from the caudex and are procumbent or decumbent in habit. Further
growth of these stems tends to be in a vertical rather than in a horizontal direc-
tion so that by the end of the season the habit is ascending or erect from an

LINUM PERENNE GROUP IN EUROPE oh

Erect Erect from an Inclined Ascending Decumbent Procumbent Reclinate
ascending base

FiGureE 6. Habit of the L. perenne group.

ascending base. This leads to a partial correlation between habit and stem length
and hence it is convenient to consider these two characters together.

From Table 2 it can be seen that L. perenne subsp. anglicum growing in tall
grass has long stems which are ascending or erect from an ascending base. Plants
from short grass or open habitats have short procumbent, decumbent or ascend-
ing stems. In cultivation in open beds with a metre between each plant, the
differences between these two forms largely disappear because the short-
stemmed plants produce considerably longer stems in cultivation (Table 2).
Small samples of L. perenne subsp. anglicum from different parts of the country
do not show any marked differences in habit when cultivated. However, cultiva-
tion of a large sample from topodeme GB2 has shown marked variation in

TABLE 2. VARIATION IN HABIT AND STEM LENGTH OF
L. PERENNE SUBSP. ANGLICUM

IN THE WILD IN CULTIVATION
Popu- Stem Stem
lation Habitat length Habit length Habit
GBI Tall grass 30-70 cm Erect from an 45-65 cm Ascending or
ascending base or erect
inclined
GBI Mown grass 20-40 Ascending or
decumbent
GB2 Tall grass 30-70 (-80) Erect froman 50-75 Ascending or
ascending base or erect
ascending
GB3 Tall grass 30-80 (-100) Erect froman
ascending base or
ascending
GBS5 Short grass 15-25 Decumbent or 60 Ascending
procumbent
GB6 _ _—s Tall grass 30-70(-90) Erect from an 55-70 Ascending or
ascending base erect
GB7 Short grass 15-30 Decumbent or
procumbent
GBl1 Short grass 10-20 Ascending or 30-50 Decumbent or
decumbent ascending
GB12 Mediumgrass 20-60 Erect from an 45-60 Decumbent or
ascending base or ascending

decumbent

222 D. J. OCKENDON

habit, indicating that there is also a genetic component to the observed differences
in habit. 89 plants had decumbent or ascending stems, while 15 had stems which
were erect from an ascending base or completely erect. The habit of individual
plants may vary from season to season. Five plants which were procumbent or
decumbent in habit in 1966 were ascending in 1967.

In the field L. Jeonii is usually strikingly smaller than L. perenne subsp. angli-
cum but in cultivation this difference in stem length largely disappears (Table 3).
There appears to be a correlation between stem length and soil depth in L.
leonii. The smallest plants are invariably found in very shallow soils and large
plants generally occur in much deeper soils. Large and small plants can occur
in mixed populations as at Bourges (topodeme F9), where there are numerous
small plants with stems 5-20 cm growing in soil 4~7 cm deep and a few larger
plants with stems 25-35 cm growing in soil 15-20 cm deep. The correlation be-
tween stem length and soil depth has been confirmed by cultivation. Plants of
L. leonii F4 grown in 10 cm pots produce stems up to 20 cm but when planted
out in beds the stem length increases to 60 cm.

In L. leonii, both in the wild and in cultivation, short stems are usually erect
but longer stems are usually ascending or reclinate. An exception is L. leonii F2,
in which the stems are always procumbent. The unusual habit of this topodeme
is presumably genetically determined.

Topodemes F17 and F25 of L. perenne subsp. perenne behave similarly in
cultivation and in the field, having erect stems 40-80 cm long. Topodeme 351/
65 from Bavaria is intermediate in habit and stem length between L. perenne
subsp. perenne F17 and subsp. alpinum 80A/63. The stems of 351/65 are 30-45 cm
long, thinner, more numerous and more flexuous than those of subsp. perenne
F17, and in all these respects tend towards subsp. alpinum.

In cultivation the stem length of L. perenne subsp. alpinum 80A/63 from Haute
Savoie reaches 45 cm compared with a maximum of about 25 cm in the wild.
The long stems produced in cultivation differ from those of L. perenne subsp.
perenne in being much branched below and in having long axillary shoots which
overtop the inflorescences. Because of the short growing season in its natural
habitats, a great development of axillary shoots does not normally occur in L.

TABLE 3. STEM LENGTH OF L. LEONII

Cultivated in beds

Population Wild (maximum values)
Fl 4-15 (-20) cm
F2 7-30 50 cm
F3 30-35 60
F4 3-10 60
F8 5-20 (-25)
F9 5-20 (-35) 45
F21 4-15 (-25) 55)
F27 20-40 60
F28 25-55
F43 30-50 90
F44 7-18

329C/65 60

LINUM PERENNE GROUP IN EUROPE 223

perenne subsp. alpinum in the wild. In cultivation, plants from the western Alps
have a low compact habit with decumbent spreading, flexuous stems. Plants
from the Pyrenees have more erect stems but retain a compact bushy habit. The
bushy habit of L. perenne subsp. alpinum in cultivation is not normally seen in
the wild. L. perenne subspp. extraaxillare and montanum share the typical bushy
habit of subsp. alpinum when cultivated, but tend to have somewhat longer stems
and inflorescence branches. .

STEM LEAFINESS

In cultivation the alpine taxa L. perenne subspp. alpinum, extraaxillare and
montanum commonly produce characteristic densely leafy sterile shoots with
squarrose or deflexed leaves. Such shoots are not normally produced by the
lowland subspecies in cultivation, but can occasionally occur when plants are
growing vigorously under moist conditions. Furthermore, the densely leafy stems
of the alpine taxa are often absent in the wild, especially when the plants are
growing amongst tall vegetation. Although stem leafiness is a conspicuous charac-
ter, which is sometimes useful for the purpose of recognition, it is generally too
phenotypically plastic to be used diagnostically.

LEAF WIDTH

Leaf width is the main character used to distinguish L. perenne subsp. extra-
axillare from subsp. alpinum, and broad-leaved Romanian plants (L. trinervium)

£2) SP

: 4 L. leonii

$29 eSATA ERNEST ene eee

F 10 REESE APE

F15 : L. austriacum subsp. co/linum

ij L. austriacum subsp. austriacum

3| L. perenne subsp. perenne

| L. perenne subsp. anglicum

F12 ee EET TE — L. perenne subsp. montanum

L. perenne
F 36 SS a :
subsp. a/pinum
F 4] RANE SEPP ree
F 42 SERS TS SO
(5) — a Le eee
subsp. extraaxillare

Li ee ere ee Ss ee ee ee

2 3 4 5mm

_

Ficure 7. Leaf width of the Linum perenne group showing mean, range (thin line) and stand-
ard deviation (thick line) for population samples. For sources of material see Appendix.

224 D. J. OCKENDON

of L. austriacum subsp. austriacum from the rest. Leaves on the branched
part of the stem are commonly wider than those on the unbranched part
and therefore a leaf from the middle of the stem was selected for measure-
ment. The results in Fig. 7 are for samples of 20 stems collected in the wild.
The amount of variation in leaf width in a single population ranges from very
little in L. leonii F21 to a large amount in L. perenne subsp. alpinum F42. The
leaves of L. austriacum and L. leonii tend to be a little narrower than those of
L. perenne but there is a large amount of overlap. The sample with the narrowest
leaves (L. leonii F21) also has the shortest stems, and it is not uncommon to
find that large plants have wider leaves than small plants from the same popula-
tion.

The greatest variation in leaf width occurs in L. perenne subsp. alpinum,
especially in certain Pyrenean topodemes such as F42, which show almost as
much variation in this character as the L. perenne group as a whole. Topodemes
of this subspecies from the Alps have moderately narrow leaves similar to those
of L. perenne subspp. montanum and perenne. L. perenne subsp. extraaxillare
seldom has leaves less than 2 mm wide but does not produce leaves as wide as
the widest produced by some Pyrenean topodemes of subsp. alpinum.

Leaf width shows some phenotypic plasticity. A few plants of topodeme F36
of L. perenne subsp. alpinum growing in partial shade had leaf widths of 2:5—
4-5 mm compared with a range of 1-5-4-0 mm for the rest of the population.
In cultivation several subspecies showed greater leaf widths than those found in
wild material. This is partly a question of the greater vigour of plants in cul-
tivation.

SEPAL SIZE AND SHAPE

The sepal dimorphism of the L. perenne group is highly characteristic. The two
inner sepals have a much wider colourless scarious margin than the two outer
sepals (Fig. 8). The fifth sepal is asymmetrical, with a wide margin on one side
and a narrow margin on the outer. The green parts of all five sepals have much
the same shape, but because of the differences in the widths of the margins, the
outer sepals are lanceolate or ovate-lanceolate, while the inner are ovate-
lanceolate, ovate or obovate.

The variation in sepal size and shape is not easy to analyse because of the
complex of characters involved. It is imperative that outer sepals be compared
with outer and inner with inner. The only sepal character which can be recorded
with precision directly from herbarium material is sepal length. For more

1mm

scarious margin

Outer Middle Inner

FiGure 8. Sepal dimorphism of the Linum perenne group.

LINUM PERENNE GROUP IN EUROPE 225

er en L. perenne subsp. perenne
SS L. austriacum subsp. austriacum
rc L. austriacum subsp. collinum
ie ee L. perenne subsp. anglicum
rr L. leonii
ee ene pr L. perenne subsp. montanum
$$ rr rc L. perenne subsp. a/pinum

ee L. perenne subsp. extraaxillare

PE ES A [ne SA a ES A OO CO RETO
4 5 6mm

FiGurE 9. Length of inner sepals of the Linum perenne group showing mean, range (thin line)
and standard deviation (thick line) for each taxon. For sources of material see Appendix.

detailed studies, sepals were dissected from the flower, mounted and examined
under a binocular microscope.

The montane subspecies L. perenne subspp. alpinum and extraaxillare tend
to have longer sepals than the lowland taxa, especially L. perenne subsp. perenne
and L. austriacum subsp. austriacum (Fig. 9), and this has been used as a diag-
nostic character. However, the ranges for the different taxa overlap so much
that this character is not very reliable. There is considerable variation in sepal
length within populations. The range found for 10 plants of topodeme GB1 of
L. perenne subsp. anglicum (4-1-5-8 mm) almost equals the total range for the
subspecies.

The difference in length between the outer and inner sepals can be used to
separate subspp. anglicum and perenne from the other subspecies of L. perenne
(Fig. 10). In L. perenne subspp. alpinum, extraaxillare and montanum the outer
and inner sepals are almost equal in length, never differing by more than 0-4 mm.

qa L. perenne subsp. a/pinum
SLD L. perenne subsp. extraaxillare
CED L. perenne subsp. montanum
Se L. leenti

TED L. austriacum subsp. austriacum
ETE L. austriacum subsp. col/linum

Per ese neste) L. perenne subsp. perenne

qs | /perenne subsp. anglicum

0.4 0.8 1.2mm

oO

FiGcure 10. Difference in length between the outer and inner sepals, showing range for each
taxon. For sources of material see Appendix.

226 D. J. OCKENDON

In L. perenne subspp. anglicum and perenne the inner sepals exceed the outer by
0:5-1-4 mm. The difference in length between the sepals can usually be seen in
herbarium material. Sepal shape can be expressed crudely in terms of length/
breadth ratios but these do not give a much better separation between the sub-
species than sepal length alone. The potential taxonomic value of sepal shape and
size in the L. perenne group has not been realised despite detailed examination
of these characters.

PETAL SIZE
The petals of the L. perenne groups are broadly obovate or obovate-lanceolate
with a very obtuse or slightly emarginate apex. They taper gradually to a short
yellow, pubescent claw. They are flimsy and fugacious. In hot weather few petais
remain on the plant after midday. A new set of flowers opens each morning.
Petals were collected in the morning and spread out carefully before pressing.
Petal length shows rather less intrapopulation variation than petal width and
therefore was chosen as a possible taxonomic character. Fig. 11 shows that there
is continuous variation in the petal length of the L. perenne group. L. leonii has
the shortest petals and could be distinguished from L. perenne subspp. anglicum
and alpinum by its petal length, but otherwise this character has little taxo-

FA3 Soe L. leonii

L. austriacum subsp. austriacum

| L. perenne subsp. perenne

F 10 S$ — L. austriacum subsp. collinum

GB I ee re L. perenne subsp. anglicum

F 36 es L. perenne subsp. a/pinum

8 12 16 20 24mm

Ficure 11. Petal length in material from the wild, showing mean, range and standard devia-
tion. For sources of material see Appendix.

LINUM PERENNE GROUP IN EUROPE Qa.

F2 a
F9 ~_ L. leonii
DD SS -
FO — _— L. austriacum subsp. collinum
125B/65 enc ———- — L. austriacum subsp. austriacum
351/65 asa pe —_ L. perenne subsp. perenne
75/65 —a— = L. perenne subsp. extraaxillare
F25 SS
F25 ee EE SSS L. perenne subsp. perenne

1255/65 a
301B/65 eo L. austriacum subsp. co/linum
244A/65 SS L. perenne subsp. montanum
GB2 oS

B82 SS —_—

a L. perenne subsp. anglicum

10 14 18 22 mm

FiGurE 12. Petal length of cultivated material (solid figures) compared with that of wild
material (open figures). For sources of material see Appendix.

nomic value. It is noteworthy that L. Jeonii, the only European member of the
group which is not absolutely dependent on cross-pollination by insects, has the
smallest petals.

The pattern of variation in petal length shown by the cultivated material is
similar to that of the wild material, but if anything the differences between the
taxa are less clear in cultivation (Fig. 12). In cultivation, the petal length of
L. leonii remains shorter than that of L. perenne subsp. anglicum, indicating that
there is a genetic basis to this difference. However, in the three cases in which
direct comparisons can be made, the petal length of cultivated material is
appreciably greater than that of wild material from the same source. This
suggests that petal length is phenotypically plastic and is partially correlated
with plant size.

PETAL COLOUR

Petal colour varies from pale blue to deep bright blue and has been used as a
diagnostic character to distinguish L. perenne from L. austriacum, and L. alpinum
from L. julicum (Hayek 1909). Occasional white-flowered variants occur in the
wild and a single plant of L. perenne subsp. perenne 351/65 from Bavaria had
deep purple flowers.

Petal colour is not a particularly good taxonomic character in the L. perenne
group because it is difficult to measure accurately and because the colour fades

Cc

228 D. J. OCKENDON

L. perenne
subsp.
alpinum

L. perenne | L. perenne
subsp. subsp.
extraaxillare| perenne

131F/63
75/65
32 O

Methyl violet
Hyacinth blue
Lobelia blue
Gentian blue

FiGureE 13. Petal colour as measured with the Horticultural Colour Chart. Each block diagram
represents a single population. Four hues are recognised and each of these can exist at a range
of intensities from 0 (most intense) to 3 (least intense). Squares in the bottom right of each
diagram represent dark blue petals, while squares in the top left represent pale mauvish petals.
For sources of material see Appendix.

L. perenne | L. perenne
subsp. subsp.
anglicum | montanum

L. leonii

subsp
austriacum

subsp.
collinum

—

190E/65

Intensity

on exposure to bright light. The colour was measured in terms of hue and
intensity by means of the Horticultural Colour Chart and the results are sum-
marised in Fig. 13. This shows that L. leonii and L. austriacum tend to have
darker, bluer petals than L. perenne. Petal colour in L. Jeonii is fairly constant
but is much more variable in L. perenne subsp. alpinum, ranging from deep blue
in 80A/63 to pale mauvish blue in F41. The tetraploids L. perenne subspp.
anglicum and montanum resemble subsp. perenne in having paler, mauvish petals.
Petal colour is not sufficiently reliable to be a diagnostic character in the L.
perenne group, but taxa which have been recognised on other grounds may show
some differences in petal colour.

LINUM PERENNE GROUP IN EUROPE 229

CAPSULE AND SEED SIZE

Each segment of the capsule is filled by a single seed. As seed and capsule length
are closely correlated there is little to be gained by measuring both characters.
Capsule length has the advantage that it can be measured directly on herbarium
sheets, but the disadvantage that it is less easy to measure accurately than seed
length. These characters should be measured on mature seeds or capsules. Popu-
lation samples tend to show more variation in capsule length than in seed length:

subsp. anglicum GB1 Mean Range Standard deviation
100 capsules 6:16 mm 5-0-7:5 0-47
100 seeds 4-71 mm 4-0-5°3 0-23

For these reasons seed length was chosen for detailed study rather than capsule
length.

3298465 ———
331065
190E/65 ia
1584/65 eA
35165 Es

L. austriacum subsp. austriacum

| L. perenne subsp. perenne

33 ea ) |
L. austriacum subsp. co/linum

3018/65 Ts
(35 ——

FA7 Ene L. perenne subsp. a/pinum

F51 — << L. perenne subsp. extraaxi/lare
2 =
329C/65 =e
£43 = L. leonii

CE)2 JERS RTE acc
GB] ees EE Rca
104D46 Sone NES
y L. perenne subsp. montanum
2444/65 a

2.6 3.4 4.2 5.0 5.8mm

FicureE 14. Seed length, showing mean, standard deviation and range for population samples
of 50 seeds. For sources of material see Appendix.

| L. perenne subsp. anglicum

230 D. J. OCKENDON

Variation in seed length in the L. perenne group is virtually continuous, but
the amount of variation within populations is relatively small and does not
obscure the differences between populations (Fig. 14). Seed length is fairly con-
stant in L. austriacum subsp. austriacum and L. leonii but varies considerably in
L. perenne subsp. alpinum, and to an even greater extent in L. austriacum subsp.
collinum. This parallels the marked variation between populations of L. austria-
cum subsp. collinum in other characters such as stem length and sepal shape. Seed
length is less phenotypically plastic than most of the other morphological
characters and is not correlated with overall plant size. Thus L. leonii, which
has the shortest stem length, has relatively large seeds, while L. perenne subsp.
perenne has long stems and small seeds.

Seed length can be used to distinguish L. austriacum subsp. austriacum from
L. perenne subsp. anglicum, but other characters are easier to use for this. Cap-
sule length is used in the key to separate the two species of L. austriacum, but
as can be seen from Fig. 14, L. austriacum subsp. collinum F15 is intermediate
in seed length between subsp. austriacum and the rest of subsp. collinum. L. austria-
cum subsp. collinum is a heterogeneous subspecies and is difficult to separate
from subsp. austriacum in any simple way.

POSITION OF MATURE FRUIT PEDICEL

The difference between erect and patent or deflexed mature fruit pedicels (here
referred to simply as ‘pedicels’) has long been the chief character for distinguish-
ing L. perenne from L. austriacum. There is a complete series of pedicel positions
in the L. perenne group but it is only feasible to distinguish a few main types
(Fig. 15).

The position of the pedicel may alter considerably as the fruit matures and it
is essential to compare the pedicels of mature fruits. A striking example of this
is a specimen of L. perenne subsp. anglicum with immature fruit collected near
Stamford on 4 July 1957, which has small capsules on patent pedicels looking
just like those of L. austriacum subsp. austriacum. Specimens with mature fruit
collected from the same site on 3 August 1965 have large capsules with erect
pedicels.

Pedicel position is not noticeably altered by cultivation and there is no
evidence of appreciable phenotypic plasticity. The species which is most difficult
to recognise by its pedicels is L. /eonii, which has erecto-patent, flexuous or
patent pedicels and is intermediate between L. perenne and L. austriacum in
this respect. Normally pedicel position is constant in a single plant but there are
some exceptions. A single plant of L. perenne subsp. alpinum F41 has erecto-
patent, flexuous and patent pedicels, this variation being almost as great as that

Erect Erecto-patent Flexuous Patent Deflexed

Ficure 15. Fruiting pedicel position.

LINUM PERENNE GROUP IN EUROPE 231

of the population from which the specimen came. Pedicel position does not
correlate with any other character and is the only means of distinguishing all the
subspecies of L. perenne from those of L. austriacum.

HETEROSTYLY

Although heterostyly is a morphological character, it is in some ways better
considered as a morphological manifestation of the breeding system. The varia-
tion pattern shown by this character is totally different from that of the quanti-
tative morphological characters previously considered. In the case of heterostyly
there are two sharply contrasting character states, namely heterostyly, in which
long- and short-styled plants occur in approximately equal numbers in a popula-
tion, and homostyly, in which all plants have stigmas and stamens at about the
same height. No population of Linum containing both heterostyled and homo-
styled plants has yet been reported in Europe and the character is not phenotypic-
ally plastic. It may be difficult to classify immature or malformed flowers as long-
styled, short-styled or homostylous, but in the field there is no difficulty in
determining whether a population is heterostylous or homostylous. As the charac-
ter can be seen more easily in the field than in herbarium material, it is highly
desirable that it be recorded in the the field whenever a species of Linum is
collected. Homostyly is by far the most reliable character for distinguishing
L. leonii from L. austriacum and L. perenne. Heterostyly is an especially valuable
taxonomic character in Linum and might well be used more in the other genera
in which it occurs.

DISCUSSION

The evidence presented clearly shows that the majority of the morphological
characters vary almost continuously. The minor discontinuities which are
found tend to reflect discontinuities in the distribution or lack of adequate
samples. Almost all the characters show some phenotypic plasticity, but this is
more marked in the stem, leaf and flower characters than in seed size and pedicel
position. In no case could the variation in any character be attributed to pheno-
typic plasticity alone, but the genetically controlled differences which do occur
are often obscured by phenotypic plasticity. There is a partial correlation be-
tween the variation of some of the characters in so far that large plants tend to
have longer stems, sepals and petals and broader leaves than small plants. Seed
length is an exception to this in that it remains fairly constant despite wide varia-
tion in the size of the plant. The phenotypic plasticity of the L. perenne group
may have considerable ecological significance because the group can behave as
an opportunist coloniser of disturbed sites and often occupies unstable habitats
where conditions such as soil depth and degree of competition are liable to
change rapidly.

Differences in the amount of variation within single populations of the L.
perenne group may be associated with particular geographical areas or with the
type of breeding system. Geographically correlated differences of this type are
exemplified by some of the Pyrenean populations of L. perenne subsp. alpinum
which are exceptionally variable. This may have resulted from past hybridization
between L. perenne subsp. alpinum and L. austriacum subsp. collinum in the
Pyrenees. Artificial hybrids between these two subspecies can be made easily
(Ockendon 1968a), but there is at present nowhere where they occur sufficiently

232 D. J. OCKENDON

close together to produce natural hybrids. A correlation between the type of
breeding system and the amount of variation within a population is exemplified
by the inbreeding species L. Jeonii, which generally shows less variation within
populations than do the obligatory outbreeding species L. austriacum and
L. perenne.

A useful comparison can be made between the variation patterns of the L.
perenne group in Europe and of Hesperolinon in California. The species of
Hesperolinon are homostylous annuals and resemble the L. perenne group in
occurring in highly disjunct and very localised populations and in having a
restricted ecological tolerance (Sharsmith 1961). In contrast to the L. perenne
group, the species of Hesperolinon can be assessed with a reasonable degree of
assurance and, where two species occur sympatrically, hybridization is very
rare. In Hesperolinon most of the populations are morphologically uniform but
may be separable one from the next. The difference between the almost continu-
ous Variation in the L. perenne group and the clear discontinuities in Hesperolinon
are a reflection of the fact that the former is outbreeding (except for L. leonii)
while the latter is chiefly inbreeding.

The majority of characters which have been used in the L. perenne group are
taxonomically unreliable because of phenotypic plasticity and genetically based
variation within populations. Although statistically significant differences in
these characters could often be demonstrated between populations, these are
insufficient to assign a single plant to a particular population. Some characters
are reliable in limited areas but break down when applied to other areas. Thus
leaf width reliably separates L. perenne subsp. extraaxillare from the Tatra from
subsp. alpinum from the Alps, but breaks down when material from Romania
or the Pyrenees is considered. The morphological variation in the L. perenne
group has arisen largely by the gradual differentiation of geographically isolated
populations. This, together with widespread extinction, has given rise to the
familiar situation of a series of forms which replace one another geographically,
but in most cases the sharp geographical discontinuities are not marked by clear
morphological discontinuities. A satisfactory taxonomic treatment of the L.
perenne group has been achieved by rejecting many of the morphological charac-

TABLE 4. MAIN DIVISIONS OF THE L. PERENNE GROUP

Ploidy Pollen Pedicel
Taxa Breeding system level type position
L. leonii Homostylous 2x 3 Furrows’ Flexuous or
patent
L. austriacum Heterostylous 2x 3 Furrows Patent or
deflexed
L. perenne subspp.
alpinum, Heterostylous 2x. 3 Furrows_ Erect
extraaxillare and
perenne
L. perenne subspp.
anglicum and Heterostylous 4x 6 Pores Erect

montanum

LINUM PERENNE GROUP IN EUROPE 233

ters which were used in the past, by emphasising the sharpiy discontinuous
characters of the breeding system, chromosome number and pollen morphology,
and by ensuring that the taxa recognised are valid in terms of their distribution
and ecology. The main divisions within the group are summarised in Table 4.

ACKNOWLEDGMENTS

Financial help from the Science Research Council, Clare College, Cambridge
and the Brooks Fund is gratefully acknowledged. Thanks are due to the keepers
of herbaria who loaned the material examined in this work, to Mr J. E. Dandy
for advice on nomenclature and to Dr W. T. Stearn for help with typification.
I should like especially to thank Dr S. M. Walters who gave much help and
encouragement during the course of the work.

REFERENCES

ALEFELD, F. (1867). Uber Adenolinum Rchb. Bot. Ztg., 25: 249-255.

ASCHERSON, P. F. A. & GRAEBNER, K. O. P. P. (1914). Synopsis der mitteleuropdischen Flora,
vol. 7, pp. 194-211. Leipzig.

BEYER, R. (1898). Uber Linum leonii Schultz und einige andere Formen der Gattung Adeno-
linum. Verh. Bot. Ver. Prov. Brandenb., 40: 82-94.

BOcHER, T. W. (1946). Grassland communities of ‘Héje M@n’. Bot. Tidsskr., 48: 1-45.

CANDOLLE, A. P. DE (1824). Prodromus Systematis naturalis Regni vegetabilis, vol. 1, p. 427.
Paris.

Davis, P. H. (1967). Flora of Turkey, vol. 2, pp. 444-447. Edinburgh.

FREYN, J. F. (1876). Uber einige Pflanzen, insbesondere der Osterungar. Flora. Ost. bot. Z.,
26: 228.

GAUCKLER, K. (1964). Linum anglicum Miller —neu fiir Bayern. Ber. bayer. bot. Ges., 37:
103-104.

GRENIER, M. C. (1865). Flora de la chaine Jurassique. Paris.

GUSSONE, G. (1845). Florae siculae Synopsis, vol. 2, p. 808. Neapoli.

HANDEL-Mazzettl, H. F. (1911). Uber das Vorkommen von Linum perenne L. in Liechtenstein.
Ost. bot. Z., 61: 227.

HAYEK, A. (1907). Vorarbeiten zu einer Bane hues ceaplidehe Karte Osterreichs. IV Die
Sanntaler Alpen. Abh. Zool.-Bot. Ges. Wien, 4(2): 109.

Hayek, A. (1909). Flora von Steiermark, vol. 1, pp. 619-620. Berlin & Graz.

Hea, G. (1924). Ilustrierte Flora von Mitteleuropa, vol. 5, pp. 15—20. Miinchen.

JACQUIN, N. J. VON (1762). Enumeratio stirpium plerarumque quae sponte crescunt in agro
Vindobonensi, montibusque confinibus. Vienna.

JUEL, H. O. (1936). Joachim Burser’s Hortus siccus. Symbolae Bot. Upsal., 2: 1-187.

JUZEPCZUK, S. V. (1949). Linum, in Komarov, V. L., ed., Flora URSS, vol. 14, pp. 103-124.
Leningrad & Moscow.

LEWEJOHANN, K. (1967). Linum leonii Schultz und sein Vorkommen in Siidniedersachsen.
Flor. Rundbr. Sud-Niedersachs., 1967: 5-8.

NIESCHALK, A. & NIESCHALK, C. (1963). Linum leonii Schultz in Hessen. Hess. florist. Br.,
137: 29-32.

Nyman, C. F. (1878). Conspectus Florae Europaeae. Orebro.

OCKENDON, D. J. (1967a). Notes on the Linum perenne group. Notulae Systematicae ad
Floram Europaeam spectantes. Reprium nov. Spec. Regni. veg., 74: 20-22.

OCKENDON, D. J. (1967b). Experimental taxonomy of the Linum perenne group in Europe. Ph.D.
thesis. University of Cambridge.

OCKENDON, D. J. (1968a). Biosystematic studies in the Linum perenne group. New Phytol., 67:
787-813.

OCKENDON, D. J. (1968b). Linum perenne ssp. anglicum (Miller) Ockendon, in Biological Flora
of the British Isles. J. Ecol., 56: 871-882.

234 D. J. OCKENDON

OCKENDON, D. J. & WALTERS, S. M. (1968). Linum L., in TuTIn, T. G., et alia, ed. Flora
Europaea, vol. 2, pp. 206-211. Cambridge.

PLANCHON, J. E. (1848). Sur la famille des Linées. Hooker’s London J. Bot., 7: 173-178.

RAUSCHERT, S. (1967). Linum leonii F. W. Schultz in Thiiringen und in nérdlichen Harzvor-
land. Wiss. Z. Univ. Halle-Wittenburg, math.-nat. Kl., 16: 944-948.

REICHENBACH, H. G. L. (1837). Handbuch des natiirlichen Pflanzensystems. Dresden & Leipzig.

SHARSMITH, H. K. (1961). The genus Hesperolinon (Linaceae). Univ. Calif. Publs Bot., 32:
235-314.

STEARN, W. T. (1957). Introduction to LINNAEUS, C. Species Plantarum, a facsimile of the Ist
edition, vol. 1. London.

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Fr., 7: 509.

APPENDIX

L. austriacum subsp. austriacum

F14 Raron, Brig, Valais, Switzerland (150)*
F18 Eichstatt, Bavaria, Germany (400)
F20 Staffelberg, Staffelstein, Bavaria, Germany (200)
F24 Randersacker, Wiirzburg, Bavaria, Germany (600)

125B/65 Letea, Tulcea, Romania

158C/65 Budapest, Hungary

190E/65 Kyffhaiiser, Thiiringen, Germany
308A/65 Hungary

329B/65 Suad, Romania

331A/65 Hovblege, Mon, Denmark
331B/65 Illmitz, Burgenland, Austria

L. austriacum subsp. collinum

F10 Puy Crouel, Clermont-Ferrand, Puy-de-Dome, France (300)
Fil Cournon d’Auvergne, near Clermont-Ferrand, France (5)
F15 Mt. Spaccato, Trieste, Italy
F32 Gap, Hautes Alpes, France (20)
F35 Villefranche-de-Confluent, Pyrénées Orientales, France (400)
F38 Bellver de Cerdana, near Seo de Urgel, Spain (1000)
301B/65 Capo di Noli, Liguria, N. Italy

L. leonii
Fl Flohiment, Ardennes, France (50)
F2 Thierville, near Verdun, Meuse, France (30)
F3 Cote Quaraille, Noveant, near Metz, Moselle, France (20)
F4 Montenach, near Sierck, Moselle, France (100)
F5 St Mihiel, Meuse, France (50)
F6 Bovée sur Barboure, Meuse, France (3)
F8 Malesherbes, Loiret, France (100)
F9 Chapelle-St-Ursin, near Bourges, Cher, France (100)
F21 Bottingheim, Taubertal, Wiirttemburg, Germany
F26 Neuville-sur-Seine, near Troyes, Aube, France (2)
F28 Gamay, near Chagny, Cote d’Or, France (1000)
F29 Santannay, near Chagny, Cote d’Or, France (1000)
F43 Chaumes de Souberac, near Jarnac, Charente, France (60)

* Figures in brackets are approximate numbers of individuals at each locality.

F13
F30
F31
F33
F34
F36
F41
F42
F47
F49
F50
80/63

GBI
GB2
GB5
GB6
GB8
GB10
GB11
GB12
GB13

F351
131F/63
75/65

F12
79/65
244A/65
104D/66

Bi.
F25
351/65

LINUM PERENNE GROUP IN EUROPE

L. perenne subsp. alpinum

Plateau d’Assy, near St. Gervais, Haute Savoie, France. 900 m

Lac du Crozet, near Revel, Isére, France. 1950 m
Col de Glaize, Gap, Hautes Alpes, France. 1950 m
Mt Charance, Gap, Hautes Alpes, France. 1800 m

PB )8,

Pic de Costabonna, Prats de Mollo, Pyrénées Orientales, France. 2000 m

Val d’Eyne, Pyrénées Orientales, France. 2000 m
Valle de Aran, near Tredos, Lerida, Spain. 2000 m

Mt. Ceciré, near Luchon, Haute Garonne, France. 2000 m

Mt. Maggiorasca, near Genova, Italy. 1800 m
Bansky suchodol, Pirin Mountains, Bulgaria. 2600 m
Val Fredda, Gran Sasso, Abruzzi, Italy. 1550 m

Grenairon, near Martigny, Haute Savoie, France. 2100 m

L. perenne subsp. anglicum

Gog Magog Hills, Cambridge, England

Babraham Hall Estate, Cambridge, England
Lakenheath Warren, Suffolk, England

Great North Road, Stamford, Lincolnshire, England
Sherburn Wold, East Riding, Yorkshire, England
West Cornforth, Co. Durham, England

Tunstall Hill, Sunderland, Co. Durham

Brighouse Bay, Kirkcudbrightshire, Scotland
Crosby Ravensworth, Westmorland, England

L. perenne subsp. extraaxillare

Chalet Hvezdon, Tatra, Czechoslovakia. 1600 m
Orava, Czechoslovakia
Kosice, Czechoslovakia

L. perenne subsp. montanum

Le Rozet, Thoiry, Ain, France. 850 m
Crozet, Ain, France. 1300 m

Mt Dole, Vaud, Switzerland

Col de Crozet, Ain, France. 1500 m

L. perenne subsp. perenne

Garchinger Heide, Eching, Munich, Bavaria, Germany
Pfungstadter Diine, near Darmstadt, Hessen, Germany
Pfatter, near Regensburg, Bavaria, Germany

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The Ranunculus acris L. complex in Europe
S. M. COLES

Department of Botany, Birmingham University

ABSTRACT

The geographical variation of the Ranunculus acris L. complex within Europe has been exam-
ined. The following taxa are recognised in this present work:
Ranunculus granatensis Boiss.
R. strigulosus Schur
R. acris L. sensu stricto subsp. acris var. acris
var. villosus (Drabble) S. M. Coles, comb. et stat. nov.
var. pumilus Wahlenb.
subsp. borealis (Trautv.) Nyman
subsp. friesanus (Jord.) Rouy & Fouc. var. friesanus
var. pyrenaeus S. M. Coles, var. noy.

R. granatensis and R. strigulosus have a restricted distribution and are relatively uniform in
appearance. R. acris sensu stricto occurs over most of Europe and is a very variable plant. The
main clinal variation in R. acris is in an east-west direction. R. acris subsp. friesanus forms no
part of this variational pattern although its distribution is completely overlapped by that of
subsp. acris.

INTRODUCTION

The plants investigated in this study are those included in Ranunculus acris L. as
defined by Tutin (1964). R. acris L. sensu stricto in the present study excludes
R. acris subsp. granatensis (Boiss.) Nyman and subsp. strigulosus (Schur) Hyl.,
which are given specific status.

The R. acris complex belongs to the section Ranunculus of the subgenus
Ranunculus. It can be separated from most other European species of this
section by having a terete pedicel, patent sepals and a glabrous receptacle. The
diploid chromosome number of R. acris L. sensu stricto is 14; within the genus
a basic chromosome number of 8 is more common than one of 7. R. serbicus
Vis. (2n=28) is generally held to be a closely related species and probably merits
inclusion within the complex, but it has not been considered here.

R. acris sensu lato is common over the whole of central and northern Europe
including the British Isles, the Faeroes and Iceland. It also occurs in Greenland,
but there has been some dispute as to whether it is truly native. Herbarium
material examined from the south and east coasts of Greenland could not be
distinguished from plants which have been introduced into Spitsbergen and
N. America. The distribution of the complex is more limited in southern
Europe. It is absent from Portugal; in Spain it only occurs in the north, east and
south-east; it extends into Morocco. In Italy, material has been seen only as
far south as Lat. 42°N. In the Balkan Peninsula it extends as far south as
Albania and the Makedhonia region of Greece (Fig. 1). The distribution of the
complex extends eastwards through Siberia and a series of forms is to be found
in eastern Asia. It has been introduced into many of the temperate parts of the
world, such as New Zealand, S. Africa and N. America; in the last it has
spread in some regions with amazing rapidity (Benson 1948).

238 S. M. COLES

Details of the ecology of R. acris are given by Harper (1957). It is tolerant of
a wide range of habitats, and is widespread over most of its range. One of its
main limiting factors appears to be that of the water content of the soil. It is not
markedly calcicole or calcifuge, although it does appear to be limited by a very
low base-status.

TY

7

cA
DY
s

R. granatensis

; ZB R. acris subsp. acris
7 var. pumilus
NX R. acris subsp. acris

R. acris subsp. friesanus Tarniiiinsts

subsp. friesanus var. pyrenaeus

FicureE 1. Distribution in Europe of the taxa recognised; based only on the material examined
during this study.

Abnormalities of both the floral and vegetative parts of the plant are common.
Some abnormalities of the flower can transform characters usually held as safe
diagnostic characters of the complex. They may result, for example, in hairs
being found on the receptacle or on the achenes. Gynodimorphism commonly
occurs in the complex. All grades from functionally female to normal hermaphro-
dite flowers occur. Although there seems to be genetical control of these flower-
types, fluctuations of types within a single plant are common (Marsden-Jones &
Turrill 1929, 1935).

MATERIALS AND METHODS

This study was largely based on herbarium material from the following herbaria:
British Museum (BM), Cambridge (CGE), Copenhagen (C), Edinburgh (E),
Florence (FI), Kew (K), Leicester (LTR), Leningrad (LE), Liverpool (LIVU),
Paris (P), Stockholm (S), Vienna (W).

RANUNCULUS ACRIS L. IN EUROPE 239

The herbarium material was supplemented wherever possible by living material.
Unfortunately living plants of R. acris subsp. acris and subsp. friesanus only
could be obtained. The plants, from over fifty localities in the British Isles and
from sixty localities on the Continent, were grown for periods of at least one
year.

Characters were investigated from as many parts of the plant as possible.
The following were found to be of most value: type of rootstock, shape of leaves,
type of hair cover, and form of achenes. A detailed evaluation of characters
and of the variations found within them, is given after a summary of the recog-
nised taxa.

THE NOMENCLATURE AND SUMMARY OF THE CHARACTERISTICS
OF THE RECOGNISED TAXA

Previous names which have been used in connection with plants belonging to
this complex are very numerous (Index Kewensis lists over forty at specific level).
Many of the names appear to have been used only by their authors and their
descriptions give no clear idea of the taxon referred to, and where type material
was seen it usually showed that the names were merely referable to plants
which were parts of clines or which were showing seasonal variations. The
synonyms given here include only names which have been commonly used by
workers other than their authors. A fuller account of nomenclature has been
given by Coles (1968).

The species category has been used where there are distinct, ‘major’, morpho-
logical discontinuities. The category of subspecies has been used to cover groups
of plants from geographical regions which show ‘major’ morphological differ-
ences from other subspecies, but where there is also an array of intermediate
types present. The category of variety has been used where the taxa have a
geographical or ecological basis, but the morphological differences are not so
great.

R. GRANATENSIS Boiss., Diagn. Pl. Or. Nov., Ser. 2, vol. 3 (1): 8 (1853).
R. atlanticus Ball, J. Bot., Lond., 11: 296 (1873).
R. acris subsp. granatensis (Boiss.) Nyman, Consp., 12 (1878).
R. acris var. atlanticus (Ball) Maire, Flore de l’Afrique du Nord, vol. 11:
162 (1964).

R. STRIGULOSUS Schur, Enum. Pl. Transs., 17 (1866).
R. malocophyllus Schur, Enum. Pl. Transs., 16 (1866).
R. csatoi Schur, Verh. naturf. Ver. Briinn, 17: 46 (1876).
R. acris subsp. strigulosus (Schur) Hyl., Symb. bot. upsal., 7: 1 (1943).

R. granatensis and R. strigulosus are both very distinct from any other taxa
within the complex. R. granatensis is probably completely geographically
isolated from the rest of the complex, and no intermediates with any other forms
of the complex were seen. The plants of R. granatensis from Morocco appear
similar to those of southern Spain, but only very poor material was seen. The
distribution of R. strigulosus is overlapped by R. acris subsp. acris, but the only
intermediate plants which were seen came from a single region in south-west
Ukraine. These plants were exactly intermediate between the two supposed

240 S. M. COLES

parental types, and none were seen indicative of back-crossing to the parental
species. This interpretation of the hybrid origin of these plants is in agreement
with that of Hylander (1943).

R. granatensis and R. strigulosus are morphologically more closely related to
each other than to R. acris. They are completely distinct from one another in
floral but not in vegetative characters, although they can frequently be separated
on the latter characters alone. They both have rhizomatous rootstocks and broad-
lobed leaves with few teeth. In both species the beaks of the achenes are straight
or only curved, never hooked, but they are longer in R. granatensis than in
R. strigulosus.

R. AcRIS L., Sp. Pl., 554 (1753).

This is a very variable species which is distributed over most of Europe. All
the infraspecific taxa recognised within it are linked by intermediates which do
not appear to be of a hybrid nature.

Subsp. ACRIS
R. acris L. f. multifidus DC., Reg. Veg. Syst. Nat., 1: 278 (1817).
R. boreanus Jord., Obs. Pl. Crit., 6: 19 (1847).
R. tomophyllus Jord., Diagn., 71 (1864).

This is the most widespread subspecies and the most variable. The main cline
of variation in Europe is from west to east. Plants characteristic of the west have
long hairs, which are patent or deflexed on the lower part of the stem and on the
petioles. In the east the hairs are usually shorter and appressed. Also, in the
western plants the achenes and petals are larger. Broadly-lobed leaves are
commoner in the western plants.

Subsp. ACRIS var. VILLOSUS (Drabble) S. M. Coles, comb. et stat. nov.
R. acris var. friesanus (Jord.) Drabble forma villosus Drabble, Rep. botl Soc.
Exch. Club Br. Isl., 9: 472 (1930).

Plants of this variety occur in northern Scotland, Ireland, the Faeroes and
Iceland. They are characterised by having broadly-lobed leaves, which have
many hairs exceeding 1-2 mm on their upper surfaces. The hairs on the petioles
are long, often deflexed and usually of an orange-brown colour. One or two
plants approaching this condition have been seen from the west coast of Nor-
way, but no plants typical of this variety have been seen from the mainland of
Europe. Plants of this variety occur in the remoter parts of these islands, where
the vegetation has been relatively little affected by agriculture. It is probable
that this variety was previously more widespread but is losing its identity by
hybridising with plants of var. acris. In other areas in Britain, e.g. Teesdale,
which have been relatively little affected by modern agriculture, the plants
typically have long, deflexed hairs, and to some extent approach var. villosus.

Subsp. ACRIS var. PUMILUS Wahlenb., Flora Lapp., 159 (1812).

Var. pumilus is characterised by having leaves which are shallowly lobed and
which are glabrous in the spring. It occurs in Scandinavia, Iceland and the
Cairngorms, Scotland. It is mainly confined to mountains where it often occurs
in snow-patch habitats. Some of the plants examined from the extreme north of
Scandinavia were intermediate in character between subsp. acris var. pumilus
and subsp. borealis. Plants of these two taxa appear to form part of a stepped
cline from the north-west to the north of Europe.

RANUNCULUS ACRIS L. IN EUROPE 241

Subsp. BOREALIS (Trautv.) Nyman, Consp., 12 (1878).
R. borealis Trautv., Byull. mosk. Obshch. Isp t. Prir., 33: 72 (1860).

Plants of this subspecies are characterised by their very shallowly lobed leaves,
the petioles of which are usually covered by deflexed hairs which are always
very fine. The description of subsp. borealis in Flora Europaea (Tutin 1964)
covers both this subspecies and subsp. acris var. pumilus. Plants of subsp.
borealis have only been examined from the arctic coast of Russia, and the
nomenclature of these plants may need to be revised when the representatives
of the complex in the Urals and in Siberia are examined.

Subsp. FRIESANUS (Jord.) Rouy & Fouc., Fl. Fr., 1: 103 (1893).
R. friesanus Jord., Obs. Pl. Crit., 6: 17 (1847).
R. nemorivagus Jord., Diagn., 74 (1864).

This subspecies forms no part of the clinal variation as seen in the rest of
R. acris sensu stricto. It is the only taxon within this species which has a rhizome;
it is also characterised by having broadly lobed leaves. Its centre of distribution,
the mountains of eastern France and western Switzerland, is completely over-
lapped by that of subsp. acris. It may have been ecologically isolated in the past,
but at the present time intermediates between it and subsp. acris are very
numerous. It has been introduced into other parts of the continent, e.g. Sweden
and Austria, and here too it seems to be hybridising with subsp. acris.

Subsp. FRIESANUS var. PYRENAEUS S. M. Coles, var. nov.

Caudex rhizomatosus. Folia plerumque anguste lobata. Pili petiolares semper
adpressi, haud tenuissimi. Habitat in Pyrenaeis Montibus et in Hispania septen-
trionali et orientali.

Rootstock rhizomatous. Leaves usually narrowly lobed. Petiole-hairs always
appressed, never very fine. Found in the Pyrenees and in northern and eastern
Spain.

Holotype: Pont de Ravi, near Hospice de France, open place in damp beech
wood, France (Haute Garonne), 26. VII. 1961, E. K. Horwood (LTR).

This variety occurs over a wide geographical area, where it appears to be the
only form of R. acris sensu stricto. It is relatively constant in appearance
throughout its range. Except for its rhizomes it does not resemble specimens of
subsp. friesanus from eastern France, but it cannot be morphologically dis-
tinguished from some of the intermediate plants between subsp. friesanus and
subsp. acris which occur in eastern France. The relationships of this variety are
not clear; no plants of R. acris were seen from the region north of the Pyrenees
adjacent to the area occupied by this variety.

A taxon of uncertain status is R. steveni Andrz. ex Bess., Enum. Pl. Volhyn.,
22 (1822). The name has been much used, but with many different interpreta-
tions. Hylander (1943) suggested that the plants originally described might
have been forms intermediate between R. acris and R. strigulosus or, judging
from inadequate and doubtful type material, might only be R. acris subsp.
acris with elongated rootstocks such as occurs when a rootstock is buried.

THE BRITISH FORMS OF R. ACRIS

Only R. acris subsp. acris occurs in Britain. The general cline in morphological
characters of subsp. acris across Europe is visible even within the British Isles.

242 S. M. COLES

Subsp. acris var. villosus occurs in northern Scotland and Ireland, and var.
pumilus occurs only in the Cairngorms, Scotland.

The principal former classifications of British material were those of Town-
send (1900) and Drabble (1930). These, and the descriptions of R. acris and its
varieties in British Floras up to only a few years ago, were based on adaptations
of continental systems to British material, even though many of the continental
forms do not occur in Britain. The varieties were chiefly delimited on the charac-
ter of the rootstock and on the width of the leaf-lobes. From an examination of
herbarium material determined by Drabble and other British botanists, no
justification could be found for them to have called some rootstocks rhizo-
matous and others premorse; and the differences between the varieties in leaf-
lobe width were found to be mainly due merely to seasonal variation. Plants
collected in the spring were generally called var. boreanus or var. multifidus,
whereas those collected in the autumn were called var. steveni or var. friesanus.

KEY TO THE EUROPEAN TAXA

Descriptions of leaves refer to those produced just before the first flowering
stem of the year.

Measurements of petiole-hairs refer to the largest hairs on the petioles,
excluding the sheath region. (Abnormally large hairs of rare occurrence, for
any particular plant, are ignored.)

1. Rootstock much elongated, rhizomatous. Leaves on vegetative shoots
arranged in two ranks

2. Beaks of achenes distinctly hooked. Petiole-hairs frequently less than
0-8 mm in length (rarely exceeding 1:2 mm) often very fine (less than
0-02 mm wide) but plants with thicker hairs (up to 0-05 mm) common.

Leaves thin, usually broadly lobed with many teeth
R. acris subsp. aes
3. Leaves broadly lobed. Petiole-hairs frequently very fine, often
patent .. .. (Mainly in France and Switzerland, commonly
introduced to other countries) R. acris subsp. friesanus var. friesanus
3. Leaves narrowly lobed. Petiole-hairs never very fine, always
appressed .. Be .. (The Pyrenees and eastern Spain)
R. acris subsp. friesanus var. pyrenaeus
2. Beaks of achenes straight, or only slightly curved, never hooked. Petiole-
hairs long, exceeding 0-8 mm in length (frequently exceeding 1-2 mm)
and thick (always exceeding 0:03 mm). Leaves thick, broadly lobed with

relatively few teeth
4. Beaks of achenes 0:5-O0-:7 mm. Stamens with fewer than
20 hairs/filament .. .. (Eastern Europe) R. strigulosus
4. Beaks of achenes exceeding 0:8 mm. Stamens often with more
than 30 hairs/filament :
(South Spain and Morocco) R. uN
1. Rootstock short, premorse. Leaves on vegetative shoots produced spirally ..
R. acris L. sensu stricto (excluding subsp. friesanus)
5. Central lobe of the median leaf-segment less than 2/5 of
leaf length. Petiole-hairs deflexed, ue very fine (less
than 0:02 mm wide), white ..

(Arctic coast of Russia) R. acris 5 subsp. forentis

RANUNCULUS ACRIS L. IN EUROPE 243

5. Leaves variously divided, only rarely with the central lobe
of the median segment less than 2/5 of leaf length. Petiole-
hairs appressed, spreading or deflexed, rarely less than
0:02 mm wide ua R. acris subsp. acris

6. Leaves produced before the first flowering stem of
the year glabrous (those produced later bearing
hairs), usually shallowly lobed, never finely divided

(Scandinavia, Iceland, Scotland (Cairngorms))
R. acris subsp. acris var. pumilus
6. Leaves hairy, variously divided

7. Hairs on the upper surface of the leaves rarely
exceeding 1-2 mm. Leaves variously divided,
often very finely .. .. (Widespread over
most of Europe) R. acris subsp. acris var. acris
7. Hairs of the upper surface of the leaves frequently
exceeding 1-2 mm. Leaves never very finely

divided, lobes usually broad
(Scotland, Ireland, Iceland, Faeroes)
R. acris subsp. acris var. villosus

DISCUSSION OF CHARACTERS

THE ROOTSTOCK

The type of rootstock in this complex, whether premorse, i.e. short and compact,
or rhizomatous (Fig. 2) has previously been held as a character of primary im-
portance. This indeed was found to be justified. Rhizomatous rootstocks occur
in all plants of R. strigulosus, R. granatensis and R. acris subsp. friesanus. (How-
ever, it should be noted that in these taxa a rhizome is not always produced by
the apical bud of a plant in its first season.) Compared with R. acris subsp. acris,
which has a premorse rootstock, all these taxa have limited geographical dis-
tributions centred on a mountain range.

Although there is this clear separation of taxa on the type of rootstock, the
only real difference between a premorse and rhizomatous rootstock appears to
be in their degree of elongation and phyllotaxy. The phyllotaxy of a premorse
rootstock is spiral, whereas that of a rhizome is distichous. The phyllotaxy is not
altered in either case when rapid upward growth by extensive elongation of the
internodes is induced by inundation with sand. All grades between a premorse
rootstock and a fully-elongated rhizome occur in the array of forms inter-
mediate between R. acris subsp. acris and subsp. friesanus.

The premorse rootstocks of subsp. acris are slightly longer in the west of
Europe than in the east (Fig. 2). Unusually long rootstocks, approaching rhizo-
matous types in length, were found in some plants from the Shetland Islands.

THE LEAF

The leaf of R. acris sensu lato is palmate in shape and primarily three-lobed.
These primary lobes, or ‘segments’, may either be shortly stalked or joined for a
short distance. The three segments are further lobed, but never as far as the mid-
vein. The shape as a whole was defined by measurements made on the median
segment only. The measurements made were as follows:

D

244 S. M. COLES

FIGURE 2. Rootstocks: showing their appearance in September.
a. & b. Premorse rootstocks of R. acris subsp. acris showing differing degrees of elongation.
a. Finland, loc. 1. b. England, loc. 62a.
c. Rhizomatous rootstock of R. acris subsp. friesanus from France, loc. 39.

L length of median segment of the leaf

X distance from base of leaf to base of the free part of the median segment

Z distance from the base of the median segment to the base of the free part
of the median lobe of this segment

W width of the median lobe of the median segment, measured at the base of
the free part

These measurements are shown diagrammatically in Fig. 4.

It was found, from observations of plants in cultivation, that there is a
marked seasonal variation in leaf-shape. The successive leaves produced by a
rosette which has overwintered show increasing dissection: those produced
immediately before the first flowering stem are the most finely-dissected leaves
produced during the whole year (‘spring’ leaves). The leaves, that are pro-
duced by the buds in the axils of these ‘spring’ leaves, show a marked discon-

RANUNCULUS ACRIS L. IN EUROPE 245

FIGuRE 3. Leaf-shape.
a. Plant from England, loc. 62b: i. ‘spring’ leaf; ii. ‘autumn’ leaf.
b. Plant from Ireland, loc. 109: i. ‘spring’ leaf; ii. ‘juvenile’ leaf produced in the autumn
after transplantation.

tinuity in shape, being more broadly lobed and less deeply divided (‘autumn’
leaves) (Fig. 3). A gradual increase in dissection of the leaves then occurs until
‘spring’ leaves are produced. This seasonal variation in leaf-shape has been
almost totally overlooked by previous workers.

The comparisons of leaf-shape were restricted to the innermost ‘spring’ leaves.
Leaf-shape as recorded by the ratios X/L, Z/L and W/L is shown diagram-
matically in Fig. 4 for the whole of Europe. Figs. 5, 6, 7 and 8 show W/L
plotted against Z/L; these two ratios are those which most closely define the

2 aa A ONS} “6 “Z Ue
7L

FIGURE 5. Graphical comparison of leaf-shapes. © R. granatensis.
x R.acris subsp. acris from E. Europe. oO R. strigulosus.

246

2 . O . 5 .
3 4 5 6 7 Z (
FIGURE 6. Graphical comparison of leaf-shapes.
R. acris subsp. acris from France.
R. acris subsp. friesanus var. friesanus from France, Austria, Switzerland and Sweden.
R. acris subsp. friesanus var. friesanus in cultivation from France, loc. 39.
R. acris subsp. friesanus var. pyrenaeus from France and Spain.

@@Oo x

“2 3 “4 5 ; z
6 7 Ze

FIGURE 7. Graphical comparison of leaf-shapes.
R. acris subsp. acris var. acris from Norway and Sweden.
R. acris subsp. acris var. acris in cultivation from Sweden, loc. 23.
R. acris subsp. acris var. pumilus from Norway and Sweden.
R. acris subsp. borealis from N. Russia.

@eopx

RANUNCULUS ACRIS L. IN EUROPE 247

major characteristics of the leaf-shapes. These figures show more clearly the
range in leaf-shapes of certain groupings. Examples of actual leaf-shapes are
given in Fig. 9.

R. strigulosus and R. granatensis are both characterised by having broad-
lobed leaves. Fig. 5 shows how their range in leaf-shape is similar. On the charac-
ters of W/L and Z/L alone they appear similar in leaf-shape to some of the
western European R. acris sensu stricto; but they differ in that although they
have broad-lobed leaves the segments, especially in the case of R. granatensis,
are often only slightly fused or even stalked and the leaves, especially in the case
of R. strigulosus, have few teeth relative to their length. This latter feature also
occurs in eastern European R. acris subsp. acris, but such leaves are always
narrowly lobed. The leaves of R. granatensis and R. strigulosus are usually
thicker than those of R. acris sensu stricto, due to the former having larger cells;
this is the only anatomical difference which has been reported among the
European taxa.

R. acris subsp. friesanus is characterised by its broad-lobed leaves, which dis-
tinguish it from subsp. acris as it occurs in the surrounding districts (Fig. 6);
but leaf-shape alone would not distinguish it from all the forms of subsp. acris
that are to be found in north-west Europe. Most of the overlap in leaf-shape
shown in Fig. 6 is due to subsp. friesanus var. pyrenaeus, rather than to plants of
subsp. friesanus showing introgression with subsp. acris.

The much wider range in leaf-shapes within plants of subsp. acris from north-
west Europe is shown in Figs. 7 and 8. Fig. 7 also shows how the leaf-shape of
subsp. borealis differs from that of nearly all other groups excepting some of the
north Scandinavian subsp. acris var. pumilus.

2 3 “4 98) 6 Y/
A

Ficure 8. Graphical comparison of leaf-shapes.
R. acris subsp. acris var. acris from England and Wales.
R. acris subsp. acris var. acris from England, loc. 62b.
R. acris subsp. acris var. acris from Scotland, Iceland and The Faeroes.
R. acris subsp. acris var. pumilus from Scotland, Iceland and The Faeroes.
R. acris subsp. acris var. villosus from Scotland, Iceland and The Faeroes.

©eoprx

248 S. M. COLES

FiGurE 9. Examples of leaf-shapes.
. R. granatensis — Sierra Nevada, S. Spain. b. R. strigulosus — Transylvania, Rumania.
. acris subsp. acris from E. Europe: i. Hungary — loc. 58. ii. Austria — loc. 54.
. acris subsp. acris from W. Europe: i. England — loc. 60. ii. England — loc. 73.
. acris subsp. borealis — Vaigach Is., N. Russia.
. acris subsp. acris var. pumilus — Kiruna, Sweden.
. acris subsp. friesanus — Lyon, France.

WE ante) hee) te)
AADARA

Figure 10
STEM AND PETIOLE HAIRS

RANUNCULUS ACRIS L. IN EUROPE 249

THE STEM

A variable number of foliar organs occurs on the stem; the lower are stalked and
similar to the basal leaves, but higher up they become sessile and simpler in
form. The shapes and sizes of these organs can nearly always be correlated with
those of the basal leaves, The exceptions to this correlation occur in most plants
of subsp. borealis, subsp. acris var. pumilus and in many plants of var. villosus.
These plants have relatively broad, little-divided leaves, but the transition to
simple linear bracts is often abrupt.

The key in Flora Europaea (Tutin 1964) uses the height of the stem as one of
the main diagnostic characters separating some of the taxa. Studies of both
herbarium and cultivated material showed this to be a character of negligible
value.

THE HAIRS

Plants of the complex are generally rather hairy, except in their winter state.
Plants of R. acris subsp. acris var. pumilus are unusual in having glabrous
‘spring’ foliage. The pedicels of all plants always bear appressed hairs.

The hairs are single-celled and simple in form. Measurements of hair character-
istics were made on the petioles (above the sheathing region) and on the lower
part of the stem. The hairs were scored as appressed, spreading or deflexed.
The density of hair cover was scored using four subjective categories: very dense,
continuous, scattered, glabrous. The density was scored for ‘spring’ organs, as
it usually increases on the ‘autumn’ organs. The maximum values of hair-length
and width on the petioles were recorded (any unusually large hairs for the organ
concerned were ignored). Length and width were divided into five arbitrary
categories as follows:

Length Width

up to 0-4 mm up to 0:02 mm

over 0:4 mm to 0:8 mm over 0:02 mm to 0:03 mm
over 0:8 mm to 1:2 mm over 0:03 mm to 0:04 mm
over 1:2 mm to 1:6 mm over 0:04 mm to 0:05 mm
over 1-6 mm over 0:05 mm

Moa P
Om BWN

The results are shown in Figs. 10 and 11. In R. acris subsp. acris glabrous peti-
oles and stems are rarely found except in subsp. acris var. pumilus. Glabrous stems
occur mainly in the plants from areas with a continental climate or from northern
mountains, whereas hairiness is associated with more oceanic climates and
southern European mountains. Other trends are for longer hairs in the west
rather than in the east of Europe, and for deflexed hairs to be rarely found
except in the western oceanic region. Throughout the entire range of subsp. acris
a wide range of hair-widths occurs and no geographical trends of variation are
apparent, except in Scandinavia where montane plants always have relatively
narrow hairs. Only very narrow hairs occur on plants of subsp. borealis.

In R. acris subsp. friesanus the range in length and types of hairs is similar to
that of western European plants of subsp. acris, except that in var. friesanus
very narrow hairs are common, and in all the plants examined of var. pyrenaeus
the hairs were appressed.

R. strigulosus and R. granatensis are alike in always having very long, wide
hairs.

250 S. M. COLES

E
D
HAIR T b ‘| 3
LENGTH’
Al
4) 246) fiers

: HAIR WIDTH
R. acris. subsps. acris
and borealis

l Plantin cultivation

R. acris subsp. friesanus

FE R. strigulosus

R. granatensis

FiGureE 11. Petiole-hairs, lengths and widths.

The lengths of the hairs on the upper surfaces of the leaves are generally simi-
lar to those on the petioles; however R. acris subsp. acris var. villosus has hairs
on the upper surface which are commonly over 1-2 mm in length, a character
not found in other members of subsp. acris, even when such hairs commonly
occur on the petioles.

In most cases the colour of the hairs appears to be closely correlated with the
hair widths. The colours from the wider to the narrower hair-types range from
an orangy-brown, through increasingly paler shades of fawn to white.

THE FLOWER

On the sepals of R. acris sensu stricto hairs of different lengths are more or less
evenly distributed. In both R. strigulosus and R. granatensis the longest hairs
are usually restricted to the upper two-thirds of the sepal. The zone lacking

RANUNCULUS ACRIS L. IN EUROPE 251

long hairs is usually more distinct in R. granatensis, but sepal hair-type does not
form a reliable diagnostic character separating R. strigulosus from R. granatensis.

Measurements of shape and size of petals were made on those from the
earliest flowers on a cyme, since flower-size decreases later. The commonest
petal-shape is broadly obovate. The only real exceptions to this are found in
subsp. acris var. pumilus and subsp. borealis, where petals varying from very
narrow to very broad occasionally occur. The shapes and sizes of the nectary-
scales show great variation even from single localities; short scales tend to pre-
dominate in subsp. borealis and subsp. acris var. pumilus, but not to the exclusion
of longer types. The petal-colour appears the same throughout the complex:
the occurrence of brown-veined petals in subsp. borealis (Tutin 1964) is not
upheld.

Most variation in petal-length is found in subsp. acris, where short petals
occur in the east of Europe and longer ones in the west. Fig. 12 shows histo-
grams of petal-length for groups of cultivated plants. The petal-lengths of subsp.
friesanus are similar to western subsp. acris, whereas those of R. strigulosus and
R. granatensis are similar to those of eastern subsp. acris.

Hylander (1943) reported the presence of hairs on the filaments of both R. gra-
natensis and R. strigulosus, and stated that it thus served as a clear diagnostic
character between these taxa and the rest of the complex. This is not the case, as
hairs are commonly found on the filaments of eastern European R. acris subsp.
acris. R. granatensis, however, is distinct from the rest of the complex in having

SUBSP. FRIESANUS Ee
Raa

SUBSP. ACRI/S

from

Finland &

Russia

- = aE

SUBSP. ACRIS
from Austria
& Hungary

7 ' 8 9 LOR eal peel S Geel Ate el oy eee lOmim
LENGTH OF PETAL

SUBSP. ACR/S
from

Sweden, Denmark
& Britain

15

10 NO. OF
PLANTS
5

FiGurE 12. Histograms of petal-length for groups of cultivated R. acris s.s.

<a ie
Core eg a eae aa
CS OC oa

2mm.

FiGurE 13. Achenes.
a. R. granatensis — from two plants. b. R. strigulosus — from two plants.
c. & d. R. acris subsp. acris from W. Europe, c. England —two plants from loc. 60.
d. Sweden — two plants from loc. 23. e. & f. R. acris subsp. acris from E. Europe,
e. Hungary—loc. 59. f. Finland—loc.5. g. R.acris subsp. friesanus from France —loc. 41.

a greater number of hairs, at least thirty on some filaments of the flower; else-
where it seldom exceeds twenty.

It was found that the achenes of R. strigulosus and R. granatensis are distinct
from those of R. acris sensu stricto in having long, straight or only curving
beaks, whereas in all the subspecies of R. acris the beaks are hooked, or very
short. R. granatensis and R. strigulosus are distinct from one another in the
lengths of their beaks: those of R. granatensis are in the range of 0-9-1:2 mm,
whereas those of R. strigulosus are only 0-5-O0-7 mm in length.

Much variation occurs in the shape and size of the achenes of R. acris sensu
stricto. Variation in shape is often great even in plants from the same locality
(Fig. 13). In general the achenes are smaller in the east of Europe and larger in
the west. Achenes of subsp. friesanus are similar in size-range to those of western
subsp. acris. Only immature achenes of subsp. borealis were seen; they appeared
similar to those of western subsp. acris.

CHROMOSOMES AND BREEDING EXPERIMENTS

The chromosomes of R. acris sensu stricto have had more attention than have
those of most species of Ranunculus. All of the more recently published counts
of the chromosome number are of 2m = 14. In the earlier counts most of the
variations found in the chromosome number were reported by two workers
only, Senjaninova and Sorokin. Comprehensive lists of these and other earlier
counts are given by Gregory (1941) and Coonen (1939).

No chromosome counts have been reported for R. strigulosus and R. grana-
tensis. Pollen-grain size and stomatal diameter were examined in herbarium
material and were found in most cases to be greater than those in R. acris sensu

— a a a ee ee ee eE—eEeeEeEeEeEeEeeEEeEeEEEeEeEeEe—eEeEeEeEeEeeeeeeee eee ee

RANUNCULUS ACRIS L. IN EUROPE 253

stricto. Although no definite conclusions can be drawn from these measure-
ments they suggest that the first two species may possibly be polyploids.

Breeding experiments were carried out using R. acris subsp. acris and subsp.
friesanus from a series of different localities. Compatibility of selfing and occur-
rence of apomixis were tested. The plants which were used were grown in a
greenhouse and the flowers were individually covered with cellophane bags. In
these tests, out of more than 3,500 carpels from emasculated buds only 29 seeds
were set and of the 4,500 carpels which had been selfed only 3 seeds were set.
For a number of reasons, the seed which was set was most probably due to con-
tamination. It is highly probable, therefore that all plants of R. acris sensu stricto
are obligate outbreeders. This was previously not thought to be the case (Mars-
den-Jones & Turrill 1935, 1952; James & Clapham 1935). Preliminary tests were
made which suggest that the incompatibility mechanism is gametophytic.

The interfertility of plants from different regions and of different morphological
types was tested by means of a series of test crosses. These were between plants
from differing localities using mainly pollen from plants from localities 5 (Fin-
land), 50 (Austria) and 39 (subsp. friesanus) (see Appendix 1 for localities). The
results are given in Table 1. All the plants tested were found to be freely inter-
fertile and the seed set between geographically remote and morphologically
dissimilar plants was generally at least as good as that between morphologically
similar plants from the same country. The percentage of seed set generally
depended on carpel fertility rather than on the origin or fertility percentage of
the pollen.

The fertility of the ‘hybrid’ F, plants was tested by examination of the meiotic
divisions of the pollen-mother-cells. However, irregularities in the pairing of the
chromosomes were found as frequently in the parental plants as in the hybrids.
An estimate of apparent fertility of the pollen of the parent and hybrids was
made by staining the pollen in cotton blue. Even parental plants of normal
appearance were found commonly to have a percentage of good pollen as low as
20°% and seldom more than 80%. The following list gives the percentages of
good pollen in the first flower produced on a stem which had flowers of normal
appearance (each value refers to a separate cultivated plant).

Loc. 62a (England) : 56, 76, 59, 83, 56, 46, 60, 75, 70, 54%
Loc. 62b (England) : 40, 24, 18, 27, 28%
Loc. 33 (Germany) : 47, 42, 35, 39, 21%
Loc. 39 (France) : 91,89, 70, 73, 67%
Woes22 (Sweden) = 79;53;, 80,79; 757%

The first flower was always taken, as the percentage of good pollen fluctuates
greatly among different flowers of the same plant; even percentages as measured
above can vary greatly from season to season.

DISCUSSION AND CONCLUSIONS

Measurements of characters were made on organs produced at apparently the
same season and at the same position on the plant, so that they would be
comparable. This restricted the number of characters that could be measured
on much of the herbarium material. Seasonal variation in leaf-shape had been
ignored by most of the previous workers on this complex.

The presence or absence of a rhizome was found to be a reliable diagnostic

254

S. M. COLES

TABLE 1. TEST CROSSES BETWEEN CULTIVATED PLANTS

Locality
numbers

Ain! BI] w

62b

32

33

39

41
40

5
79; 68

54; 96
76

92; 96
54; 91

OF WILD ORIGIN

Results expressed as percentage seed set per flower.

3 Parents
50 39 Lind 4. 6, 24 22. 62b 32 335741840
82;91; 76 61; 67; 95 92 94 94
69;61;60 85; 63 85
46
93; 90 96; 92
54; 86 So 2 oT, 34 22
87; 79; 61 70; 73; 64 29
94; 29;50 94; 29; 28
61 43; 68
90; 79 78; 91
88; 90 TBI L2
TAZ OS) 91367
90;77;65 43; 47 59
65; 67 US 7/3
Oils 32 12; 0; 6
L727 17
182 142 35, 293 19 94 77 70 90 91 85
23; 94; 83 83 87 61
92; 88; 87
48; 82 81; 90 53
TZ
84; 73 94; 89 85
14)
28 323 29; 10; 47 38
21
63; 84;91 98;95;80 74 90
91;89;79 46 1D)
86
9225:935,24 40.45 23 66
69
36; 14; 0
0; 68; 63
54; 65 39; 48 28
48;74;41 57; 54; 56 70
85 WSO US 64

RANUNCULUS ACRIS L. IN EUROPE 255

character. Leaf-shape was found to be diagnostic for most taxa, although it was
very variable even within plants from a single locality. Hair-type was similarly
found to be very variable in some localities, but most taxa were characterised
by hairs of a particular length and width. Differences in floral characters were
found between the three species, but within R. acris sensu stricto no taxa could
be delimited on such characters.

Chromosome number is constant within R. acris sensu stricto (2n=14). All
plants within R. acris sensu stricto appear freely interfertile, but information as
to the fertility of plants resulting from such crosses, compared with that of the
parental plants, would require a detailed knowledge of factors controlling
femaleness. Although femaleness is so common it would appear disadvantageous
to the species, which is an obligate outbreeder. It is suggested that there is a
possibility that R. granatensis and R. strigulosus may be polyploids, but no living
material of these two species was available.

All the rhizomatous taxa within the complex appear to have evolved in the
mountainous regions of southern Europe. R. granatensis and R. strigulosus
show so many features in common that presumably they must have come from
a common stock. R. acris subsp. friesanus, which also has rhizomes, shows no
close relationship to either of these species.

An array of intermediate plants were found between the different morpho-
logical forms within R. acris sensu stricto. This can be attributed to the facts that
all the plants of this species appear to be obligate outbreeders, there appear to
be no sterility barriers between any forms of the species, and the ecological or
geographical barriers, which must have occurred especially in the case of subsp.
friesanus, appear to be breaking down due to farming activities, which also
enable the plants to spread as weeds.

ACKNOWLEDGMENTS

I wish to thank Professor T. G. Tutin for his encouragement and advice during
this work. The work was carried out at Leicester University, during the tenure
of a Research Studentship from the Science Research Council, to whom I am
much indebted. I am most grateful to Birmingham University for a grant to
cover the cost of publication of the large maps.

I also wish to thank the curators of the herbaria, from which material was
loaned, and the many people who helped collect living material.

REFERENCES

BENSON, L. B. (1948). A treatise on the North American Ranunculi. Am. Midl. Nat., 40: 1-261.

Co es, S. M. (1968). Systematics of the Ranunculus acris L.s.l. complex in Europe. Ph.D. thesis.
University of Leicester.

_ COOoNEN, L. P. (1939). The chromosomes of Ranunculus. Am. J. Bot., 26: 49-58.

DRABBLE, E. (1930). The British forms of Ranunculus acer L. Rep. botl Soc. Exch. Club Br. Isl.,
9: 472-478.

GREGORY, W. C. (1941). Phylogenetic and cytological studies in the Ranunculaceae. Trans.
Amer. phil. Soc., 31: 443-521.

Harper, J. L. (1957). Ranunculus acris L., R. repens L., R. bulbosus L., in Biological Flora of
the British Isles. J. Ecol., 45: 289-342.

256 S. M. COLES

HyYLANpeR, N. (1943). Ranunculus acris L. ssp. friesanus (Jord.) R. & F. Symb. bot. upsal.,
7: 1-7.
JAMES, W. O. & CLAPHAM, A. R. (1935). The Biology of Flowers. Oxford.
MARSDEN-JONES, E. M. & TURRILL, W. B. (1929). Studies in Ranunculus. I. Preliminary account
of petal colour and sex in Ranunculus acris and R. bulbosus. J. Genet., 21: 168-181.
MARSDEN-JONES, E. M. & TURRILL, W. B. (1935). Studies in Ranunculus. III. Further experi-
ments concerning sex in Ranunculus acris. J. Genet., 31: 363-378.

MARSDEN-JONES, E. M. & TURRILL, W. B. (1952). Studies in Ranunculus. IV. Additional
experiments with Ranunculus bulbosus and R. acris. J. Genet., 51: 26-31.

TOWNSEND, F. (1900). Ranunculus acer L. J. Bot., Lond., 38: 379-383.

TuTIN, T. G. (1964). Ranunculus, in TuTIN, T. G., et alia, ed. Flora Europaea, vol. 1, pp. 223-
238. Cambridge.

APPENDIX I

ORIGIN OF PLANTS TO WHICH REFERENCE IS MADE

1 Espoo, Finland 40 Strasbourg, France
2 Borga, Finland 41 Massif de Hoheck, Vosges, France
3 Tampere, Finland 44 Udine, Italy
4 Punkaharju, Finland 48 Genova, Italy
5 Kuusamo, Finland 50 Graz, Austria
6 Oulu, Finland 54 Heiligenblut, Austria
8 Leningrad, Russia 58 Budapest, Hungary
22 Landskrona, Sweden 59 Dorog, Budapest, Hungary
23 Goteborg, Sweden 60 Church Stretton, Shropshire,
24 Varnamo, Sweden England
25 Stockholm, Sweden 62 Cawston, Warwickshire, England
32 Gransee, E. Germany 73 Middleton-in-Teesdale, Durham,
33 Karlmarxstadt, E. Germany England
39 Mairon-sur-Béze, Dijon, France 109 L. Mask, Co. Mayo, Ireland

APPENDIX 2
EXAMPLES OF HERBARIUM MATERIAL OF THE RECOGNISED TAXA

R. strigulosus Schur (Fig. 14A)
In pratis Transsilvan., D. J. Schur (W)
Transsilvanica, Hermannstadt, 12. VI. 1906, J. Barth (W)
Tilalmas prope Toroczko, Transsilvanica, VII. 1883, J. de Csatdé (P)
Kolozsvar, comit Kolozs, 14. V. 1916, M. Péterfi (C)
R. granatensis Boiss. (Fig. 14B)
Sierra Nevada, sunny bank by the bridge over R. Genil..., 10. VI. 1926, A. J.
Wilmott (BM)
In Valle fluv Feint, Nevada, 29. VI. 1876, M. Winkler (C, W)
Sierra de Segura, above Rio Segura near La Toba, edge of water channel, 27. VI.
1955, V. H. Heywood (LTR, LIVU)
Sierra Nevada, lot. humidus graminos in Dehesa de S. Gerorimo et valle fi.
Monothil, 9. VIII. 1879, Huter (CGE, C, E, S)
R. acris L. subsp. acris var. acris (Fig. 15A)
Beaminster, Dorset, 2. VI. 1913, J. M. Roper (CGE)
Austria Inferior, In pratis ad ‘Mauer’ prope ‘Wien’, VI. 1903, E. v. Halacsy (W)
Rossica, Archangelsk extra oppidum..., 13. VII. 1911, S. J. Enander (S)

RANUNCULUS ACRIS L. IN EUROPE 257)

R. acris subsp. acris var. pumilus Wahlenb. (Fig. 16A)
Lycksele Lappmark, Tarna, Norra Storfjallet, Dalaive, 31. VII. 1944, H. Lenand-
der (S)
Torne Lappmark, Jukkasjarvi parish, Mt. Atjaktjakko, 5. VII. 1947, H. Smith (E)
Corrie-an-Lochan, East side of Braeriach, E. Inverness, 12. VIII. 1898, E. S.
Marshall (CGE)
R. acris subsp. acris var. villosus (Drabble) S. M. Coles (Fig. 15B)
Melvich, W. Sutherland, 15. IX. 1897, W. A. Shoolbred (BM, CGE, LTR)
Coast near Rockfield, E. Ross, 19. VII. 1890, E. S. Marshall (CGE)
Near the summit of Langa, Harris, 14. VIII. 1841, - (CGE)
R. acris subsp. borealis (Trautv.) Nyman (Fig. 16B)
Novaya Zemlya, Matotschin Schar, 30. VII. 1908, R. Niemann (CGE)
Waigats Is., Dolga Bay, 1-10. VII. 1897, H. W. Fielden (CGE)
Waigatsch, sinus Warnek, 11. VIII. 1902, O. Ekstam (S)
R. acris subsp. friesanus (Jord.) Rouy & Fouc. var. friesanus (Fig. 17A)
Lyon a Chanelay, A. Jordan (CGE)
Forét du Rhin-du Bois, Cher, 2. VI. 1855, A. Déséglise (P)
Near Archettes, Forét d’Epinal, Vosges, 30. VII. 1963, E. K. Horwood (LTR)
R. acris subsp. friesanus var. pyrenaeus 8S. M. Coles (Fig. 17B)
Prov. Teruel. Albarracin, lieux humides et herbeux, sur le calcaire 1,300 m, VI.
1894, E. Reverchon (BM, E, P, S)
Guipuzcoa. Gabo de Higuer, V. 1895, M. Gandoger (C)

258 S. M. COLES

iy

4 | ik

A B
FIGURE 14
A. R. strigulosus Schur B. RR. granatensis Boiss.
[In pratis fertilibus, Transsil- [Sierra de Castril....-—. VII.
van...., —. VI.-, D. J. Schur 1903, E. Reverchon (E).]

(W).]

RANUNCULUS ACRIS L. IN EUROPE 259

B
A
FIGURE 15
A. R. acris L. subsp. acris var. acris B. R. acris L. subsp. acris var. villosus
[D’Angers, 18. VI. 1853, (Drabble) S. M. Coles [Damp ground
A. Jordan (C).] sloping to a small rocky bay,

Nr. Melvich, W. Sutherland,
Ref. No. 1864, 15. VII. 1897,
E. S. Marshall (CGE).]

260 S. M. COLES

B
FIGURE 16
A. R. acris subsp. acris var. pumilus B. R. acris subsp. borealis (Trautv.)
Wahlenb. [Lule Lappmark, Nyman [Novaja Semlja, Matot-
Kvikkjokks Kapellfors. .. ., schkin Scharr., 20. VIII. 1891,
23. VII. 1943, S. Selander & O. A. Ekstam (S).]

N. Dahlbeck (S).]

RANUNCULUS ACRIS L. IN EUROPE 261

FIGURE 17
A. R. acris subsp. friesanus B. R.acris subsp. friesanus var.
(Jord.) Rouy & Fouc. var. pyrenaeus S. M. Coles [Nr.
friesanus [Lyon a Chanelay, Hospice de France, Haute
Herb. Jordan (CGE).] Garonne. ..., 26. VII.

1961, E. K. Horwood
(LTR).]

TEST arg ie SE
4) pola Liv. aleve
SOK cori en tae
HF 98 po baronet)
bow A ad bee!
(AED

Watsonia, 8, 263-276 (1971). 263

Studies on British Poppies 4. Some aspects of variability
in the British species of Papaver and their relation to
breeding mechanisms and ecology

S. ROGERS
Department of Biology, Queen Elizabeth College, London

ABSTRACT

The patterns of intra-specific variation in the British species of Papaver have been studied
using mainly characters of the ripe capsules. The results, considered in relation to the breeding
mechanisms of the species, show that in the outbreeding P. rhoeas variation is primarily
between individuals and the variation within populations is generally such that any variation
between them is not significant. In the self-compatible species, P. dubium and P. lecogii, on the
other hand, the variation between populations is generally highly significant. Some populations
nevertheless show more internal variation than others and this appears to parallel a variation
in the degree of inbreeding.

INTRODUCTION

The five species used in this study fall into two sections of the genus as classified
by Fedde (1909), the section Orthorhoeades including P. rhoeas, P. dubium and
P. lecogii whilst P. argemone and P. hybridum are included in the Argemono-
rhoeades.

In many respects the species are all similar, being annual plants inhabiting
open communities such as annual crops, waste ground and recently disturbed
roadside verges. Yet, in the south of Britain at least, only P. rhoeas is universally
abundant; P. dubium is only locally abundant, whilst the other three species
are very localised in their distribution.

The existence of large populations of P. rhoeas focusses attention upon its
variability, especially in obvious characters such as flower colour, foliage colour
and form, and capsule shape. This has resulted in the description of numerous
varieties. Thus Fedde’s list (1909) comprises twenty varieties and six sub-
varieties, based on foliage form, petal colour, the nature of the indumentum,
the form of the stigmatic disc and the habit of growth. There is little information
regarding variability in the other species. Fedde lists nine varieties of P. dubium
based mainly on foliage characters but also including P. dubium var. lecogii
(Lamotte) Fedde, a taxon now reinstated as a species. In his description Fedde
points out the essential difference in capsule form between his var. /ecogii and
P. dubium. Thus, of the latter he wrote: ‘capsula ... ad basim paulatim attenu-
ata’, whilst of P. lecogii he said: ‘ad basim plus minusve subito rotundato-
attenuata’. Fedde’s seven varieties of P. argemone include two distinguished by
small size and non-incised foliage. Plants of this type are not uncommon in
corn crops and it may be that this growth habit results more from habitat con-
ditions than from genetic differences; the other varieties are based on the
development of the capsule setae, the indumentum, and the entire or laciniate
nature of the petals. This latter variant also occurs in other members of the
genus e.g. the Shirley section of P. rhoeas, and P. somniferum. There are only one
or two records for each of the five varieties of P. hybridum described by Fedde

264 S. ROGERS

and none of these is from British material. The taxonomic literature thus sup-
ports field observation in suggesting that P. rhoeas is far more variable than the
other species.

In attempting to assess this variability and correlate it with the reproductive
biology of the species, attention was confined to the capsules as providing
characters which could readily be measured and recorded. Data relevant to
reproductive biology are being considered in another paper in this series but
their significance in relation to the variability patterns found will be discussed
here.

METHODS

The shape of the capsule was expressed in terms of
(1) the length/breadth ratio.
(2) an angle expressing the contraction of the capsule base, referred to hereafter

as the base angle. Length and breadth were measured directly using callipers.
The base angle was obtained indirectly from a silhouette projected at a standard

FIGuRE 1. Capsule Measurements
(a) Length/breadth ratio —all species. L = length from top of flower scar to apex of disc;
B = capsule diameter (calliper measurement). (b) and (c) Base angles; (b) in short-capsuled
species and (c) in long-capsuled species. A-B diameter of top of flower scar. C-D diameter
5 mm (E) above A-B. C’ D’ diameter 10 mm (E’) above A-B. x = base angle. All measure-
ments at 3 diameters magnification.

STUDIES ON BRITISH POPPIES 265

magnification of 3 diameters on to graph paper. The lower measurement taken
in all species was that of the diameter of the top of the flower scar (A—B in
Fig. 1). For the short-capsuled species the second measurement (C—D) was
taken 0-5 cm (E) above this, but in the long-capsuled species it was found neces-
sary to employ the diameter 10 mm above the top of the flower scar to ensure that
the maximum width of the capsule was recorded. The angle was obtained by
calculation from the formula

X = 2 tan7? —— =

RESULTS

(1) CAPSULE VARIATION WITHIN THE SPECIES

The data are summarised in Table 1 and statistical analyses were carried out
using the F-test so that within sample-variation and between-sample variation
could be compared (Tables 3a & 3b).

(a) PAPAVER RHOEAS. Capsule shape in P. rhoeas shows a wide range
of variation both in length/breadth ratio and base angle. The results expressed
graphically in Fig. 2a and Fig. 5e are based on ‘three capsule means’ for every
plant sampled during the investigation. Statistical analysis of the within and
between-plant variation was performed using five-capsule-per-plant samples from
thirty randomly selected plants in a single population. The analysis was repeated
for four populations from widely separated localities in southern England. In
each case, the analysis showed the between-plant variation to be highly signifi-
cant (P < 0-001 or 0-01 in one case) in spite of the considerable phenotypic
variation within individuals (Table 3a).

a) Papaver rhoeas b) Papaver dubium

_ 20 MeN
LENT E \
TEN
i! SN
sae
1 i

° Frequency

2:0 2°5 3:0 3-5

Length breadth ratio

FiGcurE 2. Frequency distribution of capsule length/breadth ratios for three species of Papaver.
Continuous line: all data; broken lines: sample populations. Ordinates are % frequencies.

266 S. ROGERS
TABLE 1. SUMMARY OF CAPSULE ANALYSIS RESULTS
SPECIES INDIVIDUALS

Normal range Maximum
Minimum Maximum Range of variability range

P. rhoeas fed 2:9 1-8 0-2-0-5 0-7
L/B P. hybridum L372 2:0 0:8 0-2-0:3 0-63
Ratios P. dubium 2-3 4:5 22 0-35-0:9 1:47
P. lecogii 2:0 3) | (121 0-35-0-65 0-74
P. argemone 2:6 4°5 1:9 0-4-1-0 les
P. rhoeas 10 105 95 10-30 47
Base P. hybridum 49 95 46 12-30 44
Angles P. dubium 4 42 38 7-22 36
P. lecogii 19 52 35 8-16 24
P. argemone 4 37°6 33-6 11-23 31

To anticipate the criticism that variation within the individual is not fairly
represented in a five capsule sample, the same analysis was carried out using
ten and fifteen capsule samples from twenty plants of known and different origin
grown in a botanic garden. Between-plant variation reached the same level of
significance and the five capsule sample was accordingly taken as representing
adequately the variation within the individual. Smaller individual samples are
in any case preferable since relatively few plants in wild populations produce
as many as ten fully matured capsules.

By bulking the individual data for each of the four populations used to obtain

TABLE 2. CAPSULE MEASUREMENTS OF SEED PARENTS
AND PROGENY

SEED PARENT FAMILY
Inbreeding/
L/B Ratio Base L/B Base Outbreeding
Angle Ratio Angle Ratio
1 1-49-1-74 71-82 1-15-2-25 33-98 0
P. rhoeas 2 1-65-1-78 60-69 1:25-2:45 18-83 0
3 2:18-2:33 10-33 1:55-2:95 18-83 0
4 1-:2-1-48 66-95 1-15-2-55 20-103 0
P.dubium 1 3-0-3 -62 8-19 2:55-3-75 5-40 0-56/1
a 2:7-3-2 12-28 2:55-3:65 4-27:°5 7 Jes) |
P. lecogqii — 2-25-2-7 24-41 2:15-3:05 21-5-47-5 1:18/1
Inbreeding/outbreeding ratio = Se
g g = (T=S)
where S = mean number of potentially viable seeds following unaided self

pollination
T = mean number of potentially viable seeds following open pollination

STUDIES ON BRITISH POPPIES 267
TABLE 3 (a) VARIATION WITHIN POPULATIONS

Variance L/B BA L/B BA L/B BA L/B BA L/B BA
Estimates 1 2 3 4 5
between
P. rhoeas mneividel 1:21 1557 0:0833 489-2 0-067 519:3 0:203 914-2
within
individuals 0-23 129 0-032 99:09 0-006 69-6 0:005 396-9
F= 5-1* 12-06* 2:59* 4:94* 11:16* 7-46* 36-9* 2°3*
between
P. dubium individuals 0:1609 62 0-1998 49-7 0-:1186 37:09 0-116 96:27 0-:3110 59-04
within
individuals 0:0297 17:7 0:1613 36:40 0:045 25:02 0-035 8:07 0-054 23-94
F= 5-41* 3-6* 1:23 0:13 2-6f 1-48 3-31 11-9 5:78T 2:64t
between
P. lecogii individuals 0-:0700 39-4 0:0412 17:96 0:086 99-27
within
individuals 0-0265 45-0 0:024 20:02 0:0224 15-66
F= 2-6* 0:87 1-71 0:89 3:83 6:33*
between
P.argemone_ individuals 0-137 44-4
within
individuals 0-109 610
= 1:256 0-073
between
P.hybridum individuals 0-049 171
within
individuals 0-016 99-4
F= 3-06 1:72
* significant P= 0-1% + significant P = 1%

(b) VARIATION BETWEEN POPULATIONS

Variance Estimates L/B ratio Base Angle

P. rhoeas Between populations 0-0146 1576
(4 populations) Within populations 0:0423 247

a 0-3379 6:38T
P. dubium Between populations 0-4546 54:5
(5 populations) Within populations 0-0496 13-2

ls 9-16* 4-12*
P. lecogii Between populations 0:2584 38-5
(3 populations) Within populations 0-0155 10-8

i 16-67* 3-56°

* significant P = 0-1% } significant P=1% ° significant P= 5%

the five capsule samples analyses, it was possible to use the F-test to compare
within- and between-population variation. In respect of length/breadth ratio
the within-population variation was such that a significant between-population
variation was not obtained.

(b) PAPAVER DUBIUM. As in P. rhoeas there is much phenotypic varia-
tion within the individual in the expression of the capsule characters, but unlike
that species, large populations of P. dubium do not generally embrace the entire
range of variation of the species. However, the length/breadth ratio results
shown graphically in Fig. 2b indicate that different populations vary in this
respect, some showing a relatively high degree of uniformity and giving high
frequencies over a relatively narrow range of variation, while others seem much
more variable and hence yield low frequencies over a wider range of variation.

Analysis of the data shows that the within-individual variation and between-
individual variation do not differ significantly in a population of the former

268 S. ROGERS

type, whereas in one of the latter type the between-individual variation is signi-
ficantly greater than the phenotypic plasticity (Table 3a). The implication of
this would seem to be that populations of the latter type are genetically more
heterozygous than those of the former type and such variations in population
structure may be related to intraspecific variation in the breeding mechanism.

In spite of the differing amounts of variability within populations, there is
nevertheless a significant between-population variation in both characters
(Table 3b).

(c) PAPAVER LECOQII. This species has the smallest range of variation of the
long-capsuled species, both in length/breadth ratios and base angles. The range of
variation in both characters overlaps that of P. dubium; but the modal frequencies
of the two species are quite widely separated and when both characters are con-
sidered there is no difficulty in assigning capsule samples to the correct species
(Fig. 3).

Modal frequencies
P. dubium P. lecogii
Length/breadth ratio 3-0-—3-2 2°6-2:7
Base angle 25 - 30 35-40

40)

*, Frequency
i)
Lo)
——+ 4]

3°5 4:0 4:5 10) 20 40 60
Length breadth ratio Base angles

Ficure 3. Papaver dubium and P. lecoqgii: Comparison of capsule characters. Continuous line:
P. dubium; broken line: P. lecogii. Ordinates are % frequencies.

Phenotypic variability within the individual in the length/breadth ratio of the
capsule is much less than in P. dubium, and in some populations the between-
individual variation is significant whereas in others it is not (Table 3a). The
variation between populations is highly significant (Table 3b) and while a
proportion of the individuals from one population will invariably fall within the
variation range of another, yet each is distinct, at least in respect of this particular
character. The same is not true, however, of the capsule base shape, where an
inherently small range of variation in the species coupled with much phenotypic
plasticity tend to preclude the observation of any significant difference between
individuals or populations.
(d) COMPARISON OF PAPAVER RHOEAS, P. DUBIUM AND P. LECOQII

The differences between these three species are most readily appreciated by .
comparing the numerical and graphical results obtained from families of known

a) Papaver rhoeas

Frequency

o/
£0

c) Papaver lecogii

4074
0 304
fe) 20
—
) 10
2:0 2:5 3-0
CC
fo)
20 40 60
Length breadth ratio Base angles

FiGcure 4. Comparison of the variability in capsules of families of plants from known seed
parents.

The two families of P. rhoeas correspond to families 1 (broken line) and 3 (continuous line)

in Table 2. In P. dubium, family 1 in Table 2 is represented by the unbroken line and family 2

by the broken line. The horizontal lines represent the range of character variation in the seed
parent. Ordinates are % frequencies.

270 S. ROGERS

seed origin. Each family of twenty plants was raised in outdoor plots from a
sample of the seed from a single open-pollinated capsule.

In Fig. 4 the data from samples of 15 capsules/plant from 2 families of P. rhoeas,
2 of P. dubium and 1 of P. lecoqii are compared with the 3 capsule means of
their respective seed parents. Table 2 gives the summarised capsule data for
each family, together with that for two other families of P. rhoeas and the
inbreeding/outbreeding ratio for each family.

In P. rhoeas the salient features are the wide range of phenotypic variation in
the progeny relative to that within the known parent and the efficiency of the
barrier to self-fertilization. These features, together with the significant between-
plant variation establish beyond doubt the genetic heterozygosity of the species.

In both the long-capsuled species, by contrast, the capsule variability of the
progeny does not greatly exceed that of the known parent, and this pointer
towards a measure of genetic homozygosity in these species would seem to be
reinforced by the results of breeding experiments, which show a proportion of the
seed set to be self-fertilized. The difference between the degree of inbreeding in
the two families of P. dubium has been discussed previously, and while the striking
uniformity of family 2 correlates with its high inbreeding/outbreeding ratio, too
much significance should not be attached to a single result.

(e) PAPAVER ARGEMONE AND PAPAVER HYBRIDUM

Phenotypic variation within the individuals of P. argemone in the one large
population examined was such that variation between them was not significant.
Individuals from this population were subsequently shown to be strongly in-
breeding so that the results, as far as they go, are consistent. No other wild
population of sufficient size was obtained to permit of the variation between
populations being analysed.

Individual plants of P. hybridum also show considerable variation in capsule
shape. However, conflicting results were obtained from the only wild population
analysed. In length/breadth ratio the variation between individuals was signifi-
cant in spite of individual phenotypic variability but in base angle the variation
between individuals was not significant. It is obvious that more information is
required for this species before attempting to relate its variation-pattern to its
intensive inbreeding.

(2) GENERAL CONSIDERATIONS

It is apparent from this study that the long narrow type of capsule, characteristic
of P. dubium, P. lecogii and P. argemone, is more variable in length/breadth
ratio than the short, broad type of P. rhoeas and P. hybridum. Conversely, the
more abruptly contracted capsule base of the short-capsuled species is more
variable than the tapering form of the long-capsuled species. Since the base angle
is inherently less variable in the P. dubium type of capsule, and variation within
the individual may represent an appreciable part of the total species variation,
results for this character do not show the same clearly defined differences as are
obtained with the larger angles and bigger total variation of the P. rhoeas type
of capsule base. In all species, the expression of the individual genotype is sub-
ject to appreciable phenotypic modification and the significance of inter- and
intra-specific variation can, therefore, only be considered in relation to that
within the individual.

Expressed graphically both the length/breadth ratio and base angle data corre-
spond closely to normal curves in all the species, suggesting that capsule form,

STUDIES ON BRITISH POPPIES 271

in common with other variations of a quantitative nature, is polygenically
determined.

In the short-capsuled species observation suggests a correlation between the
length/breadth ratio and the base angle, a high ratio being associated with a
low angle and conversely. This association of the characters is less apparent in
the long-capsuled species.

(3) SOME OTHER ASPECTS OF VARIABILITY IN PAPAVER RHOEAS

(a) PEDICEL HAIRS
The two forms of multicellular pedicel hair occurring in the genus Papaver are
illustrated in Fig. 5. In the spreading form each hair has a cushion-like swollen
base and stands out stiffly at approximately a right-angle to the stem. In the
adpressed or strigose form the hair is bent through a right-angle just above its
point of origin so that it lies parallel to the stem, often pressed closely to it. An
intermediate form ‘semi-adpressed’ also occurs in which the hair base is ad-
pressed but the tip stands outwards instead of lying close to the pedicel. This
form has been observed in P. nudicaule by Fabergé (1943) and in P. rhoeas by
Turrill (1946). The adpressed forms in P. rhoeas have been variously designated
taxonomically as P. strigosum (Boenn.) Schur, P. x strigosum (Boenn.) Schur
and P. rhoeas f. strigosum Boenn. Field scoring shows that the frequency of
the ‘adpressed’ character is always low but that it varies from one population
and district to another. The highest incidence observed was 8%, compared with
a mean of 2:95%, and this was encountered in one of the most southerly popu-
lations examined. Over the relatively small south-north distances involved in
this survey, however, it was not possible to ascertain whether the incidence of
adpressed hairs declines in frequency northwards as suggested by Turrill (Joc.
cit.).

Families raised from ‘adpressed’ plants pollinated naturally in the field show
an approximate ratio of 3 spreading: 1 adpressed. This would point to ‘adpressed’
being a simple recessive as suggested by previous authors (Rasmuson 1920), and
also indicates that in populations where it does occur a considerable number
of spreading hair plants are heterozygous for the gene since all the seed is, of
necessity, cross-fertilized. Some further information relevant to the inheritance
of the character was obtained from hybrids between P. rhoeas and P. dubium.
The latter species invariably has adpressed hairs. P. rhoeas (adpressed) x P.
dubium, and the reciprocal cross yielded progeny all with adpressed hairs. P.
rhoeas (spreading) x P. dubium yielded equal numbers of spreading and ad-
pressed. This is in partial agreement with the results obtained by Rasmuson
(1920) and interpreted by him as indicating the dominance of spreading hairs in
P. rhoeas.

Fig. 5e shows that there is no significant difference in capsule characters
between spreading- and adpressed-hair plants, and this may be taken as further
evidence in favour of the view that the adpressed variant is a simple allelic one
which does not merit taxonomic rank.

(b) PETAL BLOTCH
Taxonomic descriptions of P. rhoeas usually imply that a black blotch at the
base of each petal is a frequent occurrence. Data collected from forty populations
in twelve vice-counties of England show that only approximately one-fifth of
the plants have blotched petals. Two hundred and fifty-four plants from 27
populations were scored for both presence and size of blotch and in only just

PUP? S. ROGERS

over 2% was the blotch found to be large and conspicuous. Both Newton (1929)
and Philp (1933b) associate the development of the blotch with a dominant gene
B, the recessive of which produces a white blotch, yellow anthers, white or tinged
filaments, white stigmatic rays and a light-coloured seed testa. This recessive
has been seen only once in all the wild populations examined though it is
characteristic of the cultivated Shirley poppy. The other phenotypic characters
which Philp and Newton ascribe to B, namely dark purple filaments and
anthers, purple rays and dark rays and dark seed coats, are all present in the
plants lacking the petal blotch.

(c) : (d)

(e)
>
oO
c
oO
Ss 1.0 1.5 2.0 2.5 30
= Length/breadth ratio

Base angle

FiGure 5. Papaver rhoeas: Variation in pedicel hairs.
(a) Adpressed pedicel hairs (x 7) and (b) detail of hair base (x 115); (c) spreading pedicel
hairs (x 7) and (d) detail of hair base ( x 115); (e) capsule characters in spreading (continuous
line) and adpressed (broken line) hair plants. Ordinates are % frequencies.

STUDIES ON BRITISH POPPIES 273

30 30
20 20
10 10
{0} ce)
6 if ksh © HO. Wi A ABs AS iS 10) ] 2 3 4 5 6
Mean stigma number per plant Range in stigma number per plant

FiGure 6. Papaver rhoeas: Variation in stigma ray number (data from all plants sampled).
Ordinates are % frequencies.

(c) THE NUMBER OF STIGMATIC RAYS

The expression of this character varies within as well as between individuals.
Using a sample of three capsules per plant the mean stigma number varied from
6 to 14 with the highest frequencies between 9 and 11. Less than 10% of the
plants had the same number of rays on the three capsules examined, a variation
of 1 or 2 being the most frequent. The maximum within-plant variation observed
was 6 and on this basis the between-plant variation is significant (P < 0-001)
(Fig. 6). It would seem, therefore, that stigmatic ray, and hence placenta
number has a genetic basis, though this is subject to considerable phenotypic
modification. The expected association between ray number and capsule size is
by no means invariable; small capsules may have numerous rays and large ones
relatively few. It is not known to what extent stigma, and hence placenta, number
might affect the reproductive potential of a capsule. A larger number of placentae
would suggest a greater seed-bearing surface within the ovary and possibly,
therefore, more seeds, but this would need to be considered in relation to capsule
SIZe.

DISCUSSION

(1) VARIATION PATTERNS AND BREEDING MECHANISM

The species of Papaver in Britain are seen to exhibit two fundamentally different
patterns of variation. One, exemplified by P. rhoeas, presents obvious visual
differences between individuals within a population, whilst comparison of large
populations shows no statistically significant difference between them, at least
in respect of the characters investigated. In the second pattern, exemplified by
P. dubium and P. lecogii, visual individuality is less apparent, and in the charac-
ters studied inter-individual variation reaches significance within some popula-
tions, but not within others. Populations, however, do differ significantly from
one another. The overall picture thus resembles that described by Baker (1953)
in Armeria maritima, where the subsp. maritima showed variation between
individuals while in the subsp. californica populations were much more uniform
internally but differed significantly from one another. This was found to be

274 S. ROGERS

correlated with breeding mechanism, subsp. maritima being self-incompatible
while subsp. californica is self-compatible.

There seems to be a fairly close parallel between the situation in these two
species and that described by Imam & Allard (1965) in the wild oat, Avena fatua.
Like the poppies, A. fatua is an annual species and an opportunist colonizer
of bare ground and annual arable crops. It has frequently been reiterated that
success in such situations requires a high degree of adaptation, hence gene com-
binations conferring this are maintained by self-compatibility, rigorous inbreed-
ing and low recombination (Grant 1958, Baker 1959). While the premium for
the opportunist colonizer is on adaptations such as rapid germination and
establishment, a short vegetative phase and a high reproductive capacity, the
flexibility that is demanded of it is the ability to mature successfully regardless of
seasonal and climatic fluctuations, macro- and micro-edaphic variation and
interspecific competition. Some degree of heterozygosity would thus also appear
to be an advantage.

In Avena fatua this is achieved by a flexibility of the genetic system in which
inbreeding maintains a fairly high level of homozygosity but a low level of out-
crossing permits of recombination and diversity of genotypes. The resemblance
between P. dubium and A. fatua goes further, in that the degree of inbreeding in
P. dubium varies from one population to another as does the mean amount of
outcrossing in the Avena populations studied by Imam & Allard (1965).

It seems likely from the variation studies that the pattern of breeding behav-
iour in P. /ecogii is similar.

P. rhoeas on the other hand is rigorously outbreeding and there is no evidence
of any breakdown in its incompatibility mechanism. There is a high degree of
genetic heterozygosity which is obvious in characters such as petal colour, petal
blotch and stigmatic ray number. Such a species would seem to be less well
equipped than a self-compatible one to maintain those combinations of genes
conferring adaptive advantage. A priori, therefore, one might expect it to be less
successful as a colonizing species, but in reality it is the only one of the five
native species which, in the absence of the selective weedkiller, becomes a
major weed.

Rigorous outbreeding implies chromosomal heterozygosity but this need not
imply an absolute genic heterozygosity. Selection pressure against a less well
adapted recessive homozygote would tend to reduce the frequency of the reces-
sive allele in populations (MacArthur & Connell 1966) and would also favour
a low mutation rate from the dominant habitat-adapted allele. Most phenotypes
would consequently approximate to the optimum for such a character while
maintaining a much higher level of heterozygosity in those characters subjected
to less rigorous selection.

Compared with Avena fatua, the reproductive capacity of P. rhoeas is enormous
and the proportion of the seed output which produces the next generation of
plants is correspondingly small. There is thus an enormous surplus of genotypes
and the small proportion of obviously ill-adapted phenotypes that occurs in
natural populations is unlikely to have any significant effect on ultimate popula-
tion size.

(2) ECOLOGY

The genetic heterozygosity of P. rhoeas would appear to enable it to exploit a
much wider range of habitats than any of the other species. P. lecogii, P. arge-

STUDIES ON BRITISH POPPIES 275

mone and P. hybridum are confined to calcareous soils, and P. dubium (while
showing no pH restriction on light-textured soils) does not colonize heavy soils.
Both P. hybridum and P. lecoqii seem to be reaching the northern limit of their
distribution in the British Isles, scarcely extending beyond the southern half of
the country (Perring & Walters 1962, Perring & Sell 1968). Both have only a
short period of inherent seed dormancy and the seedlings are consequently
vulnerable to both winter mortality and destruction by autumn or early spring
cultivations. The other three species generally have a longer period of inherent
dormancy (Harper 1960), resulting in spring germination. This may allow their
further extension northwards into central and north-west Scotland and also into
Scandinavia (Mowat & Walters 1964). Little is known regarding seed longevity
as a factor in survival other than a reported viability of five years for P. rhoeas
(Brenchley & Warrington 1933).

(3) THE POPPIES AS WEEDS

The poppies show many of the characters of the ideal weed summarised by
Baker (1965). It seems unlikely that the ecological adaptability of P. rhoeas is
endowed by a general purpose genotype of the type he postulates as being
possessed by many successful weeds, since it has none of the necessary genetic
attributes — a history of hybridization, polyploidy, complex heterozygosity,
agamospermy or self-compatibility. The long-capsuled species, while apparently
having the more favourable genetic systems for the opportunist coloniser, do
not seem to have acquired the ideal combination of genes to turn them into
aggressive weeds under British conditions.

It would seem that there is no one ideal genetic system for the aggressive weed ;
some achieve success by means of rigorous inbreeding, some by a flexible mix-
ture of in- and outbreeding and at least a few by rigorous outbreeding coupled
with a high reproductive capacity.

A marked phenotypic plasticity is also a feature of the the poppies. It is
apparent in the variation found within individual plants (as in the capsule
samples) but also, more significantly, in the plastic response of a diversity of
different genotypes to density (Harper 1961). Such plasticity permits individual
response to the immediate situation. Presumably it is genetically controlled and
as such would be expected to have a strong selective advantage in unstable
environments.

ACKNOWLEDGMENTS

The work described in these papers formed part of a thesis accepted for the degree
of Ph.D. in the University of London. The help of Dr M. A. P. Madge of
Royal Holloway College is gratefully acknowledged.

REFERENCES

BAKER, H. G. (1953). Race formation and reproductive method in flowering plants. Symp.
Soc. exp. Biol., 7: 114-145.

BAKER, H. G. (1959). Reproductive methods as factors in speciation in flowering plants. Cold
Spring Harbor Symposium, 24: 117-190.

BAKER, H. G. (1965). Characteristics and modes of origin of weeds, in BAKER, H. G. & STEB-
BINS, G. L., ed. The genetics of colonizing species. New York and London.

FE

276 S. ROGERS

BRENCHLEY, W. & WARRINGTON, K. (1933). The weed seed population of arable soil. II. Influ-
ence of crop, soil, and methods of cultivation upon the relative abundance of viable seeds.
J. Ecol., 21: 103-127.

FABERGE, A. C. (1943). Genetics of the Scapiflora section of Papaver. II. The garden Iceland
poppy. J. Genet., 44: 366-385.

FEDDE, F. (1909). Papaveraceae, in ENGLER, A., ed. Das Pflanzenreich, vol. 40, pp. 366-385.

GRANT, V. (1958). The regulation of recombination in plants. Cold Spring Harbor Symposium,
23: 337-360.

Harper, J. L. (1960). Factors controlling plant numbers, in HARPER, J. L., ed. The Biology of
Weeds. Oxford.

Harper, J. L. (1961). Approaches to the study of plant competition. Symp. Soc. exp. Biol., 15:
1-39.

Imam, A. G. & ALLARD, R. W. (1965). Population studies in predominately self-pollinated
species. IV. Genetics, 51: 49-62.

MACARTHUR, R. & CONNELL, J. (1966). The biology of populations. London.

Mowat, A. B. & WALTERS, S. M. (1964). Papaver, in TuTIN, T. G., et alia, ed. Flora Europaea,
vol. 1. Cambridge.

NEWTON, W. C. F. (1929). The inheritance of flower colour in Papaver rhoeas and related forms.
J. Genet., 21: 389-404.

PERRING, F. H. & SELL, P. D., ed. (1968). Critical supplement to the Atlas of the British Flora.
London.

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

PHILP, J. (1933a). The genetics of Papaver commutatum and its hybrids with P. rhoeas. J. Genet.,
28: 169-174.

PHILP, J. (1933b). The genetics of Papaver rhoeas and related forms. J. Genet., 28: 175-203.

Rasmuson, H. (1920). Uber einige genetische Versuche mit Papaver rhoeas und Papaver
laevigatum. Hereditas, 1: 107-115.

TURRILL, W. B. (1946). British Plant Life. London.

WincE, O. (1932). Experiments with Papaver rhoeas f. strigosum Boenn. Trudy Lab. Genet., 9:
115-120.

Watsonia, 8, 277-288 (1971). QTT

A cytotaxonomic study of Lamiastrum galeobdolon
(L.) Ehrend. & Polatschek in Britain

S. WEGMULLER
Nidau, Switzerland

ABSTRACT

An account is given of the distribution and variation of the diploid and tetraploid subspecies
of Lamiastrum galeobdolon (L.) Ehrend. & Polatschek in Britain. The tetraploid subspecies is
the common plant; the diploid has been found in two localities only, both in N. Lincolnshire.
The taxonomic distinction between the two subspecies is shown to be difficult, and the discrim-
inatory value of several characters is considered in detail.

INTRODUCTION

Until recently, the Yellow Archangel was classified either as a species of the
Linnaean genus Lamium under the name Lamium galeobdolon (L.) Nath. or in the
genus Galeobdolon Adanson (1763) as G. luteum Hudson. Both of these treat-
ments are to be found in the numerous Floras of Britain. Lindley (1835),
Babington (1843), Irvine (1858) and the authors of the older county Floras
adopt the former treatment, while the latter course is followed by Hudson (1778),
Withering (1830), Deakin (1845), Clapham, Tutin & Warburg (1962) and the
authors of the more recent county Floras.

On the evidence of new research into Bentham’s Sectio IV Galeobdolon
(Adans.) Benth. (Bentham 1832-36, 1848), Polatschek (1966) came to the con-
clusion that the treatment of this plant as a monotypic genus was justified. He
pointed out that the generic name Galeobdolon was invalid, being a nomen super-
fiuum, since Heister in 1759 had validly published the name Lamiastrum in the
Enumeratio of Fabricius under the heading of the pre-Linnaean genus Galeobdolon
Dill. The new generic name Lamiastrum Heister ex Fabr. with the single species
Lamiastrum galeobdolon (L.) Ehrend. & Polatschek was used in the second
edition of the Excursion Flora of the British Isles by Clapham, Tutin & Warburg
(1968).

According to previous studies, and in particular to more recent cytotaxonomic
investigations in northern and central Europe (Turesson 1938, Polya 1949,
Gutermann 1962, Gadella & Kliphuis 1963, Dersch 1964, Polatschek 1966),
the collective species Lamiastrum galeobdolon includes at least three taxa, one
tetraploid and two diploid, which are defined as subsp. montanum (Pers.)
Ehrend. & Polatschek, subsp. galeobdolon and subsp. flavidum (Herm.) Ehrend.
& Polatschek respectively.

In Britain, the Yellow Archangel extends over wide areas of England and
Wales (Perring & Walters 1962). It is rare in Cornwall and near the west coast
of Wales and in the East it is absent from Breckland and Fenland. The northern
limit of its unbroken distribution is in a line from the Humber to Morecambe
Bay. Several isolated localities in southern Scotland recorded in the Flora of the
British Isles (Clapham, Tutin & Warburg 1962) are considered as ‘doubtfully

native’.

278 S. WEGMULLER

During a preliminary investigation carried out in the Botany School, Cam-
bridge, partly as a class exercise for the Part II Course in 1967 and 1968, some
British material of known wild origin was brought into cultivation and exam-
ined cytologically. Several tetraploids were found, and two samples, kindly
collected locally by Mrs M. and Miss A. Smith of Willoughby, near Alford,
Lincolnshire, were found to be diploid. This material provided the basis for the
present study, which aims to examine the different forms of the collective species
Lamiastrum galeobdolon in England and Wales, to determine their chromosome
numbers, to assess the extent of variation in the characters of the taxa, and to
establish whether fundamental morphological differences are related to the
ploidy level. During the summer of 1969 plants collected in England and Wales
were grown in the experimental area of the University Botanic Garden at
Cambridge. Chromosome counts were prepared as follows: the root-tips were
pretreated for 3 or 4 hours with a saturated solution of 1-Bromo-Naphthalene,
fixed in alcohol—acetic acid ‘glacial’ 3:1, hydrolyzed for 7-9 minutes in 10%
HCl at a temperature of 60°C, then stained in aceto-carmine and finally squashed.
The voucher specimens of the plants examined have been placed in the Herb-
arium of the Cambridge Botany School (CGE), and stocks of living material
are held in the University Botanic Garden, Cambridge.

TAXONOMY AND NOMENCLATURE

Persoon (1795) was the first to differentiate two Yellow Archangels when he
described alongside his Pollichia vulgaris a new species, Pollichia montanum.
Later, he regarded them as varieties (1807). Babington (1843) in his Manual of
British Botany under Lamium galeobdolon described two different taxa, these
being ‘Galeobdolon montanum Reich.’ and ‘Galeobdolon luteum Reich.’, but he
added that he had not seen the latter in Britain — ‘in our plant the lower leaves
are coarsely and even doubly serrate’. Reichenbach (1830-32) had mentioned
both taxa in his Flora germanica excursoria and referred especially to subsp.
montanum. Druce (1927) remarks correctly in The Flora of Oxfordshire: ‘Our
plant is not Galeobdolon luteum Reichb., but Galeobdolon montanum Reichb.,
with small bracts.’ On the whole, most authors of British Floras have not taken
into account the division of the species.

Turesson (1938) recognised two different ploidy levels in Lamiastrum galeob-
dolon and equated these with the two taxa. Dersch (1964) came to the conclu-
sion that the two taxa could only be given the rank of subspecies since variation
in different characters shows overlapping. Polatschek (1966) agreed with these
conclusions and clarified the nomenclature.

CHROMOSOME COUNTS

Numerous chromosome counts have been published for both subspecies from
central and northern Europe. These are as follows:

subsp. galeobdolon: 2n = 2x = 18

Turesson (1938), Southern Sweden and Latvia; Polya (1949), Hungary; Dersch
(1964), Germany.

subsp. montanum: 2n = 4x = 36

Jérgensen (1927), unlocalised; Turesson (1938), Munich; Gutermann (1962),
Southern Germany; Gadella & Kliphuis (1963), Netherlands; Dersch (1964),
Germany; Polatschek (1966), Austria and Czechoslovakia.

LAMIASTRUM GALEOBDOLON IN BRITAIN 279

Polatschek (1966) in a survey provides more detailed information about his
individual localities.

The chromosome counts (see Appendix) of plants from England and Wales
include both diploids and tetraploids and agree with those already published.

Figure | shows the localities of the plants examined. In spite of intensive investi-
gations, diploid plants could only be found in two localities, bothin north Lincoln-
shire. Plants from localities further west in the same vice-county are tetraploid.

Special attention was paid to the Lamiastrum localities in Surrey and West
Kent on greensand areas, since Britton (1926) mentions the two varieties grow-
ing side by side in Surrey. However Britton incorrectly interpreted Persoon’s

@ LAMIASTRUM
GALEOBDOLON
subsp. MONTANUM (4x)

O LAMIASTRUM

GALEOBDOLON 8} a x
b % z)

subsp. GALEOBDOLON (2x) /_ s
F 1D AXLE

MATERIAL fe (ge SS LA

CYTOLOGICALLY ’ p ZO

INVESTIGATED 7! —z Ly a as

KM: Sif A S

ic

FicureE 1. Distribution of cytologically investigated material of Lamiastrum galeobdolon in
England and Wales.

280 S. WEGMULLER

Stomatal length in ~m
WwW
ro)
ro)

1.0 1.5 2.0 25 3.0 3:5 40
Length/breadth/ratio of upper bracts

FicureE 2. Scatter diagram of stomatal length and the length/breadth ratio of the upper bracts.
Key

Lamiastrum galeobdolom subsp. galeobdolon (2x) ..

subsp. montanum (4x)

No. of flowers per whorl up to 8

more than 8

Margin of the upper bracts crenate or
crenate-serrate

serrate-crenate
serrate

Stolon leaf-base shape truncate ..
cordate

rounded or
shortly attenuate ..

7+ CO 00 OF

taxa, and the material collected in this area proved to be uniformly tetraploid.

When collecting the specimens, special attention was also paid to plants with
intermediate characters, but no triploid hybrids could be identified. This observ-
ation agrees with the findings of Dersch (1964).

MORPHOLOGY

Plants of the two different ploidy levels were examined carefully, concentrating
on the following morphological characters: number of whorls per inflorescence;
number of flowers per whorl; shape and margin of bracts and stolon leaves;

LAMIASTRUM GALEOBDOLON IN BRITAIN 281

hairiness of the stem; size of flowers; length of pollen and stomata. An attempt
was made to establish as far as possible the degree of variation of the individual
characters. Tables 1-3 show, that as in the results of Dersch (1964), single charac-
ters cannot be used to distinguish the two subspecies.

TABLE 1. INFLORESCENCE CHARACTERS

Number of whorls per Maximum number of
inflorescence flowers per whorl
5 or 5 or 7 or more
Z 3 4. more 6 8 than 8

subsp. galeobdolon | 3-7 59-3 22 14-8 51-9 44-9 3°7
me = 1S

subsp. montanum 2:7 15-2 33-9 48:2 8-9 14:3 76°8
27,— 4x = 36

Values expressed as a percentage of 27 plants from 5 populations of subsp. galeob-
dolon and of 112 plants from 63 populations of subsp. montanum. All material
collected in the field.

POLLEN SIZE
The average pollen-length derived from 100 measurements of 12 plants of both
taxa gave the following results:

subsp. galeobdolon (2x) 32°31 — 34:82 um

subsp. montanum (4x) 33-46 — 38-35 um

A separation of the two subspecies on the basis of the pollen size is not poss-
ible, a result which agrees with that of Polatschek (1966).

STOMATAL LENGTH

The lengths of 50 stomata from the uppermost pair of stem leaves and the lowest
pair of bracts were measured and the averages calculated.

subsp. galeobdolon subsp. montanum
Number of plants examined 30 60
Number of averages (nj, n.) 30 60
Range of averages 23°48 — 29-66 um 28-77 — 37:59 um
Means (x, Xs) 26-118 »m 31-237 pm
Variance (c,” , o,”) 2-583 2-293
Standard deviation (6}, o,) 1-607 1-514
Range within o 24-51 — 27-72 pm 29-72 — 32:75 um
Range within 2c 22-90 — 29-33 um 28:21 — 34-26 um
Difference of the means (X. — X;) 5-119 um
Standard deviation of the

difference of means (6,) 0-352

The difference between the two means is significant.

The averages of stomatal length have been used along with the length/breadth
ratio of upper bracts (Table 2) to construct a scatter diagram (Fig. 2).

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LAMIASTRUM GALEOBDOLON IN BRITAIN

283

The morphology of the two subspecies is compared in the following Table.

TABLE 4. COMPARISON OF THE CHARACTERS OF BOTH
SUBSPECIES IN BRITAIN

habit

hairiness of the stem

stolon leaves:
shape

margin

upper bracts:
shape

margin

number of whorls per
inflorescence

number of flowers per
whorl

length of flowers

(measured from the base

of calyx)

stomatal length
(see p. 281)

subsp. galeobdolon (2x)

plant usually 15-45 cm

stem, especially at the base,
with deflexed, appressed
hairs mainly on the angles

orbicular or ovate; base
truncate or cordate

regularly crenate or crenate-
serrate; terminal tooth
rounded

ovate; 1/b ratio 1-0—2-0

regularly crenate or bicrenate
or serrate; often deeply
incised

2-4 (5)
up to 8

17-21 mm

22-9-29-3 pm

subsp. montanum (4x)

plant, more vigorous
20-60 cm

stem, especially at the base,
with deflexed scattered hairs
on the flat areas and on the
angles

ovate, rarely orbicular;
base shortly attenuate,
rounded or cordate

irregularly serrate-crenate
to serrate; terminal tooth
acute

narrowly or broadly lance-
olate, rarely ovate; 1/b ratio
(1 -:2—)1-8-3-5

serrate or biserrate or
serrate-crenate; teeth
usually widely separated ;
terminal tooth long and
acute

(3) 4-7(-10)
more than 8
18-25 mm

28-2-34-3 um

Many intermediate forms occur (Fig. 3). Poorly developed plants from tetra-
ploid populations can often look very similar to the diploid plants, especially
when the bracts are crenate. On the other hand, diploid plants with sharply
serrate subtending bracts can resemble tetraploids. Thus, although a positive
decision on the basis of a combination of characters is usually possible there
exist cases which demand measurements of stomatal length or even an analysis

of chromosome numbers for a precise identification.

284 S. WEGMULLER

FIGURE 3. Variation in shape and marginal teeth of upper bracts.

A. Range of examples of subsp. galeobdolon.
B. Range of examples of subsp. montanum.

HABITAT

Lamiastrum galeobdolon sensu lato is a plant of woods and hedges. Subsp. mon-
tanum favours rich, deep soils above a great variety of underlying rocks. It is
widespread in Britain in beech woods, especially over limestone, and in oak-ash
woods, where it is frequently found in association with Mercurialis perennis,
Endymion non-scriptus, Allium ursinum, Anemone nemorosa, Arum maculatum,
Primula vulgaris, Sanicula europaea, Ajuga reptans, Asperula odorata and Cir-
caea lutetiana. It is also able to grow on moderately acid soils in Castanea sativa
coppices and sometimes colonizes disturbed places. In woods where subsp. mon-
tanum is rare, it is usually found at the eastern or northern margin.

Subsp. galeobdolon also flourishes on deep soils in north Lincolnshire. The
few populations studied do not allow valid assertions as to its ecological re-
quirements. Therefore, only a few of the observations made on these popula-
tions are given here. The plants of the different populations had already flowered
by May 13th and had almost ceased to bloom by May 3lst. By the beginning of
July, two populations were found to be dying off, as they were completely
overgrown, by Pteridium aquilinum and Epilobium angustifolium in one case at
the edge of the wood and by Rubus in the other. It appears that, on account of
their early and quick development, the plants of subsp. galeobdolon are able to
occupy habitats which later support other species. Turesson (1930) established
that subsp. galeobdolon flowers earlier than subsp. montanum.

DISTRIBUTION

In the Herbaria of the British Museum (Natural History) (BM) in London and
of the Royal Botanic Gardens, Kew (K), specimens of both subspecies have been
seen from the following countries:

LAMIASTRUM GALEOBDOLON IN BRITAIN 285

subsp. galeobdolon: southern Sweden; southern Finland; Estonia; western
European Russia; Poland; Denmark; central Germany;
Czechoslovakia; Hungary, Ukraine; Romania; Yugo-
slavia; Bulgaria; Greece (Macedonia) and European
Turkey.

subsp. montanum: central Germany; Holland; Belgium; northern, central,
western and south-west France (Pyrenees); north-east
Spain (Pyrenees); Switzerland; Austria; Czechoslovakia;
southern Poland; Hungary; Yugoslavia (Bosnia, Herze-
govina); Greece (Macedonia) and Italy (Apennines).

In Britain, subsp. galeobdolon could only be found in north Lincolnshire.
This distribution in eastern England recalls that of a few other species occurring
both in Scandinavia and Continental Europe, e.g. Maianthemum bifolium (L.)
Schmidt. Subsp. montanum is the widely distributed Yellow Archangel of
Britain.

LAMIASTRUM GALEOBDOLON SUBSP. FLAVIDUM

This Yellow Archangel without stolons was described by Hermann (1958) and
later cytotaxonomically examined by Gutermann (1962), Dersch (1964) and
Polatschek (1966). The plant is diploid, the number of chromosomes being
2n=2x=18. For purpose of comparison two plants of subsp. flavidum were
grown in the University Botanic Garden at Cambridge and examined (see
Appendix). The chromosome number was confirmed on this material. The
following stomatal and pollen measurements were obtained:

Pollen diameter Stomatal length
Coll. No. (n=200) (n=200)
149-67 31-84 pm 24-93 wm
246A-67 32-30 um 25-75 wm

According to Polatschek, pollen measurements enable the separation of the
diploid subsp. flavidum and the tetraploid subsp. montanum. Measurements of
stomatal length produced averages which lie within the range of subsp. galeob-
dolon.

The typical characters of this subspecies are as follows: pliant without stolons,
bracts ovate to lanceolate, sharply serrate or biserrate, some of the teeth promin-
ently curved, the lowest bracts more than twice as long as the internode below,
number of flowers in a whorl more than 10, length of flowers 14-17 mm. Subsp.
flavidum colonizes scree slopes and other open habitats from the montane up to
the subalpine level. The distribution area extends from the southern Central Alps
to the Eastern Alps, and south to the Apennines and the northern mountains of
Yugoslavia (Polatschek 1966). In the Herbaria (BM, K) mentioned above,
plants of subsp. flavidum have been seen from the following localities:

Central Alps Italy Valle di Formazza
Austria Gschnitztal
Eastern Alps Austria K4arnten, Steiermark

Yugoslavia Herzegovina

286 S. WEGMULLER

CROSSINGS

Dersch (1964) pointed to the possibility that subsp. montanum could be con-
sidered as an allotetraploid descendant of subsp. galeobdolon and subsp. flavidum,
since it possesses characters of both subspecies. The presence of stolons and the
greater length of flowers are perhaps from subsp. galeobdolon, the number of
whorls, the number of flowers per whorl and also the prominent curved
teeth of the bracts could be derived from subsp. flavidum. Polatschek (1966)
supported this hypothesis by his scatter diagram, in which he revealed the inter-
mediate position of subsp. montanum. The range of subsp. montanum lies
geographically and altitudinally for the most part between the areas of subsp.
galeobdolon and subsp. flavidum.

In the course of the summer of 1969 crossings were made in the Cambridge
Botanic Garden between the two diploid subspecies and also the tetraploid,
but only a few seeds were produced. Germination tests carried out have so far
been unsuccessful. Even seeds pre-treated under refrigeration did not germinate,
so that no results are yet available.

CONCLUSIONS

1. A cytotaxonomic study of the collective species Lamiastrum galeobdolon in
England and Wales showed two subspecies with different ploidy levels.

2. The previously published chromosome numbers for subsp. galeobdolon
(2n=2x=18) and subsp. montanum (2n=4x=36) were confirmed. Triploid
hybrids were not found.

3. A separation of the subspecies is possible by a combination of characters,
including averages of stomatal length.

4. Subsp. galeobdolon was only found in north Lincolnshire. Subsp. montanum
is the common Yellow Archangel in Britain.

ACKNOWLEDGMENTS

This present study was made during a six months study period from April to
September 1969 at the Botany School of the University of Cambridge. I am
grateful to Professor P. W. Brian for providing laboratory facilities. 1 am greatly
indebted to Dr S. M. Walters, who suggested this topic and encouraged me by
his lively interest. With N. G. Marchant I have had many discussions on taxo-
nomic problems and methods. For his helpful comments on the manuscript I
am most grateful. I should also like to thank those people who have kindly col-
lected living specimens and the Biological Records Centre, Monks Wood Experi-
mental Station, Abbots Ripton, Huntingdon for preparing Fig. 1 from my data.
My thanks go also to the Swiss National Fund, which by a financial grant made
this period of study possible.

REFERENCES

ADANSON, M. (1763). Familles des Plantes. Paris.

BABINGTON, C. C. (1843). Manual of British Botany. London.

BENTHAM, G. (1832-1836). Labiatarum Genera et Species. London.

BENTHAM, G. (1848). Labiatae, in DE CANDOLLE, A. P. Prodromus Systematis Naturalis... ,
vol. 12, pp. 27-603. Paris.

LAMIASTRUM GALEOBDOLON IN BRITAIN 287

BRITTON, C. E. (1926). Notes on some minor varieties of British plants. J. Bot., Lond., 64:
327-328.

CLAPHAM, A. R., TuTIN, T. G. & WARBURG, E. F. (1962). Flora of the British Isles, 2nd ed.
Cambridge.

CLAPHAM, A. R., TuTIN, T. G. & WARBURG, E. F. (1968). Excursion Flora of the British Isles,
2nd ed. Cambridge.

DEAKIN, R. (1845). Florigraphia Britannica. London.

Derscu, G. (1964). Zur Cytologie und Taxonomie der Goldnessel (Lamium galeobdolon (L.)
L.). Ber. dt. bot. Ges., 76: 351-359.

Druce, G. C. (1927). The Flora of Oxfordshire. Oxford.

GADELLA, T. W. J. & KLIPHuIs, E. (1963). Chromosome numbers of flowering plants in the
Netherlands. Acta bot. neerl., 12: 195-230.

GUTERMANN, W. (1962). Diploides Lamium galeobdolon (s./.) in Bayern (vorlaufige Mitteilung).
Ber. bayer. bot. Ges., 35: 43-45.

HERMANN, F. (1958). Lamium flavidum nov. spec. in den Bayerischen Alpen. Ber. bayer. bot.
Ges., 32: 145-146.

Hupson, W. (1778). Flora Anglica, 2nd ed. London.

Irving, A. (1858). The illustrated handbook of the British Plants. London.

JORGENSEN, C. A. (1927). Cytological and experimental studies in the genus Lamium. Hereditas
9: 126-136.

LINDLEY, J. (1835). A Synopsis of The British Flora. London.

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

Persoon, C. H. (1795). Botanische Beobachtungen aus einem Brief des Hr. Persoon an den
Herausgeber. Ann. Bot. (Usteri), 14: 33-39.

Persoon, C. H. (1807). Synopsis plantarum, vol. 2. Paris.

POLATSCHEK, A. (1966). Cytotaxonomische Beitrage zur Flora der Ostalpenlander, 2. Ost. bot.
Z., 113: 102-112.

Po.yA, L. (1949). Chromosome numbers of some Hungarian plants. Acta Geobot. Hung., 6:
124-137.

REICHENBACH, H. G. L. (1830-1832). Flora germanica excursoria. Leipzig.

TURESSON, G. (1930). The selective effect of climate upon the plant species. Hereditas, 14: 110-
Weil

TURESSON, G. (1938). Chromosome stability in Linnean species. LantbrHégsk. Annlr, 5:
405-416.

WITHERING, W. (1830). A Systematic Arrangement of British Plants. London.

APPENDIX

ORIGIN OF MATERIAL

Lamiastrum galeobdolon (L.) Ehrend. & Polatschek
subsp. galeobdolon 2n = 18
v.c. 54 North Lincoln

Welton High Wood

Population 1, mixed Quercus robur and GR 53/454699 No. 31A-69
Acer pseudoplatanus wood No. 31B-69

Population 2, mixed forest, under Acer
pseudoplatanus GR 53/455700 No. 32A-69
Population 3, edge of wood under Preridium GR 53/456700 No. 33A-69
aquilinum and Epilobium angustifolium No. 33C—69
Population 4, mixed forest, by track GR 53/455701 No. 138H-68
Burwell — Muckton, roadside GR 53/370801 No. 71C—69

subsp. flavidum (Herm.) Ehrend. & Polatschek 2n = 18
Austria

Karnten, Pl6ckenpass (‘locus classicus’), cultivated in the No. 149-67

Botanic Garden at Munich

288 S. WEGMULLER

Yugoslavia
Slovenija, Kamniske Alpe, limestone scrub c 1000 m No. 246A-67
on east side of Krvavec

subsp. montanum (Pers.) Ehrend. & Polatschek 2n = 36

The chromosomes of 72 specimens of this subspecies, all from England and Wales,
have been counted. Their distribution from the following vice-counties is shown in
Fig. 1.

Vice-counties 2, 3, 6, 8, 9, 12, 15-17, 19, 20, 22-34, 39-41, 44, 47, 50, 51, 53, 54, 57,
59, 60, 62

Details of these localities have been given to the Biological Records Centre, Monks
Wood Experimental Station, Abbots Ripton, Huntingdon.

Watsonia, 8, 289-291 (1971). 289

Frequencies of pin and thrum plants in a wild population
of the Cowslip, Primula veris L.

D. R. LEES
Genetics Laboratory, Department of Zoology, Oxford

ABSTRACT

The heterostylous dimorphism in the genus Primula is discussed. A population of the cowslip
Primula veris on stable duneland at Strathy in Sutherland contained an excess of the thrum
form. It is suggested that this may be due to an advantage of the heterozygous thrums over
pins in some microclimates.

The heterostylous dimorphism in the genus Primula is well known. Two types of
flower exist, always borne on separate plants. In the ‘pin’ form, the anthers are
placed half-way down the corolla-tube, the mouth of which is occupied by the
stigma, borne on the end of a long style. In the ‘thrum’ flowers, the positions of
the sexual organs are reversed, the anthers being at the top of the corolla-tube
while the stigma is half-way down it, since the style is relatively short.

Darwin (1877) recognised that this dimorphism mechanically favours out-
breeding since insects tend to transfer pollen to stigmas at the same level as the
stamens. Yet more important in achieving outbreeding in this genus are the
physiological distinctions between the pin and thrum forms, which result in
pollen being largely incompatible on its own kind of stigma. The basis of this
incompatibility is differential pollen-tube growth (Tseng 1937, Lewis 1942).

Pollen from thrum anthers can germinate on a thrum stigma but rarely
penetrates its surface, so that fertilisation is not achieved. As Ford (1964) has
pointed out, this incompatibility occurs whether the pollen is genetically pin
or thrum and is an example of Mendelian inheritance delayed one generation.
Pin pollen readily germinates on and penetrates the surface of a pin stigma, but
pollen-tube growth is then slow compared with that of the thrum form, which
in these circumstances will always reach the ovules first. Thus of the two
‘illegitimate’ crosses, thrum x thrum probably never succeeds in wild popula-
tions while pin x pin regularly does so, but only in the absence of thrum pollen.

The genetic control of the pin/thrum heterostyly in Primula is by means of a
supergene the components of which control both physiological and structural
characteristics of pin and thrum flowers (Ernst 1928). The various loci compris-
ing this supergene are closely linked and normally behave as if they were a pair
of alleles (S and s), with the thrum character dominant to pin (Bateson &
Gregory 1905).

As a result of the incompatibility mechanisms mentioned above, thrum homo-
zygotes probably never occur and pin selfing is restricted to special circumstances
(where thrum plants are absent). Thus the breeding system gives rise to a series
of backcrosses (Ss x ss) with pin and thrum seed being produced in approx-
imate equality. In wild populations of Primula vulgaris Huds., the species most
studied in the genus, equality of pin and thrum plants is seldom found and a
slight excess of pin is the norm, probably owing to its capacity for self-fertilisa-

290

D. R. LEES

tion (Crosby 1949, Ford 1964). Excesses of thrums have however also been
found (Haldane 1938).

The report presented here concerns a large population of the cowslip, Primula
veris L., from stable duneland at Strathy in Sutherland. Four samples of plants
were collected along a transect running down the SW. side of a SE./NW. cleft
in the dunes. Its length was about 70 yards, dropping 60 feet from the top sample
A to the bottom sample D. Care was taken to collect only one flower per plant.
Details of the results are given in Table 1.

TABLE 1.

Site

A

DETAILS OF SAMPLING AREAS AND OF SAMPLES OF

PRIMULA VERIS OBTAINED FROM STRATHY, SUTHERLAND

Approx.
slope Associated plant species Pin
of dunes

205 Agropyron junceiforme 60
(A. & D. Love) A. & D. Love
Bellis perennis L.
Festuca rubra L.
Plantago lanceolata L.
Thalictrum minus L.
Ditrichum flexicaule (Schwaegr.)
Hampe

30° Agropyron junceiforme 67
(A. & D. Love) A. & D. Love
Bellis perennis L.
Festuca rubra L.
Galium verum L.
Heracleum sphondylium L.

40° Plantago lanceolata L. 44

Vicia sepium L.

Hylocomium splendens (Hedw.)
LB Ser Mee

Pseudoscleropodium purum (Hedw.)
Fleisch.

Rhytidiadelphus squarrosus (Hedw.)
Warnst.

<a Achillea millefolium L. 80
Agropyron junceiforme
(A. & D. Love) A. & D. Love
Agrostis stolonifera L.
Centaurea nigra L.
Festuca rubra L.
Heracleum sphondylium L.
Plantago lanceolata L.
Sanguisorba officinalis L.
Tussilago farfara L.
Vicia sepium L.
Equisetum arvense L.
Hylocomium splendens (Hedw.)
B., S. & G.

Thrum Total
93 153
71 138
75 119
81 161

7%

Thrum

60°8

50°3

PIN AND THRUM PLANTS IN A WILD POPULATION OF THE COWSLIP 291

The samples are homogeneous (x73, = 7:04, P > 0-05) and show a significant
departure from the expected equality of pin and thrum, there being an excess of
thrum in all of them (x74) = 8:32, P < 0-01). Style-length in the pin flowers
was variable throughout and in some the style was so short that these were
initially scored as short homostyles.

A wild sample of 522 Primula veris plants collected by Darwin (1877) contained
281 thrums and 241 pins, but he does not mention the locality where they were
found. It is possible that the excess of thrums in this population and in that at
Strathy is due to an advantage of the heterozygous thrums over pins. The
relatively large excess of thrums in areas A and C at Strathy could reflect an
increased advantage of the heterozygotes in the microclimates of these areas;
both areas A and C had a sparse cover of vegetation whereas B and D had a
thicker cover. Fixed dunes in general tend to have an extreme and locally very
variable climate (Salisbury 1952). In a quite different organism, the polymorphic
land snail Cepaea nemoralis L., rapid changes in the frequency of colour morphs
over short distances have been recorded on duneland (Cain 1968). This has been
related to its complex topography, producing microclimates (e.g. cold hollows)
within which some morphs are better able to survive than others.

An attempt will be made to sample cowslip populations from as many areas
of Great Britain as possible to establish whether an excess of thrum plants is
normal in wild populations. Such a survey might also show if wild homostyle
colonies exist in the cowslip, as in the primrose.

ACKNOWLEDGMENTS

My thanks are due to Professor E. B. Ford, F.R.S., for reading this paper in
typescript and making many helpful suggestions, and to Dr J. G. Duckett for
his help in identifying flowering plants and bryophytes from Strathy.

REFERENCES

BATESON, W. & GREGORY, R. P. (1905). On the inheritance of heterostylism in Primula. Proc.
R. Soc., Series B, 76: 581-586.

CaIn, A. J. (1968). Studies on Cepaea. V. Sand-dune populations of Cepaea nemoralis (L.).
Phil. Trans. R. Soc. Series B, 253: 499-517.

Crossy, J. L. (1949). Selection of an unfavourable gene complex. Evolution, 3: 212-230.

Darwin, C. (1877). The different forms of flowers on plants of the same species. London.

Ernst, A. (1928). Zur Vererbung der morphologischen Heterostyliemerkmale. Ber. dt. bot.
Ges., 46: 573-588.

Forp, E. B. (1964). Ecological Genetics. London.

HALDANE, J. B. S. (1938). Heterostylism in natural populations of the primrose Primula acaulis.
Biometrika, 30: 196-198.

Lewis, D. (1942). The physiology of incompatibility in plants. I. The effect of temperature.
Proc. R. Soc., Series B, 131: 13-26.

SALISBURY, E. J. (1952). Downs and Dunes. Their plant life and its environment. London.

TSENG, T. (1937). Pollen tube competition in Primula sinensis. J. Genet., 35: 289-300.

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Watsonia, 8, 293-301 (1971). 293

Short Notes

69. RHYNCHOSINAPIS — The Worcestershire records.

From 1873 to 1876 a species first identified as Brassica cheiranthos Vill. was found
‘in considerable plenty’ at Sutton Common, Kidderminster by J. Fraser, J. H. Thompson
and W. Mathews (Rea 1897). The plant was later identified by Mathews (1899) as B.
monensis L. (Rhynchosinapis monensis (L.) Dandy). The record and identification were
accepted by Amphlett & Rea (1909), Druce (1932) and Perring & Walters (1962).

In 1965 I examined the material in the herbarium of the Hastings Museum, Worcester
(WOS). The specimens, collected at Sutton Common in June and July 1873, October
1874 and June 1876, were in a poor state of preservation, but the following character-
istics could be ascertained: Stems branched, with a few simple setae on their lower
parts; simple hairs present on petioles and basal leaves and on the sepals, which slightly
exceed the pedicels in length; cauline leaves with segments narrower than those of
rosette leaves; siliquae glabrous, 6:5—9-5 cm long, with beaks 11 mm without seeds.

The specimens are, therefore, of the alien species Rhynchosinapis cheiranthos (Vill.)
Dandy and not of the endemic R. monensis. It has already been pointed out (Kiernan
1966) that the material bears no resemblance to Sisymbrium orientale L. (S. columnae
Jacq.), to which species it was referred by Rea (1921).

In view of these findings, it is possible that other inland records for R. monensis
might be erroneous.

REFERENCES

AMPHLETT, J. & REA, C. (1909). The Botany of Worcestershire. Birmingham.

Druce, G. C. (1932). The Comital Flora of the British Isles. Arbroath.

KIERNAN, J. A. (1966). Notes on the Flora of Worcestershire. I. Additions to the flora since
1909. Proc. Bgham nat. Hist. phil. Soc., 20: 19-37.

MATHEws, W. (1899). List of Midland County plants 1849-1884. Trans. Worcs. Nat. Club, 2:
55-102.

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

REA, C., ed. (1897). Trans. Worcs. Nat. Club, 1: 217, 225, 226.

REA, C. (1921). Appendix to the Botany of Worcestershire. Trans. Worcs. Nat. Club, 7 (Suppl.):
ile

J. A. KIERNAN

115/15. HyPERICUM CANADENSE L. — In England.

In a note in a recent issue of Watsonia (8: 90, (1970)) R. D. Meikle suggests that this
species is most probably a recent arrival in Europe. Further localities in the vicinity
of the original one by Lough Mask may indicate that it is spreading there, and the
discovery within such a short time of scattered localities on the Continent as well as
Ireland is consistent with the plant being a recent introduction. The Eccles Hotel at
Glengariff, Co. Cork is frequented by botanists and it is possible that the seeds of
H. canadense were conveyed there on footwear or the tyres of a car which had travelled
from Lough Mask.

It has now been found in England. In 1967 Mr Roy Lancaster sent me a specimen
found as a weed in a seed-bed in Hillier & Sons’ nursery at Ampfield, S. Hants., v.c. 11,
which has now been confirmed by N. Robson (Herb. Lousley). It grew in ground sown
earlier in the year with seed of Cedrus atlantica from an Italian source, but H. canadense
is not known in Italy and this is unlikely to have been its origin. It did not persist.
However it was conveyed to the nursery at Ampfield and this would seem further
evidence that human agency is involved.

J. E. LOUSLEY

294 SHORT NOTES

237/2. CRASSULA AQUATICA (L.) Schonl. — New to Scotland.

Crassula aquatica, hitherto recorded in Britain only from Adel Dam, Yorkshire, and
now apparently extinct there, was found, on August 21st 1969, growing on mud over-
lying stones by fresh water in v.c. 97, Westerness. The site was obviously subject to
periodic inundation and the plant, which, on the date of discovery, showed both
unopened flowers and fruit in the form of four neatly arranged little carpels in the leaf
axils, occupied a strip about six yards by one yard, parallel to the water.

Associated plants were very few and scattered: Juncus articulatus, Juncus bufonius
and Polygonum sp. were also on the mud, while Litorella uniflora and Alisma plantago-
aquatica grew in the adjacent water.

The status of this species in Britain is obviously still under the doubt indicated in
Flora of the British Isles (Clapham, Tutin & Warburg 1962). It occurs in Iceland and
Norway and its tiny seeds seem more fitted than most to arrive with mud on migrant
waterfowl. Nonetheless Scotland fits neatly into its continental northern distribution
and the lochs and rivers of north-west Scotland must offer hundreds of miles of under-
botanised shoreline where this inconspicuous annual may have found a suitable
habitat.

The plant was collected for identification without its uniqueness being realised and
thanks are due to Mary McCallum-Webster, who paid a later visit to the site, for con-
firming and amplifying some of the original observations.

Specimens are now in the herbaria of the Royal Botanic Garden, Edinburgh, the
British Museum, Kew and Cambridge.

ALAN J. SOUTER

NAUFRAGA BALEARICA Constance & Cannon — Refound.

Work on Flora Europaea (Tutin, T. G. et al. 1964, 1968) has encouraged the study in
detail of some of the botanically lesser-known areas of Europe and, perhaps surpris-
ingly, the Balearic Islands have been found to be of special interest. In fact the rapid
spread of tourism and the resulting urbanisation have underlined the need for study,
pointing the lack of readily available information and the loss of valuable, unexplored
land to building.

The proof of the interest of the Balearic flora was confirmed by the discovery in
1962 of a genus new to science. On 22 July 1962, Professor J. Duvigneaud found at
Cala San Vicente, Mallorca, a plant which he did not recognise. He was still unable to
name it when he returned to Belgium and therefore he sent it to Professor T. G.
Tutin, Chairman of the Editorial Committee of Flora Europaea, who forwarded it to
Mr J. F. M. Cannon, an acknowledged expert on European Umbelliferae.

Mr Cannon investigated the plant thoroughly and then consulted Professor Lincoln
C. Constance, an expert on American Umbelliferae; together they decided the plant
was representative of a genus new to science and it was described and named Naufraga
balearica (Constance & Cannon 1967), the description being based on this one collec-
tion. It was noted as having affinities with New Zealand and South American Umbelli-
ferae; this apparently startling relationship however was noted for other Balearic
plants by Dr H. Knoche (Knoche 1921) fifty years ago.

In 1966 I went to Mallorca to collect specimens for the British Museum (Natural
History) and to study the flora in the field in connection with work I have undertaken
on Balearic plants. In relation to this, Naufraga had little importance; its habitat and
locality were known, it had been reliably determined and its nomenclature was uncom-
plicated. But its curiosity value and the need to have further material available to
herbarium botanists made it worth collecting and a desultory search was made for it,
but it was not found.

During 1967 and 1968 four other British botanists joined the search and in 1968 my
husband and I made an intensive search but also failed. The affinities of Naufraga and
its elusiveness were now increasing its importance.

SHORT NOTES 295

In 1969 a series of errors and confusions resulted in another holiday being spent in
the Balearic Islands, and at Cala San Vicente. The proximity of the Naufraga locality
led inevitably to another attempt to find it. Careful re-appraisal of the maps and the
lure of the shaded, north-facing cliffs of Punta des Covas Blancas, with masses of
Laurentia tenella (Biv.) A.DC. and Erodium reichardii (Murray) DC., resulted in a
scrambled climb around the shoulder of the cliffs and down a steep talus slope which
ended in a 300 foot drop into the sea. Here Naufraga balearica was growing in pro-
fusion on fine talus in an almost inaccessible habitat.

Material was collected and prepared for a number of herbaria, and a living plant was
sent to the Royal Botanic Gardens, Kew, where it is now in cultivation. A chromosome
count has been made and further cytological studies will be undertaken.

In Mallorca the plant may well be more widespread on shaded, precipitous maritime
cliffs on the north-west of the island ; but attempts to study the cliff faces from land were
abortive and approach from the sea seems to be the only practical way of investigating
these areas successfully.

The type specimen, second collection, cultivated plant, maps and photographs of
the type-locality were shown at the Exhibition Meeting of the Society on 29 November
1969.

REFERENCES

CONSTANCE, L. C. & CANNON, J. F. M. (1967). Naufraga...a new genus of Umbelliferae
from Mallorca. Reprium nov. Spec. Regni veg., 74: 1-4.

KNocHE, H. (1921-1923). Flora Balearica. Montpellier.

TuTin, T. G., et alia, ed. (1964, 1968). Flora Europaea, vols. 1 & 2. Cambridge.

L. F. FERGUSON

590/1. MAIANTHEMUM BIFOLIUM (L.) Schmidt — Its status in the British Isles.

It is strange that such a conspicuous plant as the May Lily was not recorded for Norfolk
until 1955 (Petch & Swann 1962), when it was found in a wood at Swanton Novers.
Dr Francis Rose later estimated the colony to contain 50,000 plants ‘certainly native
in this ancient relic of old forest’ (in lit.). Its abundance suggests it must not only be
long-established but also the largest colony in the British Isles. Both in the work cited
and in their recent Flora of Norfolk (1968) these authors have considered the species
to be doubtfully native.

Although it is often difficult to distinguish with any certainty between natural and
semi-natural woodland, there is no doubt that the wood at Swanton Novers is a relic
of old forest, with records of management going back to the early thirteenth century.
Today it still has good stands of deciduous trees including Tilia cordata and the native
oak with hazel scrub but the inevitable planting of the more rapidly maturing conifers is
being carried out.

Although it could be said that the abundance of the May Lily gives the appearance
of it being a native plant, neither Smith (1828) nor Stillingfleet (1761), reputable and
reliable recorders of the Norfolk flora, make any reference to the species.

The plant is well adapted to spreading, both by reason of its conspicuous red berry
containing one or two seeds and also its slender rhizomes. According to Ridley (1930)
its fruit forms part of the food of the magpie, a by no means uncommon bird in this
county. Although many suitable habitats abound in Norfolk, it remains exclusive to
Swanton Novers. For these reasons it is probably more accurate to regard it as one of
the many naturalised aliens, ‘introduced species which are naturalised in natural or
semi-natural habitats’ (Lousley 1953).

Elsewhere in the British Isles, it has had a somewhat chequered history. A note
possibly by Newman in The Phytologist (1: 520 (1843)) states that Loudon gives the

296 SHORT NOTES

year 1596 as the date of its introduction into Britain. Jackson (1913) gives the history
of the plant in England and he considered its Continental distribution is entirely in
favour of its being a native species in the south of England. His conclusion is somewhat
invalidated by the fact that of the eight counties mentioned, namely Lancashire,
Northumberland, Yorkshire, Lincolnshire, Durham, Bedfordshire, Middlesex and
Oxfordshire, the majority cannot be regarded as southern English counties.

Its past and present distribution given by the Atlas of the British Flora (Perring &
Walters 1962), supplemented by information kindly provided by Mr Douglas Kent
and Dr Perring, is as follows:

v.c. 21, Middlesex. ‘Maianthemum was recorded first by Hunter, steward at Caen
Wood Towers, from Caen (Ken) Wood in J. J. Park’s Topography and Natural History
of Hampstead (1813). I have seen specimens collected there in 1829, 1840, 1845 and
1861. Trimen and Dyer in their Flora of Middlesex (1869, p. 277) refer to ‘“‘a patch of
about twenty square yards on an eminence under the shade of a very large beech in
the enclosure of the Ken Wood grounds near its south-east angle.’”’. Jackson and H.
Boyd Watt recorded it as plentiful in 1912, and from 1917 it was observed annually
by H. S. Redgrove; a good patch in flower 1917; scarce in 1922 and did not flower; a
single plant which did not flower, 1924. Redgrove thought the severe drought in 1921
was responsible for its extinction: other people have claimed it was destroyed by path-
making. It has been variously claimed as a native or denizen — my view is certainly
the latter, and most certainly an introduction. In 1933 it was reintroduced at Ken
Wood by J. S. L. Gilmour (J. Bot., Lond., 71: 168) and in 1945 was reported by R.S. R.
Fitter as naturalised and increasing. I saw it there several times after the war — the
last time in 1953’ (Kent, in Jit.).

v.c. 23, Oxfordshire. There is a specimen in the Oxford Herbarium collected by Druce
in June, 1892, and an unconfirmed record of 1900 between Thame and Wheatley.
v.c. 30, Bedfordshire. In 1833 a doubtful record from Apsley Wood, but Dony (1952)
was unable to confirm this.

v.c. 54, N. Lincolnshire. Recorded in 1895 at Turnby (Tumby), where it still persists.
Jackson (1913) suggests it is introduced.

v.c. 59, S. Lancashire. Gerard (1597) records it from Dinckley Wood, Aundernesse and
Harwood. Savidge, Heywood & Gordon (1963) state it is extinct.

v.c. 62, N. E. Yorkshire. According to Jackson (1913) it was first discovered by Breby
and Reynolds in its well-known station, Forge Valley near Scarborough, in 1857. Here
the plant has shown a steady increase in recent years, “between 90 and 100 flowers in
May, 1945’ (Garton 1945).

v.c. 65, N. Yorkshire. Recorded by Gerard (1597) from ‘. .. Howgill, Yorkshire, in a
field next Todberry.’ This record is included by Lees (1888) in his Flora of West York-
shire. Although the locality is in the West Riding it is actually in v.c. 65 and not 64.
This record is generally regarded as an error.

v.c. 66, Durham. Recorded by Oliver (1896) in a plantation at Hunstanworth where it
still persists.

v.c. 67, S. Northumberland. There is an unsubstantiated record from near Blanchland
but Dr G. A. Swan considers that the species is not in this vice-county.

v.c. 68, N. Northumberland. First found in 1843 by the Rev. Oswald Head in one of
the woods at Howick, close to Earl Grey’s garden, and Watson (1849) in his Cybele
Britannica considered the plant ‘Incognit or Alien’. Oliver (1896) refers to a further
record from Rothbury.

v.c. 83, Midlothian. There is a specimen in the Cambridge Herbarium collected by
T. Cowan, Junr, in 1909 from Carlowrie in West Lothian.

The foregoing evidence shows that it has been occasionally introduced and, although
formerly far from uncommon (Parkinson 1640), it has considerably diminished in
frequency with the exceptions of Norfolk and north-east Yorkshire. British woodlands
have been much reduced in acreage by felling with the consequent loss of the character-
istic flora. A survey of the literature has shown conflicting views concerning its status.

SHORT NOTES 297

In his summary, Jackson (1913) quotes de Candolle’s note (1855) to the effect that,
contrary to both Babington (1847) and Watson (1849), who considered the plant to be
an alien, the French botanist had no doubt at all that it was a good British native.

In an attempt to assess its status here it is necessary to take into account its distribu-
tion and ecology outside the British Isles. In France, Boreau (1857), Coste (1906) and
Fournier (1961) refer it to woods and pastures, particularly in the mountainous districts.
It is certainly abundant in the mountainous woods of Switzerland. Clapham, Tutin
& Warburg (1962) give its European distribution as ‘Europe from Scandinavia and
N. Russia to N., E. and C. France, N.E. Spain, N. Apennines, Bosnia and S. Russia’.
I can find no correlation between this presumably Eurasian element and any of the
British stations, particularly the largest colony extant in Norfolk.

In conclusion, therefore, there is so much suspicion on historical grounds, sugges-
tions of introduction, and slender ecological correlation, that a more accurate estima-
tion would be that in Britain the May Lily is a naturalised alien.

REFERENCES

BABINGTON, C. C. (1847). Manual of British Botany, 2nd ed., p. 324. London.

BoreEAu, A. (1857). Flore du Centre de la France, 3rd ed., p. 616. Paris.

CANDOLLE, A. P. DE (1855). Géographie botanique raisonée, p. 696. Paris.

CLAPHAM, A. R., TUTIN, T. G. & WARBURG, E. F. (1962). Flora of the British Isles, 2nd ed.,
p. 968. Cambridge.

Coste, H. J. (1906). Flore descriptive et illustrée de la France, vol. 3, p. 350. Paris.

Dony, J. G. (1952). Flora of Bedfordshire, p. 394. Luton.

FOurRNIER, P. (1961). Les quatres Flores de France, p. 179. Paris.

GarTON, G. A. (1945). The Naturalist, 1945: 111.

GERARD, J. (1597). The Herbal, p. 330. London.

JACKSON, A. B. (1913). Maianthemum bifolium Schmidt in England. J. Bot., Lond., 51: 202-208.

Legs, F. A. (1888). Flora of West Yorkshire, p. 443. London.

LousLey, J. E., in Lous.ey, J. E., ed. (1953). The Changing Flora of Britain, p. 156. London.

OLIVER, D. (1896). Maianthemum bifolium in Durham. J. Bot., Lond., 34: 431.

PARKINSON, J. (1640). Theatrum Botanicum, p. 505. London.

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

Petcu, C. P. & SWANN, E. L. (1962). West Norfolk Plants Today. Proc. bot. Soc. Br. Isl., 4.
Supplement.

Petcu, C. P. & SWANN, E. L. (1968). Flora of Norfolk, p. 226. Norwich.

RID_ey, H. N. (1930). The Dispersal of Plants throughout the World, p. 457. Ashford.

SAVIDGE, J. P.,. HEyYwoop, V. H. & GorDon, V. (1963). Travis’s Flora of South Lancashire,
p. 297. Liverpool.

SMITH, J. E. (1828). The English Flora, vol. 2. London.

STILLINGELEET, B. (1761). The Calendar of Flora. London.

Watson, H. C. (1849). Cybele Britannica, vol. 2, p. 465. London.

E. L. SWANN

668/1. SIEGLINGIA DECUMBENS (L.) Bernh. — Pulvini of palea.

The palea, the characteristic scale of the grass flower, is usually a thin two-veined
structure with margins folded inwards to form uniformly sharp keels which are fre-
quently ciliolate. Seldom does the palea exhibit striking characters useful in identification
— unlike its opposing scale, the lemma — and consequently it is often overlooked in keys
and descriptions.

However, in the heath-grass, Sieglingia decumbens, common in moorland, the keels
of the palea are considerably enlarged and progressively thickened towards the base
(Fig. 1). These tissue-swellings, or pulvini, are soft-textured, well-rounded below, and
flesh-coloured or colourless in fresh plants. In dry specimens they become pale orange

298 SHORT NOTES

and shrivelled but on wetting soon regain their natural colour and turgescence. Intern-
ally they are composed of relatively large thin-walled cells bounded by a poorly defined
epidermis.
These distinctive pulvini were first noticed in a plant grown at Ottawa, Canada,
transplanted from Louisburg, Nova Scotia, where Sieglingia thrives in peaty coastal
turf as a local introduction, possibly dating back to the time of early French coloniza-
tion. The plant was of the usual cleistogamous (apomictic) form with reduced anthers
and abortive lodicules. Examination of numerous specimens from throughout the
native range of the species, including a chasmogamous specimen from Austria, revealed
a consistent occurrence of the paleal pulvini (although these are not described or clearly
depicted in any of the numerous published works so far searched). I consider, therefore,
that these swellings are an inherent feature of S. decumbens, not pathological or abnor-
mal structures, even though they may have superficial resemblance to rust pustules
especially when dry. Their function is not immediately apparent, although their
activity may serve to loosen the ripe floret in the absence of a well-developed awn
such as members of the closely related genus Danthonia possess.

Indeed, the relationship of Sieglingia to Danthonia is so close, that the propriety of
maintaining the former as a monotypic genus has long been questioned (Hubbard
1948). When De Candolle created the genus Danthonia in 1805 he even referred Festuca
decumbens L. to it, apparently unaware that this was already the type of Sieglingia of
Bernhardi of 1800. The choice has largely rested on whether the central tooth of the
lemma in Sieglingia represents an abbreviated awn or not, other features being com-
parable with Danthonia. The recent action taken to conserve Danthonia (Baum 1968;
Moore et al. 1970) would mean that the now familar name Sieglingia would be lost to
Danthonia, if considered congeneric. The prominent feature of paleal pulvini noticed
here supports the maintenance of Sieglingia as a distinct genus.

2mm

---"-
-==
ee

M_ ZZ

Ficure 1. Sieglingia decumbens. A. Complete floret in adaxial view, the lemma flattened out.
B. Palea (diagrammatic) showing the pulvini on lower portion of keels, ciliate upper portion
of keel, and infolded margins (dashed lines). C. Sections of palea at levels indicated, showing

position of vein (circle) and thin-walled tissue (not stippled) of turgid pulvini. (Drawn from
Dore & Marchant 23253 - DAO.)

SHORT NOTES 299

REFERENCES

Baum, B. R. (1968). Proposal to conserve the generic name 280. Danthonia (Gramineae).
Taxon, 17: 444-446.

HUBBARD, C. E. (1948). Gramineae, in HUTCHINSON, J., British Flowering Plants, pp. 284-348.
London.

Moore, H. E., STAFLEU, F. A. & Voss, E. G. (1970). XI International Botanical Congress:
final mail vote and Congress action on nomenclatural proposals. Taxon, 19: 43-51.

W. G. Dore

Plant Research Institute Contribution No. 729, Canada Department of Agriculture,
Ottawa 3.

671/2 x r. LOLIUM MULTIFLORUM Lam. x L. RIGIDUM Gaud.-— At Blackmoor, N.
Hants., v.c. 12.

The following natural interspecific hybrids involving different species of Lolium have
been listed in the botanical literature (Ascherson & Graebner 1902, Ullmann 1936,
Terrell 1966): Lolium multiflorum x perenne (L. x hybridum Hausskn.), L. multi-
florum x remotum, L. multiflorum x rigidum, L. perenne x rigidum and L. perenne x
temulentum. Of these, only L. multiflorum x perenne has hitherto been reported from
Britain (Dandy 1958). When examining British material of Lolium at the British Museum
(Natural History), Dr E. E. Terrell, of Beltsville, Maryland (U.S.A.), identified a
specimen collected by J. E. Lousley on October 29, 1960, at Blackmoor, N. Hants.
(v.c. 12) and distributed in his Wool Adventive Flora of Britain under the name L.
rigidum (No. W/1651), as L. multiflorum x rigidum, which has not been previously
recorded from Britain.

L. multiflorum x rigidum is intermediate in several respects between its putative
parental species, and is characterized by the following features.

_Annual or biennial, about 60 cm high. Culms tufted, geniculate in the lower part,
erect in the upper, somewhat stout, branched at the base, usually 3-noded, scaberulous
below the spike, otherwise glabrous, smooth; nodes tinged with purple. Leaves green,
glabrous; sheaths rounded on the back; ligules about 1 mm long, membranous; leaf-
blades 10-25 cm long, 3—7 mm wide, scaberulous above, rolled in the young shoot,
with short, spreading auricles. Spike 20—25 cm long, rather stout, erect or slightly nod-
ding, compressed; axis scaberulous. Spikelets 1-5—2-8 cm long, 9—11-flowered, sessile,
with their edges in hollows of the axis, oblong-lanceolate, breaking up at maturity
below each lemma. Lower glume present only in the terminal spikelet, similar in shape
and size to the upper; upper glume 14-17 mm long, narrowly oblong, 5—7-nerved,
obtuse. Lemmas 5—8 mm long, lanceolate-oblong, 5-nerved, tipped with a short, straight
awn 1—2 mm long. Paleas as long as the lemmas, 2-keeled, minutely ciliate towards the
tip. Rhachillas slightly scaberulous. Anthers about 4 mm long.

This hybrid differs from L. multiflorum mainly in having a longer upper glume and
a much shorter awn of the lemma, and from L. rigidum in having a shorter upper glume
and a short-awned lemma.

The results obtained by Jenkin et al. (Jenkin 1931, 1935, 1954a, b, c, d, e, 1955; Jenkin
& Thomas 1938, 1939) in crosses among the three cross-pollinated and largely self-
sterile species, L. multiflorum, L. perenne and L. rigidum, indicate that they are strongly
interfertile. Crosses between these species with self-pollinated species, such as L. Joli-
aceum, L. temulentum and L. remotum, produced sterile hybrids.

REFERENCES

ASCHERSON, P. F. A. & GRAEBNER, K. O. P. P. (1902). Synopsis der Mitteleuropdischen Flora,
vol. 2, pp.759-760. Berlin.
Danpy, J. E. (1958). List of British Vascular Plants, p. 158. London.

300 SHORT NOTES

JENKIN, T. J. (1931). The interfertility of Lolium perenne and L. perenne var. multiflorum. Bull.
Welsh PI. Breed. Stn, ser. H, 12: 121-125.

JENKIN, T. J. (1935). Interspecific and intergeneric hybrids in herbage grasses. II. Lolium perenne
x L. temulentum. J. Genet., 31: 379-411.

JENKIN, T. J. (1954a). Interspecific and intergeneric hybrids in herbage grasses. IV. Lolium
rigidum et alia. J. Genet., 52: 239-251.

JENKIN, T. J. (1954b). Interspecific and intergeneric hybrids in herbage grasses. V. Lolium
rigidum sens. ampl. with the other Lolium species. J. Genet., 52: 252-281.

JENKIN, T. J. (1954c). Interspecific and intergeneric hybrids in herbage grasses. VI. Lolium
italicum A.Br. intercrossed with other Lolium types. J. Genet., 52: 282-299.

JENKIN, T. J. (1954d). Interspecific and intergeneric hybrids in herbage grasses. VII. Lolium
perenne L. with other Lolium species. J. Genet., 52: 300-317.

JENKIN, T. J. (1954e). Interspecific and intergeneric hybrids in herbage grasses. VIII. Lolium
loliaceum, Lolium remotum and Lolium temulentum, with references to ‘Lolium canadense’.
J. Genet., 52: 318-331.

JENKIN, T. J. (1955). Interspecific and intergeneric hybrids in herbage grasses. XVIII. Various
crosses including Lolium rigidum sens. ampl. with L. temulentum and L. loliaceum with
Festuca pratensis and with F. arundinacea. J. Genet., 53: 467-486.

JENKIN, T. J., & THomas, P. T. (1938). The breeding affinities and cytology of Lolium species.
J. Bot., Lond., 76: 10-12.

JENKIN, T. J. & THomas, P. T. (1939). Interspecific and intergeneric hybrids in herbage grasses.

Ill. Lolium loliaceum x Lolium rigidum. J. Genet., 37: 255-286.

TERRELL, E. E. (1966). Taxonomic implications of genetics in Ryegrass (Lolium). Bot. Rev., 32:
138-164.

ULLMANN, W. (1936). Natural hybridization of Grass species and genera. Herb. Rev., 4: 105-

i A. MELDERIS

683/p. BROMUS PUMPELIANUS — An introduced species, at Walthamstow, S. Essex, v.c. 18.

A Brome found by J. Whisher on 23rd September, 1967, at Walthamstow, S. Essex
(v.c. 18), growing in large quantities on a bank of the reservoir, proved to be Bromus
pumpelianus Scribn., which is a polymorphic N. American species, occurring in mead-
ows and on grassy slopes from Colorado to S. Dakota, Idaho, Yukon and Alaska.
It is closely related to B. inermis Leyss., a native of northern and central Europe and of
temperate Asia, extending eastwards to China. As a fodder plant, B. inermis has been
introduced in the British Isles (also in N. America), where it has become naturalised in
widely scattered localities, especially on sandy and stony soils. The following are details
of the Walthamstow specimen:

A perennial, 50-100 cm high, forming loose tufts, spreading by long wiry rhizomes.
Culms erect, stout, unbranched, minutely pubescent below the nodes. Leaves dark
green; sheaths tubular, splitting in the upper part, striate, smooth or minutely pubescent
towards the base; ligules short; leaf-blades about 20-25 cm long, 3-5-9 mm wide,
scaberulous on the upper surface, glabrous, smooth beneath, flat, except for the involute,
pungent tip, auricled at the base. Panicle 10-20 cm long, more or less dense, green;
branches 5—10 cm long, erect, divided; the lower in pairs, the upper 3—5 at each node;
axis glabrous. Spikelets about 30 mm long, narrowly lanceolate to narrowly oblong,
somewhat compressed, S—10-flowered, breaking up below each lemma. Glumes per-
sistent, glabrous, keeled, unequal; the lower 5-7 mm long, narrowly lanceolate, 1-
nerved, finely pointed; the upper 7-9 mm long, oblong-lanceolate, 3-nerved, obtuse.
Lemmas 9-12 mm long, 7-nerved, minutely pubescent towards the base and along the
margins, tipped with a fine straight awn 2-3 mm long. Paleas shorter than the lemmas,
2-keeled, with shortly ciliate keels. Rhachillas pubescent. Anthers 4:5—5:5 mm long.

B. pumpelianus differs from B. inermis in having a lemma which is more or less
pubescent along the margins and across the back (densely and long-hairy all over in
forms of the northern parts of N. America), tipped with a short awn 2-3 mm long. In
B. inermis the lemma is glabrous, smooth or slightly scaberulous, with an obtuse tip,

awnless or mucronate. A. MELDERIS

SHORT NOTES 301

PERSISTENCE OF SILENE MARITIMA AND OTHER PLANTS ON AN INLAND TIP IN SOUTH
LANCASHIRE, V.C. 59.

In 1942, imported sand was deposited on an ash-tip at Leigh, South Lancashire which
I had first known in 1938. The site lies about thirty miles inland. Several new plants
appeared on the sand in 1943 and a watch was kept on these until 1948, after which the
site was not visited until 1968.

In the intervening years, part of the ash-tip and a large part of the adjoining railway
complex had been removed. Despite this, the sand was still visible in places in 1968
and, on and around it, all but one of the adventives of 25 years earlier were still
present. Status changes were as follows:

Cakile maritima. 1943, several large plants. 1947, five, very small, sickly plants.

1968, not refound.

Diplotaxis muralis. 1943, several, robust plants. 1948, a few, very small but healthy

plants. 1968, ten, small, fertile specimens.

Erodium cicutarium subsp. cicutarium. 1943-1948, some fluctuation due to tramp-

ling, usually about ten moderate plants. 1968, twelve tiny plants among taller

vegetation.

Linaria dalmatica. 1943-1948, 1968, fairly constant, five or six large and as many

small, unbranched flowering plants.

Silene maritima. 1943-1948, constant at four perennial plants. 1968, 25 large mats,

now also on the surrounding ashes.

Ononis repens. 1943-1948, increased and spread to the ashes. 1968, still common

but now discontinuous over a wider area.

Saponaria officinalis. 1943-1948, 1968, very similar in status and history to Ononis

repens.

Anthyllis vulneraria. 1943 and 1944, common on the sand. 1948, apparently gone.

1968, common on and near the sand.

The persistence of several of these species and the increase of Silene maritima at this
inland, lowland site is unusual.

The plant community of the main tip, which is well-trodden, appeared to have
changed little between 1938 and 1968, the main components being:

very common — Epilobium angustifolium, Artemisia vulgaris and Conium maculatum

common — Reseda luteola, Melilotus officinalis and M. alba

frequent — Oenothera biennis, Potentilla norvegica and Silene vulgaris

T. EDMUNDSON

TWO ADDITIONS TO THE FLORA OF FOULA, ZETLAND, V.c. 112.

Since the publication of ‘Notes on the Flora and Vegetation of Foula, Zetland (v.c.
112)’ (Hawksworth 1969), it has been possible to confirm two further species for the
island:

19/1. Cystopteris fragilis (L.) Bernh. in shelter of Old Red Sandstone boulders Hamna-
fjeld, Foula, 1965, D. L. Hawksworth; under a boulder, Hamnafjeld, Foula, 1968,
W. Scott and R. C. Palmer.

192/17. Trifolium hybridum L. near Ham, introduced with corn, 1965, D. L. Hawks-
worth, confirmed by T. G. Tutin and A. O. Chater, LTR.

REFERENCE

HAwkswonrtH, D. L. (1969). Notes on the Flora and Vegetation of Foula, Zetland (v.c. 112).

Proc. bot. Soc. Br. Isl., 7: 537-547.
D. L. HAWKSWoRTH

302 Watsonia, 8, 302-312 (1971).

Plant Records

Records for publication should be submitted in the form shown below to the appropriate
vice-county Recorder and not direct to the Editors.

Records are arranged in the order given in the List of British Vascular Plants (L.B.V.P.) by
J. E. Dandy (1958) and in his subsequent revision (Watsonia, 7: 157-178 (1969)). Herbarium
abbreviations are those used by D. H. Kent (1957).

The following signs are used:

* before the record: to indicate a new vice-county record.
+ before the L.B.V.P. 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. 9, Dorset: Pilsdon Pen, GR 31/41.01. D. R.
Seaward, 1967, field record. Ist record this century. 27, Norfolk: Winterton, GR 63/49.
20. Small colony in dune slacks. F. W. Simpson, 1963, herb. E. L. Swann; Lenwade,
GR 63/11.18. Marshy area. M. J. Wigginton, 1969, herb. E. L. Swann. Ist post-1930
records.

4/1. EQUISETUM HYEMALE L. *42, Brecon: near Llanbedr, GR 32/24.19. In beech
litter in damp wood. M. Porter, 1968, NMW. (Nature Wales, 11:138 (1969)).

4/4. EQUISETUM VARIEGATUM Schleich. ex Weber & Mohr *107, E. Sutherland:
south-west side of Loch Sail an Ruathair, Benmore Forest, GR 29/33.14. On a small
limestone outcrop. U. K. Duncan, 1969, herb. U.K.D.

4/9 x 5. EQUISETUM X LITORALE Kiihlew. ex Rupr. *45, Pembroke: Merry
Vale, St David’s, GR 12/74.24. Moist ground at roadside. S. G. Harrison, 1969, NMW.
(Nature Wales, 12: 37 (1970)).

15/6. ASPLENIUM VIRIDE Huds. *103, Mid Ebudes (Watsonia, 8: 147 (1970)).

19/1. CYSTOPTERIS FRAGILIS (L.) Bernh. *112, Zetland (see Short Notes p. 301).
Ist definite record.

21/3. DRYOPTERIS ABBREVIATA (DC.) Newm. 104, N. Ebudes: Coire Uaigneich,
Blaven, Skye, GR 18/53.21. On scree. M. McCallum Webster, 1966, herb. C. W.
Murray. 2nd record. *111, Orkney: Ward Hill, Rousay, GR N31/38.30. Rocks at
north end, 600 ft. E. R. Bullard, 1968, BM, conf. A. C. Jermy.

21/6. DRYOPTERIS CARTHUSIANA (Vill.) H. P. Fuchs 47, Montgomery: Dolanog,
GR 33/07.11. In bog. D. E. M. Paish, 1967, NMW. 2nd record.
24/3. THELYPTERIS PHEGOPTERIS (L.) Slosson *45, Pembroke: Craig-y-cwm,

Foel Cwmcerwyn, Mynydd Prescelly. Large colony on wet face of disused quarry,
1550 ft. T. A. W. Davis (69/1229), 1969, BM, NMW. (Nature Wales, 12: 37 (1970)).
*111, Orkney: Nowt Bield, Ward Hill, Hoy, GR N31/23.01. Below rocks and in
heather, 250 ft. E. R. Bullard, 1969, E. 1st definite record.

28/1. BOTRYCHIUM LUNARIA (L.) Sw. 80, Roxburgh: Drinkstone Hill, GR 36/48.
18. Pasture, 1000 ft. R. W. M. Corner, 1967, field record. Ist post-1930 record.

29/1. OPHIOGLOSSUM VULGATUM L. 104, N. Ebudes: valley of Allt nan Leac,
Skye, GR 18/59.18. In turf and limestone gryke. C. W. Murray, 1966, herb. C.W.M.
Ist record for Skye.

43/1. ANEMONE NEMOROSA L. H6, Waterford: west of Ballynaclogh cross roads,

3 miles north-west of Tramore, GR 02/46. Recently cleared scrub. I. K. Ferguson,
1961, field record. 2nd record. (U/r. Nat. J., 16: 94 (1968)).

PLANT RECORDS 303

45/1. CLEMATIS VITALBA L. *+103, Mid Ebudes (Watsonia, 8: 147 (1970)).

46/21. RANUNCULUS TRICHOPHYLLUS Chaix *104, N. Ebudes: stream between
Lock Fada and Loch Leathan, Skye, GR 18/49.50. J. Macdonald, 1968, herb. C. W.
Murray.

57/1. CERATOPHYLLUM DEMERSUM L. *43, Radnor: Pen Cerrig Lake, near Builth,
GR 32/04.54. A. C. Powell, 1969, field record, conf. G. Ellis. (Nature Wales, 12:36
(1970)). *67, S. Northumberland: pond near High West House, Ponteland, GR
45/14.73. 250 ft. G. A. & M. Swan, 1969, herb. G. A. Swan, conf. J. E. Dandy.

58/6. PAPAVER SOMNIFERUM L. *103, Mid Ebudes (Watsonia, 8: 147 (1970)).

61/1. GLAUCIUM FLAVUM Crantz H6, Waterford: Bunmahon, GR 02/36. Sand
dunes. I. K. Ferguson, 1961, field record. Ist post-1930 record. (/r. Nat. J., 16: 94
(1968)).

77/1. CAKILE MARITIMA Scop. 104, N. Ebudes: Glen Brittle beach, Skye, GR
18/40.20. A. McG. Stirling, 1954. Ist record for Skye.

92/1. LOBULARIA MARITIMA (L.) Desv. *80, Roxburgh: shingle of R. Tweed,
Melrose, GR 36/53. M. McCallum Webster, 1969, E.
95/1. EROPHILA VERNA (L.) Chevall. H6, Waterford: Tramore, GR 02/46. Sand
dunes. I. K. Ferguson, field record. 2nd record. Ur. Nat. J., 16:94 (1968)).
$97/7. CARDAMINE RAPHANIFOLIA Pourtr. *33, E. Gloucs. (Watsonia, 8: 86 (1970)).
+98/3. BARBAREA INTERMEDIA Bor. *103, Mid Ebudes (Watsonia, 8: 147 (1970)).
*104, N. Ebudes: Portree, Skye, GR 18/48.43. Garden weed. C. W. Murray, 1963,
herb. C.W.M.

102/5. RORIPPA AMPHIBIA (L.) Bess. *47, Montgomery: Stalloe Pool, GR 32/22.
98. H. Webster, 1969, NMW. (Nature Wales, 12: 38 (1970)).

113/5. VIOLA REICHENBACHIANA Jord. ex Bor. *42, Brecon: Cwm Claisfer, GR
32/14.16, and Dyffryn Crawnon, GR 32/09.15. On carboniferous limestone. M. Porter,
1968, both NMW. (Nature Wales, 11: 138 (1969)). Ist and 2nd records.

113/7. VIOLA LACTEA Sm. *H16, W. Galway: near Costello school, GR 94/00.
G. Mackie, 1969, herb. G.M., conf. D. H. Valentine.

7115/4. HyPERICUM CALYCINUM L. *103, Mid Ebudes (Watsonia, 8: 148 (1970)).

123/1. SILENE VULGARIS (Moench) Garcke *103, Mid Ebudes (Watsonia, 8: 148
(1970)).

124/2. LYCHNIS VISCARIA L. 80, Roxburgh: Minto Crags, GR 36/58.20. R. W. M.
Corner, 1966, field record. Ist post-1930 record.

131/5. CERASTIUM ARCTICUM Lange 104, N. Ebudes: Sgurr nan Gillean, Skye,
GR 18/47.25. R. M. Murray, 1966, herb. C. W. Murray. Confirmation of the only
record (pre-1930) for the v.c.

131/11. CERASTIUM PUMILUM Curt. *29, Cambridge: Cambridge station car park,
GR 52/46.57. A. P. Conolly, 1969, CGE, det. P. D. Sell.

133/3. STELLARIA PALLIDA (Dumort.) Piré *46, Cardigan,: near Ynys Las, GR
22/60.93. Rabbit scrapes in old dunes. P. M. Benoit, 1970, NMW. *107, E. Sutherland:
Brora, GR 29/90. Sand dunes in golf links. M. McCallum Webster, 1969, CGE.

136/3. SAGINA MARITIMA Don 103, Mid Ebudes (Watsonia, 8: 148 (1970)).

154/14. CHENOPODIUM RUBRUM L. *47, Montgomery: Trewern, GR 33/28.11.
Garden weed. W. F. Jary, 1969. (Nature Wales, 12: 38 (1970)).

159/1. SALSOLA KALI L. 104, N. Ebudes: Camus Ban, Portree, Skye, GR 18/49.
42. N. Dennis, 1963, herb. C. W. Murray. New to Skye.

160/5. SALICORNIA PUSILLA Woods H6, Waterford: Back Strand, Tramore, GR
02/56; White House Bank near Dungarvan, GR 02/25. Both I. K. Ferguson, field
records. Ur. Nat. J., 16: 95 (1968)). 2nd and 3rd records.

304 PLANT RECORDS
168/15. GERANIUM LUCIDUM L. 104, N. Ebudes: south-east of Torvaig, Skye,
GR 18/50.44. On wall. C. W. Murray, 1961, herb. C.W.M. Ist record for Skye.

+171/3. IMPATIENS PARVIFLORA DC. *43, Radnor: Boultibrook, near Presteigne,
GR 32/30.65. On rubble of lane verge. K. Lomax, 1969, field record, det. A. C. Powell.
(Nature Wales, 12: 36 (1970)).

179/1. RHAMNUS CATHARTICUS L. *+103, Mid Ebudes (Watsonia, 8: 148 (1970)).

192/4. TRIFOLIUM MEDIUM L. *48, Merioneth: railway embankment between
Towyn and Aberdovey, GR 22/59. K. M. Stevens, 1969, NMW. Ist definite record.

4192/17. TRIFOLIUM HYBRIDUM L. 103, Mid Ebudes (Watsonia, 8: 148 (1970)).

206/10. VICIA SYLVATICA L. 79, Selkirk: Buck Cleuch, Clearburn, GR 36/33.14.
950 ft. R. W. M. Corner, 1968, field record. Ist post-1930 record.

206/17. VICIA BITHYNICA (L.) L. *32, Northants.: Finedon ironstone pits, GR
42/91.71. Grassed-over quarry slope. E. M. Cooke, 1968, field record.

207/2. LATHYRUS NISSOLIA L. *+42, Brecon: disused railway line near Llan-
santffraed, GR 32/12.23. On embankment, presumably introduced. M. Porter, 1968,
NMW. (Nature Wales, 11: 138 (1969)).

207/6. LATHYRUS SYLVESTRIS L. *103, Mid Ebudes (Watsonia, 8: 149 (1970)).

207/10. LATHYRUS JAPONICUS Willd. *68, Cheviot: Birling Links, GR 46/25.07.
G. A. & M. Swan, 1969, herb. G.A.S.

212/12. POTENTILLA CRANTZII (Crantz) G. Beck 97, Westerness (Argyll): Ben
Iadain, GR 17/65. Steep south-facing rocks. A. A. Slack, 1966, field record. 2nd record.

212/14. POTENTILLA ANGLICA Laichard. *105, West Ross: Eilean Mheadhonach,
Crowlin Islands, GR 18/68.35. A. A. Slack, 1967, herb. A.A.S.

+235/11. SEDUM REFLEXUM L. 80, Roxburgh: Minto Crags, GR 36/58.20. R. W.
M. Corner, 1966, field record. Ist post-1930 record.

235/12. SEDUM VILLOSUM L. *97, Westerness (Argyll): Ben Iadain, GR 17/65.
Damp hollow in summit plateau. A. A. Slack, 1966, field record.

237/2. CRASSULA AQUATICA (L.) Schonl. *97, Westerness (see Short Notes
p. 294).
239/15. SAXIFRAGA HYPNOIDES L. *80, Roxburgh: north-east side of Ewenshope

Fell, GR 35/37.99. Beside burn, 1400 ft. R. W. M. Corner, 1969, field record; Twistle-
hope and Chapel Grain Burns, GR 35/49. 1000-1250 ft. R. W. M. Corner, 1970, field
record. 2nd record.

+241/1 ToLMIEA MENzIESI (Pursh) Torr. & Gray *79, Roxburgh: by River
Ettrick, GR 36/27.14. In Salix scrub. R. W. M. Corner, 1968, K, det. R. D. Meikle.
The record for v.c. 79 in Proc. bot. Soc. Br. Isl., 5: 350 (1964) is actually in v.c. 80.
*103, Mid Ebudes (Watsonia, 8: 149 (1970)).

+254/6. EPILOBIUM ADENOCAULON Hausskn. *51, Flint: south-east shore of Llyn
Helyg, GR 33/17. Wooded ground. P. M. Benoit, 1969, NMW.

254/12. EPILOBIUM ALSINIFOLIUM Vill. *67, S. Northumberland: Todd’s Sike,
near Allenheads, GR 35/83.45. 1650 ft; Tyne near Kirkhaugh, GR 35/64. On river
gravels, 800 ft. Both G. A. & M. Swan, 1969, herb. G.A.S., conf. T. D. Pennington.
Ist and 2nd records respectively.

+254/13. EPILOBIUM NERTERIOIDES Cunn. 43, Radnor: Water-break-its-neck,
New Radnor, GR 32/18.60. M. Jenkins, 1968, field record, conf. A. C. Powell. (Nature
Wales, 11: 139 (1969)). *104, N. Ebudes: Skerinish road, Skye, GR 18/41.50. In quarry.
J. Raven, c. 1951, field record. 1958 specimen in herb. C. W. Murray. 111, Orkney:
Wideford Hill, St Ola, GR N30/41.11. Roadside, 740 ft. E. R. Bullard, 1968, field
record. Ist Mainland record.

PLANT RECORDS 305

261/1. HIPPURIS VULGARIS L. *104, N. Ebudes: north of Digg, Skye, GR 18/47.
70. Lochan by shore. C. W. Murray, 1965, herb. C.W.M.

262/2. CALLITRICHE PLATYCARPA Kiitz. 48, Merioneth and *49, Caernarvon:
R. Glaslyn near Pont Croesor, GR 23/54. P. M. Benoit, 1969, NMW. 2nd record for
v.c. 48.

276/1. MyYRRHIS ODORATA (L.) Scop. 43, Radnor: near Fron Rocks, GR 32/19.
76. In wet ditch. A. C. Powell, 1969, field record. 2nd record. (Nature Wales, 12: 36
(1970)). *45, Pembroke: Caldey Island, GR 21/14.96. Waste ground. F. Stewart-
Liberty, 1969, NMW (T. A. W. Davis 69/1191). (Nature Wales, 12: 38 (1970)).

301/1. AETHUSA CYNAPIUM L. 103, Mid Ebudes (Watsonia, 8: 150 (1970)).

7311/2. HiERACIUM MANTEGAZZIANUM Somm. & Levier *103, Mid Ebudes
(Watsonia, 8: 150 (1970).

320/2. POLYGONUM RAI Bab. 68, Cheviotland: Holy Island, GR 46/12.41. G. A.
& M. Swan, 1969, herb. G.A.S., conf. B. T. Stiles. 2nd record. 104, N. Ebudes: Glen
Brittle beach, Skye, GR 18/40.20. C. W. Murray, 1966, herb. C.W.M. Confirmation
of the only (1916) Skye record.

+320/18, 20-22. POLYGONUM BALDSCHUANICUM Regel, P. SACHALINENSE F. Schmidt,
P. POLYSTACHYUM Wall. ex Meisn. and P. CAMPANULATUM Hook. f. *103, Mid
Ebudes (Watsonia, 8: 150 (1970)).

325/14. RUMEX SANGUINEUS L. *104, N. Ebudes: North Fearns, Raasay, GR
18/59.35. Edge of wood above shore. C. W. Murray, 1969, herb. C.W.M.

325/18. RUMEX MARITIMUS L. 47, Montgomery: Hen Flash, Forden, GR 33/23.
00. V. J. Macnair, 1968, NMW. (Nature Wales, 11: 140 (1969)). 2nd record.

333/1. MyYRICA GALE L. H6, Waterford: between Annestown and Dunhill, GR
02/36. Marshy ground. I. K. Ferguson (889), 1963, TCD. Ur. Nat. J., 16: 95 (1968)).
2nd record.

343/20. SALIX MYRSINITES L. 104, N. Ebudes: Sgurr Mor, Skye, GR 18/44.70.
Rock ledge. H. J. B. Birks, 1968, herb. C. W. Murray. Ist record for Skye.

358/4. VACCINIUM OXYCOCCUS L. *79, Selkirk: west end of Alemoor Loch, GR
36/38.14. Sphagnum bogs, 900 ft. 1961; Pot Loch, GR 36/37.18. 1965. Both records
R. W. M. Corner, field records. Ist and 2nd records respectively. 103, Mid Ebudes
(Watsonia, 8: 150 (1970)).

359/1. PyROLA MINOR L. 57, Derby: Ashwood Dale near Buxton, GR 43/08.72.
Bottom of limestone scree. K. M. Hollick, 1969, field record, det. A. R. Clapham.
Probably only surviving colony in v.c.

359/2 PYROLA MEDIA Sw. 104, N. Ebudes: Tote, GR 18/42.49; Skerinish, GR.
18/41.50. Several stations in moorland. J. Raven, c 1951, field records, later specimens
herb. C. W. Murray. Ist and 2nd records for Skye.

360/1. ORTHILIA SECUNDA (L.) House *103, Mid Ebudes (Watsonia, 8: 150
(1970)).

361/1. MONESES UNIFLORA (L.) A. Gray 88, Mid Perth: Black Wood of Ran-
noch, GR 27/55. A. W. Punter, 1969, field record. Ist post-1930 record.

367/3. PRIMULA VERIS L. H6, Waterford: Ballykinsella, between Waterford and
Tramore, GR 02/56; Pickardstown House near Tramore, GR 02/46. Both I. K. Fergu-
son, field records. (/r. Nat. J., 16: 95 (1968)). 2nd and 3rd records.

370/3. LYSIMACHIA VULGARIS L. 103, Mid Ebudes (Watsonia, 8: 150 (1970)).
4370/5. LYSIMACHIA PUNCTATA L. *103, Mid Ebudes (Watsonia, 8: 150 (1970)).

372/4. ANAGALLIS MINIMA (L.) E. H. L. Krause 104, N. Ebudes: Varkasaig,
Skye, GR 18/25.42. Damp gravel by shore. C. W. Murray, 1961, herb. C.W.M. Ist
record for Skye.

306 PLANT RECORDS

374/1 SAMOLUS VALERANDI L. 104, N. Ebudes: mouth of Allt nan Leac, Skye,
GR 18/58.18. Damp ground. C. W. Murray, 1968, field record. 2nd record.

402/1. MERTENSIA MARITIMA (L.) Gray 104, N. Ebudes: shingle near Cleat
Skerry, Ardroag, Skye, GR 18/2.4. J. K. Morton, 1969, herb. C. W. Murray. Ist
record for Skye.

+406/2. CALYSTEGIA PULCHRA Brummitt & Heywood *42, Brecon: Llangynidr,
GR 32/15.21. Roadside hedge. M. Porter, 1968, NMW. (Nature Wales, 11: 138 (1969)).

*406/3. CALYSTEGIA SILVATICA (Kit.) Griseb. *42, Brecon: Llansantffraed, GR
32/12.23; Brecon, GR 32/05.27. Hedgerows. Both M. Porter, 1969, NMW. (Nature
Wales, 12: 35 (1970)). Ist and 2nd records.

+409/1. LycrtUM BARBARUM L. *103, Mid Ebudes (Watsonia, 8: 150 (1970)).

410/1. ATROPA BELLA-DONNA L. *58, Cheshire: Malpas churchyard, GR 33/48.
47. M. W. Cullen, 1969, field record.

+420/2. LINARIA PURPUREA (L.) Mill. 79, Selkirk: shingle of R. Tweed, Gala-
shiels, GR 36/43. M. McCallum Webster, 1962, field record. 2nd record.

420/3 x 4. LINARIA xX SEPIUM Allman 48, Merioneth: near Penmaenpool, GR
23/61. Disused railway line, one plant growing with parents. P.M. Benoit, 1969. 2nd
record.

421/1. CHAENORHINUM MINUS (L.) Lange 79, Selkirk: shingle of R. Tweed,
Galashiels, GR 36/43. M. McCallum Webster, 1966, E. 2nd record.

+425/1 x c. MIMULUS GUTTATUS DC. x CUPREUS Dombrain 103, Mid Ebudes
(Watsonia, 8: 151 (1970)).

+425/3. MIMULUS MOSCHATUS Dougl. ex Lindl. 103, Mid Ebudes (Watsonia, 8:
151 (1970)).

430/2. VERONICA ANAGALLIS-AQUATICA L. *42, Brecon: along R. Wye near
Llanigon, GR 32/20.40. Abundant in wet ditches. H. Soan, 1968, NMW. (Nature Wales,
11: 139 (1969)).

430/22. VERONICA POLITA Fr. 79, Selkirk: shingle of R. Tweed, Galashiels, GR
36/43. M. McCallum Webster, 1969, E. Ist post-1930 record.

432/1. PEDICULARIS PALUSTRIS L. H6, Waterford: Islandtarsney, west of Tra-
more. Marsh. I. K. Ferguson (2094), 1967, BM. (Ur. Nat. J., 16: 95 (1968)). 2nd record.

432/2. PEDICULARIS SYLVATICA L. subsp. HIBERNICA D. A. Webb *103, Mid
Ebudes (Watsonia, 8: 151 (1970)).

433/1. RHINANTHUS SEROTINUS (Schonh.) Oborny 17, Surrey: Chipstead Valley,
GR 51/27.57. Chalky meadow. S. B. Smith & E. J. Clement, 1968, SLBI, conf. D. J.
Hambler. 2nd record.

439/1. LATHRAEA SQUAMARIA L. 106, East Ross: by AY, 1 mile north of Dingwall,
GR 28/56. On calcareous bank, growing on Sambucus. A. Langton, 1969, E. 2nd
record.

442/1. UTRICULARIA VULGARIS L. H6, Waterford: south-east of Savagetown
cross roads, west of Dunhill, GR 02/26. I. K. Ferguson (354), 1962, TCD. (Ur. Nat. J.,
16: 95 (1968)). This and others are the 2nd and subsequent records.

442/4. UTRICULARIA MINOR L. *80, Roxburgh: Blackpool Moss, near Selkirk,
GR 36/51.29; Gattonside Moss, GR 36/54.36. Both 800 ft, R. W. M. Corner, 1956,
field records. Ist and 2nd records respectively.

445/2. MENTHA PULEGIUM L. *42, Brecon: near Llyswen, GR 32/11.37. Damp
hollows beside moorland pool. M. Porter, 1968, NMW. (Nature Wales, 11: 139 (1969)).

445/4 x 3. MENTHA X VERTICILLATA L. 103, Mid Ebudes (Watsonia, 8: 151
(1970)).

PLANT RECORDS 307

461/1. LAMIASTRUM GALEOBDOLON (L.) Ehrend. & Polatsch. *{67,S. Northumber-
land: near Longwitton Hall, GR 45/08.88. Naturalised in a wood. T. Trevelyan, 1938,
later specimen herb. G. A. Swan.

467/1. GLECHOMA HEDERACEA L. *104, N. Ebudes: Dunvegan Castle estate
woodland, Skye, GR 18/24.49. D. G. Campbell, 1959, herb. C. W. Murray; Kilmarie,
Strathaird, Skye, GR 18/55.17. Estate woodland. C. W. Murray, 1966, herb. C.W.M.
2nd record.

473/1. LirTTORELLA UNIFLORA (L.) Aschers. H6, Waterford: Carrickavrantry
Lake, west of Tramore, GR 02/46. I. K. Ferguson. Ur. Nat. J., 16: 96 (1968)). 2nd
record.

481/1. SHERARDIA ARVENSIS L. 97, Westerness: Ardtoe sand dunes, GR 17/67.
M. McCallum Webster, 1969, field record. 2nd record.

485/2. GALIUM BOREALE L. *79, Selkirk: Wilson Burn, Alemoor, GR 36/38.15.
950 ft. R. W. M. Corner, 1960, field record; ravine at Buck Cleuch, Clearburn, GR
36/33.14. R. W. M. Corner, 1968, field record. 2nd record.

485/3. GALIUM MOLLUGO L. *104, N. Ebudes: Edinbane, Skye, GR 18/34.50.
Hayfield, possibly introduced. M. Macdonald, 1968, herb. C. W. Murray.

485/7. GALIUM STERNERI Ehrend. *67, S. Northumberland: S. Tyne near Knars-
dale, GR 35/67.53. River gravels at 650 ft. 1968; near Low Burnfoot, GR 35/67.60,
450 ft. 1969. 2nd record. Both G. A. & M. Swan, herb. G.A.S., conf. K. M. Goodway.

485/10. GALIUM ULIGINOSUM L. 103, Mid Ebudes (Watsonia, 8: 151 (1970)).
7487/3. SAMBUCUS RACEMOSA L. *103, Mid Ebudes (Watsonia, 8: 151 (1970)).

494/1. VALERIANELLA LOCUSTA (L.) Betcke 97, Westerness: Ardtoe, Ardna-
murchan, GR 17/67. Sandy ground by shore. M. McCallum Webster, 1969, E. 2nd
record. 104, N. Ebudes: Ord. Sleat, Skye, GR 18/61.13. Roadside. C. W. Murray, 1967,
herb. C.W.M. Ist record for Skye. H6, Waterford: Bunmahon, GR 02/36. Roadside
wall. I. K. Ferguson (2117), 1968, BM. Ur. Nat. J., 16: 96 (1968)). 2nd record.

494/2. VALERIANELLA CARINATA Lois. *47, Montgomery: roadside between
Llandyssil and Montgomery, GR 32/21.94. O. Vaughan, 1968, NMW. (Nature Wales,
11: 140 (1969)).

502/1. BIDENS CERNUA L. 98, Argyll: by Loch Illiter, Luing, GR 17/71. A. A.
Slack, 1968, field record. Ist post-1930 record; either confirmation of Ist or a 2nd
record.

4503/1. GALINSOGA PARVIFLORA Cav. *29, Cambridge: west of Gamlingay, GR
52/22.51. Arable field. S. Richardson, 1969, CGE. 32, Northants.: Great Billing, GR
42/81.61. Gravel pits. G. M. Gent, 1968, field record. 2nd record.

4503/2. GALINSOGA CILIATA (Raf.) Blake *32, Northants.: Wollaston, GR
42/91.71. Waste ground. S. L. M. Karley, 1969, field record.
506/2 x 1. SENECIO X OSTENFELDIL Druce. *103, Mid Ebudes (Watsonia, 8:

151 (1970)).

7506/4. SENECIO SQUALIDUS L. *94, Banff: just west of Buckpool station, GR
38/46. On railway ballast. A. J. Sowter, 1969, field record.

_ 3506/6. SENECIO SYLVATICUS L. H6, Waterford: Kilfarrasy, west of Tramore, GR
02/46. Cultivated ground on cliff tops. I. K. Ferguson (935), 1963, TCD. Ur. Nat. J.,
16: 96 (1968)). Ist post-1930 record.

7509/2. PETASITES ALBUS (L.) Gaertn. *103, Mid Ebudes (Watsonia, 8: 152
(1970)).

7509/4. PETASITES FRAGRANS (Vill.) C. Presl *103, Mid Ebudes (Watsonia, 8:
152 (1970)).

H

308 PLANT RECORDS

7526/1. ANTHEMIS TINCTORIA L. *80, Roxburgh: Newtown St Boswells, GR
36/53. Shingle of R. Tweed, probably introduced with wool. M. McCallum Webster,
1969, E.

540/8. CIRSIUM DISSECTUM (L.) Hill *48, Merioneth: near Maentwrog, GR 23/
63. In Sphagnum-Eriophorum angustifolium bog. W. I. Jones, 1969, NMW. Ist definite
record.

7557/3. CICERBITA MACROPHYLLA (Willd.) Wallr. *103, Mid Ebudes (Watsonia,
8: 152 (1970)).

558/1/18. HIERACIUM LINGULATUM Backh. *104, N. Ebudes: lower slopes of
Blaven, Skye, GR 18/52. A. McG. Stirling, 1966, det. Sell & West.

558/1/38. HIERACIUM AMPLIATUM (W. R. Linton) A. Ley 104, N. Ebudes:
Suardal limestone near Loch Cill Criosd, Skye, GR 18/62. A. McG. Stirling, 1966,
det. Sell & West. 2nd record.

558/1/39. HIERACIUM PETROCHARIS (E. F. Linton) W. R. Linton *104, N.
Ebudes: Tote, Skye, GR 18/41.50. Rocky knoll. C. W. Murray, 1967, CGE, det. Sell &
West.

558/1/40. HIiERACIUM LANGWELLENSE F. J. Hanb. 104, N. Ebudes: Quirang, Skye,
GR 18/46. A. McG. Stirling, 1968, det. Sell & West. 2nd record.

558/1/43. HIERACIUM IRICUM Fr. 104, N. Ebudes: Bearreraig, Skye, GR 18/55.
On cliff. C. W. Murray, 1966, herb. C.W.M., det. Sell & West. 2nd record.

558/1/50. HIERACIUM NITIDUM Backh. *104, N. Ebudes: Skudiburgh, Skye, GR
18/37.64. Cliff ledge. C. W. Murray, 1963, herb. C.W.M., det. Sell & West.

558/1/91. HIERACIUM CYMBIFOLIUM Purchas *104, North Ebudes: between
Fearns and Hallaig, Raasay, GR 18/53. Limestone cliffs; near Brae, Raasay, GR 18/54.
Limestone gorge. Both A. McG. Stirling, 1969, CGE, det. Sell & West.

558/1/108. HiERACIUM PICTORUM E. F. Linton *104, N. Ebudes: lower slopes
of Blaven, Skye, GR 18/52; between Elgol and Camusunary, Skye, GR 18/51. Both A.
McG. Stirling, 1966, det. Sell & West. Ist and 2nd records respectively.

558/1/109. HiERACIUM POLLINARIOIDES Pugsl. *104, N. Ebudes: between Fearns
and Hallaig, Raasay, GR 18/53. Limestone cliffs; rocks by Loch Eadar de Bhaile,
Raasay, GR 18/54. Both A. McG. Stirling, 1969, CGE, det. Sell & West. Ist and 2nd
records respectively.

558/1/118. HiERACIUM OISTOPHYLLUM Pugsl. *51, Flint: between Rhydymwyn
and Leet Valley, GR 33/16. W. S. Laverock, 1901, LIV; Nant Alyn, T. Edmondson,
1969, herb. J. N. Mills. The latter probably refers to the same locality; both det.
J. N. Mills.

558/1/125. HiERACIUM AURANTIFLORUM Pugsl. *62, N.E. Yorks.: Hawnby, GR
44/59. Frequent on woods, walls and rocks. Boult, LIV, det. J. N. Mills.

558/1/136. HizRACIUM CAESIOMURORUM Lindeb. *104, N. Ebudes: between
Fearns and Hallaig, Raasay, GR 18/53. A. McG. Stirling, 1969, CGE, det. Sell & West.

558/1/138. HIERACIUM CRAVONIENSE (F. J. Hanb.) Roffey *104, N. Ebudes:
near Loch Cill, Criosd, Skye, GR 18/62. By burn. A. McG. Stirling, 1966, det. Sell &
West; south of Kilmarie, Skye, GR 18/55.16. Rocks above shore. C. W. Murray, 1967,
herb. C.W.M., det. Sell & West. 2nd record.

558/1/143. HiERACIUM CALEDONICUM F. J. Hanb. *51, Flint: banks by R. Alyn,
14 miles south of Rhydmwyn, GR 33/26. R. Brown, 1894, LIV; Nant Alyn, GR 33/16.
In disused limestone quarry. T. Edmondson, 1969, herb. J. N. Mills. 2nd record. Both
det. J. N. Mills. *70, Cumberland: rocks by Piers Ghyll, GR 35/20. 1900-2100 ft. J. N.
Mills, 1969, herb. C. West.

PLANT RECORDS 309

558/1/157. HIERACIUM DIAPHANOIDES Lindeb. *104, N. Ebudes: Druim an Aon-
ach, Raasay, GR 18/54. A. McG. Stirling, 1969, CGE, det. Sell & West.
558/1/188. HIERACIUM UIGINSKYENSE Pugsl. *69, Westmorland (N. Lancs.):

Seathwaite, Dunnerdale, GR 34/23.96. J. N. Mills (69/108), 1969, herb. J.N.M.

558/1/204. HIERACIUM CALCARICOLA (F. J. Hanb.) Roffey *64, Mid-W. Yorks.:
rocks by R. Wharfe below Appletreewick, GR 44/05.59. J. N. & J. R. J. Mills (69/117,
118), 1969, herb. J. N. Mills.

558/1/219. HIERACIUM PERPROPINQUUM (Zahn) Druce *104, N. Ebudes: wood-
land at Kilbride House, Skye, GR 18/58.20. C. W. Murray, 1967, CGE, det. Sell &
West.

558/1/m. HIERACIUM MIRANDUM Sell & C. West *69, Westmorland (N. Lancs.):
by small brook in Dunnerdale, GR 34/23.96. J. N. Mills (69/107), 1969, herb. J.N.M.
4th locality for the species.

558/2/3. HIERACIUM LACTUCELLA Wallr. *64, Mid-W. Yorks.: at the foot of the
Ingleborough hills, GR 34/77. — , 1818, LIV, det. J. N. Mills. 1st definite record. *69,
Westmorland: (No locality). 1814, LIV, det. J. N. Mills.

558/2/9. HIERACIUM PRAEALTOUM Vill. ex Gochnat *43, Radnor: Knucklas, GR
32/27. On a wall. J. R. Palmer, 1967, field record, det. C. West as subsp. praealtum.
(Nature Wales, 11; 30 (1968)).

561/1. BALDELLIA RANUNCULOIDES (L.) Parl. *104, N. Ebudes: Loch Cill Criosd,
Skye, GR 18/61.20. By outflow. N. Dennis, 1963, herb. C. W. Murray; edge of Loch
Airidh na Saorach, GR 18/68.20. C. W. Murray, 1968, herb. C.W.M. 2nd record.
H6, Waterford: by Carrickavrantry Lake, west of Tramore, GR 02/46. I. K. Ferguson
(2091), 1967, BM. Confirmation of the only (pre-1930) record. Ur. Nat. J., 16:96 (1968)).

576/2. ZOSTERA ANGUSTIFOLIA (Hornem.) Reichb. *103, Mid Ebudes (Watsonia,
8: 152 (1970)). *H6, Waterford: Back Strand, Tramore, GR 02/00.56. I. K. Ferguson
(2261), 1968, BM, DBN, LTR, det. J. E. Dandy & T. G. Tutin. Ur. Nat. J., 16: 176
(1969)).

576/3. ZOSTERA NOLTIT Hornem. *H6, Waterford: Back Strand, Tramore, GR
02/00.56. I. K. Ferguson (2262), 1968, BM, det. J. E. Dandy & T. G. Tutin. Ur. Nat. J.,
16: 176 (1969)).

577/3. POTAMOGETON COLORATUS Hornem. *79, Selkirk: Selkirk racecourse,
GR 36/49.27. Moss at 850 ft. D. J. Bellamy & J. Edmondson, 1968, BM, conf. J. E.
Dandy. )

577/8. POTAMOGETON PRAELONGUS Wulf. 48, Merioneth: the westernmost Llynau
Crogenen, GR 23/61. P. M. Benoit, 1969, NMW. *107, E. Sutherland: Loch Sail an
Ruathair, Benmore Forest, GR 29/33.14. U. K. Duncan, 1969, BM, conf. J. E. Dandy.

579/2. RUPPIA MARITIMA L. 103, Mid Ebudes (Watsonia, 8: 152 (1970)).

603/1. PARIS QUADRIFOLIA L. *104; N. Ebudes: Suardal, Skye, GR 18/61.19.
Limestone gryke above marble quarries. J. C. Brownlie, 1961, herb. C. W. Murray;
Coille Gairellach, Skye, GR 18/60.19. Grykes and swallow holes. C. W. Murray,
1962, herb. C.W.M. 2nd record.

7605/2. JUNCUS TENUIS Willd. *51, Flint: track on south-east side of Llyn
Helyg, GR 33/17. P. M. Benoit, 1969, NMW.

605/10 x 9. JUNCUS CONGLOMERATUS L. x EFFUSUS L. *45, Pembroke: Amble-
ston Common, GR 12/99.25. Wet moorland. N. D. Simpson, 1969, NMW (T. A. W.
Davis, 69/1231). (Nature Wales, 12: 38 (1970)).

605/17. JUNCUS SUBNODULOSUS Schrank *68, Cheviotland: near Cornhill, GR
36/85.38. G. A. & M. Swan, herb. G.A.S., conf. A. Melderis.

310 PLANT RECORDS

605/20. JUNCUS ALPINOARTICULATUS Chaix *79, Selkirk: shores of Akermoor
Loch, GR 36/41.21. Calcareous flushes, 1050 ft. R. W. M. Corner, 1968, BM, conf.
G. A. Matthews; Tima Water, GR 36/28.11. Sandy banks, 850 ft. R. W. M. Corner,
1969, herb. R.W.M.C, 2nd record. *80, Roxburgh: Murder Moss, near Selkirk, GR
36/50.28. 800 ft. R. W. M. Corner, 1968, BM, conf. G. A. Matthews. 2nd record.

607/3. ALLIUM SCORODOPRASUM. L. *80, Roxburgh: St Boswells, GR 36/58.30.
Established for more than 30 years in railway cutting. R. W. M. Corner, 1956, field
record.

625/6. EPIPACTIS PHYLLANTHES G.E.Sm. *28, W. Norfolk: Santon, GR 52/81.88.
Among ivy in conifer plantation. A. J. Davison, 1969, herb. E. L. Swann, conf. D. P.
Young as var. vectensis (T. & T. A. Stephenson) D. P. Young.

627/1. SPIRANTHES SPIRALIS (L.) Chevall. H6, Waterford: Ballatin, near Tramore,
GR 02/46. I. K. Ferguson, 1967, field record. Ur. Nat. J., 16: 96 (1968)). 1st post-1930
record.

628/2. LISTERA CORDATA (L.) R.Br. 79, Selkirk: south-west end of Akermoor
Loch, GR 36/40.20. R. W. M. Corner, 1966, field record. Ist post-1930 record.

643/3. DACTYLORHIZA INCARNATA (L.) So6 _*47, Montgomery: Llyn Mawr,
near Llanwnog, GR 32/00.97. R. Lovegrove, 1969, field record.
643/5. DACTYLORHIZA PURPURELLA (T. & T. A. Stephenson) Sod *42, Brecon:

near Llanwrtyd Wells, GR 22/88.41. Meadows by brook. W. J. H. Price & M. Porter,
1969, NMW. (Nature Wales, 12: 36 (1970)).

649/1. ARUM MACULATUM L. *+103, Mid Ebudes (Watsonia, 8: 152 (1970)).

652/2. SPARGANIUM EMERSUM Rehm. *H6, Waterford: shores of Carrickavrantry
Lake, GR 02/46. I. K. Ferguson (2086), 1967, BM, det. C. D. K. Cook. (Ir. Nat. J.,
16: 96 (1968)).

652/4. SPARGANIUM MINIMUM Wallr. *42, Brecon: Traeth Mawr bog, Brecon
Beacons, GR 22/96.25. M. Porter, 1969, NMW. (Nature Wales, 12:36 (1970)). H6,
Waterford: Ballin Lough, near Kill, GR 02/36. G. C. Argent, 1966, LANC. Ist
post-1930 record.

654/3. ERIOPHORUM LATIFOLIUM Hoppe *79, Selkirk: south-east of Akermoor
Loch, GR 36/41.20. Basic flush, 1050 ft. R. W. M. Corner, 1968, field record; north
side of Wedder Law, Ettrick, GR 36/26.14. Basic flushes. R. W. M. Corner, 1969,
field record. 2nd record.

656/3. ELEOCHARIS QUINQUEFLORA (F. X. Hartmann) Schwartz 79, Selkirk:
south side of Akermoor Loch, GR 36/40.20. Basic flushes, 1050 ft. R. W. M. Corner,
1966, field record. 1st post-1930 record.

660/1. RHYNCHOSPORA ALBA (L.) Vahl 47, Montgomery: near Glanmerin Lake,
Machynlleth, GR 22/75.99. D. E. M. Paish, 1967, field record. 2nd record.
661/1. CLADIUM MARISCUS (L.) Pohl *42, Brecon: Traeth Mawr bog, Brecon

Beacons, GR 22/96.25. 1100 ft. M. Porter, 1969, NMW. (Nature Wales, 12: 36 (1970)).
104, N. Ebudes: pool south of Brochel road, Raasay, GR 18/57.45. C. W. Murray,
1969, herb. C. W. Murray. 2nd record.

663/13. CAREX CAPILLARIS L. 96, Easterness: Coire Coulavie, Mam Sodhail, GR
28/11. 24. Basic flush at 3000 ft. R. W. M. Corner, 1961, field record. Major extension
in V.C.

663/15. CAREX PSEUDOCYPERUS L. *42, Brecon: marsh near R. Usk at Pencelli,
GR 32/09.24. M. Porter, 1968, NMW. (Nature Wales, 11: 139 (1969)).

PLANT RECORDS 311

663/17. CAREX VESICARIA L. 51, Flint: shore of Llyn Helyg, GR 33/17. 1 plant.
P. M. Benoit, 1969, NMW. 2nd record. *103, Mid Ebudes (Watsonia, 8: 153 (1970)).

663/20. CAREX RIPARIA Curt. H6, Waterford: marsh between Annestown and
Dunhill, GR 02/46. I. K. Ferguson (347), 1962, TCD. (/r. Nat. J., 16: 97 (1968)).
1st post-1930 record.

663/23. CAREX STRIGOSA Huds. *42, Brecon: near Llyswen, GR 32/14.37. Damp
woodland path. M. Porter, 1968, NMW. (Nature Wales, 11: 139 (1969)). *51, Flint:
Whitford Wood near Mostyn, GR 33/17. With C. sylvatica by side of track. P. M.
Benoit, 1969, NMW.

663/48. CAREX AQUATILIS Wahlenb. 79, Selkirk: confluence of Ettrick and Tima
Waters, GR 36/27.14. 750 ft. R. W. M. Corner, 1969, K, conf. S. S. Hooper. Ist post-
1930 record.

663/54 x 71. CAREX X BOENNINGHAUSIANA Weihe *67, S. Northumberland:
Tod Burn, GR 45/11.94. G. A. & M. Swan, 1969, herb. G.A.S., conf. A. Melderis.

663/55. CAREX APPROPINQUATA Schumach. *80, Roxburgh (Proc. bot. Soc. Br.
Tsl., 7: 562 (1969)).

663/62. CAREX DIVISA Huds. *68, Cheviotland: Holy Island, GR 46/09.43. G. A.
& M. Swan, 1964, herb. G.A.S., conf. A. C. Jermy. 1st definite record.

663/64. CAREX MARITIMA Gunn. 111, Orkney: Wideford Hill, St Ola, GR N30/
41.11. By tarmac road, 741 ft, probably introduced. E. R. Bullard, 1968, field record.
Highest record in the British Isles.

663/67. CAREX SPICATA Huds. *67, S. Northumberland: Maglin Burn, GR
45/10.98. 300 ft. G. A. & M. Swan, 1969, herb. G. A. Swan, conf. A. Melderis.

663/78. CAREX PAUCIFLORA Lightf. 104, N. Ebudes: bog by Loch Meodal,
Sleat, Skye, GR 18/65.11. A. McG. Stirling, 1966. 1st record for Skye.

669/2. GLYCERIA PLICATA Fr. H6, Waterford: near Tramore old water works,
GR 02/46. I. K. Ferguson (325), 1962, TCD. 2nd record. Ur. Nat. J., 16: 97 (1968)).

669/4. GLYCERIA MAXIMA (Hartm.) Holmberg *42, Brecon: near Llyswen, GR
32/14.37. Margin of pool. H. Soan & M. Porter, 1968, NMW. (Nature Wales, 11: 139
(1969)).

670/6 x 672/2. FESTUCA RUBRA L. X VULPIA BROMOIDES (L.) Gray #75018.
Suffolk: Shingle Street, Hollesley Bay, GR 62/34. 5 plants in thin sand over shingle.
P. J. O. Trist, 1969, K, det. C. E. Hubbard. Probably 2nd British record.

670/10. FESTUCA VIVIPARA (L.) Sm. *47, Montgomery: Cerrig-y-Duon, GR 33/
05.30. 2000 ft. A. Bull, 1966, herb. A.B. *67, S. Northumberland: Gilderdale, GR
35/69.47. On limestone crags, 850 ft and basalt by stream nearby. G. A. & M. Swan,
1969, herb. G.A.S. Ist definite record.

674/1. CATAPODIUM RIGIDUM (L.) C. E. Hubbard *42, Brecon: Cwm Clydach,
GR 32/22.12. Disused limestone quarries. M. Porter, 1968, NMW. (Nature Wales,
12: 36 (1970)).

676/9. POA COMPRESSA L. *103, Mid Ebudes (Watsonia, 8: 153 (1970)).
677/1. CATABROSA AQUATICA (L.) Beauv. 104, N. Ebudes: mouth of Lealt river,
Skye, 18/52. 60. H. J. B. Birks, 1969. 1st record for Skye.

681/2. MELICA NUTANS L. *79, Selkirk: by Tima Water below Meerless, GR
36/28.11. R. W. M. Corner, 1969, field record. 104, N. Ebudes: Tokavaig wood, Skye,
GR 18/61.12. A. McG. Stirling, 1968, herb. C. W. Murray. 2nd record.

700/1. CALAMAGROSTIS EPIGEJOS (L.) Roth *48, Merioneth: between Towyn
and Aberdovey, GR 22/59. Marshy ground. K. M. Stevens, 1969, NMW. Ist definite
record. 104, N. Ebudes: Tokavaig wood, Skye, GR 18/61.12. A. McG. Stirling, 1968
herb. C. W. Murray. Ist record for Skye.

*H

312 PLANT RECORDS

701/4. AGROSTIS GIGANTEA Roth 50, Denbigh: Rhyd-y-Crenau Field Centre,
Betws-y-Coed, GR 23/85. Garden weed. P. M. Benoit, 1969, NMW. Ist record for
over 100 years.

701/5 x 3. AGROSTIS STOLONIFERA L. x A. TENUIS Sibth. *28, W. Norfolk:
Cranberry Farm, Hockham, GR 52/93.93. P. M. Benoit & E. L. Swann, 1968; Derby
Fen, Gromston, GR 53/70.20. E. L. Swann, 1969. 2nd record.

707/5. PHLEUM ARENARIUM L. *103, Mid Ebudes (Watsonia, 8: 153 (1970)).

708/4. ALOPECURUS AEQUALIS Sobol. *42, Brecon: near Llyswen, GR 32/11.37.
Margin of moorland pool. M. Porter, 1968, NMW. (Nature Wales, 12: 36 (1970)).

Watsonia, 8, 313-322 (1971). 313

Book Reviews

Flore de la Belgique, du Nord de la France et des régions voisines. Edited by W. Mullen-
ders. Pp. xliv + 756, with 1 table, 1 folding map, 25 sets of line drawings and 14 text-
figures. Ed, Desoer, Liége. 1967 Price 420 B.Frs.

This new Flora of Belgium and northern France embraces a traditionally accepted
phytogeographical territory. In many respects it reads like a modern edition of Themis-
tocle Lestiboudois’s Botanographie Belgique, ou Flore du nord de la France, et de la
Belgique proprement dite (1827). However, the new Flora confines itself to the vascular
plants, and works from the family to the species by means of simple, but adequate
keys. In the case of critical taxa, these are accompanied by clear line drawings illus-
trating important diagnostic characters. In addition to the taxonomic description of each
genus, the Flora includes detailed floristic and geographical observations on most
species. Particular emphasis is laid on the vernacular names of species, their season of
flowering, life-form, ecology and geographical distribution. Subspecies and hybrids
receive rather arbitrary treatment.

An introductory analysis of the phytogeographical regions involved in the territory
covered is less than adequate, and the folding map is too complicated and poorly
reproduced. At the end of the Flora there is a well-illustrated glossary. The index is
satisfactory, and there are some blank pages for personal notes.

I am less than happy with the section on the collection and preservation of speci-
mens. I should have preferred one on the conservation of the flora. In my experience,
the herbarium botanist in France and Belgium has little enough regard already for the
preservation of the flora. I have too often seen the careless collection of rarities, such
as Osmunda regalis and the micro-endemic Biscutella neustriaca. A cautionary note
should be appended with rare species.

In some instances, the geographical notes on species may be found wanting. For
example, Globularia vulgaris is recorded as ‘doubtful’ for the district ‘Picardo-Braban-
con’. I know it at a number of sites in the Somme valley east of Amiens. Eriophorum
gracile is recorded for three districts, but not for ‘Maritime’, where it grows near St
Aubin (Pas-de-Calais). Aceras anthropophorum, Carex diandra, C. appropinquata and
Carex lasiocarpa should also be recorded in ‘Maritime’. Unfortunately, these minor
errors are too frequent.

The Flora will probably find a niche alongside the herbarium and in the study. It is
not quite an excursion Flora and is far from being a great reference work; but for
general reference purposes it may prove useful. I would recommend it as an introduc-
tion to the flora of Belgium and Northern France. The phytogeographical information
is of particular value in this respect. However, with the publication of the Flora
Europaea, some of the nomenclature and taxonomy will become obsolete very soon.

PuiLtre A. STOTT

A List of the Flowering Plants and Ferns of Carmarthenshire. R. F. May. Pp. 88.
West Wales Naturalists’ Trust Ltd. Obtainable from the Hon. Gen. Sec., West Wales
Naturalists’ Trust, 4 Victoria Place, Haverfordwest, Pembs. Price 50p. post free.

This booklet admirably fulfils its aim of collating the published records for this Flora-
less county, and many previously unpublished records seem also to have been included.
It lists all native and many alien species, brief ecological notes are given for each, and
localities, with references to the recorders, are given for the less common species. The
total number of species covered is given as 1117. Short introductory chapters on

314 BOOK REVIEWS

topography, plant distribution and the history of botanical exploration give practical
and detailed information, and Appendices 1 and 2, with details of records of over
70 unlocalised, lost or extinct species, should prove an irresistible inducement to further
activity. There is a useful list of the localities referred to, with grid references, and a
geological map which is unfortunately poorly reproduced.

The older records seem to have been thoroughly covered, with the exception of those
of Evans (1804), who mentioned 10 species of interest. Evans’s record of Asplenium
septentrionale is the only one for the county, for example, and his record of Spergularia
rubra from Ferryside pre-dates the cited Barker record by nearly a century. In other
cases old records known to the author (for example, Ray’s Juncus acutus from
Kidwelly, mentioned in the introduction) are omitted in favour of more recent ones from
the same locality; both together would have been more useful. Of more recent records,
that of Epilobium nerterioides is a notable omission (cf. Hyde & Wade 1957).

The author’s concept of nativeness is at the other extreme from that of Benoit &
Richards (1963), and a comparative reading of the lists for Merioneth and Carmarthen
from this angle is a puzzling experience. May unfortunately does not define ‘introduced’
as precisely as Benoit & Richards; he seems to follow Clapham, Tutin & Warburg
(1962), whose statement that a species is native implies that it is so in some part of
Britain, but not necessarily throughout its whole range, and in cases of doubt, such as
Mercurialis annua and Myrrhis, credits them with native status. Since it is to local
Floras that one turns for evidence on the nativeness or otherwise of a species, and since
this evidence may be only temporarily observable, a more independent and reasoned
approach would have been welcome.

The author modestly hopes that his list ‘will save much time if at any future date a
more scientific flora is ever contemplated.’ It must be added that it is just such lists as
his that become one’s inseparable field companion and are most likely to lead to
increased interest in a local Flora.

REFERENCES
BENoIT, P. M. & RICHARDS, M. (1963). A Contribution to the Flora of Merioneth, 2nd ed.
Haverfordwest.
CLAPHAM, A. R., TuTin, T. G. & WARBURG, E. F. (1962). Flora of the British Isles, 2nd ed.
Cambridge.

Evans, J. (1804). Letters written during a Tour through South Wales. London.
Hype, H. A. & WapeE, A. E. (1957). Welsh Flowering Plants, 2nd ed. Cardiff.

A. O. CHATER

The Victorian Fern Craze, a history of pteridomania. David Elliston Allen. Pp. 85,
illustrated in black-and-white. Hutchinson, London. 1969. Price £1-05.

For over 30 years I have lived close to ferns and fern men — and to fern books. The
books have a great fascination, with their wealth of information and the enthusiasm
of the many authors who churned them out in vast numbers during the latter half of
the 19th century and gave a very vivid picture of the ‘fern craze’. I thought I knew a
lot about its history until I read this book by David Elliston Allen who has skilfully
captured the picture of the period and made it live in his pages. We feel the excitement
of the hunt for new fern varieties, bad ones and good ones, and their subsequent
development in garden and greenhouse, and above all we feel the excited urge for more
and more information and the avid seizing of each new book as it poured from the
presses. The author is to be congratulated on the very successful outcome of the large
amount of labour and research which has gone into this production, bringing into
small compass so much new and interesting information.

The subject matter would not seem to lend itself to the making of an interesting and
successful book and we would expect it to be a dry recital of many facts. The first
stimulus to the fern cult was the discovery of reliable spore-raising methods, which

BOOK REVIEWS 315

encouraged the growing of many exotic ferns and launched the fern trade. Next came
the Wardian Case, which provided the growing conditions for easy culture, and this
was followed by an awakening interest in our British ferns and their varieties. Then
came the books in ever-increasing numbers until the end of the century, when the
‘craze’ was fast declining. The story is written by Mr Allen in a graphic manner which
holds our interest, and sums up the whole period, bringing its fevered craziness into
true perspective.

To the present-day pteridologist the book is a revelation-—and a warning! The
wind of commonsense blows through it. Ferns are still an absorbing subject with the
power to obsess, and with today’s revival of interest in them we need the antidote
provided by Mr Allen, who signposts the dangers and the absurdities which can still
beset us, and helps us to preserve a balanced outlook on ferns.

This delightful small book should be read by everyone interested in ferns. It is
excellently produced by Hutchinson, and has several fine black-and-white illustrations
taken from the literature of the period. It is certainly very good value for 21s.

J. W. Dyce

Flowering Plants, Origin and Dispersal. Armen Takhtajan. Pp. x + 310. Oliver &
Boyd, Edinburgh. 1969. Price £2-50.

Although this book is basically an English translation of The origin of Angiospermous
plants, 2nd ed. (Moscow, 1961, in Russian), the text has been revised to such an
extent that it is in fact a new work. As such it is particularly useful, as it provides a
key (both by text and bibliography) to works in Russian on the topics under considera-
tion — topics that seem to have provoked as much discussion in that language as they
have in other European ones. Those of us who do not have a fluent reading knowledge
of Russian must therefore be grateful to Mr Charles Jeffrey for his most readable
English translation.

Takhtajan approaches his subject from many different angles and succeeds in form-
ing a picture — orthodox in many respects, but none the less fascinating — of a group
of plants originating at some time well before the Cretaceous from some as yet unknown
Pteridosperm with primitive secondary xylem and bisexual strobili. He explains the
‘gap’ in the fossil record in terms of population genetics, i.e. of small transitional
populations in which evolution could operate at an accelerated rate, and repeatedly
emphasises the evolutionary importance of neoteny (e.g. the concept that the flower is
a neotenic form of the ancestral Gymnosperm strobilus).

The first flowering plants are depicted as woody, evergreen and possibly pachycaul.
As each character is examined in detail and the evidence considered, a picture is gradu-
ally formed of a plant with many Magnolialean characters but more primitive in some
respects (e.g. with a perianth of bracts). The viewpoint is very similar to that given by
Cronquist in The evolution and classification of flowering plants, which was recently
reviewed in this Journal, i.e. it is based on the ideas of Bessey and Arber & Parkin that
the Magnolia type of flower is primitive and basic to the whole of the Angiospermae.
Little consideration, if any, is given to more recent theories that cast doubt on the
validity of the orthodox view, although these can often provide more convincing
explanations than can the classical theory. Thus, while Takhtajan finds it difficult to
account for the occurrence of both extrorse and introrse anther dehiscence in primitive
families, and therefore thinks lateral dehiscence to be primitive, Melville’s Gonophyll
Theory provides a simple explanation: where the ‘foliar’ stamens are not regarded as
primitive but as derived from a ‘typical’ stamen with its subtending foliar organ. If
this is so, the ‘typical’ stamen with filament and anther has not gone through a ‘foliar’
stage, and the differences in dehiscence were fixed before the foliar type evolved.

After an interesting and illuminating discussion of primitive families containing
‘living fossils’, the author indicates (i) how the various apetalous Dicotyledons are

316 BOOK REVIEWS

thought to have been derived from different families in which petals are present and
then (ii) how the Monocotyledons, in his view, are derived monophyletically from the
Nymphaeales (without discussing the serious difficulties involved in this theory). Chap-
ters then follow on the time and place of origin of the Angiosperms and their sudden
apparent evolutionary radiation, in which a tropical, possibly S.E. Asian origin, well
pre-Cretaceous in time and a subsequent radiation into temperate regions are sug-
gested. This radiation is thought to have involved migration by means of land bridges,
and the words ‘Continental Drift’ are not even mentioned.

To the main text are added two appendices, of which the first is a useful synopsis
of Takhtajan’s own classification of flowering plants, published in Russian in 1967,
and the second gives the floristic regions of the world with the endemic and otherwise
prominent families and genera of each.

All in all this is a book that will fascinate anyone interested in the evolution and
distribution of the Angiosperms. With due allowance for the author’s occasional
dogmatic tendencies, it should also be a useful university text-book.

N. K. B. RoBsoN

Wild Flowers of Greece. C. N. Goulimis. Edited by W. T. Stearn. Pp. 212, with
104 colour plates by N. A. Goulandris. The Goulandris Botanical Museum, Kifissia,
Greece. 1968. Price £26.

This handsome volume is a worthy and fitting memorial to one of Greece’s outstanding
amateur botanists of the last two decades, Constantine N. Goulimis, who died in 1963.
Like his British predecessor, the amateur botanist Shirley Clifford Atchley, he left an
uncompleted manuscript of his work, which in this instance has been edited and com-
pleted by William T. Stearn of the British Museum. Both Atchley’s Wild Flowers of
Attica, and Goulimis’s Wild Flowers of Greece have therefore benefited from the
hand of a professional botanist before publication.

Dr Goulimis was by profession a lawyer, and one of the leading lawyers of Greece,
and at the same time he was for 44 years the legal adviser to the British Embassy in
Athens. It was comparatively late in life—in his 60th year — that his interest in the
Greek flora suddenly flourished, and from 1946 until his death in 1963 he made num-
erous journeys into remote parts of Greece and to the Islands, adding not only 230
new species to the Greek list, but discovering a number of species new to science,
including Anthyllis serpentinicola, Campanula goulimyi, Inula serpentinica, Linaria
hellenica, Linum goulimyi, Crocus goulimyi, Silene goulimyi and Tulipa goulimyi. This
is surely unprecedented in the twentieth century: a self-trained amateur starting his
botanical life at sixty and discovering in Europe so many species new to science,
particularly when his terrain had been worked botanically by such great European
botanists as Tournefort, Sibthorp, Boissier, Halacsy, Hayek, Turrill and K. H.
Rechinger, to mention but a few.

The richness of the Greek flora compares only with that of the Iberian Peninsula in
Europe. Greece, though a quarter the size, has probably one and a third as many
species of flowering plants again as France, for example. Goulimis estimated that
when the more remote areas had been fully explored ‘the number of 6,000 will be
considerably exceeded’. He must have known the Greek flora better than anyone of
his time. He describes how he has made more than 220 expeditions, great and small,
in search of flowers, and has climbed more than 70 Greek mountains, often on more
than one occasion, on his explorations. In his fascinating introduction to the book
he describes some of his favourite hunting grounds and the plants he has discovered.
For my own part, I wish that this section were many times longer and more detailed,
for here lies the heart of the matter, and Dr Goulimis’s real contribution to our
knowledge of probably the most interesting flora in Europe.

In 1954 Dr Goulimis began to plan a series of illustrated volumes on the Greek flora

BOOK REVIEWS 317

in collaboration with Mrs Niki Goulandris, who painted the plants as he brought
them in from the field, sometimes illustrating new, unnamed species. Her fine water-
colours of these plants, so ably reproduced in Athens, give a great distinction to this
volume. There is a total of 110 species reproduced in full colour plates, selected from
a wide range of families.

Widespread and familar lowland species, such as Epilobium angustifolium, Rosa
canina, Globularia alypum and Euphorbia characias subsp. wulfenii, stand page to page
with such new and rare species as Crocus goulimyi, Fritillaria davisii, Staehelina arborea,
Centaurea lactucifolia and Campanula formanekiana. With each species there are
short notes on its distribution in Greece, in particular listing those localities where
they have been discovered by Dr Goulimis. Notes on their medicinal and other ‘uses’,
both ancient and modern, are included, as well as their habitats and vernacular names.
Only new species are described in detail. It is with the selection of species that my main
criticism lies; there seems no reason for the odd assembly of rare and common plants,
unless as I suspect, they have been chosen to supplement the ten volumes of Sibthorp
and Smith’s Flora Graeca and the magnificent paintings of Ferdinand Bauer.

Few, alas, will be fortunate enough to use Wild Flowers of Greece in conjunction
with the senior volumes. Be that as it may, Dr Goulimis’s volume is indeed a landmark
in Greek botany — the first book of any stature from the pen of a Greek botanist to be
published in recent years. It may well prove to be more than this, for it is also the first
publication of the Goulandris Botanical Museum, founded in 1963 by Mr and Mrs
Angelos Goulandris as a centre for the study of the Greek flora. It has set a fine
example of both scholarship and production, and it is a worthy beginning to what we
all hope will be a long series of works by contemporary Greek botanists.

O, POLUNIN

Plant Anatomy. A. Fahn. Pp. viii + 534. Pergamon Press, London. 1967.
Priceres-i 5:

During the last 20 or 30 years British students have had to rely on a number of Ameri-
can text books of plant anatomy. However excellent such works are, it is a pleasure
to welcome this new book which presents the student with a somewhat different
approach and outlook. Professor Fahn’s book was written originally in Hebrew and
has been revised during translation for this English edition.

The first half of the book deals with the various kinds of cells and tissues which are
to be found in the vascular plants (Tracheophyta). A clear and concise chapter on the
cell is followed by a chapter on meristems and apices, which is an attractive presenta-
tion of histological and dynamic aspects including an evaluation of recent research
and citing about 100 references.

Seven chapters — Parenchyma, Collenchyma, Sclerenchyma, Xylem, Phloem, Lacti-
cifers, Epidermis - make up the section on Mature Tissues. They contain a great
amount of detailed information, clearly presented and well illustrated on structure and
development with considerable emphasis on economic and evolutionary considerations.
There is, for example, a valuable discussion on the evolutionary trends in the develop-
ment of tracheary elements in various angiosperm groups. The chapter on the epider-
mis, with its detailed consideration of stomatal types, trichomes and other surface
appendages, is of special interest to taxonomists.

The next two sections are entitled ‘Primary Vegetative Body of the Plant’ (three
chapters) and ‘Secondary Body of the Plant’ (five chapters). Here again there are various
features worth drawing attention to such as the account of adaptation to desert and
saline habitats shown by stems, and an example of a key for the identification of trees
and shrubs using features of the secondary xylem.

The last section of the main text, which is concerned with the reproductive organs,
comprises a long and detailed chapter on the flower and two shorter chapters on the
fruit and the seed. That on the flower covers such topics as ontogeny, vascularization

318 BOOK REVIEWS

(which is particularly well treated), phylogeny, structure and development of the
stamen, pollen grain development, phylogeny and histology of the carpel, sporogenesis,
an extensive survey and classification of nectaries, and the formation of endosperm
and the embryo. It contains a great amount of information, most of it up to date,
although the section on pollen grains, especially with regard to the pollen wall, is some-
what superficial, and makes no references to research published during the last 10—
15 years.

In the chapter on the fruits it is good to see a distinction made between the achene
(e.g. Ranunculus) and the cypsela (of Compositae). Most recent anatomical texts treat
the fruits of Compositae as achenes although, as Fahn points out, they are formed
from an inferior ovary and surrounded by other floral tissues in addition to the ovary
wall and are therefore false fruits (pseudocarps).

There is a glossary of some 400 terms, as well as indices to authors and subjects.

Professor Fahn has written a valuable and enjoyable book which should be widely
used by undergraduates in British Universities. Many of the examples and illustrations
are different from those they will normally encounter in American texts, making a
refreshing change. The translation by Mrs Sybil Broido-Altman reads very well. The
book is well produced and printed on coated paper, the printing having been done in
Hungary. It is good value at 75s.

V. H. HEYwoop

Die Farnpflanzen Zentraleuropas. K. and H. Rasbach (photographs) and O. Wilmanns
(text). Pp. 295. Quelle & Meyer, Heidelberg. 1968. Price DM 38.

This is one of the most beautiful botanical books I have ever had the pleasure of
reviewing. A comprehensive introduction covers the life-cycle and evolution of ferns
and ‘fern-allies’. There is also a section on hybridisation and the biogeography of the
group. However, valuable as Dr Wilmanns’ account is, it is the systematic section,
which comprises about three-quarters of the book, which will undoubtedly prove the
major attraction of the work. Seventy-seven of the ninety Pteridophytes of Central
Europe are illustrated by superb black and white full-page photographs, taken in such
a way that the essential characteristics of each species are caught. That they can be
compared to pictures taken in this country by Michael Proctor ought to be a sufficient
indication of their standard. By paying particular attention to lighting and back-
grounds the Rasbachs have produced some startling effects: you can almost feel the
rigid texture of Asplenium fissum (p. 137), whilst the picture of Ophioglossum vulgatum
(p. 165) suddenly makes the resemblance to a snake remarkably plain. The three
moonworts, Botrychium lunaria, B. matricariifolium and B. virginianum, are so life-
like that they seem to be growing on the page. Probably most successful of all are the
species of Lycopodium sensu lato. The clear distinction between the sporophylls of
Diphasium complanatum and D. issleri is vividly brought out, which makes the absence
of mature sporophytes of this complex in Britain even more frustrating.

The plants are arranged according to a broad habitat classification: cliffs, scree,
alder woods, beech woods, open water and so on, and each habitat is introduced by
a typical landscape including one or more of the species to be illustrated later in detail.
Whilst this does help to relate species to habitat in a telling way, it does detract from
the book as a work of reference. It is aggravating to have the genus Equisetum scattered
in at least three places. The habitats and the systematic material could have been
separated without in any way reducing the value of the former.

There is a synopsis of classification which closely follows Flora Europaea in its treat-
ment, but analysis shows that the thirteen species for which there are no illustrations
are not mentioned elsewhere. It is unfortunate that lack of a picture should also mean
lack of a description, as it tempts the term ‘Kaffee-tisch,’ which would be a gross under-
estimate of the full value of this book.

F. H. PERRING

BOOK REVIEWS 319

A Guide to the Naming of Plants: with special reference to heathers. David McClintock.
Pp. 38. The Heather Society. 1969. Price 324p

Mr McClintock is one of the few botanists to have successfully bridged the gap that
exists between botanical theory and horticultural practice. His knowledge of both
spheres and of the problems and misunderstandings which have long prevented their
complete agreement is succinctly but sympathetically demonstrated in this little book-
let.

It is 17 years since the first edition of the International Code of Nomenclature for
Cultivated Plants was published and today, with the fourth revised edition now
available, I doubt that many copies are to be found on the bookshelves of gardeners.
Certainly few nurserymen possess a copy, and of those that do only a handful have
attempted to put its rules and regulations into practice.

Nurserymen and gardeners are normally much too busy growing plants to be
bothered with codes of nomenclature and the likes, but it is these people, the producers
of new cultivars, the hybridists and the mysterious men with an eye for a ‘good ’un’
who must benefit from the Code in the long run. As far as practising gardeners are
concerned the Code fails in that it assumes too much, and this is where Mr McClin-
tock’s question and answer method of explaining the basic facts is so very much more
digestible. He succeeds in putting across in a simple yet accurate manner important
aspects of what is to many people a most confusing subject.

The booklet is divided into three parts. The first sets out the chief principles on which
the correct naming of plants is based; the second exemplifies these in the naming of
hardy heathers (which include Calluna, Erica, Daboecia, etc.); and the third lists
recommendations of the Heather Society on certain cultivar names.

The type is clear and the sections arranged in a simple, uncluttered manner; this, plus
an excellent index, makes the booklet easy to refer to and enjoyable to read.

I have only one minor criticism. I do not agree with the coining of new English
names for plants which have no available established one. Our friends the Americans
are particularly fond of this unfortunate albeit amusing pastime. Established English
names have been with us a long time and their origins are often part of a rich heritage
of folk-lore and history, but to invent a new English name for a species perhaps already
well known by its botanical name seems pointless. It gives the idle gardeners amongst
us an unnecessary alternative. Examples of these names in the booklet are thankfully
few but include ‘Balkan Heath’ (Bruckenthalia spiculifolia), “Winter Heath’ (Erica
carnea), ‘Williams’ Heath’ (Erica x williamsii) and ‘Menziesia’ (Phyllodoce caerulea).
The last is particularly unfortunate in view of the existence of the genus Menziesia,
several species of which are cultivated in British gardens. I must emphasise that this
is only a personal criticism and in no way detracts from the value of the booklet to
the user, be he botanist, gardener or layman.

The Heather Society is to be congratulated on its foresight in commissioning the
work and in choosing such a ‘natural’ author.

Roy LANCASTER

The Oxford Book of Food Plants. S. G. Harrison, G. B. Masefield & Michael Wallis.
Pp. viii + 206. Oxford University Press, London. 1969. Price £2-75.

This is another volume of the successful series of Oxford books on plants and animals.
Like its companion volumes on Wild Flowers, Garden Flowers and Flowerless Plants
it is an exceptionally fine book, and there is certainly a need for it. In an age of tech-
nological advancement where people become, to an alarming degree, ignorant of the
natural resources on which their lives depend, books of this kind serve as educational
tools of great importance.

The main purpose of this work is to depict and describe the plants which provide
food for the human race. On 87 colour plates about 420 useful plants (species, varieties
or cultivars) have been magnificently illustrated by B. E. Nicholson. (Incidentally,

320 BOOK REVIEWS

here is one more proof that even today the artist’s rendering of plants is far superior to
anything which colour photography has to offer.) Special emphasis has been given to
the organs of plants which are actually consumed as food, but in many instances the
whole plant has been illustrated. There are not many books on the market in which the
same high degree of colour fidelity has been achieved. Each description is printed
facing the relevant illustration, which is convenient for the reader.

For each plant, information is given about its geographical distribution, origin,
morphology and value as a nutrient. Also the most important particulars about both
cultivation and processing methods are given. The length of the description and the
amount of information it contains varies according to the importance of the plant.

Although one does not expect a cookery book, in some instances the reader may
expect to be told more about the usefulness of certain plants. The reviewer was very
unhappy that even this book did not enlighten him on why Monstera deliciosa deserves
its specific epithet. For goodness’ sake, how is it eaten? To stay with this plant, is the
statement that it is ‘the only genus of plants which has natural holes in the leaves’
really correct ?

The arrangement follows a classification of foods rather than a botanical one, begin-
ning with cereal crops and followed by oil-producing plants, nuts and legumes; spices,
salad plants, leaf vegetables and root crops form the final sections. General topics
of interest are treated in three concluding chapters (the domestication, spread and
uses of food plants).

An illustrated glossary of botanical terms at the beginning of the book will assist
the less initiated user of the work. The figure given for the term bipinnate is not quite
correct since the pinnae are not completely divided, but it would be pedantic to argue
about such minor mistakes in a book of this nature.

In conclusion, this delightful book can be thoroughly recommended to anybody,
be he just curious, a lover of plants or, like the reviewer, interested in all possible

aspects of ‘gastrosophy’. EDMUND LAUNERT

Nightshades. The paradoxical plants. Charles B. Heiser, Jr. Pp. 200. W. H. Freeman &
Co., San Francisco. Price $5.95.

Too scarce are those who both know their subject expertly and can, or do, write or
broadcast well about it. Consequently much that is published or heard is by those
who can talk or write (or the publishers think they can), and know enough to get by.
Perhaps this is the lesser evil, when we see how heavily and obscurely some specialists
express themselves. But in this book we have a specialist who can, and does, write well.
Professor Heiser, we are told, has published over 50 scientific papers (although I am
never sure at what stage any given piece of writing qualifies for this designation) and
the bibliography includes some of his on Solanums. Internal evidence too shows his
first-hand knowledge and investigations into their family.

There are nine chapters, each devoted to one or a group of Solanaceous plants,
such as Peppers (Capsicum spp.), Potatoes, Egg Plants, Tomatoes, the Wonderberry,
Mandrakes, Jimsonweed (anglice Thorn Apple, for this is an all-American book),
Henbane, Tobacco, Lulos and Pepinos (know what they are?), and some garden
annuals. The author discusses their history, uses, composition, variation, even their
nomenclature, and covers widely each plant in a way to be understood by the general
reader. Even the more expert might well learn from what he relates, sometimes with
dry humour — ‘The superior ovary — this refers to the position of the ovary, not its
quality’. Most of the taxa discussed are American, in origin at any rate. But this is no
drawback to the lessons to be learnt from this book, perhaps particularly in the story
of the Wonderberry; and ‘certain principles first delineated in Datura and other
‘worthless’ plants have been of tremendous importance in the understanding of all
plants, including those of great direct importance to Man’. Not all is botanical, for
example the history of cigarettes, but this fairly broadens the interest. I regret only that

BOOK REVIEWS 371

the author did not discuss some plants (e.g. Jerusalem Cherry) more, or include others
such as Nicandra, or Woody Nightshade (the origin of the name Nightshade remains
a mystery); and that it should be out-priced, at any rate to sell well in the British
market — but would it not make an excellent paper-back ?

Davip MCCLINTOCK

The Lowland Grasslands of Co. Limerick. A. M. O’Sullivan. Pp. 57 + 6 plates and
3 maps. An Foras Taluntais, Irish Vegetation Studies 2, Dublin. 1968. Price 25p.

This is a detailed analysis of grassland communities based upon 110 semi-random
samplings made over a wide area of lowland County Limerick, and conducted accord-
ing to the methods of the Ziirich-Montpellier system of phytosociology.

All the communities enumerated are considered to belong to the major class of
European lowland grasslands, the Molinio-Arrhenatheretea, within which two orders,
three alliances, four associations (Lolio-Cynosuretum, Centaureo-Cynosuretum, Sene-
cioni-Juncetum acutiflori, Junco acutiflori-Molinietum) and three sub-associations of
Centaureo-Cynosuretum are recognised. The associations and sub-associations are in
turn surveyed from the point of view of floristic composition and variation, phenology,
distribution and ecology, management and improvement, and the findings are set out
briefly in a final summary.

The job is very competently done, and the conclusions may be very useful in grassland
research, but I cannot help feeling that Mr O’Sullivan would have done better had he
been freed from the intolerable restrictions of the phytosociological straitjacket, and
allowed to roam the verdant acres of Limerick guided by his own eyes and native
commonsense.

Plant societies are of absorbing interest, and phytosociology will one day engage
the sedulous attention of our best botanical intellects; but the Ziirich-Montpellier
line went astray when it hitched its wagons to that grand old puffer we call formal
taxonomy.

R. D. MEIKLE

Flowers of Europe. Oleg Polunin. Pp. 662, 192 pp. colour photographs, 50 plates of
line drawings, and text figures. Oxford University Press, London. 1969. £4-20.

It is easy to be unkind about this book, and to point out that 2,600 descriptions, how-
ever judiciously supplemented by line drawings and colour photographs, cannot con-
ceivably provide fully informative guidance to the vast, varied and intricate flora of
Europe. Five large volumes, without illustrations, are not likely to accomplish this to
everyone’s satisfaction. Mr Polunin’s book is not intended for the expert, but for that
much-appealed-to class of person, the intelligent tourist, who will find in these pages
as much botanical information as, or possibly more than, he is likely to be willing
to digest.

From the viewpoint of book-production, author, publisher and printer are alike
to be congratulated. The colour photographs are mostly of the highest quality and
well reproduced, though I am still not convinced that photography is a satisfactory
substitute for line drawings. Photographs, however good, too often fail to reveal those
very features which are essential for accurate determination, and which an accom-
plished botanical artist will take care to stress. Some of the ‘close-ups’ in this volume
(e.g. 70 Osyris alba, 363 Eruca vesicaria, 1215 Scrophularia scorodonia, 1216 S. hoppii,
1282 Plantago media) afford virtually no indication of the general appearance of the
plant, while others (e.g. 669 Euphorbia dendroides, 1193 Verbascum thapsiforme) are
taken at a distance which lends enchantment to the view, but precious little in the way
of taxonomic data. The text figures are, on the other hand, uniformly lucid, accurate
and helpful.

The text, in a book of this kind, might almost be said to be a minor consideration. The

322 BOOK REVIEWS

author has made few concessions to the novice. Descriptions are terse and technical,
with the odd appended note of economic, medical or general interest to assuage the
general aridity. I have not tested the keys to families and genera, but they look prac-
tical enough, though, again, an entry such as ‘Valves rounded at the back; median
nectaries absent; lateral nectaries prismatic’ may test the patience of the tourist, what-
ever his I.Q., to somewhere near breaking-point. The species are not individually
keyed, probably not a serious omission where there are group headings and abundant
illustrations as well as descriptions. So many species have had to be passed over in
silence that a key, at this level, might be more misleading than helpful.

In brief, it is a handsome book, at a very reasonable price, but it tries to cram too
much into one nutshell; and the nutshell may well have become too heavy for those
who travel light in permanent fear of the excess baggage exactions inexorably demanded
by the grim-faced servants of the aircraft corporations.

R. D. MEIKLE

Photosynthesis. Isaac Asimov. Pp. 193. George Allen & Unwin, London. 1970.
Pricereieioe

This book is a popular but by no means superficial account of the process upon which
virtually all life at the present day ultimately depends. In fact, ‘Photosynthesis’ is too
narrow a title, because the authors consider the whole of the carbon cycle — both the
energy-fixing process of photosynthesis and the energy-releasing process of respiration.
Dr Asimov, a chemist by training, is well known within his own field as a biochemist,
as well as enjoying a considerable reputation as a scientific populariser and science-
fiction writer. The present book, written in a readable and uninhibited style, is evidently
aimed at the intelligent layman. The essential facts and concepts are lucidly set out,
at a pace, and with a degree of background explanation, which seem nicely judged for
a reader who may have only limited scientific knowledge — though it would be idle to
pretend that a completely non-scientific reader is likely to find this an easy book. The
author has achieved more than merely a sound elementary exposition of our present
understanding of the biochemistry of the carbon cycle. He has written with a sense of
the wider relevance of the mechanisms and processes he describes, and of the human and
social origins and ends of science. Only a few points left me a little uneasy. I am sure
the author could have contrived to explain more lucidly and specifically why the
heat of combustion of glucose (673 kcal/mole, p. 61) is different from the standard
free-energy change for the same reaction (686 kcal/mole, p. 67). Also, I know that I
(as a non-biochemist) am not alone in finding the molecular changes of the Krebs and
Calvin cycles peculiarly fascinating, and I should have liked rather more detail here.
However, these are small criticisms of an admirable piece of popular scientific writing,
and the reader whose interest is aroused will be able to pursue these and other questions
elsewhere.

M. C. F. PROCTOR

Index Kewensis Plantarum Phanerogamarum. Supplementum XIV (1961-1965). Com-
piled at the Herbarium of the Royal Botanic Gardens, Kew, under the direction of
Sir George Taylor. Pp. 149. Clarendon Press Oxford: University Press, London.
1970. Price £6-50.

The latest supplement of Index Kewensis makes this invaluable work complete as far
as the end of 1965. ‘Complete’ is possibly an over-statement because, like its pre-
decessors, this part includes some names that were omitted from previous supple-
ments. It is nevertheless to all intents and purposes a complete index to the descrip-
tions of all Phanerogamous species published from 1961 to 1965 (inclusive). Would
that the same information were available for subspecies and varieties.

N. K. B. ROBSON

Watsonia, 8, 323-338 (1971). 323

Reports

REE Ok OF THE COUNCIL FOR 1969

(This report and the audited accounts relate to the calendar year 1969.
Figures in parentheses are the 1968 ones, for comparison. It is a reprint
of the report circulated to members in April 1970)

A number of important events have occurred in the Society’s history in the year 1969.
The last numbers of Proceedings and Watsonia appeared before their amalgamation
as the ‘new Watsonia’; the Treasurer has launched the Covenant scheme; and a
professional agent has been appointed to handle the sales of publications.

The membership figures have held at a reasonably steady level during the year with
119 (128) members joining the Society, and a loss in membership of 136 (89) by reason
of death, resignation or the operation of Rule 28; thus the year ended with a member-
ship total of 1716 (1733) representing a net loss of 17 (net gain 39). These results are
not as discouraging as it might at first seem, considering the effect of the increase in
the subscription rate, a factor which has inevitably accounted for some of the loss in
numbers. It is with great regret that we record the deaths of Rev. W. Keble Martin and
Mr G. C. Brown who had been a member of the Society since 1911.

The accounts of the Society are appended on pages 328 to 329. Despite the increase
in the rate of subscription for Ordinary members by 10s. to £2, expenditure for the
year exceeds income, and there is a deficit on General Income and Expenditure Account
of £22. Although they have received several reminders, about 125 members have still
not paid the additional 10s. due for 1969; if these dues had been received, the deficit
would have been covered.

Whilst expenditure generally continues to rise in accordance with forecast, certain
exceptional items were incurred during the past year. In particular, the charge for
journals covers approximately four parts of either Watsonia or Proceedings, whereas
in future the annual charge will relate to two parts only of the new Watsonia plus one
issue of the supplement Abstracts from Literature. Other exceptional items are shown
in the accounts but also general expenses have been unusually high because of the
heavy stationery and postage costs (at higher rates than heretofore) incurred in remind-
ing members of the increased rate of subscription.

In November, the Hon. Treasurer wrote to all Ordinary members inviting them to
enter into Covenants in respect of their annual subscriptions to the Society. The response
has been gratifying; by the end of the year well over 200 members had signed Coven-
ants. Thus we can expect to recover more than £300 p.a. in future in respect of income
tax. It is hoped that any member who is in a position to do so, and who has not already
done so, will sign and send to the Treasurer a covenant form. (Further copies can be
obtained from the Treasurer if required.)

Proceeds from sales of journals exceeded the figures of the previous year, this being
a reflection of the increased number of parts published. Sales proceeds, royalties re-
ceived and expenditure relating to other publications sponsored by the Society are set
out in the account of the Publications Fund. Sales of British Sedges have continued
at a satisfactory rate and justified the printing of a further 500 copies before the type
was broken up; stocks at the end of the year amounted to just over 500 copies and it
seems likely that these will be exhausted within the next year or two.

With effect from Ist January 1970, the Society has appointed E. W. Classey Limited
(Natural History Publishers, Booksellers and Literary Agents) of 353 Hanworth Road,
Hampton, Middlesex, to act as their agents for the sale of their journals and publica-
tions (other than those handled by other publishers). This change has been made partly

374 REPORTS

to relieve the over-burdened voluntary officers who have undertaken this work in the
past and partly because it is hoped that, in time, the organisation and publicity of a
professional organisation will help to increase sales. In the early years, however, the
net receipts by the Society will decrease by reason of the charges which E. W. Classey
Limited must necessarily make for their services. The thanks of the Society are due to
D. H. Kent for handling all orders for our publications during the year and in many
previous years.

Mention must be made of the working parties set up from Council. That set up to
report on a ‘List of recommended English names of wild plants’ has prepared a very
full draft which it is hoped will be published in the near future. The party investigating
proposed collaboration with European botanical societies is continuing its discussions.

THE MEETINGS COMMITTEE arranged as the Society’s main event of the year, a very success-
ful conference in September on “The Flora of a Changing Britain’. This was attended
by some 150 delegates and held in Imperial College by kind permission of Professor
C. P. Whittingham. The Society is much indebted to Dr D. H. Dalby for his help in
organising this conference which consisted of two days of lectures and discussion and
concluded with an enjoyable excursion on the Sunday to the Forestry Commission’s
lands at Alice Holt, Hants and the Surrey Naturalists’ Trust’s Reserve at Thursley
Common.

The Annual General Meeting was held in conjunction with a meeting of the North-
west Region at Merlewood Research Station, Grange-over-Sands by kind permission
of the director, Mr J. N. A. Jeffers. Thirty-six members were present to hear a series of
lectures relating to the flora of the North-west Region and in the evening Mr S. Jeeves
of the Council for the Protection of Rural England gave an excellent illustrated lecture.
A field excursion to Roanhead Dunes and Roudsea Wood National Nature Reserve
was held on the Sunday.

In addition to the two above-mentioned field excursions the Committee arranged
six field meetings all of which were well supported, none having less than 20 members
present and the average attendance being 35. These meetings were at Guernsey, Mid-
hurst in Sussex (with the Sussex Naturalists’ Trust), Flatford Mull, Leicestershire, East
Norfolk (with the Norfolk and Norwich Naturalists’ Society) and London (with the
London Natural History Society). The Midland Region meeting at which a repre-
sentative was elected to fill the vacancy on Council was held in conjunction with the
Leicestershire field meeting.

A successful one-day conference entitled ‘Hedges and Local History’ arranged by
the Society and the Standing Conference for Local History was held in June at Monks
Wood Experimental Station by kind permission of the director, Dr K. Mellanby. Our
Patron wrote expressing her interest in the conference and extending greetings to the
Society.

The Committee also organised the Annual Exhibition Meeting in November, held
as usual in the Department of Botany, British Museum (Natural History) by kind per-
mission of the Keeper of Botany, Mr R. Ross. Thirty-five exhibits of a very varied
nature were staged and there was a good attendance of members and guests. Sixty-eight
stayed for the Reception afterwards; an encouraging number considering the very
inclement weather, a heavy and unusually early snowstorm.

THE JUNIOR ACTIVITIES SUB-COMMITTEE arranged six field meetings in Britain in 1969,
four of which were cancelled due to lack of support, these being those at Edinburgh,
Perthshire, Kidderminster and Oxford; the other two (S. Essex and Northumberland)
had a total of nine participants. This very poor response has resulted in the number of
Junior day field meetings planned for the coming year being greatly reduced. Steps
are being taken to arrange in future years more joint meetings with other bodies
organising field activities for juniors, and residential meetings at home and abroad.

REPORTS 325

THE DEVELOPMENT AND RULES COMMITTEE had been in some doubt about its continued
usefulness and on a recommendation from Council the Committee met during the year
and discussed its function and future. It was finally decided with great regret that the
present Committee should be disbanded and a Co-ordinating Committee set up of the
officers of the Society (excluding the editors), the secretaries of the permanent working
committees and a member of Council to act as Secretary. The new committee is to
prepare business for Council, consider policy on general matters and make recommend-
ations to Council for consideration. J. F. M. Cannon has agreed to act as secretary of
the Co-ordinating Committee.

THE PUBLICATIONS COMMITTEE has given considerable thought to the future of the
Society’s periodical publications and has reported that it can see no justification for
continuing to have two periodicals. The needs of today could be served more effectively
by one journal, an amalgamation of the present two. This new journal will appear
twice a year and will contain all that has in the past been included in the two publica-
tions with the exception of Abstracts from Literature which will be published annually
as a supplement. The year saw preparations for this amalgamation in the appearance
of the last part of Proceedings and three parts of Watsonia.

Plans for the Society to publish companion volumes to British Sedges on other
difficult groups have been under discussion and R. D. Meikle has agreed to write one
on Willows.

Agreement has been reached between the Society and the Academic Press to publish
jointly a book relating to British hybrids under the editorship of Dr C. A. Stace.

Arrangements have been made for a Membership list to be published separately from
Watsonia.

THE CONSERVATION COMMITTEE had a very busy year, once again acting as a useful
‘watchdog’ on threats to and conservation of the British flora. The Society is now
recognised as the national botanical society concerned with advising onconservation
and it is represented on the Society for the Promotion of Nature Reserves Conservation
Liaison Committee by the President, Dr S. R. J. Woodell and P. J. Wanstall. The
Society has co-operated with local conservation trusts and natural history societies. It
has functioned in bringing together the various bodies concerned to consider the threat
to the Lady’s Slipper Orchid in Yorkshire; it has made representations concerning the
Monkey Orchid in Kent; the removal of limestone in Westmorland; the threat to
Braunton Burrows in Devon and Pembrey in Carmarthen, to mention but some of its
activities.

As part of its contribution to “The Countryside in 1970’ the Committee has prepared
a ‘Code of Conduct’ to serve as a guide for members of the Society. It has advised upon
and given its approval to a booklet being prepared by D. McClintock on ‘Wild flowers
you may pick’ designed for schools.

The sponsor of the Wild Plant Protection Bill, which the representatives of the
Society played a leading part in framing, failed to obtain an early place in the House
of Commons’ ballot. The Working Party are having the Bill re-drafted by a professional
parliamentary draftsman, the cost of which, up to a maximum of £500, is generously
being met by a grant from the World Wildlife Fund (B.N.A.). It is hoped to introduce
the Bill in the House of Lords in 1970.

Mrs A. Quinn (née Folan) from the National Institute for Physical Planning and
Construction Research and a member of the Irish Regional Committee has been
actively concerned with conservation problems in Ireland and has agreed to join the
Committee.

The President represented the Society at a banquet given by the Lord Mayor of
London at Guildhall to inaugurate European Conservation Year 1970.

é

326 REPORTS

THE RECORDS COMMITTEE has continued with its long-term task of improving the
Society’s whole system of record-collection by supplying to further Recorders who re-
quested them, sets of index drawers and cards. The Carnegie United Kingdom Trust
has made a generous contribution towards the cost of these index drawers. The Society
has provided voucher labels to recorders to attach to herbarium specimens which sup-
port published records and, in an effort to promote better understanding among
recorders and museum curators of herbarium management, a ‘Guide to Herbarium
Practice’ by J. W. Franks has been sent to each vice-county recorder. A new format
for ‘Plant Records’ has been designed for use in the merged Watsonia and Proceedings.

‘Network research’ has continued to flourish with a big demand still this spring for
cards for the Symphytum survey. Holly and Mistletoe surveys have now been launched
and further projects are being planned.

The Committee has continued to keep under close review the progress of the many
local Floras known to be in preparation and has provided specialist advice. The pro-
duction of a ‘‘ Guide for Sussex Flora Writers”, an expanded version of the ‘‘ Blue
Book’’, prepared by P. C. Hall has been undertaken as it is felt that this would be
useful to many other local Flora writers despite being primarily intended for Sussex.

Mention may be made that the report of the Local Flora Writers’ Conference at
Aberystwyth (1968) has come out during the year and has been circulated to vice-
county recorders. A Recorders’ Conference is to take place in Edinburgh in September
of this year (1970).

The Society was represented at the Conference of National Biological Societies where
the setting up of regional recording centres, a code of conduct and joint recording
meetings were considered.

Of the three Regional Committees—

THE COMMITTEE FOR THE STUDY OF THE SCOTTISH FLORA included eighteen field meetings
in its customary very full programme. Three meetings, including two junior ones, had
to be cancelled (see under Junior Activities) through lack of support. The fifteen which
took place were at St. Abb’s Head, Pease Dean (Berwickshire), Ardnamurchan, Girvan,
the rivers Tummel and Tay, Glen Prosen, Isle of Raasay, Braes of Balquhidder, Shet-
land, Handa and Durness, Northumberland, Skinflats (Stirlingshire), Dalmahoy Hill,
Penicuik Woods and Lawers. In addition the Annual Exhibition Meeting was held
jointly with the Botanical Society of Edinburgh at the Royal Botanic Garden, Edin-
burgh, by kind permission of the Regius Keeper, Dr H. R. Fletcher.

THE WELSH REGIONAL COMMITTEE arranged four field meetings which were generally
well attended and attracted a number of members from England. These were in Pem-
brokeshire, Montgomeryshire, Carmarthenshire and Brianne, Rhandirmwyn. The
Region staged a comprehensive exhibit at the Council for the Protection of Rural
Wales Conference at Aberystwyth in March.

The Annual General Meeting was held in Bangor, where Dr W. S. Lacey and Dr
Ball gave talks on the calcicole-calcifuge concept and R. H. Roberts on problems in
fern species. A field excursion in Anglesey next day excellently illustrated these talks
which will appear in the Welsh Regional Bulletin in 1970.

THE IRISH REGIONAL COMMITTEE arranged an extended field meeting based at Fermoy
in Co. Cork and a one day meeting at Lough Ross in Co. Armagh. The newsletter did
not appear during 1969 but it is hoped an issue will appear during 1970. The Annual
General Meeting and Exhibition took place in October at the School of Botany,
Trinity College, Dublin, by kind permission of Professor W. A. Watts.

WARBURG MEMORIAL FUND. Donations to the Warburg Memorial Fund now amount
to £563. The sum of £516 has been invested in 300 units of the Equities Investment
Fund for Charities. The Rules of the Fund, as agreed between the British Bryological
Society and the B.S.B.I. are set out in an appendix to this Report. It is hoped to make
the first award from the Fund in 1970.

REPORTS 327

The Council thanks the many friends of the Society who have helped during the year
to further its progress and also all those members who have taken an active part in
organising its affairs, on committees, as representatives on other bodies, and in other
ways. In particular, thanks must be expressed to the authorities of University College,
London, the Nature Conservancy and the Linnean Society of London for allowing the
regular use of their premises for Council or committee meetings; to Imperial College
of Science and Technology, London, for allowing the Conference to be held there; and
to the Trustees of the British Museum (Natural History) for the use of their Lecture
Hall and Conversazione Room in connection with the Annual Exhibition Meeting.
Finally, we thank our Honorary Auditors, Messrs Price Waterhouse & Co., for their
continuing services.

E. MILNE-REDHEAD, President
I. K. FERGUSON, Hon. General Secretary

17th March, 1970

328 REPORTS
BALANCE SHEET AS AT 31st DECEMBER, 1969

1968 1968
General Fund: Investments:
Balance Ist aps 44% Defence
1969 . £3,482 12 0 Bonds .. -- £500 0 0
Add:Premium on 5% National
conversion of evelopment
Defence Bonds. . 60 0 O Bonds . 1,000 0 0O
—————— 54% National
3,542 12 0 Development
Less: Excess of Bonds 500
expenditure over 6% British Sav-
income for year. . 218) *5 ings Bonds .. 1,000 0
£3,483 ——_— £3,520 13 7 10% Local
4,446 Publications Fund ae .. 4,784 17 11 Authority
40 Benevolent Fund ak 39 11 O Bonds .- 1,000 0 O
2,405 Creditors for Printing and The Equities
General Expenses if .. 1,641 18 6 Investment
171 Subscriptions Received in Fund for
Advance ae ae <0 180 0 O Charities—
1,500 units
(Market
value—£2,737) 2,789 1 3
£7,430 ————— £6,789 1 3
Cash at bank:
Deposit Account 3,000 0 O
Current Account 377 19 9
3,115 —————._ 3,377 19 9
£10,545 £10,167 1 O £10,545 £10,167 1 0

J. C. GARDINER,
Hon. Treasurer

In our opinion the above Balance Sheet and annexed General Income and Expenditure Account and summary of
Publications Fund transactions give a true and fair view of the state of the Society’s affairs at 31st December 1969,
and of its income and expenditure for the year ended on that date

27th February 1970 PRICE WATERHOUSE & CO.,
Chartered Accountants,
Hon. Auditors

GENERAL INCOME AND EXPENDITURE ACCOUNT

FOR THE YEAR ENDED 31ST DECEMBER, 1969

1968 1968

Cost of Printing and Distributing: £2,484 Subscriptions received .. sesh 5)

Proceedings, 597 Dividends and interest received . "621 6 5
Vol VII, Part3 Proceeds from Sales:
(balance) and Proceedings .. £184 12 10
Part4 .. .. £1,314 10 6 Watsonia we 456 1 5

Watsonia, 467 ———-. 640 14 3
Vol VII, Parts 2 785 Excess of expenditure over income
and 3 and part charged to General Fund es 21S

provision for
Vol VIII, Part 1 1,985 8 6
Storage and distri-
bution of back
numbers se Te & 2
£3,309 —————— £3,372 7 2
297 Printing and Distributing Annual
Report and Notices of Field

and Indoor Meetings .. 311 13 3
67 Expenses of Council and Com-
mittee Meetings VW 2S

101 Travelling Expenses of "Officers,

Secretaries and Members

attending Meetings .. 113 2 0
299 General printing, stationery, post-

ages, telephone and petty

expenses 348 3 8
71 Expenses of Regional Organisa-

tion, including Card Indexes

for Recorders, less grant 34 33 13 8

119 Res Research printing and
ostage 136 E16

52 Field ana Indoor "Meetings, ex-
penses less fees received : (3 By 5)
— Expensesre Covenants .. she 65 4 9

— Donation to Linnean Society
Appeal = ae ae 20 0 O
£4,315 £4,481 4 1 £4,315 £4,481 4 1

REPORTS 329
PUBLICATIONS FUND

TRANSACTIONS IN 1969

1968 1968
£1,253 British Sedges — Royalties and £3,437 Balance from 1968 ee .. £4,446 1 2
printing additional copies .. £240 6 10 Sales:
60 Storage and distribution, printing 1,621 British Sedges .. £329 11 O
price lists and sundry expenses S83} 17/- © 60 Conference
Balance to 1970: Reports a S019
General. . soe sss) 7/ Il 48 Other
Atlas Royalties publications .. 13 Gens
less out-goings 2,479 10 0 ———._ 372 19 _‘'5
4,446 —————_ 4, 784 17 11 Royalties received:

142 +~3 Atlas of the

British Flora. . 105 10 6
417 Critical Supple-

ment to the

Atlas .. a 96 6 0
7 Conference
Report No 9 44 11 0
27. Index to Botanical
Monographs .. 13 14 2
——§\— 260 1 8
£5,759 £5,079 2 3 £5,759 £51079) 2793

TEESDALE DEFENCE COMMITTEE REPORT

After further delays, the Charity Commissioners ultimately decided to register the
Fund as a charity. It was expected that this satisfactory outcome of the considerable
efforts made, and legal expenditure incurred, in order to persuade the Commissioners
to reach this conclusion, would result in the Inland Revenue withdrawing the provisional
tax assessments made on the Fund. The Committee would then have been able to dis-
charge outstanding liabilities and hand over the balance of the Fund to the Northum-
berland and Durham Naturalists’ Trust, in accordance with their undertaking. Regret-
tably, the Inland Revenue informed the Committee that they were not satisfied that the
registration of the Fund as a charity necessarily precluded them from continuing to
treat the Fund as taxable. Further delays thus arose, but within the last few weeks the
Inland Revenue have decided not to pursue their claim and it has, therefore, now been
possible to hand over to the Northumberland and Durham Naturalists’ Trust the
remaining balance on the Fund, amounting to £1,903 17s. 3d.

J. C. GARDINER, Hon. Treasurer
March, 1970

ANNUAL GENERAL MEETING, MAY 91TH, 1970

The Annual General Meeting of the Society was held in the Lecture Theatre at the
Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey, on Saturday,
9th May, 1970 at 12 noon. .

Mr E. Milne-Redhead (President) was in the Chair and 58 members were present.

The Minutes of the last Annual General Meeting, as printed in Watsonia, Vol. 8,
Part 1, pages 81-82, were adopted on the motion of Mr D. McClintock, seconded by
Mrs B. H. S. Russell after the sentence at the foot of page 81 commencing ‘Mrs M.
Briggs was proposed by the Chairman for re-election as Honorary Membership Sec-
retary’ had been altered to read “Meetings Secretary’.

REPORT OF COUNCIL

The Report together with the Accounts for the year 1969 had been circulated to all
members. There being no queries, the Report was adopted on the motion of Mr M. H.
Mannering, seconded by Mr P. J. Wanstall.

330 REPORTS

ELECTION OF OFFICERS

The Council had nominated Professor D. H. Valentine for re-election as a Vice-
President and this was carried unanimously. Dr I. K. Ferguson had been nominated for
re-election as Honorary General Secretary and he was re-elected unanimously. The
President thanked the Honorary General Secretary for his work throughout the pre-
vious year.

Mr J. C. Gardiner had been nominated for re-election as Honorary Treasurer. The
President spoke of the invaluable work Mr Gardiner had done for the Society and he
said that this was the last year in which Mr Gardiner was prepared to be nominated to
serve as Treasurer. The Society was confronted with the great difficulty during the
coming year of finding a new Treasurer. Mr Gardiner’s re-election was carried unani-
mously.

Mr E. F. Greenwood, Dr G. Halliday, Dr M. C. F. Proctor and Dr N.K.B. Robson
were Unanimously re-elected as Honorary Editors. The President voiced the Society’s
thanks to the hard-working Editors.

Mrs M. Briggs was proposed by the President for re-election as Honorary Meetings
Secretary following Council’s nomination and the meeting unanimously carried the
proposal. The President thanked Mrs Briggs for her wonderful work in organising
meetings for the Society.

Council had nominated Mrs J. G. Dony for re-election as Honorary Membership
Secretary and the President in proposing her re-election said that the name of Dony
had long been associated with the Society and he congratulated Mrs Dony on the
recently published membership list. The proposal was carried unanimously.

ELECTION OF NEW MEMBERS TO COUNCIL

In accordance with Rule 10, Mr R. S. R. Fitter, Dr S. M. Walters and Professor V. H.
Heywood retired. Three nominations had been received: Dr D. M. Moore, Mrs S. C.
Holland and Dr D. J. Ockendon and these were elected, their seniority for purposes of
Rule 10 beind decided (in the order shown above) by lots drawn by the President.

ELECTION OF HONORARY MEMBERS

Council had nominated Professor J. L. van Soest and Mr E. C. Wallace as Honorary
Members. Mr Milne-Redhead said that Professor van Soest was an expert on the
genus Taraxacum and had done much to further our knowledge of British Dandelions.
Professor van Soest’s election was carried unanimously. In proposing that the Meeting
approve Mr Wallace’s nomination the President said Mr Wallace had given invaluable
service to the Society over many years in compiling Plant Records and he had an im-
mense knowledge of the British flora. The election was carried unanimously and greeted
with loud applause.

AMENDMENTS TO THE SOCIETY'S RULES

The President said, that as explained in the Annual Report, the Development and
Rules Committee had now been replaced by a Co-ordinating Committee and it was
necessary to amend the Society’s Rules.
The following amendments were approved.
Rule 4: alter ‘(a)’ to read ‘Co-ordinating Committee’
Schedule II (a): alter to read ‘Co-ordinating Committee’ and add ‘Prepare business
for Council; consider policy on general matters and make recommendations to
Council for consideration’.

ELECTION OF HONORARY AUDITORS

Council recommended the re-election of Messrs Price, Waterhouse & Co., who had
kindly agreed to continue this work on an honorary basis. Their election was duly

REPORTS 331

carried, and the President in expressing the Society’s gratitude for their services said
how fortunate the BSBI were in having such a distinguished firm as its auditors.
Mr P. C. Hall proposed a special vote of thanks to Messrs Price, Waterhouse & Co..,
seconded by Mr D. McClintock.
There being no further business, the Meeting closed at 12.35 p.m.

I. K. FERGUSON

SOUTH-EAST REGION MEETING, 1970

A meeting of members resident in the South-East Region was held in the Kew Lecture
Theatre at the Jodrell Laboratory, Royal Botanic Gardens, Kew, on Saturday, 9th
May, 1970, following the Annual General Meeting of the Society.
The President was asked to take the Chair. The main business was to elect a Regional
Representative to serve on the Council of the Society in place of Mr P. J. O. Trist
who retired in accordance with Rule 9. The President thanked the retiring Repre-
sentative for his four years service on the Society’s Council.
The only nomination for Regional Representative received had been Mr D. E. Allen.
Mr Allen’s election was proposed by Mr R. M. Burton, seconded by Mr P. J. Wanstall.
There being no other business the meeting closed.

I. K. FERGUSON

COMMITTEE FOR THE STUDY OF
THE SCOTTISH FLORA

FIFTEENTH ANNUAL REPORT TO 31ST DECEMBER 1969

The Committee met twice in Perth and arranged an Exhibition Meeting at the Royal
Botanic Garden, Edinburgh (by kind permission of the Regius Keeper). Eighteen field
meetings were included in the Scottish programme for the year.

The Exhibition Meeting was held on Saturday Ist November, 1969 and about seventy
people were present. The following exhibits were shown: new Scottish vice-county
records, Hypericum montanum for v.c. 85 (G. H. Ballantyne), Crassula aquatica for
v.c. 96 and also new to Scotland (A. Souter) and Zannichellia palustris for v.c. 72 (Mrs
O. Stewart); Epilobium species and hybrids (hirsutum x montanum (Roxburgh), mon-
tanum X roseum, montanum x obscurum, adenocaulon x montanum, adenocaulon x
roseum and adenocaulon x obscurum (Fife)) and Salix hybrids (alba x fragilis, alba x
stipularis, caprea Xx viminalis, cinerea x viminalis and aurita x cinerea) (G. H. Ballan-
tyne); Bryophyte mapping at Roslin (Dr D. Chamberlain); Rubi, Taraxaca and Scirpus
cespitosus forms (Miss U. K. Duncan); a survey of Scottish Rubi (E. S. Edees and
Miss C. W. Muirhead); lichens from Iceland (Miss F. Kupicha); herbarium specimens
from and map of Inverness-shire, ‘the Cinderella of the Scottish Counties’ (A. A. P.
Slack); specimens from Stubai, Austria in the Hilda Wilson Herbarium, University of
Aberdeen (Mrs A. H. Sommerville); water colours of flowers of southern Scotland
(Mrs O. M. Stewart); ‘introduction to the sections of the genus Hieracium’ (A. McG.
Stirling); living specimens of wool adventives (Miss M. McC. Webster).

After tea Mr D. M. Henderson gave a talk on his travels in Scandinavia. Following
a buffet supper in the University of Edinburgh Staff Club colour transparencies were
shewn by B. S. Brookes, Miss E. R. T. Conacher, B. W. Ribbons, G. Rodway, A. McG.
Stirling and Miss M. McC. Webster on C.S.S.F. field meetings, J. Anthony on Suther-
land plants, Miss E. Bullard on Orkney plants, J. Grant Roger and Mrs A. H. Sommer-
ville on plants of the Austrian Tirol, Mrs B. Burbridge on plants of the Dolomites
and Canary Islands, B. Burbridge on botanical illustrations and Mrs H. Salzen on
Ontario woodlands.

I

332 REPORTS

Of the eighteen field meetings in the programme, three were cancelled (one to Border
S.S.S.I.s and two junior meetings to Craiglockhart Dell and Birks of Aberfeldy); one,
to Glen Prosen, was arranged by the Alpine Botanical Secretary of the Botanical Society
of Edinburgh and three, to Dalmahoy Hill, Penicuik Woods and Ben Lawers, by the
Cryptogamic Society of Scotland (for reports of these see Trans. bot. Soc. Edinb., 41,
part 1); the other meetings were held at St Abbs (7 June) led by Dr E. Conway (40);
at Pease Dean, Berwickshire (8 June) led by J. Cousens (12); at Ardnamurchan (14—
16 June) led by Miss A. Burns (5); at Byne Hill, Girvan, Ayrshire (15 June) led by H. A.
McAllister (19); on shingle beds of the Tummel and Tay (21-22 June) led by A. W.
Robson (4); on the Isle of Raasay (28 June—5S July) led by Mrs C. Murray (8); at the
Braes of Balquhidder (12 July) led by A. McG. Stirling (13); in Shetland (1-9 August)
led by W. Scott and R. C. Palmer (6); on Handa Island (2 August) and at Durness,
Sutherland (3 August) led by B. S. Brookes (5); to Holy Island, Northumberland
(30 August) led by Dr J. Richards (5) and at Skinflats (3 August) led by Miss E. P.
Beattie (12). The attendance figures are given in brackets. Reports of these meetings
appeared in Watsonia, 8 (2): 189-194.

Miss E. P. Beattie resigned from the Committee in April and hence ceased to be
B.S.B.I. Scottish Regional Representative. With the agreement of the Council of the
Botanical Society of the British Isles the Committee appointed its Chairman to be
Regional Representative and Mr A. A. P. Slack to fill the vacancy in the B.S.B.I.
Membership of the Committee. The eight elected members appointed Miss E. P. Beattie
and Mr A. W. Robson to serve for the calendar year 1970. Mr R. Mackechnie was
appointed Chairman, Mr B. W. Ribbons, Acting Honorary Secretary, Mrs A. H.
Sommerville, Honorary Treasurer, Miss E. P. Beattie, Honorary Field Secretary and
Mr R. Mackechnie, member of B.S.B.I. Records Committee for 1970. The Committee
was represented at B.S.B.I. Council by Miss Beattie on 11 March and by Mr Ribbons
on 29 October (it was unrepresented on 21 May) and at B.S.B.I. Records Committee
by Miss Beattie on 3 March and by Mr Ribbons on 20 October (it was unrepresented
on 3 June).

The list of Recorders for Scottish vice-counties was reviewed and recommendations
were made to the B.S.B.I. A Sub-committee was appointed in March to plan a pro-
gramme of exploration of the flora of Inverness-shire.

The accounts showed a deficit of £17.13.9 and this amount has been refunded in
equal sums by the two sponsoring societies.

The Committee wishes to express thanks to Miss Beattie for her work as Honorary
Secretary since 1966, to Dr Pritchard for his work as Honorary Treasurer since 1966,
to the officers of the sponsoring societies, to the leaders of the field meetings and to
other members and friends who helped the Committee during the year.

B. W. RIBBONS

EXHIBITION MEETING, 1969

An Exhibition Meeting was held at the Department of Botany, British Museum
(Natural History), London, S.W.7, on Saturday 29th November 1969 from 12 noon
to 5.30 p.m.

SALIX AND EPILOBIUM SPECIES AND HYBRIDS FROM
LOWLAND FIFE (V.C. 85)

During 1967 some Salix material was gathered in lowland south Fife and sent to
R. D. Meikle for identification. This confirmed the presence of the following taxa:
S. alba, S. alba x fragilis, S. fragilis (including vars. decipiens and russelliana), S. pur-
purea, S. daphnoides, S. viminalis, S. x stipularis, S. calodendron, S. caprea ssp. caprea,
S. caprea x viminalis, S. cinerea ssp. atrocinerea, S. cinerea x viminalis, S. aurita, S.
aurita x cinerea, and S. repens ssp. repens. Specimens of most of these taxa were
exhibited.

REPORTS 333

Previous records of Epilobium in v.c. 85 indicate that E. roseum and E. obscurum
have increased during the last few decades. E. adenocaulon, first noted in 1959, has
spread rapidly during the past ten years and may soon become as widespread as E.
angustifolium. Specimens of these species were exhibited along with material collected
in 1968, which had been sent to T. D. Pennington for identification. From this the
following hybrids were determined: E. montanum x roseum, E. montanum x obscurum,
E. adenocaulon x montanum, E. adenocaulon x roseum, and £E. adenocaulon x ?
obscurum. In addition, E. hirsutum x montanum was shown from v.c. 80 (Roxburgh).

G. H. BALLANTYNE

SOME OFTEN CONFUSED PLANTS

The plants most often confused are not necessarily the most closely allied species but
those with an overall superficial similarity. This similarity itself leads to misidentifica-
tions but the confusion is often amplified by careless collecting, labelling, and use of
keys and descriptions and casual comparison with authentically named material.
Herbarium specimens and notes demonstrated some of these points and indicated
some characters not used often enough:
Lepidium campestre (L.) R.Br. and L. heterophyllum Benth.; anthers yellow or purple
respectively.
Arenaria serpyllifolia L. and A. leptoclados (Reichb.) Guss.; need careful examination
of several small differences.
Lysimachia vulgaris L. and L. punctata L.; calyx-margins reddish or green respectively.
Veronica montana L. and V. chamaedrys L.; stems evenly pubescent or strongly
bifarious respectively.
Veronica polita Fr. and V. agrestis L.; sepals ovate, acute or oblong, obtuse respec-
tively.
Galium palustre L. and G. uliginosum L.; leaves blunt or mucronate respectively.
Scirpus setaceus L. and S. cernuus Vahl; fruits longitudinally ribbed or smooth
respectively.
E. B. BANGERTER & L. F. FERGUSON

NARDURUS, VULPIA OR FESTUCA

Nardurus and Vulpia are genera which have been segregated from Festuca, but which
have very uncertain circumscriptions. The exhibit consisted of herbarium sheets of the
three main species of Nardurus and of six representative species of Vulpia. They were
chosen to demonstrate the range of variation of Vulpia and Nardurus and the fallibility
of all characters which have been used to separate these genera from each other and
from Festuca. Nardurus maritimus, for instance, appears to be more similar to certain
species of Vulpia than to N. salzmannii; and one or two species of Vulpia (e.g. V. setacea)
possess the perennial habit of Festuca rather than the annual habit of Vulpia.

B. Batts, H. Jones & C. A. STACE

STEREOSCAN STUDIES ON SEED MORPHOLOGY

In an attempt to distinguish between fossil seeds of Arenaria ciliata and A. norvegica,
the seed surface characters have been examined by means of the scanning electron
microscope. For each species, seeds from localities throughout their ranges were
examined and in addition a 40,000 year old seed of A. ciliata from East Anglia, from
a last glaciation fossil flora.

At a magnification of 800 x , the details of surface sculpture can be seen to vary from
one population of a species to another. A possible distinction between the two species
may be found in the ornament on the cells. The fossil seed closely resembled recent
seeds of A. ciliata from Ireland.

334 REPORTS

Micrographs were exhibited of seed surface of Stellaria pallida, S. media and S.
neglecta at various magnifications. In these seeds magnification up to 6000x shows
interesting differences in the finer ornament between the species.

F. G. BELL

A STUDY OF THE AERODYNAMICS OF WINGED SEEDS

This exhibit, by M. Stone and R. Dudzinski of Harrow County Grammar School,
showed the results of experiments carried out on the rate of fall of various samaras.
Specimens of seeds, leaves of trees and graphs on which results were tabulated were
shown.

CHILDREN’S CENTRE CLUB, BRITISH MUSEUM (NATURAL HIsTorRy)

YET MORE ABOUT MULL

The exhibit represented an interim progress report on the first three years of the British
Museum Mull Survey Project. The number of species of vascular plants now known is
almost twice that recorded by George Ross in his detailed list of 1877. The total number
of records amassed up to the end of 1968 was approximately 32,000, a number that
bears comparison with the 117,000 records used by Dr J. G. Dony for his Flora of
Hertfordshire, an area nearly twice as large as Mull. Specimens were exhibited to illus-
trate new vice-county records, records new to the Mull part of v.c. 103 and important
confirmations of old records localised only as ‘Mull’. A fern hybrid new to science,
Dryopteris x pseudoabbreviata Jermy (D. aemula x abbreviata), was discovered during
one of the survey visits to the island. Important contributions to the survey have been
made by many people outside the museum and members of the B.S.B.I. have been
prominent among those who have offered help.

DEPARTMENT OF BOTANY, BRITISH MUSEUM (NATURAL HIsTory)

WATERCOLOUR DRAWINGS OF BRITISH PLANTS BY THE
LATE MRS WINIFRED BUSSEY

Mrs Bussey had been a flower lover and a painter all her life, but at the advanced age
of 71 she began painting wild flowers and trees as a record of the English countryside
and of the plants that had sprung up in London as a result of the wartime bombing.
In all, she collected and painted 440 specimens. Her work has been widely exhibited in
libraries around the London area and at the shows of the Royal Horticultural Society.

G. BUSSEY

TEN SPECIES OF TARAXACUM FROM SCOTLAND

The following species, all determined by Dr A. J. Richards, were exhibited: Taraxacum
brachyglossum (v.c. 90), T. cyanolepis (v.c. 106), T. drucei (v.c. 90), T. faeroense (V.C.
106), T. fulvum (v.c. 90), T. gotlandicum (v.c. 90), T. lacistophyllum (v.c. 106), T. platy-
glossum (v.c. 90), T. spectabile (v.c. 90), T. unguilobum (v.c. 88).

U. K. DUNCAN

SOME INTERESTING CO. WATERFORD PLANTS

Large areas of Ireland remain relatively unbotanised in comparison with much of
England and Wales and there is a need for considerably more recording not only of
critical groups but also of well-known species. Co. Waterford (v.c. H6) in south-
eastern Ireland is an area which has been remarkably neglected by botanists; in fact,
apart from two or three well-known areas it has been almost ignored.

REPORTS 335

The exhibit showed some herbarium specimens, together with dot maps showing the
distribution of each species in the British Isles, of new, confirmed pre-1930, and other
interesting records that have recently been made in Co. Waterford.

REFERENCES

FERGUSON, I. K. (1968). Notes on the flora of Co. Waterford. Ir. Nat. J., 16: 94-97.
FERGUSON, I. K. & SCANNELL, M. J. P. (1969). Zostera in Co. Waterford. Ir. Nat. J., 16: 176-
Nis
I, K. FERGUSON

MORECAMBE BAY—NORTHERN AND SOUTHERN ELEMENTS
IN THE FLORA

Within Morecambe Bay a number of maritime species are found close to the northern
or southern limits of their range in the British Isles. The reasons for the geographical
limitations of these species are not known but may vary from species to species. As they
near the limit of their range they exhibit strong niche preferences, being less generally
distributed throughout the maritime zone than they are further north or south, and show
a particular susceptibility to biotic and disturbance factors such as grazing.

Notes on the ecology and distribution maps were presented of seven species with
southern distributions (Halimione portulacoides, Agropyron pungens, Parapholis strigosa,
Limonium humile, L. binervosum, Adiantum capillus-veneris and the non-coastal Spiran-
thes spiralis) and three with northern distributions (Eleocharis quinqueflora, Blysmus
rufus and Mertensia maritima).

A. J. GRAY

THE GUERNSEY FLORA IN 1969

The rich flora of Guernsey has been surprisingly neglected and unexplored during the
present century. 1969 gave fantastic proof of this, for something like 20 plants were
found new to the island. The precise number depends on assessments, some still to be
made, of status, permanence, identification, etc. Most of these finds were made on the
visit by the Society in April, despite a season backward by 4—6 weeks, a report on which
has already appeared (Watsonia, 8: 185-186 (1970)). Species not recorded then, or
seen since, include — Lunaria annua, Salicornia europaea, Epilobium nerterioides and
Ipheion uniflorum.

In addition, second stations were seen for other plants, e.g. Adiantum and Equisetum
telmateia; new places found for plants almost as rare in the island, e.g. Trachelium
caeruleum, Veronica catenata and Myriophyllum alterniflorum; or not seen for many
years, such as Galeopsis tetrahit; or confirmed identifications made, such as Salix
calodendron.

Some vouchers of the year were exhibited. Not available was the remarkable new
large Romulea rosea var. australis, specimens of which were on their way back from
determination in South Africa.

D. McCLINTOCK

CHENOPODIUM PUMILIO-—A KENTISH OBITUARY?

Chenopodium pumilio R.Br. is one of three similar small-leaved, usually prostrate,
Australian species of Goosefoot, all of which have occurred in Kent. The other two are
C. carinatum R.Br. and C. cristatum (F.v.M.) F.v.M.

It differs from the other two by lacking any keel or wing on the sepals. C. carinatum

336 REPORTS

has a well marked wing-like keel, hence its Latin trivial; C. cristatum has a crest of
fringed processes on its keel, hence cristatum. In addition there is the hybrid between
the last two, C. x bontei Aellen. All these have been found in shoddy.

Not only is C. pumilio the commonest, it seems to be the only one capable of natural-
ising itself. This it has done in plenty at Borthwick Bank south of Edinburgh; did (still
does?) at Frensham, Surrey; and has done its best to do likewise at Borough Green
and Wrotham Station, W. Kent, since at least about 1960. It has been there every year,
in varying quantity, in the sidings, chiefly between the rails nearest to the east end of
the down platform. This was despite regular clearing — the station often received an
award for the best-kept station. It was a remarkable case of persistence in an alien
country, undeterred by relentless discouragement.

There it still was in 1969; and a voucher specimen (now at Maidstone) was shown.
But earlier in the year much of its area was submerged beneath asphalt for a new car
park, and the rest of the siding is to be altered. It remains to be seen whether the seeds
it must have so liberally shed enable it to survive. It deserves to.

D. MCCLINTOCK

SOME SCOTTISH HAWKWEEDS

The exhibit comprised twenty-three herbarium sheets of Hieracium species collected
during the past four years, mainly in west and north Scotland, and all determined by
P. D. Sell and Dr C. West.

Most of the Sections of the genus were represented, and the species shown ranged
from H. shoolbredii E. S. Marshall and H. pictorum E. F. Linton, both widespread and
common in the area, to the rare local endemics H. dovrense Fries and H. pollinarioides
Pugsl. Recently described species were represented by H. nigrisquamum P. D. Sell &
C. West and H. solum P. D. Sell & C. West. An as yet undescribed species of the Sec-
tion Alpina was also shown as representative of the considerable number of such
undetermined taxa which have been shown to occur throughout the area.

A. McG. STIRLING

SPECIES OF LOLIUM AND THEIR HYBRIDS IN BRITAIN

The exhibit consisted of: (1) species occurring in Britain: Lolium perenne L. (indi-
genous), L. multiflorum Lam., L. rigidum Gaud. (L. strictum Presl), L. temulentum
L., L. remotum Schrank (L. linicolum A.Br.), L. loliaceum (Bory & Chaub.) Hand.-
Mazz. (L. subulatum Vis.) and L. persicum Boiss. & Hohen. (all introduced, the last
three being rare casuals), accompanied by a key to species; (2) interspecific hybrids of
Lolium found in Britain, including L. multiflorum x rigidum, recently discovered in
N. Hants. (v.c. 12), (see Short Notes p. 299); (3) intergeneric hybrids, involving Lolium
and Festuca, recorded from Britain; (4) diagrams showing natural interspecific and
intergeneric (Festuca x Lolium) hybrids reported in the European botanical literature;
and (5) a selection of interspecific and intergeneric hybrids of Lolium synthesized by
T. J. Jenkin and his collaborators at the Welsh Plant Breeding Station, Aberystwyth.

A. MELDERIS

COMPUTER-GENERATED KEYS

A FORTRAN program has been written which prepares printed ready-to-use keys-
The taxa for which a key is required are described by a rectangular array of characters
and their values, some of which may be missing. Either bracketed or indented keys are
obtainable, with an optional limit on the number of characters used per lead. The key is
optimised for efficiency, and is made to be dichotomous wherever possible. Several

REPORTS 337

computer-produced keys were demonstrated. In particular, the key in Flora Europaea
Vol. 2 for 134 genera of the Umbelliferae has been regenerated by computer, and a key
for 86 potato cultivars was also shown. No previous key for the potatoes had been
available on account of the labour involved in compiling it by hand. The program
and a user manual are available on request.

R. J. PANKHURST

POLLINATION PHOTOGRAPHS

Photographs were exhibited showing visits of pollinating insects to a number of
flowers, mostly zygomorphic, including the following (name of pollinator in paren-
theses): Lamiastrum galeobdolon (Bombus agrorum), Stachys sylvatica (Bombus agrorum),
Salvia glutinosa (Bombus agrorum), Iris pseudacorus (Bombus hortorum, Rhingia cam-
pestris), Lonicera periclymenum (Plusia gamma), Listera ovata (Athous haemorrhoidalis),
Anacamptis pyramidalis (Zygaena sp., Melitaea cinxia).

M. C. F. PROCTOR

TARAXACUM IN UPPER TEESDALE

A study has been made of the occurrence and distribution of Taraxacum species in
Upper Teesdale. T. proximum Dahlst. and T. brachyglossum Dahlst. are Erythrosperma
species found locally in dry places on the limestone. Vulgaria species are found on road-
sides below 1300 ft, but are not of particular interest. In addition, there are eight
Spectabilia species. These range from common Scandinavian species (T. maculigerum,
T. spectabile, T. praestans) and common or local northern Atlantic species (T. faeroense,
T. unguilobum, T. landmarkii) to endemics, not found outside Upper Teesdale (T. pseudo-
nordstedtii, T. pseudolarssonii). In a small area of the dale, where the Sand Syke meets
the Langdon Beck, all eight species occur. A study, both phytosociological and environ-
mental, was made to determine in what way these species differ in habitat requirements.
The following tentative conclusions can be made at this stage of the study:
T. pseudonordstedtii Occurs in base-rich flushes, usually with a northerly aspect.
T. landmarkii Occurs in base-rich and very open localities.
T. faeroense Occurs in closed and species-rich base-rich communities.
T. maculigerum Occurs in basic and very well-drained localities.
T. pseudolarssonii Occurs in shaded localities above the influence of flood water,
associated with woodland species.

T. praestans Differs from the former chiefly in occurring in less shaded sites.

It is hoped that the continuation of this study after the flooding of the Cow Green
Reservoir will yield information on the biological effect in stream-side communities
near Widdybank Fell of changes in water regime resulting from the flooding.

A. J. RICHARDS

AN EXPERIMENT IN FLOWER MOUNTING

Two hundred specimens were suspended by plastic tape from a portable wooden stand
(6’ high, 3’ wide, 1’ 6” at base, weight 4lb) and mounted on good quality card by
slightly adhesive transparent sealing cover (COVERLON by Fablon).

The exhibit effectively displayed specimen protection, retention of colour and shape
for short and long-term preservation, and safe and easy transport.

Specimens included: Geum rivale (collected 1947), Primula farinosa (c. 1952), six
species collected a week earlier and mounted the same day, Carduus, Ilex, Eryngium,
ripe fruit of Viburnum opulus and leaves and inflorescence of Heracleum mantegazzia-
num.

338 REPORTS

Whether specimens are dried and pressed, or mounted and sealed in the field, effective
preservation necessitates periodical inspection to ensure freedom (for any lengthy
period) from excessive temperature, dampness, direct sunlight or tight packing of
specimen cards.

H. E. W. SELBY

HYBRIDS IN JUNCUS SECTION GENUINI

Five species of this section occur in Britain: J. effusus, J. inflexus, J. balticus, J. filiformis
and J. conglomeratus. The first three of these form hybrids (recognised by their high
or complete sterility and their intermediacy in a wide range of morphological and
anatomical characters) in all combinations: J. effusus x J. inflexus is quite widespread
though seldom common, and was first recognised in Britain in 1843; J. effusus x J.
balticus was represented by one colony, now lost through building, on the Lancashire
coast; and J. inflexus x J. balticus occurs as three distinct colonies on the Lancashire
coast. Neither of the latter two hybrids has ever been discovered elsewhere in the
world.

J. filiformis hybridises extensively with J. balticus in Scandinavia, but the two species
do not grow together in Britain.

J. conglomeratus has often been reported as forming hybrids with J. effusus and J.
inflexus, but none has ever been satisfactorily substantiated.

The two non-British species of this section found in Europe (J. jacquini and J.
arcticus) appear never to hybridise.

C. A. STACE

The following also exhibited.

E. J. CLEMENT & T. B. Ryves’_ B.S.B.I. Alien Hunt, October 1969 (see Field Meetings
1969, Watsonia, 8: 188-189 (1970))

I. K. & L. F. FERGUSON Naufraga rediscovered (see Short Notes p. 294-295)

A. N. Gipsy Botany on stamps

M. GrRigERSON' British Wild Flower Calendar
Illustrations of specimens grown at Kew

M. Hickey Analytical flower drawings from life

R. B. IvimeEy-Cook & M. C. F. Proctor A proposed new Flora of Devon

D. Marris & A. H. F. WEBBE Some British plants and others from the High-Arctic

R. Marris ‘Temperate’ Potamogeton species new to the High-Arctic

A. MELDERIS Bromus pumpellianus Scribn.; an introduced species in Britain (see
Short Notes p. 300)

Y.L. Moscati Postage stamps of botanical interest

F. H. PERRING & G. H. PETERKEN Network Research 1970

W.T.STEARN Stereoscan photographs of pollen

S. M. WALTERS & S. WEGMULLER A diploid subspecies of Lamiastrum galeobdolon

(see p. 277-288)
J.T. WISELY Various British wild plants, mostly from the B.S.B.I. Shetland meeting

From 4.30 p.m. the following members gave short lectures and showed colour slides
in the Lecture Hall.

H. J. B. BirKs Plants of the Grossglockner, Austria

J. E. LOUSLEY Flowers of the Algarve

D. Marris Some High-Arctic plants

M. C. F. Proctor Some plants and habitats of S.W. England
G. TucK Swiss Alpine plants

T. WHITE Some rarities of the Isles of Wight

Watsonia, 8, 339-344 (1971). 339

Panel of Referees and Specialists

Please read carefully before submitting specimens

In all instances it is understood that a complete plant (excluding root except in
annuals and where specified) will be submitted, together with any necessary notes
on colour, habitat and locality (not for publication without sender’s permission).
For tall herbaceous plants the inflorescence and upper leaves, a short mid-sec-
tion and radical leaves should be included, and when in doubt it is advisable
to consult the referee beforehand as to the best methods of collection and pre-
servation of material for subsequent determination. Where extensive hybridisa-
tion is known or suspected to occur, samples should be sent showing the range
of variation in the population. The abbreviations after the referees’ names are of
features and parts that are essential for accurate determination. If these are not
sent the referee may refuse to determine the specimen.

Except where specified, pressed and dried material should be sent and a note
enclosed as to whether the specimen can be retained by the referee or is to be
returned to the sender. It should be remembered that critically named material
is of greater value when located in large regional or national collections where
it is available for consultation by future workers. If the specimen is to be re-
turned, postage stamps must be included or return postage refunded on receipt.

The arrangement of the list is that of Dandy’s ‘List of British Vascular Plants’
(1958). It must be realised that the referee does not necessarily agree with the
taxonomic or nomenclatural treatment of his group in Dandy’s List or in any
other publication and that he may also qualify his identification with a query
Orne.

The referees’ names in the List are given in alphabetical order when two or
more are available for consultation. The addresses of most of the referees are to
be found in the list of members, but some referees are not members and some
others do not want material sent to the address given in the list. In these instances
the specimens should be sent to the referee c/o the institution specified.

Do not forget to enclose postage for reply
(preferably a stamped addressed envelope or stamped addressed

label for parcel)
ABBREVIATIONS
bds. =a) Duds.
ckn. = catkins.
cn. = colour notes (with dried material).
fl. = flowers.
fre. = fresh material preferred.

mfr. == nature (rit.

ml. — mature leaves.

rE = roots or underground parts.
ri. = radical leaves.

S. = seeds.

sm. = spirit material.

340 REFEREES AND SPECIALISTS

stsh. = sterile shoots.

young fruits.

Botany Department, British Museum (Nat. Hist.), London, S.W.7.

CFI = Commonwealth Forestry Bureau, Commonwealth Forestry Insti-
tute, South Parks Road, Oxford.

ze
II

FC = Forestry Commission, Forest Research Station, Alice Holt Lodge,
Wrecclesham, Farnham, Surrey.
K = The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey.
L = Genetics Department, University of Liverpool.
NMW = The Botany Department, National Museum of Wales, Cardiff.
UCD = Botany Department, University College, Dublin.
UCL = Botany Department, University College, London, W.C.1.
UCNW = Botany Department, University College of North Wales, Bangor,
Caernarvonshire.
CRITICAL SYSTEMATIC GROUPS
PTERIDOPHYTA

Filices: J. A. Crabbe (BM), mfr.

Tsoetes: A. C. Jermy (BM), mfr.

Equisetum: P. G. Taylor (K), r.

Dryopteris filix-mas agg.: H. Corley, Mrs P. J. Newbould, Dromore House,
Drumcrome, Coleraine, N. Ireland.

Dryopteris dilatata-carthusiana (spinulosa) agg.: Dr S. Walker.

SPERMATOPHYTA

RANUNCULACEAE

Ranunculus flammula: Mrs P. Candlish, r.

Ranunculus hederaceus — R. baudotii: Dr R. W. Butcher, Prof. C. D. K. Cook,
mfr.

Thalictrum: Dr R. W. Butcher, mfr., rl.

FUMARIACEAE
Fumaria: M. G. Daker (K), mfr (each gathering of fresh material packed in a
separate bag as fr. and sepals deciduous).

CRUCIFERAE

Erophila: Dr G. Smith.

Barbarea: G. A. Matthews (BM), mfr., bds., rl.
Thlaspi: Dr D. P. Young, mfr.

Cochlearia: Dr D. H. Dalby, J. J. B. Gill (L), mfr., rl.
Cardamine pratensis agg: D. E. Allen, rl.

Nasturtium: H. K. Airy Shaw (K), mfr., s.

VIOLACEAE

Viola odorata — V. palustris: Prof. D. Valentine.
Viola lutea — V. kitaibeliana: R. D. Meikle (K), cn.
POLYGALACEAE

Polygala: Dr J. McNeill, mfr., r.

HYPERICACEAE

Hypericum: Dr N. K. B. Robson (BM).

CISTACEAE: Dr M. C. F. Proctor.

REFEREES AND SPECIALISTS 341

TILIACEAE
Tilia: H. A. Hyde (NMW),.

TAMARICACEAE
Tamarix: D. R. Hunt (K).

CARYOPHYLLACEAE

Silene nutans agg.: F. N. Hepper (K).

Silene, other spp.: Dr S. M. Walters.

Dianthus: Miss S. Hooper (K), cn.

Cerastium, annual spp.: E. Milne-Redhead (K), mfr., fl.

Stellaria: P. M. Benoit, mfr.

Sagina: F. N. Hepper (K), N. Jardine, mfr., for annual plants, whether the
sepals were appressed or spreading: for perennial plants, whether the
petals exceeded the sepals.

Arenaria: Dr G. Halliday.

Spergularia: Dr J. A. Ratter, mfr., s., number of stamens.

ILLECEBRACEAE
Herniaria: Dr L. C. Frost, Dr A. Melderis (BM).

PORTULACACEAE
Montia: Dr S. M. Walters, s.

AMARANTHACEAE ,
Amaranthus: J. P. M. Brenan (K), mfr., cn.

CHENOPODIACEAE

Chenopodium: J. P. M. Brenan (K), mfr., cn., s.
Atriplex: Mrs B. A. Hulme, mfr.

Salicornia: Dr D. H. Dalby, Prof. T. G. Tutin, mfr.

MALVACEAE
Malva: Dr N. K. B. Robson (BM), mfr.

GERANIACEAE
Erodium: P. M. Benoit, mfr., cn.

OXALIDACEAE
Oxalis: Dr D. P. Young, r.

PAPILIONACEAE

Ulex: P. M. Benoit, Dr M. C. F. Proctor, M. J. Woodman, fl. or mfr.
Medicago: C. C. Townsend (K), mfr.

Anthyllis: Dr J. Cullen, cn., basal ml.

Vicia: P. F. Hunt (K), mfr.

ROSACEAE

Rubus: E. S. Edees, bds., yfr., piece from current year’s stem near apex but with
fully developed leaves.

Potentilla: Dr B. Matfield, Dr S. M. Walters,

Alchemilla: Dr M. E. Bradshaw, Dr S. M. Walters, rl.

Aphanes: Dr S. M. Walters, mfr.

Rosa: Dr R. Melville (K), mfr.

Prunus: Prof. D. A. Webb, mfr., f1.

Sorbus: P. J. M. Nethercott, mfr.

Crataegus: Prof. A. D. Bradshaw, mfr., fl.

342 REFEREES AND SPECIALISTS

CRASSULACEAE
Sedum: Dr H. J. M. Bowen, Prof. D. A. Webb, stsh.

SAXIFRAGACEAE
Saxifraga: Prof. D. A. Webb, stsh.

ONAGRACEAE
Epilobium: T. D. Pennington (CFI).
Circaea: P. M. Benoit, mfr.

CALLITRICHACEAE
Callitriche: R. D. Meikle (K), Dr J. P. Savidge, mfr.

UMBELLIFERAE (general): J. F. M. Cannon (BM), mfr., rl., r.
Apium: R. D. Meikle (K), mfr.

Oenanthe: Prof. C. D. K. Cook.

Heracleum: Dr R. K. Brummit (K).

EUPHORBIACEAE
Euphorbia uralensis — E. cyparissias: Dr T. Pritchard, A. R. Smith (K).

POLYGONACEAE

Polygonum aviculare s.l.: Dr B. T. Styles, mfr.
Polygonum (other spp.): Dr J. Timson.
Rumex: J. E. Lousley, mfr., rl.

ULMACEAE
Ulmus: Dr R. Melville (K), mature short shoots of trees, NOT sucker or
epicormic shoots.

BETULACEAE
Betula: G. A. Matthews (BM), ckn., ml.

FAGACEAE
Quercus: J. E. Cousens.

SALICACEAE
Populus: P. G. Beak (CFI), J. Jobling (FC), ckn., ml.
Salix: R. D. Meikle (K), R. C. L. Howitt, ckn., ml.

ERICACEAE
Erica: Dr P. Gay, Prof. D. A. Webb.
Vaccinium (incl. Oxycoccus): Dr N. K. B. Robson (BM).

PLUMBAGINACEAE
Limonium: Dr L. Boorman.

PRIMULACEAE
Primula: Prof. D. H. Valentine, Dr S. R. J. Woodell, fre.
Anagallis: Dr F. H. Perring, P. G. Taylor (K).

GENTIANACEAE
Centaurium: Dr A. Melderis (BM).
Gentianella: Dr N. Pritchard.

BORAGINACEAE
Symphytum: Dr G. A. Nelson, Dr F. H. Perring, A. E. Wade, r., cn.
Myosotis: A. E. Wade, D. Welch, mfr.

CONVOLVULACEAE
Calystegia: Dr R. K. Brummitt (K), C. A. Stace.

REFEREES AND SPECIALISTS 343

SCROPHULARIACEAE
Verbascum: Dr I. K. Ferguson (K), rl., cn. of all parts of plant, esp. filament
hairs.

Mimulus: R. H. Roberts.

Veronica: E. B. Bangerter (BM), Prof. J. H. Burnett (aquatic spp. only).
Rhinanthus: Miss A. 8. Crozier (UCNW), Dr D. J. Hambler.
Melampyrum: Dr A. J. E. Smith.

Euphrasia: Dr P. F. Yeo.

OROBANCHACEAE
Orobanche: Dr D. J. Hambler, fre., cn. (incl. stigma), host, sketch of corolla

shape.

LENTIBULARIACEAE
Utricularia: P. G. Taylor (K), fre. or sm.

LABIATAE (general): Dr R. M. Harley (K).

Mentha: Dr R. M. Harley (K).

Thymus: Prof. C. D. Pigott.

Calamintha and Acinos: Miss C. Whitefoord (BM).

Galeopsis tetrahit agg.: P. M. Benoit, C. C. Townsend (K), fre.

RUBIACEAE
Galium: Dr K. M. Goodway.

VALERIANACEAE
Valerianella: C. C. Townsend (K), mfr.

COMPOSITAE (gen.): C. Jeffrey (K), mfr., rl., cn.
Senecio: C. Jeffrey (K), mfr., rl., cn.

Filago: J. E. Lousley.

Gnaphalium: J. E. Lousley.

Aster: Dr P. F. Yeo, cn.

Erigeron: J. B. Marshall (BM).

Arctium: Dr F. H. Perring, Dr W. A. Sledge, mfr., rl.
Carduus: Dr W. A. Sledge, rl.

Cirsium: Dr W. A. Sledge, rl.

Solidago: D. H. Kent.

Lactuca serriola-virosa group: D. R. Hunt (K), mfr.
Hieracium: C. E. A. Andrews, rl., cn. (esp. styles), date of gathering.
Crepis: J. B. Marshall (BM), mfr., rl.

Taraxacum: Dr A. J. Richards, fl., fr.

ALISMATACEAE
Alisma: Miss P. Boyle (UCD), fre.

ZOSTERACEAE
Zostera: Prof. T. G. Tutin.

POTAMOGETONACEAE
Potamogeton: J. E. Dandy (BM), Sir George Taylor.

ZANNICHELLIACEAE
Zannichellia: J. E. Dandy (BM).

LILIACEAE
Allium: Dr W. T. Stearn (BM).

344 REFEREES AND SPECIALISTS

JUNCACEAE
Juncus: G. A. Matthews (BM).
Juncus bulbosus s.l.: P. M. Benoit.

ORCHIDACEAE: P. F. Hunt (K), fre., or sm. with cn.
Epipactis: Dr D. P. Young, fre., or sm. with cn.
Dactylorhiza: P. F. Hunt (K), P. Vermeulen, fre., or sm. with cn.

SPARGANIACEAE
Sparganium: Prof. C. D. K. Cook.

CYPERACEAE, (excl. Eleocharis): A. C. Jermy (BM), mfr., r.

Eleocharis: Dr S. M. Walters, mfr.

Carex: A. O. Chater, Miss S. Hooper (K), A. C. Jermy (BM), E. C. Wallace,
mie Ac

GRAMINEAE, esp. Festuca, Poa, Bromus, Agropyron, Puccinellia, Vulpia (not aliens):
Dr A. Melderis (BM), (esp. aliens): Dr C. E. Hubbard (K).

Glyceria: Dr M. Borrill.

Poa, Agropyron: Prof. T. G. Tutin.

Dactylis: Dr M. Borrill.

MISCELLANEOUS SUBJECTS

Biographical details of British botanists: D. H. KENT.

Climate in relation to plant distribution: Dr J. P. SAVIDGE.

Collection and preservation of botanical specimens: E. MILNE-REDHEAD (K).

Folklore: D. E. ALLEN.

Floras, foreign: Dr W. T. STEARN (BM).

Floras, local: N. D. SIMPSON.

Genetics and cytology in relation to systematics: Prof. D. H. VALENTINE.

Herbals: R. DESMOND (K).

Herbaria: D. H. KENT.

Infraspecific taxa of British phanerogams (references to descriptions and cultiva-
tion experiments): D. E. ALLEN.

Nomenclature: J. E. DANDY (BM).

Popular names: Miss C. M. Ros.

Systematic works and monographs: D. H. KENT.

Vice-County boundaries: J. E. DANDY (BM).

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INSTRUCTIONS TO CONTRIBUTORS

PAPERS and short notes concerning the systematics and distribution of British and European vascular plants
as well as topics of a more general character and wider appeal are invited from both members and non-
members of the Society. Careful attention to these notes before a paper is submitted can save the author and
editors much work, and do much to speed publication.

Papers must be submitted in final, fully-corrected form. They should be typewritten, with wide margins,
double-spaced, on one side of the paper only. If possible, two copies should be sent, of which one must be a
top copy. Authors should keep a fully-corrected carbon copy of their typescripts for reference.

Write clearly and concisely, rigorously excluding extraneous matter. It sometimes helps the clarity of
an argument to place extensive experimental or tabular data in appendices, rather than in the main text.
Technical terms are of value only as aids to clarity, precision and conciseness of expression. Take care that
they are used consistently, and defined where necessary. Ill-defined or general terms often need definition in
a particular context, and can often with advantage be replaced by something more precise, as by the ‘deme’
terminology of Gilmour and Gregor in genecology. Make certain that the main conclusions of the paper are
clearly displayed to the reader.

Avoid complicated hierarchies of headings, and check carefully the consistency of those that you use;
a table of contents, setting out the full hierarchy of headings with the MS. page numbers, is often helpful
even if it is not to be printed with the paper. Avoid footnotes as far as possible, and keep cross references
by page number to a minimum.

Tables, unless very small, should be typed on separate sheets and attached at the end of the typescript.

They should be kept within a reasonable size, and as simple in structure as possible.
Keys should be in one of the generally accepted forms; Flora of the British Isles by Clapham, Tutin and
Warburg (2nd ed., Cambridge, 1962) and Flora Europaea (Cambridge, 1964-7) provide suitable patterns.

Names of genera and species should be underlined, but any other typographical indications should be
inserted lightly in pencil. Names of vascular plants should normally follow the List of British Vascular
Plants by J. E. Dandy (British Museum (NH) and BSBI, London, 1958) and changes published in Watsonia,
7: 157-178 (1969) or Flora Europaea (Cambridge 1964— 8) and may then be cited without authorities. Other-
wise authors’ names must be cited, at least on the first occasion where they appear in the text and followed by
the ‘Dandy’ or Flora Europaea name in parenthesis. Authors must follow strictly the International Code of
Botanical Nomenclature (Utrecht, 1966). Synonyms should be cited in chronological order; misapplied or
illegitimate names should be placed in square brackets. Examples will be found in recent parts of Watsonia.

Except for citations of the place of publication of plant names, which may be given in abbreviated
form in the text, full references should be listed in alphabetical order of authors’ names at the end of the
paper; the form used for references in a recent part of Watsonia should be followed carefully. Names of
periodicals should be abbreviated as in the World List of Scientific Periodicals (4th ed., London, 1963-1965).
References to herbaria should follow the abbreviations given in British Herbaria (BSBI, London, 1958) and
Index Herbariorum Part 1 (Sth ed., [APT, Utrecht, 1964). Foreign language references should, where necessary,
be transliterated into the roman alphabet according to a recognized convention; for Cyrillic script (Russian,
etc.) the Royal Society (BS 2979: 1959) or UNESCO schemes are acceptable. Unless authors expressly
indicate that they cannot get access to the necessary literature, the editors will assume that they have checked
the correctness of all titles, abbreviations, transliterations and references.

Papers should begin with a short abstract, in the form of a piece of connected prose conveying briefly
the content of the paper, and drawing attention to new information, new names and taxa, and the main
conclusions.

Line-drawings should be boldly drawn in Indian ink on Bristol board, smooth surface white cartridge
paper of good quality, or suitable draughtsman’s tracing materials, and should normaily be suitable for
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Remember that fine detail may be lost on even the best half-tone blocks.

Legends to text-figures and plates should be typed on a separate sheet of paper and attached at the end
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If in doubt about the citation of names or references, or the presentation of illustrations or tabular
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Further useful information and advice will be found in General Notes on the Preparation of Scientific
Papers (2nd ed., Royal Society, London, 5s. 0d.).

Proofs. Authors will normally receive a galley proof. Particular care should be given to the thorough
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Twenty-five offprints are given free to authors of papers. Further copies may be obtained at the Society’s
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The Society as a body takes no responsibility for views expressed by authors of papers.

Papers and short notes should be sent to Dr G. Halliday, Dept. of Biological Sciences, The University,
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vice-county recorders.

a] Fhrend & Polatschek i in » Britain

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Book REVIEWS = .
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Report of the Council for 1969 .. —.. AG
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Officers for 1971-72

Elected at the Annual General Meeting, 8th May 1971
President, D. McClintock

Vice-Presidents, Prof. D. H. Valentine, Prof. J. Heslop-Harrison, J. E. Lousley,
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Watsonia, 8, 345-356 (1971). 345

Studies on Cytisus scoparius (L.) Link with particular
reference to the prostrate forms

J. J. B. GILL and S. WALKER

Department of Genetics, University of Liverpool

ABSTRACT

The distribution in the British Isles of the prostrate forms of Cytisus scoparius (L.) Link is
given. Cultivation experiments show that growth habit is genetically determined. Whereas in
wild populations the prostrate form at Dungeness is indistinguishable in growth habit from
subsp. maritimus (Rouy) Heywood, there are significant differences under cultivation, when the
Dungeness plants show a growth habit intermediate between subsp. scoparius and subsp.
maritimus. Preliminary results from transplant experiments are reported.

Hybridization has been effected between erect and prostrate forms. Results suggest a difference
in expression of the genes controlling growth habit in the wild and horticultural varieties of
subsp. scoparius. The chromosome number of Cyfisus scoparius, erect or prostrate, is 2n = 46,
although isolated counts of 2” = 48 may be found.

INTRODUCTION

Cytisus scoparius (L.) Link (Sarothamnus scoparius (L.) Wimmer) is a common
species throughout Western Europe. It is very variable in height and Bocher &
Larsen (1958) have described a number of races or ecotypes which are primarily
recognisable on this character.

The extreme prostrate form of the taxon, recognised as Cytisus scoparius (L.)
Link subsp. maritimus (Rouy) Heywood, is genetically distinct from the erect
subsp. scoparius. This was shown by Tutin (1953) and Bocher & Larsen (1958),
who demonstrated that subsp. maritimus maintains its prostrate habit in
cultivation. According to Tutin, subsp. maritimus is the same as Sarothamnus
scoparius subsp. prostratus (Bailey) Tutin.

Many reports of the chromosome number of Cytisus scoparius subsp.
scoparius are available but no general agreement on one particular number has
been reached. Most early authors gave a count of 2n = 48 (Kawakami 1930;
Sugiura 1931, 1936; Tscheckow 1931; Kihara, Yamamoto & Hosono 1931;
De Castro 1949; Morton 1955; Bocher & Larsen 1955). Maude (1940) reported
a count of 2n = 46. This count of 2n = 46 has been confirmed by Boécher &
Larsen (1958), Melecka (in Skalinska et alia 1964) and Gilot (1965). In their 1958
paper, B6cher & Larsen stated that their previous count of 2m = 48 (1955)
arose from the misinterpretation of two large satellites as a separate pair of
chromosomes. The chromosome number of subsp. maritimus has been recorded
as 2n = 24 (Morton 1955), 21 = 48 (Adams 1957) and 2n = 46 (Bécher &
Larsen 1958).

From 1957 to 1961 a study of Cytisus scoparius formed part of a larger
programme of experimental taxonomic and genetical investigation undertaken
at Liverpool. This work was supported by a D.S.I.R. grant made to one of us

A

346 J. J. B. GILL AND S. WALKER

(S.W.). The work on Cytisus scoparius is a long-term project but the results so
far obtained are presented here.

The distribution of the prostrate forms in the British Isles has been ascertained
and studies have been carried out on both wild and cultivated populations,
with particular reference to their growth habit, cytology and hybridization.

DISTRIBUTION AND HABITAT OF THE PROSTRATE FORMS
IN THE BRITISH ISLES

Subsp. maritimus occurs in various exposed cliff habitats in the British Isles
and populations in Caernarvonshire, Pembrokeshire, Devon (Lundy Isle),
Cornwall, Co. Cork, Co. Kerry (Valencia Isle) and the Channel Isles have been
reported (Scully 1916, Wright 1933, Rees 1942, Tutin 1953, Morton 1955, ,
Adams 1957, Bocher & Larsen 1958). A prostrate population has also been
noted at Dungeness in Kent (Hepburn 1952, Morton 1955).

Localities on the mainland of Britain and Ireland have been visited by us
and the extent of the populations determined. Information on the populations
in the Channel Isles was supplied by a colleague and that for Lundy Isle and
Valencia Isle was obtained from previously published data. A survey in 1957
of many areas along the coast of Britain, from North Wales south to Land’s
End and east to Dungeness, did not reveal any other prostrate populations.
Fig. 1 indicates the total known distribution of prostrate forms of Cytisus
scoparius in the British Isles.

The most northerly site (area 1) for subsp. maritimus is near Parwyd on the
Lleyn peninsula in North Wales, where there is a small colony at the top of
an exposed south-facing cliff extending into the close-grazed Callunetum-
grassland on the cliff-top. It is locally abundant on the cliffs in Pembrokeshire
(area 2) in South Wales, where it occurs on the south-west coast of the county
from St Ann’s Head to Eastfield Farm and occasionally as far north as Trevine.
The habitat is similar to that in North Wales with the colonies, some of which
contain more than 100 plants, extending from the exposed cliff-face back into
the vegetation on the cliff-top. On Lundy Isle, off the north coast of Devon
(area 3), it is found on the exposed western cliffs (Wright 1933). In Cornwall
(area 4) it is locally abundant near the lighthouse at Lizard Point and is also
found at Kynance Cove and Gew Graze. At Lizard Point the habitat is similar
to that described for North and South Wales. The small colony at Kynance is
on a rocky outcrop a little way back from the cliff-top whereas that at Gew
Graze, also on a rocky outcrop, is about half a mile inland. Colonies also
occur on the east side of the Lizard peninsula on the cliff-face and top south of
Carleon Cove and in Downas Valley, where in many places it grows much less
prostrate due to heavy competition.

In the Channel Isles, subsp. maritimus is locally frequent on exposed south-
west-facing cliffs associated with Calluna, Erica and Festuca at Pleimont Point
and St Martin’s Point in Guernsey (area 5) and at Grosnez Point in Jersey
(area 6).

In Co. Cork it grows at Three Castles Head and more extensively at Mizen
Head (area 7); both localities are similar in habitat to those in Wales. It is also
reported (Scully 1916) as abundant on cliffs at Clynacartan on Valencia Isle,
Co. Kerry (area 8).

A further population of prostrate-growing plants exists at Dungeness (area 9)

STUDIES ON CYTISUS SCOPARIUS 347

FiGure 1. Known distribution in the British Isles of the prostrate forms of Cytisus scoparius.

in Kent. This habitat is strikingly different from those already described. It
comprises more or less flat and very extensive shingle with large numbers of
prostrate plants of Cytisus scoparius which form the dominant vegetation.
Whether the plants in this population are the same as subsp. maritimus will be
discussed later.

EXPERIMENTAL CULTIVATION OF CYTISUS SCOPARIUS

All plants were grown from seed of known wild origin. Seed was sown in August
or September in lime-free John Innes Seed Compost. Germination was greatly
improved if the seed coat was scratched prior to sowing. The young plants were
transferred to pots in a cold frame until the following March or April. They
were then planted out in their permanent positions and maintained without
competition at the University of Liverpool Botanic Gardens, Ness, Cheshire.

GROWTH HABIT

Subsp. maritimus has been distinguished from subsp. scoparius on the basis of
height (B6cher & Larsen 1958). The degree of hairiness has also been used as a
taxonomic character in some Floras, but although subsp. maritimus often

348 J. J. B. GILL AND S. WALKER

appears more hairy we have not found this a reliable character. Variation in
hairiness of Cytisus scoparius creates a large degree of overlap between subsp.
scoparius and subsp. maritimus. We consider that a more useful and complete
measure of the variation of growth-habit within the species can be obtained from

height? =

the ratio -}. This was calculated for all the three-year-old culti-
mean breadth

B

vated plants in September 1961. Height was taken as the maximum height
above ground level of the tallest shoot, and mean breadth as the average of the
greatest and smallest diameters of the plants.

The population at Ness at that time consisted of 88 specimens of subsp.
scoparius from Britain and Europe, 65 specimens of subsp. maritimus (of which
50 originated from Pembrokeshire, eight from Cornwall, five from the Lleyn
peninsula, one from Co. Cork and one from Guernsey), and eight plants from |
Dungeness.

The ratio described above when converted to log,, values gave good separa-
tion of the two subspecies on a histogram, with the specimens from Dungeness
falling approximately in the middle of the total distribution (Fig. 2). The means
and variances of the measurements for the three types are given in Table 1.

+ 1 +1°5 +, 2-#2°3

Ficure 2. Histogram showing separation of the forms of Cytisus scoparius using measurement
of growth habit.
Solid black area = subsp. scoparius.
Shaded areas = subsp. maritimus.
Outlined, unshaded areas = Dungeness form.

STUDIES ON CYTISUS SCOPARIUS 349
TABLE 1. MEASUREMENTS OF THREE-YEAR-OLD CULTIVATED PLANTS
OF CYTISUS SCOPARIUS

H2
oe == Height Mean Breadth
B in inches in inches

Mean Variance Mean Variance Mean Variance

subsp. maritimus —0-3594 0-1690 4°8923 14-4413 41-9923 198-3086
Dungeness +0-7999 0:0689 20-1250 30:4107 59-8125 39-7813
subsp. scoparius +1-7171 0:0494 59-9318 279-7195 63-6080 258-8417

In January 1970, further measurements were made of prostrate plants in
cultivation. This sample consisted of 51 specimens of subsp. maritimus from
the various localities mentioned before and 26 plants from Dungeness. The
plants were from 3 to 12 years old and the sample therefore approximated more
in age structure to a natural population than did the first series of measurements.
The results of this second series of measurements are given in Table 2. (The
erect subsp. scoparius is so obviously different from the two prostrate forms
that it was not included in this series).

Measurements of three wild populations, one of each type, were made in
1964 and these are given in Table 3. The population of subsp. maritimus was
from Pembrokeshire and the sample contained 83 plants. The Dungeness sample

TABLE 2. MEASUREMENTS OF MIXED-AGE CULTIVATED PLANTS OF

CYTISUS SCOPARIUS
H2 .
log =— Height Mean Breadth
B in inches in inches

Mean Variance Mean Variance Mean Variance

subsp. maritimus +0-4632 0-4232 14-4314 82-9302 48-8039 138-4108
Dungeness +0-9931 0:0906 29°8077 = 21:3202 79°7692 1474-8846

TABLE 3. MEASUREMENTS OF PLANTS OF CYTISUS SCOPARIUS IN WILD

POPULATIONS
H2 ,
log = Height Mean Breadth
B in inches in inches

Mean Variance Mean Variance Mean Variance

subsp. maritimus +0-1893 0-2702 =11:5904 51-4643 52-4650 372-0832
Dungeness +0-2400 0-0832 12-6400 20-4954 84-3500 699-0530
subsp. scoparius +1-:9364 0:0190 68-4833 291-4744 52-9167 396-1880

J. J. B. GILL AND S. WALKER

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PEATE.

Chromosomes of Cytisus scoparius (all x 1600).

a. Root mitosis showing 2” = 46 including two satellited chromosomes (arrowed).

b. Meiosis showing 23 bivalents.
c. Root mitosis showing 2” = 48 including two satellited chromosomes (arrowed).

STUDIES ON CYTISUS SCOPARIUS 351

contained 100 plants and the population of subsp. scoparius came from near
Whitchurch in Shropshire, the sample comprising 60 specimens.

An analysis of the three sets of results on the prostrate forms is summarised
in Table 4,in which figures are given only for those differences which are sig-
nificant.

CYLOLOGY

Chromosome studies have been carried out on both mitotic and meiotic pre-
parations. The somatic chromosome number has been determined as 2” = 46
in 33 specimens of subsp. scoparius, 29 specimens of subsp. maritimus (from a
variety of localities) and 11 specimens from Dungeness. One specimen of subsp.
maritimus from Cornwall gave a count of 2” = 48.

Mitotic preparations were obtained from root-tips pretreated for four hours
in a saturated aqueous solution of paradichlorobenzene (Meyer 1945), fixed in
1:3 glacial acetic acid : absolute alcohol, stained by the Feulgen method and
squashed in 45% acetic acid. Meiotic preparations were obtained from pollen-
mother-cells similarly fixed and stained and squashed in iron-acetocarmine
(Belling 1926). All preparations were made permanent by the freeze-drying
technique (Conger & Fairchild 1953), modified by finally passing through a
1:1 mixture of absolute alcohol : xylene to allow the mounting of preparations
in Canada balsam. Meiosis in Cytisus scoparius takes place very early in the
development of the flower; hence buds for meiotic studies were fixed when
about 3-4 mm long and just beginning to show the colour of the petals above
the calyx.

The somatic chromosomes of Cytisus scoparius (Plates la and lc) are too
small for effective karyotyping, being about 1-1-5 um long after pretreatment
with paradichlorobenzene. Most of them are either metacentric or sub-meta-
centric and, as reported by Bécher & Larsen (1958), two chromosomes carry
satellites nearly as large as themselves (arrowed in Plates la and Ic). The
connection between these satellites and the remainder of the chromosome is
frequently very tenuous, so that mistaken counts of 2” = 48 are easily obtained.
The one plant of subsp. maritimus from Cornwall which gave a count of 2n = 48
did show clearly a pair of satellited chromosomes (Plate Ic) so that a mistaken
count in this case is unlikely.

Meiosis in all specimens examined was normal with the constant formation
of 23 bivalents at metaphase I (Plate 1b). Bocher & Larsen (1958) reported the
presence of a single univalent in some of the meiotic cells from a plant of subsp.
maritimus from Jersey and interpreted this as being indicative of the presence
of a trivalent. No evidence of multivalents has been apparent in our investigation.
Chromosome segregation was normal at both anaphase I and anaphase II.
It was not possible to examine meiosis in the plant with 48 chromosomes, as
this count was made from one of a large number of seedlings which were
discarded after roots had been fixed.

HYBRIDIZATION

The pollination mechanism of Cytisus scoparius has been described by Miller
(1883). Essentially it is an explosive mechanism which depends for its effective-
ness on the bursting of the anthers and the shedding of their pollen over the

352 J. J. B. GILL AND S. WALKER

immature style and stigma while the flower is still in the bud. The style then
elongates so that, when the flower is completely open, it is held under tension.
The tension is released suddenly when the sutures of the keel are burst by an
insect visiting the flower and the style throws a cloud of pollen above the
insect to settle on its back. The stigma rapidly describes a spiral path and, in
doing so, first strikes the insect’s back where it may pick up pollen from the
previous flower visited. It then continues in its path to strike the underside of
the insect and possibly pick up pollen deposited there by the short stamens of
its own flower.

The existence of this mechanism suggests that emasculation of the flowers
prior to the bursting of their anthers would be a necessary preliminary to
successful hybridization. All attempts to emasculate flowers however resulted in
the buds withering. This also occurred when occlusion of the stigma with a
piece of grass culm, a technique found successful in Lathyrus (Davies 1957),
was substituted for emasculation. Application of various hormone solutions in
lanolin to the pedicel following either emasculation or occlusion of the stigma
allowed only a low frequency of bud retention to be achieved.

We noted, however, that flowers which remained unexploded or which had
been exploded artificially so as to minimise damage to the stigmatic surface
set very little seed, although abundant pollen might be present on the stigma.
This suggested that some mechanical damage to the stigma is necessary before
fertilization can occur.

We therefore gently burst the sutures of the keel by vertically compressing
it between the fingers, care being taken to prevent the style from springing.
As much as possible of the pollen adhering to the style and stigma was blown off.
The stigma was rubbed with forceps to damage it and then cross-pollinated.

In order to test the efficiency of this technique a number of prostrate, yellow-
flowered (wild type) plants were pollinated with pollen from erect cultivars
(red- or pale yellow-flowered). Approximately 80% of the progeny were recog-
nised as hybrid, having the habit and flower colour of the cultivars (Table 5).

This series of hybridizations was followed by a number of crosses between
the various growth-forms using plants of known wild origin. The results of
these crosses are summarised in Table 6. In this Table, plants marked with an

TABLE 5. HYBRIDIZATIONS BETWEEN WILD PROSTRATE AND ERECT
CULTIVARS OF CYTISUS SCOPARIUS

Female parent Male parent Hybrid

Growth Growth Flower Growth Flower

habit Origin habit colour habit colour
Prostrate Guernsey ETeCt Pale yellow Erect Pale yellow
Prostrate Guernsey Erect Scarlet wing Erect Scarlet wing
Prostrate Guernsey Erect Brick red Erect Brick red
Prostrate Pembrokeshire Erect Pale yellow Erect Pale yellow
Prostrate Pembrokeshire Erect Scarlet wing Erect Scarlet wing
Prostrate Pembrokeshire Erect Brick red Erect Brick red

Prostrate Lizard Erect Brick red Erect Brick red

STUDIES ON CYTISUS SCOPARIUS 353

TABLE 6. HYBRIDIZATIONS BETWEEN WILD PROSTRATE AND WILD
ERECT FORMS OF CYTISUS SCOPARIUS

Female parent Male parent Hybrid
Grown habit Origin Growth habit Origin Growth habit

erect Guernsey prostrate Guernsey prostrate
erect Guernsey prostrate Pembrokeshire prostrate
erect Poland prostrate Pembrokeshire prostrate
erect France prostrate Cornwall prostrate
erect Switzerland prostrate Pembrokeshire prostrate

erect Guernsey prostrate Dungeness erect*
prostrate Dungeness erect Ainsdale prostrate*
prostrate Pembrokeshire erect Poland prostrate*
prostrate Pembrokeshire erect Spain prostrate*
prostrate Pembrokeshire prostrate Dungeness prostrate*

* Putative hybrids awaiting analysis.

asterisk are considered to be hybrid but their certain identification as such
must await the analysis of F, segregants.

TRANSPLANT EXPERIMENTS

In order to compare survival, resistance to exposure and growth-rate of the
erect and prostrate forms of Cytisus scoparius in a habitat characteristic of
subsp. maritimus, 80 plants, each six months old (32 of subsp. scoparius, 32 of
subsp. maritimus, and 16 from the Dungeness locality), were planted out in 1964
on a cliff-face and top near Abermawr in Pembrokeshire. They were planted in
mixed groups of five (2 of subsp. scoparius, 2 of subsp. maritimus and one
from Dungeness) to eliminate as far as possible any ecological differences
within the area of planting. Seven of these plants are still surviving and all
exhibit a prostrate growth habit. However, only five can be identified, for the
labels from two have been lost. Four of these are subsp. maritimus (none from
Dungeness) and the fifth is a low-growing specimen of subsp. scoparius from
the coast of Britanny. The habit of the two unlabelled plants will be determined
by taking cuttings from them and growing at Ness. A further 50 plants of each
of the two subspecies were planted out randomly on the cliff in July 1969 and
their establishment and growth are being regularly recorded.

Of the seven plants established on the cliff since 1964, only one has so far
flowered. This was a specimen of subsp. maritimus which flowered after five
years, but did not set any seed. This compares with a three-year period before
flowering in cultivation.

DISCUSSION

The significant differences in growth-habit between cultivated populations of
subsp. maritimus, subsp. scoparius and the population from Dungeness, all of
uniform age, show that all three are genotypically differentiated from each
other.

354 J. J. B. GILL AND S. WALKER

The maintenance of the differences in cultivated populations of mixed ages
show that the Dungeness population is made up of plants which are geno-
typically taller and broader than typical subsp. maritimus. The variance of both

2

logio ® and height however is significantly greater in the cultivated mixed-age

population of subsp. maritimus than in the similar population from Dungeness,
suggesting that genotypically the Dungeness population is the more uniform
for these characters. The reverse is true for breadth (Tables 2 and 4).

The absence of any difference other than that of variance of breadth between
the wild and cultivated (mixed-age) populations of subsp. maritimus suggests
that this subspecies is strongly canalised to a fairly extreme prostrate growth-
habit. The greater variance of breadth in the wild as opposed to the cultivated
population may be due to competition in the wild or to the greater age differ-
ences within the wild population.

The cultivated mixed-age Dungeness population was significantly taller and

2

had a significantly greater value for logo imi than did the wild population

(Tables 2, 3 and 4). This suggests that the population at Dungeness is less
rigorously adapted to its particular growth-habit than is subsp. maritimus and
so suffers some degree of kill-back of upright shoots in the wild. The absence

2

! Jab
of any differences in variance, however, of either height or log, Br between

wild and cultivated populations must be a function of all plants in the wild

suffering equal exposure and therefore all being killed back to a similar height.

As the habitat at Dungeness is very homogeneous this effect might be expected.
2

There is no difference in either mean height or mean log,, a between the

wild subsp. maritimus and the population growing at Dungeness. The action
of the environment is therefore causing the Dungeness population to mimic
2;

subsp. maritimus. The greater variance of height and of log,5 nbs in subsp.

maritimus is probably due to the extreme heterogeneity of the Pembrokeshire
habitat in comparison with that at Dungeness. The greater variance in breadth
of the Dungeness population compared with subsp. maritimus may be due to
the absence of competition on the beach at Dungeness allowing the breadth
of any specimen to be limited only by its own capacity for lateral growth. The
breadth of the specimens at Pembrokeshire is most likely governed by com-
petition with other species, in particular Calluna vulgaris.

The chromosome number of Cytisus scoparius has been determined as
2n = 46 with the exception of a single plant of subsp. maritimus from Cornwall
which gave a count of 2m = 48. This confirms the observations of Bocher &
Larsen (1958). We also agree with these authors that many of the reported
counts of 27 = 48 probably arose from the misidentification of the two large
satellites as a separate pair of chromosomes. It is extremely difficult to account
for the single reported count of 2n = 24 (Morton 1954). Not only is his count
at variance with all others, but the chromosomes which he illustrates are much
larger than those shown by Bocher & Larsen or those seen by us. The longest
chromosome illustrated by Bocher & Larsen is approximately 1-5 »m, which
agrees with our observations, whereas in Morton’s illustration the shortest

STUDIES ON CYTISUS SCOPARIUS 355

chromosome is about 1-5 um and the longest is about 4-5m. This must raise
some doubts about the correct identification of Morton’s material.

No internal barriers to hybridization are apparent between prostrate and
erect forms of Cytisus scoparius. The results of confirmed crosses between pros-
trate forms and erect cultivars are different from those between prostrate forms
and erect wild plants (Tables 5 and 6). Whereas in the former the hybrid plants
are erect, in the latter they are prostrate. This difference in the expression of
the genes controlling growth-habit between two crosses suggests that the
horticultural coloured varieties of Cytisus scoparius are of hybrid origin and
contain, as well as genes causing alternative colour development in the flower,
genes governing growth habit which do not normally occur in wild Cytisus
scoparius.

The results of the transplant experiments so far analysed show that the
selection acting against the establishment of any other ecotype of Cytisus
scoparius in the natural habitat of subsp. maritimus is extremely strong, and
causes the elimination of very large numbers of young plants even when these
are planted out at a stage which overcomes any competition for initial establish-
ment.

The evidence so far accumulated suggests that Cytisus scoparius exists as two
quite distinct growth-forms and that these erect and prostrate forms are easily
distinguishable. The prostrate type in Britain is represented by subsp. maritimus,
which occurs on various exposed sea-clifis, and by the prostrate population on
the shingle beach at Dungeness. There is no chromosomal difference between
the various forms, all having 2n = 46. The Dungeness population, although
referable in the field to subsp. maritimus, behaves differently from this sub-
species in cultivation and has thus been shown to be genetically distinct from
subsp. maritimus.

ACKNOWLEDGMENTS

The authors wish to acknowledge the help given them by Mr T. A. W. Davis
in studying the detailed distribution of Cytisus scoparius in Pembrokeshire,
by Dr P. S. Dixon in collecting specimens from the Channel Isles, and by
Dr G. Halliday in supplying information on the population at Mizen Head,
Co. Cork.

REFERENCES

ADAMS, F. W. (1957). Chromosome numbers of Sarothamnus scoparius (L.) Wimmer. Watsonia,
4: 17-18.

BELLING, J. (1926). The iron acetocarmine method of fixing and staining chromosomes. Biol.
Bull. mar. biol. Lab. Woods Hole, 50: 160-162.

BOcHER, T. W. & LarsEN, K. (1955). Chromosome studies on some European flowering
plants. Bot. Tidsskr., 52: 129.

BOcHER, T. W. & LARSEN, K. (1958). Secondary polyploidy and ecotypical differentiation in
Sarothamnus scoparius. New Phytol., 57: 311-317.

Concer, A. D. & FaircuiLp, L. M. (1953). A quick freeze method for making smear slides
permanent. Stain Technol., 28: 281-283.

Davies, A. J. S. (1957). Successful crossing in the genus Lathyrus through stylar amputation.
Nature, Lond., 180: 612.

De Castro, D. (1949). Novas nemeros de chromosomas para o genero Cytisus. Agronomia
lusit., 11: 85-89.

356 J.J. B. GILL AND S. WALKER

Gmtot, J. (1965). Contribution a l’étude cytotaxonomique des Genisteae et des Loteae.
Cellule, 65: 317-347.

HEpPBurRN, I. (1952). Flowers of the coast, pp. 54, 135. London.

KAWAKAMI, J. (1930). Chromosome numbers in Leguminosae. Tokyo Bot. Mag., 44: 319-328.

KiHARA, H., YAMAMOTO, Y. & Hosono, S. (1931). Shokobutsu Senshokutaisi no Kenkyu.
Tokyo.

MaAubpgE, P. F. (1940). Chromosome numbers in some British Plants. New Phytol., 39: 17-32.

Meyer, J. R. (1945). Prefixing with paradichlorobenzene to facilitate chromosome study.
Stain Technol., 20: 121-125.

Morton, J. K. (1955). Chromosome studies on Sarothamnus scoparius (L.) Wimmer and its
subspecies prostratus (Bailey) Tutin. New Phytol., 54: 68-69.

MULLER, H. (1883). The fertilisation of flowers. London.

Rees, J. (1942), in Plant Records. Rep. botl Soc. Exch. Club Br. Isl., 12: 273.

SCULLY, D. W. (1916). Flora of Kerry, p. 64. Dublin.

SKALINSKA, M. et alia (1964). Additions to chromosome numbers of Polish Angiosperms.
Acta Soc. Bot. Pol., 33: 45-76.

SuGIuRA, T. (1931). A list of chromosome numbers in Angiospermous Plants. Tokyo Bot. Mag.,
45: 353-355.

SuGiuRA, T. (1936). Studies on the chromosome numbers in higher plants with special reference
to cytokinesis, I. Cytologia, 7: 544-595.

TsCHECHOW, W. (1931). Karyologisch-systematische Untersuchung des Tribus Sophoreae,
Podolarieae und Genisteae. Isvest. Tomsk Otdel. Russk. Bot., 3: 121-131.

TutTin, T. G. (1953). Sarothamnus scoparius subsp. maritimus (Rouy) Ulbrich, in New
combinations in the British flora. Watsonia, 2: 297.

WRIGHT, F. R. E. (1933). Contribution to the flora of Lundy Island. J. Bot., Lond., 71, Suppl.
Deel

Watsonia, 8, 357-368 (1971). 307,

Botanical Keys Generated by Computer
R. J. PANKHURST

Computer Laboratory, University of Cambridge

INTRODUCTION

This paper describes the application of a computer program to produce botanical
keys. A brief summary of this work has already appeared (Pankhurst, 1971)
and the methods used are discussed here in detail, the techniques of computation
having been considered in earlier papers (Pankhurst 1970a, b). In all cases the
taxonomy from which the data is derived is taken as already established. The
key generating program should be thought of as an aid to the identification of
specimens, not as a method for classification.

ADVANTAGES OF THE METHOD

The generation of keys by computer offers the following advantages:

(1) The manual labour of actually constructing the key is avoided.

(2) The key is as accurate as the data on which it is based. In manual key
construction, it is very difficult to be certain that the textual descriptions of
taxa correspond exactly with a key to the same material.

(3) An attempt can be readily made to shorten the key. The strategy by
which the key is constructed allows for an effort to make the key as efficient as
possible in the following sense. The most efficient key is taken to be that which
has the shortest average path (i.e. number of leads used) to a correct identifica-
tion of all taxa. It has been shown (Osborne 1963) that the key with the shortest
average route to an identification is dichotomous with groups of taxa of equal
size falling into each lead of a pair. The construction of such a key can also
take into account the different usefulness or ease of observation of the characters
used by the key and the commonness or rarity of the taxa. Improvement of keys
in the above sense is not feasible with manually constructed keys above a
certain size without considerable effort.

(4) It is easy to edit the key. If one wishes to add or remove taxa, or to add,
remove, correct or re-arrange the characters used, then a new key is produced
just by running the computer program again. In particular, this makes it easy
to produce keys based on the same taxonomy but tailored for different needs,
as for example, keys for herbarium specimens, or keys to perennials in winter.
This flexibility is achieved by altering the relative importance (weighting) of
the characters.

PERFORMANCE OF KEYS

A key is expected first of all to be accurate, and then in addition to be efficient.
An accurate key gives a correct identification with the great majority of
specimens. The most efficient key is one that requires the least effort on the
part of the user, and the theorem of Osborne indicates how this may be done.

358 R. J. PANKHURST

A key may give a wrong identification because of a number of different types
of error, which are discussed below.

(1) The specimen may belong to a taxon which is not included in the key in
any case. This should become apparent from either definite disagreement or
an unlikely result. An unlikely result is recognised by previous experience, or
failing that, by disagreement with other details not in the key, such as habitat
or distribution. Hence, as many details as possible of each taxon should be
used in the key, in order that taxa which do not belong are seen to disagree.
However, since a key can often be constructed without using all the available
characters, some of the remaining ones are often included at the point where
a taxon keys out. This helps the key-user to check his identification.

(2) Leads in a key can be misunderstood or characters of a specimen can be
misread. If there is only one character per lead in the key, then a wrong branch
can more easily be taken. This is much more serious at the beginning of a key
than towards the end, because the error causes the key-user to consider a larger
number of taxa, none of which is the right one. As a precaution against errors
of this kind, keys which have several characters in each lead are preferred,
since there is less doubt if several characters agree. Alternatively, several keys
using different sets of characters should be tried. If we give a numerical value
to the probability of answering each lead correctly, then these probabilities
have to be multiplied together to give the probability of arriving correctly at
the end of a sequence of leads, and finally to keying out. Suppose as an example
that this probability is 0-9, and consider a key for 50 taxa. Suppose that the
key is dichotomous, in which case about 6 leads will be used as a specimen
of each taxon is identified. This is because, ideally, two taxa could be separated
by one lead, four taxa by two leads, and so on. In general, a key for N taxa
involves the use of at least n leads per taxon, where 2” is approximately equal to
N. In the above example, the expected probability of getting a correct result for
an average taxon is 0-9 to the power of 6, i.e. about one half. In such a case,
the key is as likely to be right as it is likely to be wrong. Therefore, if errors
occur in the leads of a key, the key should be as short as possible to increase the
likelihood of a right result. This is, incidentally, a reason why keys to more than
a few hundred taxa are rare; even with a high probability of answering each
lead correctly, the chance of a correct final result can be small, so that large
keys are rather impractical.

It is often stated that ‘good’ characters should be used in keys. A good charac-
ter is one which is both easy or cheap to determine and which has a high prob-
ability of being correctly read. The ease of determination might be a matter of
time spent, or the need for equipment, training, or previous experience, or the
cost of chemicals or a technician’s pay. A character will more likely be read
correctly if it is, among other things, constant, available on average specimens,
and not too technical. These aspects of the quality of a character are partly
subjective, so measurement of the cost and probability is not generally practical.
For reasons of accuracy, those characters with the highest probability will be
used first in a key, since it is at the beginning of a key that an error would
have the worst effect. For reasons of efficiency, the easiest or cheapest characters
should be used first and the others, if necessary, used later. In general, therefore,
the characters in a key encountered in successive leads are often in the order of
decreasing quality and increasing cost.

(3) Characters used in the key can be missing from the specimen. This can be

BOTANICAL KEYS GENERATED BY COMPUTER 359

due, for example, to not having a complete plant, or to deterioration after
pressing, or to collection at the wrong season. If the key has only one character
in a lead at some point, and that character is missing, it is impossible to continue.
As above, this may be avoided by having more than one character in a lead, or
by having a variety of keys using different sets of characters. If a key with
several characters per lead can be provided, then the user may use the key by
selecting whichever characters he does happen to have. This is then really a
matter of selecting one of a number of alternative keys which are expressed
in the form of one. In this case, therefore, multi-character leads may be used
as if they are single-character leads. Key-users may often select only one
character from a multi-character lead, even when all the characters are available,
in order to save effort.

Once the considerations of accuracy have been considered, one may turn to
questions of efficiency. Dichotomous keys (where the leads are in pairs) are
often preferred over polychotomous keys (where there can be more than two
leads as alternatives). This is simply because a choice between two alternatives
is easier than between many. Each lead considered adds to the effort required.
It is also easier to choose between single character leads than between multi-
character leads in which every character is considered. Reasons have been
given for preferring multi-character leads, but, unfortunately, it is generally
difficult to find sets of characters in natural groups for use in leads which give
an efficient dichotomy.

More explicitly, multi-character leads producing an efficient dichotomy are
hard to construct at the top of a key, but easy to construct at the bottom,
where they occur as lists of differences between alternative taxa. A device which
is often used to counter this difficulty is to qualify auxiliary characters with
adverbs such as ‘usually’ or ‘rarely’. This is of limited usefulness because of a
clear element of doubt in the extra characters, as for example, in the following
pair of two-character leads:

1. Leaves hairy, flowers usually white.

Leaves glabrous, flowers pink.

As has been mentioned above, it has been shown using formal mathematics
(Osborne 1963) that the key with the shortest average route to an identification
is dichotomous with groups of taxa of equal size falling into each lead of a
pair. Such an arrangement is an ideal which can only be approximated to in
practice, because of unequal distribution of characters and odd numbers of
taxa. This does however give a very useful rule of thumb for choosing an
efficient key. Notice that the ‘shortest average route’ should take into account
the frequency with which specimens are found, so that common taxa should
key out after only a few leads, and rarer ones after more. Many keys will be
seen to key out unusual taxa near the beginning. If an unusual taxon is taken
as one which differs in many of its characters from the others, it may therefore
be expected to key out after rather few leads. If it is unusual in only a few
characters, then the efficiency of the key may be reduced by making it key out
near the beginning.

From the above discussion it is evident that the choice between few or many
leads, or single or multi-character leads, is a matter of compromise. The
computer program discussed below takes the following general strategy. The
order of the characters is used to decide on how to satisfy accuracy before
efficiency. If no order is specified, then only considerations of efficiency are

360 R. J. PANKHURST

used, to try and shorten the key. If an order is specified, and if, after this has
been satisfied, there is still some freedom of choice, rules of efficiency are used.
The program will try out as many characters per lead as it is asked to consider,
but will check that the multi-character leads are reasonably efficient, and so
may still prefer single character leads. All the remaining distinctive characters
are given when a taxon keys out, since this is useful and it does not interfere
with efficiency.

PROGRAM DATA AND OPTIONS

There is no restriction on the rank of taxa which may be put in to the program.
One may equally well work with families, genera, species, subspecies, varieties
or microspecies. There is nothing in fact in the program’s design to restrict its
application to botany. The taxa are described as a rectangular matrix of taxa
in rows versus characters (features, properties, attributes) in columns, filled
with values. Both the character and value are usually phrases in natural
language: for example, Epilobium roseum has an entire stigma, which is expressed
as a character ‘stigma’ with a value ‘entire’. The choice of the characters, and
the values assigned to them, are entirely in the hands of the program user.
Two or several values can be used with each character, i.e. characters can be
two- or many-valued. A value can just as readily be qualitative (e.g. ‘long’)
as quantitative (e.g. ‘10 to 20 cm’). Values can also be missed out if they are
not known, inapplicable or too variable to be useful.

The program prefers to produce dichotomous keys where possible, but can
create polychotomous keys if necessary. This depends on what proportion of
many-valued characters are given. The user may specify the maximum number
of characters that the program is to consider in combination as each branch
of the key is formed. The characters can be given equal weights, or any sequence
of weighting that the botanist prefers. It is also possible to give weights to the
taxa, the highest weight going to the commonest, so that the program will try
to arrange for a short path to common taxa, as far as the data allow.

Either of the two common forms of key can be produced, with an option
for indentation if required. Provision is made for highly distinctive taxa to be
recognised and keyed out early, although this is deliberately restricted because
it may interfere with the process of shortening the key. If the taxa are not all
distinguishable from one another with the given data, then a partial key can be
formed, such that several taxa may key out together. The character matrix has
to be sufficiently well scored before it is possible for keys to be constructed.
This requires that, initially, all the values of at least one of the characters must
be supplied. The names of the taxa form part of the data, and these may all
be different, or some of them may be the same. In the latter case, unnecessary
key branches involving only different descriptions of the same taxon are avoided.
As an aid in checking the data, the program will print a synopsis of all taxa.
This could be useful to a Flora writer who wishes to make his text agree exactly
with his key.

METHOD OF CONSTRUCTING KEYS

It should be remarked that since computer programs can evolve very rapidly,
the following discussion is only strictly accurate at the time of writing.

BOTANICAL KEYS GENERATED BY COMPUTER 361

At the beginning of the key construction process, all the taxa and all their
characters are available for consideration. The characters which have missing
values are then temporarily dismissed, since they cannot be used without
uncertainty. The fully scored characters are then examined to see which one
will form the best division. Suppose for the moment that all the characters have
equal weighting. If one character has two different values, then this character
could be used to form a pair of leads. If we ignore taxon weighting for the
moment, then this character would be a good choice if the two groups of taxa
with the two different values are about equal in size. Other characters might
have more than two values, but could still form conveniently equally-sized
groups with the different values. Another possibility is to choose any pair of
characters and construct leads with two characters at a time. This may be
expected to give rise to more than two leads and polychotomous branching.
Then one may consider three characters at a time, and so on, up to the maximum
number of characters available. In practice, a limit is set in the data as to the
maximum number of characters in a lead to be considered. Up to this limit,
all the possible combinations of characters are examined. The choice between
different possible ways of branching the key is made by calculating which is the
best from the number of leads and the number of taxa in each group. This is
described in detail below.

Once a branch has been made, then the existing leads and a list of the taxa
belonging to each lead are remembered. The process is then repeated on each
unfinished branch of the key. Notice that this time only those taxa which belong
to the previous lead are considered, and only those characters which have not
already been used in the preceding sequence of leads. To put this another way,
some of the rows and columns of the character matrix are ignored when lower
branches of the key are created.

It was assumed above that all the characters were equally weighted. If they
have different weights, then each time a new key branch is to be made, an
unused character with the highest weight is considered. It is possible for more
than one character to have the same weight. If just one character has a higher
weight than the others, then this one is used at once to make the next branch,
and the other characters are not examined. If several characters share the
highest weight, then these are examined as described above to choose the best
way of making a new branch, and those of lower weight are ignored, unless the
higher weighted characters turn out to have missing values.

A simple method of calculation is used to choose the best way of branching;
it is based only on convenience and has no theory to back it. The result of the
calculation is arranged so that it is always positive, and would be zero in the
ideal case. The key branching which gives the smallest result is the one which
is chosen. The result has two parts. The first part depends only on the number
of leads at a branch, and is given by (K—2)*, where K is the number of leads.
A dichotomy is the case K = 2, which makes the first part of the result zero.
A polychotomy of four branches (K = 4) would give (4 — 2)? = 4, which is
larger and not so favoured.

The second part depends on how equally the taxa are distributed in groups,
and is small for groups of equal size and large for unequal groups. A numerical
example will be helpful. Suppose we have a dichotomy with 2 taxa in one group
and 8 in the other, which is a total of 10 taxa. Since there are two groups, the
ideal would be if each group were of size *s = 5. If we take the ratio of the

B

362 R. J. PANKHURST

actual sizes to the ideal we have = = -4 and 2 = 1-6. Ideally these ratios would
each be unity. The differences from one with the sign removed are 1 — -4 = -6
and 1-6 — 1 = -6 and these are added to give a result for the second part of
1:2. A dichotomy of 4 and 6 respectively gives the second part equal to -4,
and would be taken as a better grouping. This calculation depends only on
the relative number of taxa in different groups. For example, if we take 4 and
16 taxa in a group of 20, the second part of the result is still 1-2. The two parts
are added together, dichotomies with equal groups therefore giving the lowest
result.

Taxon weighting, if required, is allowed for by using the numerical value of
the weighting to be an indication of the frequency of the taxon concerned. A
common taxon will be given a high weighting. If the original example above —
consists of two common species with weighting 6 each, and 8 local species each
with weighting 1, then the effective number of species in the first group is now
6 times greater i.e. 12. The total effective number of species is now 12 + 8 = 20.
The result for the second part of the calculation will now be -4 instead of 12,
which makes it more favourable to branch the key this way. The two common
species would then be separated by one more dichotomy, whereas the eight local
species would need about three more branches. Hence the desired effect of
shortening the path through the key to the identification of common taxa is
achieved. This method of taxon weighting was derived from that of Hall (1970).

To express the above as a mathematical formula for the case of no taxon
weighting, suppose that the size of each of the K groups is n; for i = 1 up to K,

K

and that the total number of taxa is N, where N = yn. Then the next key
i=1
branch is chosen by finding the minimum value of

, — ik
| N

K
CO a>

where the vertical lines stand for the absolute value, i.e. the expression between
the lines with the sign removed. The corresponding formula for taxon weighting
is
nj K
N’

K
(K—2)?+Y

a
i= 1

r

where n; is the effective number of taxa in group i (the sum of the taxon weights)
and WN’ is the effective total number of taxa.

Once the key is complete, the program works through it again to put in the
lead numbers, both before and after each lead. The key is remembered in the
order suitable for the indented type of key. If it is indeed this form of key that
is wanted, then the leads are sorted in the order of groups of taxa ascending
size, since this gives a better printed layout. If it is the bracketted type of key
that is required, then the details of the key are re-ordered after generating the
key and before printing it. Indentation is achieved by remembering with each
lead how many previous leads preceded the current lead, and this gives the
number of print positions to allow for the indentation.

In addition to the procedure outlined above, the program makes an attempt
to key out unusual taxa earlier than the rest. This is omitted if any of the taxa
to be compared have missing values of characters. The similarity of the taxa

BOTANICAL KEYS GENERATED BY COMPUTER 363

to each other is estimated and those which are most unlike the rest may be keyed
out separately, without the usual branch-choosing procedure.

EXAMPLES

The keys printed here as examples (Figs. 1-3) cover the species of Epilobium L.
found in Great Britain. Fig. 1 is produced by giving all the characters equal
weighting, so that the program alone has decided which to use and in what
order. Fig. 2 emphasises floral features while Fig. 3 emphasises vegetative
features. The second and third keys use character weighting to achieve the
different emphasis. All three keys were produced from one computer run, and
are all indented and in one of the two possible styles. This is a small example,
as it involves only 13 species. The flexibility of the computer approach shows
well in this example. The ability of the program to produce keys based on
fewer characters and with fewer leads as compared with hand-written versions
only begins to show with the order of 50 or more taxa. The keys in all three
figures have been reproduced as far as possible in the form that the computer
prints them. There are restrictions on the letters that can be used in FORTRAN
programming, which is why the keys cannot appear in the conventional style
of printing without being re-typed.

The data used for the Epilobium keys were derived from the Flora of the
British Isles (Clapham, Tutin & Warburg 1962) and the J/lustrated Flora of
Great Britain (Butcher 1961). Certain rules of thumb were used in transcribing
the written descriptions of species to the data matrix. It is sometimes necessary
to make deductions about character values from descriptions of the form
‘species B is like A but...’. Care is needed with adverbs such as ‘usually’,
‘rarely’ and ‘sometimes’ used in describing characters. One may choose to leave
out the value of such a variable character altogether, or repeat the taxon with
different character values, or just score the ‘usual’ value. The last choice means
that a variable character is incompletely scored, and the finished key can then
fail on certain ‘unusual’ specimens. It may be convenient to make simpli-
fications, for example in Epilobium “pale purple-lilac’ was scored as ‘pink’.

It was stated that the key-generating program prefers to work with characters
which are fully scored. This gives the program a better opportunity to create a
short key. This criterion of completeness is not usually required when descrip-
tions of taxa are prepared for Floras, so that, in the Epilobium example, the
data matrix prepared from two Floras was more complete than that which
could have been prepared from either alone. The experience with key-generating
to date shows that the collection of data on which to base a key requires most
of the effort.

The earliest example of a computer-generated key was based on the micro-
species of Alchemilla vulgaris. A key to the 134 genera of the Umbelliferae in
Flora Europaea (Tutin et alia 1968) was computed from data supplied by Pro-
fessor Tutin. These data came from an intermediate stage in the preparation of
the Flora, and are somewhat incomplete. The computed key proved to have 20%
fewer leads, and was based on fewer characters, than the handwritten version.
Keys for up to 116 potato cultivars have been computed for Mr T. Webster
of the National Institute of Agricultural Botany. One such key has been
published (Webster 1969). These keys have been especially welcomed since,
on account of the effort required, no handwritten key has ever been prepared.

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BOTANICAL KEYS GENERATED BY COMPUTER 367

Other keys have been created for common species of Cortinarius, microspecies
of Rubus fruticosus in Cambridgeshire, European species of Veronica, and
species of Jurinea (Compositae). The Veronica key used fewer leads and less
than half the characters than the corresponding handwritten key.

Keys relevant to other disciplines have been computed, in particular in
zoology, geology, medicine and mechanical and electrical engineering. Further
keys, generated at other institutions which have been given copies of the
program by the author, are not discussed here.

OTHER WORK

Several efforts have been made to compute the structure of keys, although
nearly all of these are numerical calculations without any arrangement to
write out the key at the finish (discussed in Pankhurst 1970a). Research by a
variety of authors in that branch of computer science known as artificial or
machine intelligence has produced a number of papers on decision trees, which
are similar to keys (discussed in Pankhurst 1970a). None of these authors
appeared to be aware of the use of keys in biology.

The only comparable program known to me is that due to Morse (Morse
1968, Shetler et alia 1971). This is similar to mine in its aims and achievements,
but differs considerably in detail. For instance, leads are not split into pairs of
characters and values, but each character value combination is treated as an
indivisible ‘couplet’. Character weighting takes effect by being included in the
computation for choosing key branching along with the number and size of
groups of taxa.

CONCLUSIONS

Diagnostic keys, until recently always composed by hand, can now be generated
by computer. A program to do this, written in standard FORTRAN computer
language, is available from the author on request.

ACKNOWLEDGMENTS

The author is indebted to Dr S. M. Walters of the Botany School, Cambridge,
for encouragement and valuable criticism throughout this research project.

REFERENCES

BuTcuer, R. W. (1961). A New Illustrated British Flora, vol. 1. London.

CLAPHAM, A. R., TuTIN, T. G. & WARBURG, E. F. (1962). Flora of the British Isles, 2nd ed.
Cambridge.

Hatt, A. V. (1970). A computer-based system for forming identification keys. Taxon, 19:
12-18.

Morse, L. E. (1968). Construction of identification keys by computer. (Abstract). Am. J. Bot.,
SS: 73).

OsBorNE, D. V. (1963). Some aspects of the theory of dichotomous keys. New Phytol., 62:
144-160.

PANKHUuRST, R. J. (1970a). A computer program for generating diagnostic keys. Comput. J.,
13: 145-151.

PANKHuRST, R. J. (1970b). Key generation by computer. Nature, Lond., 227: 1269-1270.

368 R. J. PANKHURST

PANKHurRST, R. J. (1971). Computer-generated keys, in Exhibition Meeting, 1969. Watsonia,
8: 336-337.

SHETLER, S. G., ef alia (1971). Pilot data processing systems for floristic information, in
CuTBILL, J. L., ed. Advances in Data Processing for Biology and Geology. London.

TuTin, T. G., et alia, ed. (1968). Flora Europaea, vol. 2. Cambridge.

WEBSTER, T. (1969). Developments in the description of potato varieties, 1. Foliage. J. natn.
Inst. agric. Bot., 11: 455-475.

Watsonia, 8, 369-377 (1971). 369

Six brambles (Rzdbz) from the north Midlands
A. NEWTON
11 Kensington Gardens, Hale, Cheshire

ABSTRACT

Descriptions are given of six new species of Rubus referred to in the author’s Flora of Cheshire
(1971).

DESCRIPTIONS

1. RUBUS ACCRESCENS A. Newton, sp. nov.

Turio primo semisuberectus, in autumno accrescens et decumbens, interdum
radicans, viridis, glaber, obtuse angulatus, sulcatus, aculeis sat validis sed
sparsis, rectis, e basi longa declinatis, ad angulos dispositis.

Folia (3-4) quinata, pedata, plana, atro-viridia, superne glabrescentia, subtus
ad nervos pubescentia, tenuia; petiolus longus et pubescens, aculeolis declinatis
vix uncinatis armatus, paullo canaliculatus. Foliolum terminale oblongum vel
ellipticum vel suborbiculare, basi cordatum, apice acuminatum cuspidatumque,
margine inaequaliter serratum.

Inflorescentia undulata, brevis, subracemosa, apice truncata, ramulis infimis
adscendentibus 2-3-floris, superioribus erecto-patentibus 1—2-floris.

Rachis obtuse angulata, pilis brevibus, aculeolisque sparsis, longis, gracilibus,
rectis, declinatis vestita.

Sepala inermia, tomentosa, albo-marginata, apicibus viridibus attenuatisque,
post anthesin reflexa, fructum laxe amplectantia. Petala alba, elliptica, apice vix
emarginata. Stamina alba stylos flavo-virides vix superantia. Carpella glabra.

Stem suckering, occasionally tip-rooting, at first growing at a low angle to
the ground surface, scrambling and trailing later in the summer, green, grooved,
bluntly angular, glabrous, eglandular except for minute sessile glands. Prickles
fairly strong, + straight, declining from stout bases, fairly sparse, confined to
the angles.

Leaves (3-4) quinate pedate, dark green, the petiole long, pubescent with
sparse declining curved pricklets, slightly furrowed above; leaflets thin, glabre-
scent above, shortly hairy on the veins beneath; terminal leaflet oblong, elliptic
or suborbicular with a cordate base and cuspidate, acuminate point, the margin
sub-compound serrate.

Panicle slightly flexuose, short cylindrical, few-flowered, subracemose above,
the apex truncate, with 1-2 distant lower branches which are sharply ascending
and 2-3-flowered, subtended by ternate leaves, the upper laxer, spreading or
slightly ascending.

Rachis bluntly angled, shortly pilose, armed with long, sparse, slender,
straight, declining pricklets.

Flowers c 2:5 cm diameter. Sepals felted, white-margined, with attenuated
green tips, reflexed after flowering, become patent on the ripening fruit, the tips

370 A. NEWTON

ascending. Petals, white, pale pink in bud, elliptic, the tips sometimes erose.
Stamens white, slightly exceeding greenish-yellow styles. Carpels glabrous.

Hototyprus: Southern part of Lindow Common, Cheshire, GR 33/825.794,
2/7/1970, A. Newton no. 13A1 (BM)

Characteristic features distinguishing this bramble from Rubus opacus Focke
ex Bertram, with which it sometimes grows, are the slender, straight, declining
prickles on the panicle, slightly later flowering commencement (in Cheshire
late instead of early June), and the strong, straight growth of new stems, at a
low angle to the ground, some later scrambling and tip-rooting. This plant is
particularly characteristic of the surrounds of peat mosses and damp heaths,
persisting amongst birch scrub; it is also to be found in scrub or open woods
on gritstone rocks in the Pennine moorlands.

Associated species include Betula pubescens, Corydalis claviculata, Deschampsia
flexuosa, Dryopteris carthusiana, Dryopteris dilatata, Galium saxatile, Holcus
mollis, Molinia caerulea, Rubus opacus, R. scissus, R. selmeri, R. sprengelii, Salix
cinerea, Sorbus aucuparia.

The observed range of distribution is from North Staffordshire south of
Leek, to the Glossop district of Derbyshire, extending over the morainic
foothills to the mosses and heathland of the Eastern Cheshire Plain. Hybrids
with Rubus sprengelii have been observed, distinguished by abnormally large
leaves, the terminal intermediate in shape, and fruit almost entirely abortive.

The species should be included in Rubus section Semisuberecti W.C.R. Wats.
which includes plants intermediate in character between Suberecti and Sylvatici.

2. RUBUS ROBII (W.C.R.Wats.) A. Newton, comb. et stat. nov.
R. muenteri Marss. var. robii W.C.R.Wats., Handbook of the Rubi of
Great Britain and Ireland, 287 (1956).

syn. R. godronii Lec. & Lam. var. foliolatus Rogers & A. Ley, J. Bot., Lond.,
44: 58 (1906), pro parte, quoad spec. ex Cheshire et Yorkshire.
R. muenteri Marss. sensu Sudre, Rubi Europae, 38 (1908), quoad exsicc.
leg. A. H. Wolley-Dod, ex Larkton Lane, Cheshire. Non R. muenteri
Marss. spec. auth. in Herb. Focke, Bremen, leg. Marsson ex Wolgast,
Jagerhofer Forst, 1876.
R. larktonensis W. C. Barton, MSS. in Mus. Brit.

Watson (1956 p. 287, 1958 p. 71) states that his var. robii differs only from
‘typical R. muenteri Marss.’ in the smallness of all its parts. However, the
type specimen of var. robii is not noticeably smaller than other specimens
named R. muenteri by Watson. The plant is widespread in Staffordshire,
Cheshire, South and West Lancashire, N.W. and N.E. Yorkshire, Derbyshire
and Warwickshire; it is quite distinct from R. muenteri and is considered to
merit specific rank.

The varietal description given by Watson (1958 p. 71) under R. muenteri
needs amendment only in a few particulars: leaflets are normally five, the
base of the terminal leaflet is better described as subcordate, the predominant
shape is obovate-pentagonal-suborbicular; the petals are usually pink, some-
times pale pink. The following Latin description is given here as there is no
satisfactory one elsewhere in the literature.

Validus, eglandulosus. Turio latiarcuatus, atrorufescens, leviter pilosus, obtuse

ee re

SIX BRAMBLES FROM THE NORTH MIDLANDS 371

angulatus, canaliculatus, aculeis validis rectis vel leviter declinatis ad angulos
dispositis.

Folia (3-4) quinata subdigitata, nonnunquam imbricata, plana, utrinque
viridia, inferne pubescentia, aequaliter serrato-dentata (dentibus haud_ pro-
fundis); petioli aculeolis falcatis obsiti. Foliolum terminale plerumque longe
petiolulatum, basi subcordatum obovatum vel pentagonum vel suborbiculare,
apice breviter cuspidatum.

Inflorescentia late pyramidalis ad apicem truncata, laxa, subracemosa; ramuli
medii vix ascendentes, 3—7-flori, superiores longe pedicellati.

Rachis pilis patulis vestita, aculeolis rectis vel leviter declinatis nonnullis e
basi longa validis praedita.

Flores c 2:5 cm diameter. Sepala externe inermia, villosa, albo-marginata,
apice foliosa, sub anthesi et post anthesin reflexa. Stamina alba ad basin
roseata stylos viridescentes vel rufescentes longe superantia. Carpella glabra.

Hototypus: Yearsley Moor, N.E. Yorks., 29/7/1937, C. M. Rob (BM)

3. RUBUS WIRRALENSIS A. Newton, sp. nov.

Turio altiarcuatus, rufescens, superficiebus leviter excavatis striatisque;
aculeis validis subaequalibus leviter declinatis, aculeolis sparsis, glandulis raris et
pilis crebris munitus.

Folia (3-4) quinata, pedata, + imbricata, utrinque viridia, in apricis coriacea,
infra pubescentia; petioli aculeolis falcatis declinatisque armati. Foliolum
terminale late obovatum vel suborbiculare cuspidatum vel acuminatum,
elusdem petiolulo ter longius, simpliciter, vel ad apicem irregulariter serratum,
basi truncatum vel emarginatum. Stipulae villosae glandulis brevibus munitae.

Inflorescentia pyramidalis (foliis nonnullis quinatis ternatisque inferne
instructa, superioribus ad folium abbreviatum unum saepe redactis) racemosa,
in apricis saltem multiflora, ramuli medii multiflori, leviter adscendentes vel
primo suberecti, superiores patuli.

Rachis breviter tomentoso-villosa, aculeis tenuibus autem validis aliquot
armata, glandulis stipitatis haud multis sed in pedunculis superioribus crebrior-
ibus praedita.

Sepala tomentosa griseo-albescentia, post anthesin et sub fructu reflexa.
Petala angusta, obovata rosea. Stamina alba, ad basin rosea vix stylos superantia.
Carpella sparsim pilosa.

Stem high-arching, rufous particularly in exposure, subsulcate, faces shallowly
grooved, striate; armed with prickles often tipped yellow, strong (+ stem width),
slightly declining, almost confined to the angles; with scarce pricklets, rare
short-stalked glands and frequent short hairs.

Leaves (3-4) quinate pedate, + imbricate, green on both sides, thick in
exposure, softly pilose and slightly felted beneath, the veins prominent; petiole
prickles hooked, declining. Terminal leaflet broadly obovate or suborbicular,
rarely cuneate-based, cuspidate or acuminate, approximately three times the
length of its petiole, evenly serrate but more irregularly toothed towards the
apex, the teeth patent or directed forwards, with truncate or emarginate base.
Stipules linear, villose, shortly glandular.

Panicles pyramidal, narrowing to a truncate apex, with several ternate and
quinate leaves below, and one or two simple leaves above, compound racemose

3/2 A. NEWTON

in well-developed specimens, the median branches many-flowered, slightly or
occasionally sharply ascending at first, the uppermost spreading.

Rachis clothed with fine ashy tomentum beneath dense patent hairs, a few
slender but strong declining prickles and variable quantities of stalked glands,
shorter than or + equal to the hairs. The glands are most numerous in the
upper portion of the panicle and on the upper pedicels.

Flowers c 2-5 cm diameter. Sepals grey-white felted, reflexed after flowering
and remaining so in fruit. Petals long, narrow, obovate, deep pink. Stamens
cream, pink-based, + equal to styles. Anthers glabrous. Carpels sparingly pilose.

Ho.totypus: Hedge bank, Aldford, Cheshire, GR 33/423.581, 19/7/1970,
A. Newton no. 182A1 (BM)

This plant is abundant in Wirral on clay and sandy soils but is particularly
well-developed on the damper clays, where it forms a conspicuous element of
hedges, wood borders and thickets. It appears to be absent however from the
immediate sublittoral fringe of the north-west and north Cheshire coasts. It
exhibits some variation of leaf-size, shape and texture, leaflet-number and
strength of stem-armature due to its wide range of habitat, being equally
tolerant of damp thickets on clay and open, dry, sandy banks. It is not however
a species of the open heath. Some specimens from Wirral in MANCH are
labelled ‘R. mucronatus’ (=R. mucronifer Sudre), with which there is some
affinity, but there are also significant differences, for example glabrous anthers,
narrower, deeper pink petals, more complex and racemose panicle with less
glandular but hairier rachis and more imbricate leaves with a tendency towards
cuneate bases in some terminal leaflets.

The plant is however not confined to Wirral but extends as far as the lower
Dee Valley; specimens have also been seen from Anglesey, South Lancashire
and West Lancashire, one of which from near Knott End (see below) was said
by W. M. Rogers to be “‘in leaves somewhat intermediate between R. cinerosus
and R. mucronatus but I suppose going best under the latter”. Identical plants
have also been found recently by E. S. Edees in several places in Connemara.
The following exsiccata (inter alia) are in MANCH:

Patrick Wood, Claughton, Cheshire, —/7/1862, —-, ‘R. mucronatus’

Speke Road, Garston, South Lancs., 7/7/1872, J. H. Lewis, °“R. mucronatus
Bor, Blox.’

Egerton Road, Claughton, Cheshire, 2/9/1871, J. H. Lewis, “R. mucronatus’

Hillside, Holyhead, Anglesey, 8/10/1885, C. Bailey, ‘R. mucronatus Y

Lane by canal, Maghull, South Lancs., 5/8/1902, J. A. Wheldon, “R. cinerosus’

Near Knott End, West Lancs., —/7/1901, J. A. Wheldon, *R. mucronatus
Blox.’

Preesall, West Lancs., —/7/1902, J. A. Wheldon, ‘R. cinerosus Rog.’

The position of this species in Dandy (1958) should be within the section
Appendiculati (Genev.) Sudre close to R. mucronifer Sudre.

4. RUBUS PORPHYROCAULIS A. Newton, sp. nov.

Turio arcuato-procumbens, obtusangulus, gracilis, purpurascens, pilosus,
superficiebus planis, striatis; glandulis crebris purpureis, quam pilis brevioribus
ornatus; aculei declinati vel subfalcati, e basi longa compressi, nonnulli

SIX BRAMBLES FROM THE NORTH MIDLANDS 373

mediocres, acicularibus minimis immixti. Specimina omnia ex apricis pro-
venientia parum validiora paullo validius crebriusque armata.

Folia 3(4)-nata, plana, supra glabrescentia, subtus ciliata, atrovirentia.
aequaliter serrata. Foliola inferiora patentia vel e petiolo declivia. Foliolum
terminale ovatum vel pentagonum, breviter cuspidatum, basi truncata vel
rarius emarginata, eiusdem petiolulo quater longius.

Inflorescentia laxa, elongata, apicem versus vix decrescens, basi leviter
flexuosa, bracteis foliaceis basi ternatis, superne in foliolum unum redactis.
Ramuli patuli, 3—4-flori, ad apicem plures 1—2-flori.

Rachis obtusangula, striata, pilosa, glandulis crebrissimis, aculeis raris
falcatis praedita.

Sepala deflexa et post anthesin et sub fructu, glanduligera, attenuata. Petala
alba, mediocria, distantia. Stamina alba stylos virides vix superantia. Carpella
glabra.

Stem low arching, bluntly angular, slender, striate, purplish-red or purplish-
green in shadier places, hairy, with many purple, stalked glands, nearly all less
than or equal to the hairs, and many acicles of similar length. Prickles rare or
few, the larger nearly equal in size, as long as the stem-width, mostly on the
angles, the smaller about half the size of the larger, slanting from long com-
pressed bases. The tips of the prickles often yellowish and hooked. The density
of all the arms is higher in open-grown plants.

Leaves flat, almost uniformly ternate, rarely 4-nate, thin, dark green,
glabrescent above, lighter green but not felted, with sparse short hairs beneath.
Terminal leaflet ovate-pentagonal, evenly serrate with some patent teeth
particularly towards the apex, shortly cuspidate with a truncate or emarginate
base, about four times as long as its petiolule; basal leaflets at right-angles or
somewhat deflexed from the petiole, lobed in the lower portion when ternate.
Petioles glandular hairy and like the stem aciculate with short falcate prickles.
Stipules linear, long, hairy.

Panicle usually lax, cylindrical, the lower branches stronger and more flori-
ferous. The lower foliaceous bracts ternate, the upper two-lobed and simple.

Rachis usually wavy at the base, with dense spreading hairs, purple stalked
glands about half as long as the hairs, and slightly declining and curved, sub-
equal pricklets.

Sepals ovate, acuminate, shortly glandular, hairy, with slender points, light
grey margins, felted within, strongly reflexed after flowering and remaining so in
fruit. Petals small (1:75—2 cm), obovate, white or faintly pinkish, spaced. White
stamens barely exceeding the greenish styles. Carpels small, well-formed.

HoLotypus: Woods near Marple Sanatorium, Goyt Valley, Cheshire, GR
33/950.897, —/8/1969, A. Newton no. 223A1 (BM)

This species has been found to be locally plentiful in the Goyt, Etherow and
Tame valleys and their tributaries (N.E. Cheshire and N.W. Derbyshire around
Marple, Romiley and Glossop) in Quercus petraea oakwood on medium acid
soils. Other Rubi present in this district are: R. polyanthemus, R. calvatus,
R. selmeri, R. sprengelii, R. accrescens, R. echinatoides, R. distractiformis, R.
hylocharis, R. newbouldii, R. vestitus.

Rubus porphyrocaulis is quite distinct in habitat and physiognomy from all
these species. It has affinities with R. flexuosus Muell. & Lefév. and there are

374 A. NEWTON

specimens in MANCH collected by W. R. Linton from Dinting (22/8/1894)
and Charlesworth (23/8/1894), both in north Derbyshire, labelled ‘R. foliosus
Weihe & Nees’. The latter name has often been regarded in the past as
synonymous with R. flexuosus. Another gathering of the plant by C. Bailey
from quarry refuse between Mottram and Charlesworth (17/6/1893) was
labelled ‘R. podophyllus’ by Linton (this sheet also includes a leaf of R. newbouldii
Bab.).

Similarities to R. flexuosus lie in the deflexed, grey-felted sepals and the single
terminal ‘leaf’: it differs in more pronounced hairiness, stronger stem armature,
less zig-zag panicle rachis, purplish dark-green cast and white flowers. True
R. flexuosus is not known in Cheshire or, reliably, in Derbyshire.

5. RUBUS ADENANTHOIDES A. Newton, sp. nov.

Turio fusco-rufescens, obtuse angulatus, leviter pilosus, superficiebus planis
vel concavis, striatis; aculeis crebris e basi longa et angusta abrupte contractis,
rectis vel subfalcatis, declinatis 50°-80°, aculeolis aciculisque numerosis
inaequalibus (nonnullis glanduligeris) et glandulis paucis obsitus.

Folia 3-4(5)-nata pedata, subimbricata, profunde duplicato-crenata, subtus
leviter pilosa, viridia vel cinereo-tomentosa. Foliolum terminale angusto-
ellipticum, elliptico-obovatum vel rhomboideum, acuminatum, basi subcordata,
elusdem petiolulo 3-5—4:5 longius; foliola infima breviter pedicellata.

Inflorescentia pyramidata, ad apicem truncatum vel rotundatum decrescens,
inferne foliis ternatis, superne foliis usque 4 simplicibus subtus cinerosis ornata.
Ramuli medii ascendentes c 60°, 3—5-flori, superiores breves, patuli, cymosi,
1—3-flori, pedunculis pedicellisque brevibus ad apicem fasciculatis.

Rachis inferne vix flexuosa, pilosa, superne villosa, tomentosa; aciculis
numerosis (nonnullis glanduligeris), glandulis stipitatis inaequalibus, aculeolis
flavescentibus inaequalibus rectis vel declinatis praedita.

Sepala cinereo-tomentosa aculeolata post anthesin patentia vel leviter reflexa,
mox fructum laxe amplectantia. Petala dilute rosea, oblonga, apice emarginata.
Stamina alba stylos pallide luteos vix superantia. Carpella glabra vel parce
pilosa. Receptaculum pilosum.

Stem reddish brown, sparsely hairy, obtuse-angled, sides flat or shallowly
grooved, striate, armed with numerous light brown, yellow-tipped prickles
slanting at an angle of 50°-80°, straight or slightly falcate, abruptly contracted
from a long compressed base, frequent pricklets and acicles (some gland-
tipped) and a few short-stalked glands.

Leaves 3-4(5)-nate, pedate, petioles armed as the stem but with some longer-
stalked glands. Leaflets almost imbricate with margins deeply subcompound-
crenate, slightly hairy, green or grey felted beneath. Terminal leaflet narrow
elliptic, elliptic-obovate or rhomboidal acuminate, base subcordate, 3:5—4:5
times the length of its petiolule. Petiolules of intermediate leaflets c 1/3rd length
of that of the terminal leaflet, of basal leaflets very short.

Panicle pyramidal narrowing to a truncate or rounded apex, with several
ternate leaves below and up to four simple leaflets above, at least the uppermost
grey felted beneath. Median branches 3—5-flowered, ascending (c 60°). Apical
branches short, + patent, cymose, 1-3-flowered, with very short pedicels
giving a narrow congested top.

Rachis slightly flexuose and pilose below, villose and felted above with

SIX BRAMBLES FROM THE NORTH MIDLANDS 375

abundant acicles, some gland-tipped, glands short-and long-stalked, and num-
erous yellowish declining pricklets slightly exceeding the stem-width, straight or
slightly curved.

Flowers 2-2-5 cm diameter. Sepals armed with short pricklets, slightly
glandular, grey felted with short linear tips, at first patent or loosely reflexed
then loosely clasping the fruit. Petals pale pink, elliptic, notched at the tip.
Stamens white, slightly exceeding cream styles. Anthers glabrous. Carpels
glabrescent. Receptacle hairy.

Ho.Lotypus: Lane between Harthill and Dropping Stone House, Cheshire,
GR _ 33/50.55, 25/7/1970, A. Newton no. 225A1 (BM)

Rubus adenanthoides is one of several taxa included by Rogers (1900 p. 53)
in R. adenanthus. It is abundant in the Peckforton Hills area of West Cheshire
and is also widespread in other parts of the British Isles. I have seen the following
exsiccata:

MANCH: Below Peckforton Castle, Cheshire, 5/8/1903, A. H. Wolley-Dod,
“R. adenanthus’
CGE: Under Peckforton Castle, Cheshire, 18/9/1951, W. H. Mills, ‘R. pseuda-

denanthus’

N.E. margin of Peckforton Hills, Cheshire, 2/8/1952, W. H. Mills, ‘R. pseuda-
denanthus’

Cleadon, N.E. Yorkshire, 1865, J. G. Baker, ‘R. pallidus’

Wood below Peckforton Hill, Cheshire, 11/8/1902, A. H. Wolley-Dod,
*R. adenanthus’

Hedge W. of Bickerton sandpit, Cheshire, 29/7/1905, A. H. Wolley-Dod,
*R. adenanthus’, ‘I agree WMR’

Lane at Pennsylvania, Cheshire, 12/8/1903, A. H. Wolley- Dod, ‘R. adenanthus’ ,
‘apparently WMR’

Below Peckforton Castle, Cheshire, 3/8/1963, B. A. Miles, ‘R. adenanthus,
Rog. Hbk.’

Roundhill Wood, | mile N. of Clifton Hampden, Oxon, 6/8/1965, B. A. Miles,
“R. pseudadenanthus’

S. edge of Wood, Humphrey Head, N. Lancs., 6/8/1964, B. A. Miles, ‘R.
pseudadenanthus’

Below Gowbarrow Park, Ullswater, Cumberland, 4/8/1964, B. A. Miles,
*“R. pseudadenanthus’

Lower Ballacotter Glen, I.0.M., 6/8/1963, D. E. Allen, *? R. pseudadenanthus’

Birtley, Northumberland, Y. Massey, ‘R. pseudadenanthus’

Lane to Mountshannon Pier, L. Derg (H9), 4/9/1963, D. E. Allen

This species was certainly included in Watson’s (1958 p. 138) concept of
Rubus pseudadenanthus but unfortunately the type specimen of this taxon (BM)
comes from Roscarrock Hill, Port Isaac, Cornwall, and is a hard-ground state
of R. echinatus Lindl. The name R. pseudadenanthus must therefore be discarded.

6. RUBUS DISTRACTIFORMIS A. Newton, sp. nov.

Turio arcuato-decumbens, obtusangulus, pruinosus, pilosus, superficiebus
planis, striatis; aculei numerosi e basi angusta declinati, robustiores subaequales
ad angulos dispositi, aculeoli et aciculi et glanduli stipitati numerosi pilis
plerumque breviores.

376 A. NEWTON

Folia 3(4-5)-nata, pedata, utrinque viridia, serrato-dentata, supra glabre-
scentia, inferne pubescentia; foliolum terminale obovatum vel ellipticum,
cuspidatum, ad basin subcordatum vel emarginatum, eiusdem petiolulo ter
longius.

Inflorescentia ad apicem fere aphylla, angusta, elongata, apicem versus vix
decrescens, ramuli inferiores 3—7-flori adscendentes, superiores 1—3-flori suberecti.

Rachis villosa, hirsuta, sulcata, striata, aculeis numerosis vix declinatis,
glandulis nonnullis longis, aculeolis glanduligeris raris munita.

Sepala tomentosa, glandulis aciculisque praedita, post anthesin reflexa sed
sub fructu nonnulla subpatentia. Petala alba, angusta, distantia. Stamina alba
stylos virescentes superantia. Carpella pilosa.

Stem high arching then decumbent, obtusely angled, pruinose, hairy with
simple and stellate hairs, striate, armed with many, often yellow-tipped prickles
and pricklets slanting from narrow bases, the stronger confined to the angles,
about half the width of the stem, numerous acicles and scattered short-stalked
glands mostly shorter than the hairs.

Leaves 3(4~-5)-nate pedate, light green on both sides, glabrescent above,
pubescent beneath; terminal leaflet obovate or elliptic-obovate, serrate-dentate,
some of the larger teeth sometimes patent or recurved, cuspidate, the base
shallowly emarginate or subcordate, 2-3 times the length of its grooved petiole,
which is pilose and armed with scattered falcate pricklets and short glands.

Panicle narrow cylindrical, elongate, all branches + erect, the lowest 3-7
-flowered, the upper 1—3-flowered.

Rachis villous and hairy, grooved and striate with many scarcely declining
prickles, sparse long glands and rare gland-tipped pricklets.

Flowers c 2:5 cm diameter. Sepals felted, clothed with long and short glands
and acicles, reflexed in flower and early fruit, a few spreading on the mature
fruit. Petals white, narrow, spaced. Stamens white, exceeding the greenish styles.
Carpels hairy.

Ho.ortypus: North bank of R. Bollin, Hale, Cheshire, GR 33/793.846, 5/7/1970,
A. Newton no. 248A1 (BM)

In shade the armature is weaker and the panicles much abbreviated; the
predominant light green cast, ternate leaves often held erect from the pruinose
stem, even armature and pilose panicles are distinctive features.

This bramble is widespread over a considerable area of the western Pennines
and their foothills from the Leek (Staffs.) area to Glossop (Derbyshire) including
the valleys of the Mersey tributaries, and it extends over the morainic foothills
into the Cheshire plain as far as Northwich, Knutsford and the eastern part of
Delamere Forest.

It appears to be equally prevalent on boulder clay and marl as on Carbon-
iferous shales but it is particularly prominent on shallow soils derived from
glacial sand in the vicinity of old mosses and heathland where it is a major
member of the scrub community in woodland and hedgerows.

Specimens collected by botanists at the end of last century received a number
of names and the following exsiccata are in MANCH:

Hedge in lane near Knutsford, Cheshire, 31/8/1889, C. Bailey, ‘Pyramidalis
Kalt. teste Focke’; ‘nr. Bloxamii, J. G. Baker’, ‘like Welsh Newton plant
(R. hirsutissimus), C. C. Babington’

SIX BRAMBLES FROM THE NORTH MIDLANDS 377

Wood between Chelford and Alderley Edge, Cheshire, 20/8/1898, C. Bailey,
‘R. regillus A. Ley, form less glandular than type, W. Moyle Rogers’

Hedge in cornfield near Knutsford Heath, 6/8/1884, C. Bailey, ‘R. derasus
Mull. ?’

Knutsford Heath—Tabley Lane, 6/9/1873, F. M. Webb, ‘R. warrenii Blox.
MS’

Rushton Spencer, Staffs., 7/1890 W. H. Painter, ‘R. hystrix’

Rudyard Moor, Staffs. at 600’, 27/8/1889, W. H. Painter, ‘R. distractus
P.J.M. teste Focke’

Between Rudyard and Harracles Mill, Staffs., 8/9/1887, C. Bailey, ‘R. radula’

This plant is also referred to by Rogers (1900 p. 94) as R. distractus Muell.
and by Butcher (1961 p. 658) as R. disjunctus Muell. & Lefév.

With the exception of R. warrenii Blox. MS.,comparisons have been made with
authentic material of all these taxa from which R. distractiformis is, despite
occasional similarity in some points of detail, clearly distinct. It is nearest to
R. menkei Weihe & Nees, of which I have seen an authenticated specimen in
Babington’s herbarium (Focke’s Rubi selecti no. 81). R. distractus (to which
Focke referred the Rudyard Moor, Staffordshire plant listed above) is synony-
mous with R. menkei. This was the opinion of W. C. Barton (MSS. in Mus. Brit.)
on the basis of a specimen collected and named R. menkei by Weihe and now in
Focke’s Rubi authentici. I have seen a photograph of this specimen and also one
of Miiller’s type specimen of R. distractus from the Vosges and agree with
Barton’s view. R. distractiformis differs from R. menkei in the following respects:

Leaf shape and indentation: the terminal leaflet in R. menkei is pentagonal,
considerably incise-serrate at the apex, and abruptly acuminate.

Stem armature: the longer prickles in R. menkei are sparser and unequal.

Rachis armature: in R. menkei the prickles are more declining, less crowded
and long glands are more frequent.

The affinity with R. menkei is sufficiently close to justify its inclusion next
to that taxon in Dandy (1958) within the section Appendiculati (Genev.) Sudre.

ACKNOWLEDGMENT

I am greatly indebted to E. S. Edees for his advice and encouragement during
my Rubus studies.

REFERENCES

ButTcuHer, R. W. (1961). I/lustrated British Flora, vol. 1, p. 658. London.

Danpy, J. E. (1958). List of British Vascular Plants. London.

Newton, A. (1971). Flora of Cheshire. Chester.

Rocers, W. M. (1900). Handbook of British Rubi. London.

Watson, W. C. R. (1956). New species and combinations in the genus Rubus L. Watsonia,
3: 285-290.

Watson, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland. Cambridge.

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Watsonia, 8, 379-384 (1971). 379

Studies on Gloucestershire populations of Allium
paradoxum (Bieb.) G. Don

D. M. BARLING

Royal Agricultural College, Cirencester

ABSTRACT

Populations of Allium paradoxum in woodland, headland and grass road-verges near Barton-
bury, Cirencester, have been studied since 1948. The seasonal development of the plant is
outlined, and data are given on bulbil production, floral abnormalities, stature, leaf-dimensions,
and the density of established plants and young plants produced from bulbils.

Of a sample of 250 plants, 72° showed floral abnormalities. Only five mature seeds were
found in the course of extensive searching. Reproduction is predominantly vegetative, mainly
by bulbils.

INTRODUCTION

Allium paradoxum (Bieb.) G. Don is a perennial species of shady woods in
the Caucasus, north Persia and neighbouring parts of central Asia (Vvedenskii
1935). It has been naturalised for over a century in northern Britain, especially
in mid-Scotland. In recent years many new localities have been reported in the
south of England where the plant appears to be spreading rapidly. It has been
recorded in Gloucestershire by Riddelsdell et alia (1948) and the object of the
present work is to study populations near the junction of the Stroud and Tetbury
roads, Cirencester, particularly at Bartonbury House.

It is probable that the original centre of establishment was Bartonbury gardens,
where the species might have been introduced for decorative purposes or
naturalising in rockeries. At present, dense populations are found in the wood-
lands of the grounds and also on road-verges nearby, though they cover only
small areas. The associated flora of the woodland and verges is presented in
Table 1. The sites overlie the Great Oolite limestone and the soil pH at six sites
(woodland, road-verges, headland) lies between 6-8 and 7:3. A number of sites
have been studied since 1948 and in most subsequent years new small pop-
ulations have been recorded; two are about 1 mile away, near Chesterton bridge
and along the Fosse Way.

The spread of the species has been influenced by several factors. First, the
movement of waste material and formation of leaf-mould dumps have provided
ideal means of spreading the numerous bulbils that lie on the soil surface.
Secondly, the mowing of road-verges and the sweeping of the hay has moved
and shaken off bulbils. Thirdly, the species is frequently picked for its appear-
ance in springtime and eventually thrown away because of its overwhelming
odour. In this way bulbils are dispersed over wider areas than is normal and this
may be the cause of establishment at Chesterton. The net result has been a
gradual movement along the Stroud and Tetbury roads.

380 D. M. BARLING
TABLE 1. ASSOCIATES OF ALLIUM PARADOXUM

Bartonbury Bartonbury Fosse Way

Wood (Verges) (Verge)
Bromus ramosus re)
Dactylis glomerata a O
Festuca rubra a re)
Poa trivialis f
Poa angustifolia f re)
Arrhenatherum elatius f re)
Brachypodium pinnatum oO a-d
Brachypodium sylvaticum oO
Geranium robertianum f
Geranium lucidum ‘e)
Saponaria officinalis re)
Eranthis hyemalis O
Rubus sp. r
Hedera helix a-d
Allium vineale ’e)

d dominant, a abundant, f frequent, 0 occasional, r rare

POPULATION DENSITY AND SIZE

The population density of the Bartonbury centre is remarkable and during late
winter a thick carpet of established plants and newly germinated bulbils is
found. There are in the main wood approximately one million plants and the
density may be as high as 100/sq. ft but this varies (Table 2). If bulbil develop-
ment is impeded in the previous year by early cutting then the plant density is
not high. The number of plants which are capable of flowering is much lower
and they can easily be detected by their broader leaves and deeper bulbs. The
mortality rate amongst the bulbils is high.

TABLE 2. DENSITY OF ALLIUM PARADOXUM

Mature
Site Plants / sq.ft plants / sq.ft
Bartonbury 1 104-2 + 4-2 8-3
Bartonbury 2 96:2 + 4-2 6°5
Road-verge 1 89-3 + 3-4 4-5
Road-verge 2 120-0 + 4-7 9-0

SEASONAL DEVELOPMENT

Mature plants of A. paradoxumhavesmall spherical bulbs about 1-2 cmin diameter
lying at soil depths of up to 13 cm. There is a long dormant period between
June and October when hardly any trace of the populations can be found above
ground. However, bulbils are easily located in the surface litter. Germination
of the fleshly bulbs usually begins in October with the production of fine adventi-
tious roots. A blunt shoot emerges and on dissection is seen to consist of an
outer modified leaf with a small bud in its axil, a large leaf with another axillary

GLOUCESTERSHIRE POPULATIONS OF ALLIUM PARADOXUM 381

bud and a triquetrous stem with the inflorescence enclosed in a spathe. The
modified leaf appears above ground and then the large linear leaf elongates.
During early January the buds are suitable for the study of PMC meiosis.
During February inflorescence buds are easily found and dissected out and both
bulbils and flowers can be found within the spathe. In March the leaves are
longer, the apex unrolls to reveal a linear blade and the closed inflorescences
are just beginning to appear on the strong triquetrous scapes.

During late March and April the inflorescences are fully emerged, the pedicels
and bulbils clustered at the base being subtended by the spathe.

The form of the inflorescence in the Bartonbury populations is very irregular.
It usually consists of a spathe subtending several bulbils and a single white
flower (occasionally two) on a slender pedicel. Variations recorded consist of
the following:

a. Bulbils only.

b. Pedicel bearing bulbils and no flowers.

c. Pedicel bearing bulbils with another pedicel extending from these bearing a

flower.

d. Wing-like expansions of the pedicels.

The many-flowered umbel as described by Vvedenskii (1935) has not been
found in the Bartonbury populations.

Various Diptera have been observed on the flowers; seed-setting is however
poor. The aerial parts gradually wither and the bulbils are scattered over the
adjacent area; the plants eventually die back completely during late May and
June.

The small plants developed from recently dispersed bulbils are different from
those already described. They have very small bulbs that lie on or just beneath
the soil surface and occasionally produce a short scape with a few bulbils. They
occur in very high densities, have a high mortality rate and rarely flower.

When established plants begin to die back in the autumn the following devel-
opment occurs. The base of the linear leaf swells and its axillary bud develops
to perpetuate growth and produce the bud containing the aerial parts of the
following year. The axillary bud of the modified leaf dies or develops into a
small offset on the outside of the swollen linear leaf-base.

PLANT CHARACTERS

The other characters studied were those of flower-structure, leaf-length and
width, stem-height, bulbil and seed production in established plants (Tables
3 and 5) and cytology.

The normal structure of the hypogynous flower is a perianth of six white
segments, six stamens adnate to these, and a trilobed ovary with trilobed
stigma. The following variations are common:

a. Deficiencies of one unit in all parts giving P2+2, A2+2, G(2) with a

bilobed stigma.

b. Deficiency in perianth and androecium, with stigmas and style imperfect

or absent.

c. Proliferation to give additional stamens and ovary.

d. Perianth segments with one or two of the anthers adnate to the apical

margin.

e. Bulbils within the flowers.

382 D. M. BARLING

The abnormalities of individual parts were scored and the percentage of
abnormal flowers calculated; the results are presented in Table 3.

It can be seen that abnormalities occur in all the floral parts and in 72% of
the flowering plants examined. A non-random selection of 10 two-flowered
plants was scored for inflorescence and floral variation (Table 4).

TABLE 3. PERCENTAGE ABNORMALITY OF INDIVIDUAL
FLOWER PARTS

Flower Part Number abnormal ’% abnormal
Gynoecium (whole) 119 47-60
Stigmas 97 38-76
Style 2 0-81
Carpels 45 17-89
Androecium 146 58-64
Perianth 129 51°82

Total normal plants 70
Total abnormal plants 180 72:00

Total plants sampled 250

TABLE 4. THE INFLORESCENCE CHARACTERS OF TWO-FLOWERED

PLANTS
Plant no.
Character 1 D 3 4 5 6 7 8 9 10

No. of pedicels 5 Z 2 2 3 2 2 D Z »
No. of bulbil

groups 4 2 2 2 3 2 y 2 Z 2

No. of bulbils/

group $4342 10:20 9252) 89 OF73352) 99 CUTTY EZ Soe earl
2:5

No. of perianth

parts 6:5. 6:5. 6:4. 6:33.. 6:5 6:4. 6:5. 6:45556:Geemore
No. of carpels 3:2 3:72 3:2 3:2 3:2 3:2 3:2) See omore
No. of styles 1:1 1S 1:1 1:1 et 1:1 1:1 gil Pei Let
No. of stigmas 3:2... 3:2 9° Be2 Sth 322 She 32" 8 ee Ome
No: of stamens’ 6:4 6:5 6:2 633" 6:5" (6:4 6:5 76:4 3G -OmmmoeZ

Leaf-length was determined by measuring the distance from the point of
unrolling of the leaf to the apex and this gave a mean length of 31-92 + 0-44 cm.
Leaf-width measurement is greatest near the middle and gave a mean of 1:53 +
0-09 cm.

Stem-length was taken as that length between the emergence of the scape
from the leaf to the point of origin of the spathe. This gave a mean of 21:8 +
0-41 cm.

As no seed was recorded in the sample plants an extensive search was made
for seed-producing plants as it had been observed that ovules were developing
in most ovaries. Seed has been found on five plants though only one seed in
each case. The seed has a black, rough testa. Offsets have been recorded and

GLOUCESTERSHIRE POPULATIONS OF ALLIUM PARADOXUM 383

TABLE 5. LEAF-LENGTH AND WIDTH, STEM-LENGTH AND
REPRODUCTIVE CAPACITY

Character Sample size Mean
Leaf-length 200 30:25 + 0:60 cm
Leaf-width 2000 1-55 + 0:09 cm
Stem-length 100 21-80 + 0-41 cm
Bulbils / plant 7 6:03 + 0:18
Flowers / plant Wa, 1-00
Seeds / plant 200 0°0

although no extensive data are available it appears that their contribution to the
reproductive capacity is small. The effective reproduction of A. paradoxum is
therefore by bulbils of which, at Bartonbury, the annual production is in the
region of 47:8 bulbils per sq ft.

The chromosome number of the plants was studied in pollen-mother-cells, in
pollen-mitosis and in root-tips, all prepared by the Feulgen technique. The
chromosome number is 2n = 16. At meiosis there was abundant evidence for a
chromosome inversion as bridges were recorded in the first and second divisions.
Fragments were recorded and these were easily seen in pollen grains as well.
Data and photographs have been exhibited (Barling 1958).

DISCUSSION

The observations at Bartonbury and adjacent areas show that the species is well
established and that bulbils are the main means of reproduction. This has
resulted in high plant densities and plants of varying size, only some of which
flower. It is likely that the spring-growth reduces shade and aerial competition
from its associates in both woodland and road-side swards and contributes to
successful colonisation.

The umbel of numerous flowers described by Vvedenskii (1935) has not been
found and the poor flowering recorded could be due to bulbil competition
during plant development. Gustafsson (1946) considered that in European
material the bulbils become more vigorous as flowers are reduced. Bulbil
competition probably influences the high percentage of abnormal flowers as
the variation found between flowers of the same plant indicates that it is not
due to hereditary differences.

Gustafsson (1946, 1947a b) listed three types of embryo-sac degeneration
associated with seed sterility in Allium and stated that A. paradoxum forms
slightly developed embryos that never mature. However a small number of
fully developed seeds have been found at Bartonbury. The chromosome inversion
found will have some effect on fertility but is unlikely to be the main cause of
poor seed production. Levan (1937) found that seeds were set if bulbils were
removed from bulbiferous Allium but no artificial removal has been done in
the present study. Gustafsson (1947b p. 296) stated that seed sterility may
arise automatically, not owing to any special lethal factor in the gametes or
zygote but to bulbil competition for nutrients, and this probably occurs in the
Bartonbury populations.

The general importance of bulbils in the Gloucestershire populations con-
trasts with Vvedenskii’s (1935) description “‘often with bulbils’” which suggests

384 D. M. BARLING

that they are usually absent from Russian plants. It is possible that the dominant
role of vegetative reproduction and the poor flower production and sexual
reproduction in Gloucestershire are responses to changing ecological conditions
as the species migrated westward, though more detailed data from its whole
range would be needed to clarify this point.

REFERENCES

BARLING, D. M. (1958). Reproduction and cytology of Allium paradoxum. B.S.B.1. Exhibition
Meeting 1957. Proc. bot. Soc. Br. Isl., 3: 85.

GUSTAFSSON, A. (1946). Apomixis in higher plants I. The mechanism of apomixis. Lunds Univ.
Arsskr., 42 (3): 1-66.

GUSTAFSSON, A. (1947a). Apomixis in higher plants IJ. The causal aspects of apomixis. Lunds
Univ. Arsskr., 43 (2): 71-178.

GUSTAFSSON, A. (1947b). Apomixis in higher plants III. Biotype and species formation. Lunds
Univ. Arsskr., 44 (2): 183-370.

LEVAN, A. (1937). Cytological studies in the A/lium paniculatum group. Hereditas, 23: 317-370.

RIDDELSDELL, H. J., HEDLEY, G. W. & PRicE, W. R. (1948). Flora of Gloucestershire. Cheltenham.

VVEDENSKH, A. I. (1935). Allium, in KoMARov, V. L. ed., Flora U.R.S.S., vol. 4, pp. 254-255.
Leningrad.

Watsonia, 8, 385-389 (1971). 385

Observations on Phyteuma tenerum R. Schulz
in England

M. KOVANDA

Czechoslovak Academy of Sciences, Botanical Institute, Pruhonice

ABSTRACT

The taxonomic relationships of English populations of Phyteuma tenerum are reviewed in
terms of morphological, cytological, ecological and phytogeographical criteria. Morphological
differences are shown to be slight but associated with distinct ecological requirements and dis-
tribution. Chromosome counts indicate that n = 11 in plants from two English localities. Some
preliminary comments are made on the structure of the P. orbiculare complex as a whole.

INTRODUCTION

In 1904, the genus Phyteuma L. was monographed by Schulz who proposed the
name P. tenerum to cover a major segment of the Linnaean species P. orbiculare,
defined by the following characters: stem densely leafy; basal leaves with
prominent lateral veins; involucral bracts triangular-lanceolate, acute, much
shorter than the inflorescence; ovary mostly 2-locular; widely distributed in
western Europe. The new species was subdivided into two geographical sub-
species: subsp. ibericum R. Schulz (having basal leaves elliptic, crenate, obtuse;
cauline leaves + subbracteiform; stigmas sometimes 3; occurring in N.E.
Spain), and subsp. anglicum R. Schulz (with basal leaves lanceolate or ovate-
lanceolate, serrate, acute; cauline leaves decreasing in size but distinctly foliar;
stigmas mostly 2; occupying the rest of the range of the species). Unlike some
other species described by Schulz in his monograph, which have been strongly
criticized and dropped into synonymy (see Kunz 1940), P. tenerum has always
been recognized as a distinct taxonomic entity. The present study was prompted
by a need for a more critical understanding of that species and by an interest in
the P. orbiculare complex as a whole. It is also hoped that the insight gained
may ultimately contribute to a better taxonomic arrangement of the entire
genus. The evidence reported here was obtained through study of herbarium
material from the major British herbaria and during field observations con-
ducted in several localities in the south of England in the late summer of 1969;
field studies have also been carried out on central European populations of
P. orbiculare. Chromosome counts were made on cultivated plants of known
wild origin.

MORPHOLOGY

The English populations of P. tenerum appear fairly homogeneous and uniform
in morphology. Nonetheless, there is a certain amount of variation in some
characters, even those believed to be important from the taxonomic viewpoint.
These characters will be briefly discussed below.

(1) The shape of the basal leaves has been widely used in the taxonomic
subdivision of the genus, and Schulz (1904) made it one of the main criteria

386 M. KOVANDA

in delimiting both species and infraspecific taxa. It is indeed characteristic of
most species, but the differences are not readily apparent unless leaves of equal
maturity are compared. This is technically most difficult and, with herbarium
specimens, virtually impossible since the oldest leaves, which are most important
for comparison, are seldom present at flowering time when the plants are usually
collected and dried. The shape of the leaves gradually changes as the plant
develops, and the later leaves bear little or no resemblance to the earlier ones,
those present during and after flowering being the least characteristic. Super-
ficial observation may be quite misleading. In many local populations of
P. orbiculare the shape of the radical leaves is of paramount importance as a
taxonomic guide, and P. tenerum is likely to follow this pattern of differentiation.
Unfortunately, Schulz does not appear to have been aware of this particular
seasonal variation, and the characteristics given by him in keys and descriptions
should be re-examined carefully.

(2) The prominence of the nerves of the basal leaves is difficult to assess and
hardly relevant taxonomically. It may depend on the condition of the plant,
time of collection, the way the plants have been dried and so forth. No specific
differences could be found in the available material.

(3) The leafiness of the stem varies greatly in P. tenerum, as it does in P.
orbiculare. English plants often have a densely leafy stem, as required by
Schulz’s diagnosis, but individuals with but four leaves do occur. Also, the
cauline leaves are generally shorter than is usual in P. orbiculare. A statistical
study might perhaps reveal a significant difference, but for diagnostic purposes
these characters are almost useless.

(4) The shape of the involucral bracts proved to be a truly diagnostic character
in both P. orbiculare and P. tenerum. In the first they are conspicuous, often
foliar, ovate to lanceolate, always distinctly acuminate (sometime even mucron-
ate), and generally equalling or exceeding the flowers. The bracts of P. tenerum
differ in being tiny, triangular to narrowly lanceolate, acute (not acuminate),
and invariably shorter than mature flowers. In both species the bracts vary
from entire to repand, and in P. orbiculare they are sometimes finely serrate.

(5) The number of carpels is fairly constant in most Phyteuma species. It is
either two (e.g. P. nigrum F. W. Schmidt, P. michelii All. and P. charmeiii Vill.)
or three (P. hemisphaericum L., P. globulariifolium Sternb. & Hoppe, P.
scheuchzeri All.), and deviations, if any, are rare. In P. orbiculare this variation
is considerable, and the ratio of 2- and 3-locular ovaries seems to vary from
one locality to another. In the English P. tenerum the former clearly predominate
but 3-locular ovaries are far from rare.

CHROMOSOME NUMBER

In recent years, chromosome numbers have been reported for many Phyteuma
species (e.g. Favarger 1953, Contandriopoulos 1962, Ochlewska 1965, Gadella
1966, Polatschek 1966). These studies have clearly demonstrated the existence
of a dysploid series within the genus Phyteuma. Contrary to the situation in
other genera of Campanulaceae, polyploidy does not seem to be involved,
except perhaps occasionally in P. spicatum L. (Ochlewska 1965). In P. orbiculare
alone, the following numbers have been found: 2n = 22 (Ochlewska in Skalinska
et alia 1964, Ochlewska 1965), 2n = 22 + O0-2B (Polatschek 1966), 2n=24
(Mattick in Tischler 1950, Baksay 1956, Contandriopoulos 1962, Gadella 1966),

PHYTEUMA TENERUM IN ENGLAND 387

2n = 26 (Sugiura 1942, Baksay 1956). As far as P. tenerum is concerned, the
extensive chromosome study by Mlle Contandriopoulos has included one
count made on a collection of this species from the vicinity of Mainz (““Mayence’’)
in West Germany. The author reports n = 12 for that sample and, having found
the same number in P. orbiculare, she concludes that the series Orbiculata as a
whole is characterized by n = 12. No cytological work seems to have been done
on English material.

Considering the striking karyological diversity encountered throughout the
genus, it seemed desirable to examine material of P. tenerum from another part
of the distribution area. The counts reported here were obtained during studies
of meiosis in pollen-mother-cells, flower buds being fixed in 1:3 acetic alcohol
for 24 hours and stored in 70% alcohol until examined three months later.
Squashes were made in lacto-propionic orceine. For convenience, fixations
were made from plants grown from seed in the University Botanic Garden at
Cambridge. The localities of the collections were as follows:

Easton Hill, Bishop’s Cannings, North Wiltshire (v.c. 7), leg. D. E. Coombe.
By chalk pit, Erringham Downs, north of Shoreham-on-Sea, West Sussex
(v.c. 13), GR 51/212.102, leg. A. Moppett, 9/1965.

In both samples a number of meiotic divisions have been studied, and in all
of them invariably 11 bivalents were found. In addition, in the Easton Hill
plant 27 = 22 could be counted in somatic mitoses in the young corolla tissue.

Thus the number ascertained for English material differs from that reported
by Contandriopoulos (1962). It may well be that P. tenerum is represented by
more than one cytotype.

It should however be pointed out that, in 6 of 13 Phyteuma species discussed
by Mlle Contandriopoulos, the chromosome numbers given in the table (p.
265) and in the legend differ from those seen in the figures, and one cannot,
therefore, be quite certain whether or not the number n = 12, reported for
P. tenerum, actually belongs to this species.

GEOGRAPHICAL DISTRIBUTION AND ECOLOGY

P. tenerum is an Atlantic species with a well-defined area embracing, with many
gaps, most of France (except for the Mediterranean region and, oddly enough,
much of the Massif Central) and extending to north-eastern Spain, south-western
Germany and southern England (for distribution maps see Schulz 1904 and
Contandriopoulos 1962). The distribution in England has recently been mapped
by Perring & Walters (1962). Its northern limit is shown as northern Wiltshire
but in the Kew Herbarium there are two interesting sheets of P. tenerum
collected by anonymous persons in 1792 and 1833, from ‘Manchester’ and
‘Chalky banks, Broughton, near Manchester’ respectively, which suggest that it
might occur farther to the north. Unfortunately, there is no additional evidence
to support these records; I have failed also to find a confirmation of them in
any Flora, either local or general.

Throughout its range, P. tenerum clearly prefers calcium-rich soils. It is not
surprising, therefore, to discover that in England this species is closely associated
with chalk vegetation. The species is especially characteristic of the Downs,
where it is locally abundant on south-facing slopes, avoiding, however, sites

388 M. KOVANDA

dominated by tall grasses. Unlike continental populations, which are commonly
found on rocks (e.g. Liger 1952), the English P. tenerum seems to be restricted to
grassland communities. Tansley (1949) lists it (as P. orbiculare) among so-called
exclusive species, i.e. those either confined, or nearly confined, to chalk grass-
lands in England. This strong ecological specialization of P. tenerum is very
noticeable inasmuch as P. orbiculare, its closest relative and immediate pro-
genitor, is largely indifferent to soil and occurs over a wide range of habitats,
from lowland peat bogs to alpine rocks.

Data concerning flowering periods indicate an interesting difference between
the species. Flowering begins in lowland populations of P. orbiculare in late
May or early June and seldom exceeds two weeks but starts a month later and
continues until October in populations of P. tenerum.

TAXONOMY

It is obvious that the evidence presented here challenges, more than it supports,
the specific status of P. tenerum. There is a degree of morphological differentia-
tion combined with a fairly clear geographical separation and differences in
ecological preferences and flowering time, but most of the morphological
characters tend to intergrade, often making a clear-cut separation quite difficult.
Forms intermediate between P. orbiculare and P. tenerum have been reported
from the Vosges and adjoining parts of Germany (Schulz 1904), and Kunz
(1940), in a somewhat derogatory manner, describes difficulties in identifying
material from an area where the ranges of both taxa overlap.

The occurrence of 11 bivalents at meiosis casts little light on the evolutionary
status and taxonomic rank of P. tenerum. The same chromosome number
occurs also in such widely different species as P. nigrum (Gadella 1966, Kovanda
in Léve 1970) and P. spicatum (Ochlewska 1965, Gadella 1966, Polatschek 1966).
The earlier count, m = 12, has been reported for, in addition to P. orbiculare,
P. betonicifolium Vill. (Favarger 1953, Contandriopoulos 1962, Gadella 1966,
Polatschek 1966), P. spicatum (Contandriopoulos 1962), P. zahlbruckneri Vest
(Gadella 1966, Polatschek 1966) and other species.

It has long been known that variation within the P. orbiculare complex is
enormous, yet how much is individual and how much is of taxonomic impor-
tance remains a vexing problem. Schulz (1904) tried to cope with this variation
by defining, rather vaguely, six subspecies, which, however, fail to give a true
picture of the diversity encountered in natural populations. Possibly P. orbiculare
sensu lato represents a single ecospecies with more or less well-defined geo-
graphical ecotypes ranging from Spain and southern England to the Ukraine,
but it seems premature to attempt any major taxonomic realignments within
that complex. It is suggested, therefore, that P. tenerum continues to be recog-
nized as a distinct species until comparable data are available for a representative
series of races of P. orbiculare.

ACKNOWLEDGMENTS

This account is based on work carried out during the tenure of a Flora Europaea
bursarship spent mainly at Kew in 1969-1970. I wish to express my gratitude

PHYTEUMA TENERUM IN ENGLAND 389

to Dr S. M. Walters, Botany School, Cambridge, for encouragement and
placing living material at my disposal. To Mr S. G. Shetler, Department of
Botany, Smithsonian Institution, Washington D.C., I am indebted for extending
to me the facilities to make the cytological study.

REFERENCES

Baksay, L. (1956). Cytotaxonomical studies on the flora of Hungary. Annis hist.-nat. Mus.
natn. hung., ser. n., 7: 321-334.

CONTANDRIOPOULOS, J. (1962). Recherches sur la flore endémique de la Corse et sur ses
origines. Annls Fac. Sci. Marseille, 32: 1-354.

FAVARGER, C. (1953). Notes de caryologie alpine, 2. Bull. Soc. neuchdtel. Sci. Nat., 76: 133-169.

GADELLA, T. W. J. (1966). Some notes on the delimitation of genera in the Campanulaceae,
1 & 2. Proc. ned. Akad. Wet., 69: 502-508.

Kunz, H. (1940). Bemerkungen zu einigen schweizerischen Phyteuma-Sippen. Ber. schweiz.
bot. Ges., 50: 363-378.

Licer, J. (1952). Etudes sur la végétation des falaises calcaires de la Basse-Seine. Bull. Soc.
Amis Sci. nat. Mus. Rouen, 88: 17-54.

Love, A., ed. (1970). IOPB Chromosome number reports, 25. Taxon, 19: 102-113.

OCHLEWSKA, M. (1965). Cytological studies in the genus Phyteuma L. Acta biol. cracov., sér.
bot., 8: 135-145.

PERRING, F. H. & Walters, S. M., ed. (1962). Atlas of the British Flora. London. ”

POLATSCHEK, A. (1965). Cytotaxonomische Beitrage zur Flora der Ostalpenlander, 2. Ost.
bot. Z., 113: 101-147.

SCHULZ, R. (1904). Monographische Bearbeitung der Gattung Phyteuma. Geisenheim a. Rhein.

SKALINSKA, M. et alia (1964). Additions to chromosome numbers of Polish Angiosperms.
Acta Soc. Bot. Pol., 33: 45-76.

SuGiurRA, T. (1942). A list of chromosome numbers in angiospermous plants, 8. Proc. imp.
acad. Japan, 18: 30.

TANSLEY, A. G. (1949). The British Islands and their Vegetation, 2nd ed., vol. 2, p. 539.
Cambridge.

TISCHLER, G. (1950). Die Chromosomenzahlen der Gefasspflanzen Mitteleuropas.’s Gravenhage.

Watsonia, 8, 391-393 (1971). 391

The Manchester Poplar
C. A. STACE

Botany Department, University of Manchester

It is well known that the great majority of ‘Black Poplars’ in this country are
not referable to Populus nigra L., a native of central and southern Europe and
probably of parts of England, but to horticultural hybrids between this species
and P. deltoides Marsh. from eastern North America (= P. x canadensis
Moench). This hybrid exists as at least a dozen variants, resulting from crosses
between different strains or races of the two species and from extensive back-
crossing. A good account of these trees is given by Peace (1952). Apparently
by far the most common plant in Britain is var. serotina (Hartig) Rehd. (often
called P. x serotina Hartig), which exists only in the male state. Gilbert-Carter
(1936) gives a good impression of the differences between this taxon and P. nigra.

There is, in horticultural circles, a tree known as ‘Manchester Poplar’, which
according to most authorities is P. nigra var. betulifolia Pursh. Bugula (1967)
concluded that this variety is the only one native in England and western France.
He recognised it as P. nigra subsp. betulifolia (Pursh) W. Wettst.; it differs from
the more south-eastern subsp. nigra in having pubescent young twigs, petioles
and inflorescences. Henry (1914) came to a similar conclusion to Bugula, and
gave very useful notes on the identification of planted Black Poplars.

Because of the identity of the English native Black Poplar and the horti-
cultural Manchester Poplar, the natural distribution of the former has probably
become clouded by planting. The native area of P. nigra in Britain is at any rate
uncertain. Warburg (1952) suggests it is from Essex and Lincoln to Gloucester
and Shropshire, and Edlin (1956) says ‘mainly in eastern England, though it
is also common on the Cheshire Plain.’ Edlin provides a coloured plate of a
good Cheshire specimen. Mr A. Newton, who has just completed a new Flora
of Cheshire (1971), informs me, on the other hand, that he considers P. nigra
to be decidedly scarce in the county, and nowhere undoubtedly native.

Hadfield (1957), writing about P. nigra, comments that it ‘stands smoke well
and was popular in the industrial north—hence its name of Manchester Poplar’.
McClintock (1966), discussing a rather different topic, states that “Manchester
Poplar is a name for the true Black Poplar P. nigra, which is said to withstand
the soot and filth of Manchester’.

For the past few years I have attempted to ascertain the correctness or other-
wise of some of the above statements. The commonest roadside trees in Man-
chester are ash, elm, lime, sycamore and plane. Poplars are commoner in parks
and similar places, and in a great many cases these are P. x canadensis. It was
soon found, however, that the Manchester Poplar does still exist in Manchester,
and that in some areas of the conurbation it is by far the commonest poplar.
It is mainly found in the older (i.e. mostly inner) suburbs, being scarcer in the
city centre and in the postwar suburbs. The examples I have observed closest to
the centre of Manchester are in Great Ancoats and Adair Streets (Manchester
postal districts 4 and 1).

392 Cc. A. STACE

Large numbers are to be found to the south of the city centre (districts 13,
14, 19 and 20, and in various parts of Stockport); and again to the north towards
Bolton, Bury and Rochdale (districts 8, 9 and 10, and in parts of Oldham,
Middleton, Prestwich, Pendlebury and Kearsley). Particularly good stands
have been observed in Cringle Fields Park (district 19), in Hollywood Park
(Stockport), in Queens Park and the east side of Heaton Park (district 9), and
especially in Broadhurst Park and St Joseph’s Cemetery (districts 9 and 10).
In districts 9 and 10 (notably Lightbourne Road and Victoria Avenue East)
many roadside trees are Manchester Poplars. Detailed searches to the east and
west have not been made, but specimens have been seen in Ashton-under-Lyne.

Even in areas where these trees are relatively plentiful they are, however,
rather locally distributed. Thus in Cringle Fields Park most of the trees are
Manchester Poplars, but in Ladybarn Park, about 1 km west, they are scarce,
although there are in the latter place plenty of examples of P. x canadensis
and P. nigra var. italica Duroi (Lombardy Poplar). There is, however, in the
British Museum (BM) a specimen of P. nigra var. betulifolia collected by R. S.
Adamson in 1915 from Mauldeth Road Station (which adjoins Ladybarn Park).

Most of the Manchester Poplars appear to me to be between 30 and 100 years
old, though it is difficult to make an accurate assessment and the City Parks
Department has no records which would help. Several of the parts of Man-
chester in which the trees occur are slum-clearance zones (e.g. in district 13
near the University), and it is pleasing to see, in recently cleared areas, specimen
trees of Manchester Poplars (amongst other species) preserved for incorporation
in the future development of the site. Preservation of existing trees seems almost
their only hope of salvation in Manchester, for I am told by the Parks Depart-
ment that they are extremely rarely planted nowadays, being largely replaced
by other variants of the complex, notably “P. eugenei Simon-Louis’, which can
easily withstand the now much cleaner atmosphere. Manchester Poplars are,
however, recommended in the C.P.R.E.’s 1970 Cheshire Tree Campaign
booklet for planting ‘in difficult areas, e.g. wet, badly drained, overgrown or
generally neglected’.

Whether the newer hybrids are more or less attractive than P. nigra is, of
course, a matter of personal taste. Gilbert-Carter found the heavily bossed
trunks of the latter ‘grotesque’, but I much prefer their arching branches and
rounded overall outlines (rather similar to an ash) to the exotic shapes of the
various cultivars and hybrids. It might also be argued that there are too few
native English trees which are widely used for town planting. One advantage
of using certain variants of P. x canadensis (e.g. var. serotina and P. eugenei)
for horticultural purposes is that they exist only as males, and thus councils
are spared the expense of clearing up the often extremely copious deposits of
fruiting catkins in early summer. Hayfever is perhaps a less obvious consequence
of this policy. One might presume that P. nigra, being a natural taxon, exists
in roughly equal numbers of each sex. Of about one hundred planted trees
examined in April 1970, all were, however, males. It seems clear, therefore, that
horticultural material of Manchester Poplars is propagated solely or largely by
vegetative means.

REFERENCES

BUGULA, W. (1967). Systematyska eurozjatyckich topoli z grupy Populus nigra. Arb. Kornick.
Roczn., 12: 45-219 (Abstracted in Proc. bot. Soc. Br. Isles, 7: 429 (1968)).

THE MANCHESTER POPLAR 393

Eputn, H. L. (1956). Trees, Woods and Man, p. 206. London.

GILBERT-CARTER, H. (1936). British Trees and Shrubs, pp. 49-50. Oxford.

HADFIELD, M. (1957). British Trees, p. 150. London.

Henry, A. (1914). The Black Poplars. Gdnrs’ Chron., 56(1): 1-2; 56(3): 46-47; 56(4) 66-68.

McCunTock, D. (1966). Companion to Flowers, p. 193. London.

NewrTon, A. (1971). Flora of Cheshire. Chester.

Peace, T. R. (1952). Poplars. Forestry Commission Bulletin 19.

WARBURG, E. F. (1962). Populus, in CLAPHAM, A. R., TUTIN, T. G. & WARBURG, E. F. Flora
of the British Isles, 2nd ed., pp. 578-581. Cambridge.

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Watsonia, 8, 395-402 (1971). 395

Short Notes

88/4. COCHLEARIA SCOTICA DrRuUCE—Does it exist in northern Scotland?

The British endemic, Cochlearia scotica, is stated to have a chromosome number of
2n = 14 (Gairdner 1939). As such it resembles the arctic species of the genus: C.
arctica Schlecht., C. groenlandica L. and C. oblongifolia DC. C. scotica, although
consistently recognised by recent workers (Clapham 1952, 1962; Chater & Heywood
1964), is stated to be easily and frequently confused with dwarf specimens of C.
officinalis L. This latter species is in all characters a much larger plant than C. scotica,
but in exposed conditions may become so dwarfed as to resemble C. scotica. The
following are said to be the most consistent points of difference between the two
species:

C. scotica C. officinalis
Basal leaves usually truncate at base Basal leaves usually cordate at base
Lowest stem-leaves sometimes stalked Lowest stem-leaves not stalked
Stem-leaves clasping the stem Stem-leaves not clasping the stem

The author has sampled a large number of populations from northern Scotland
and the Scottish Isles which were phenotypically identifiable as C. scotica. Most proved
to have a chromosome number of 2n = 24 with many of the populations containing
B chromosomes (Table I). Unfortunately, it was not possible to ascertain the origin
of Gairdner’s material, so no direct check on its validity was possible. This does not,
of course, deny the existence of C. scotica as a rare diploid entity in northern Britain,
but must cast doubts on its supposed distribution.

TABLE 1. CHROMOSOME COUNTS OF SPECIMENS REFERABLE TO C. SCOTICA
(all counts based on at least five specimens, except where stated)

Locality Chromosome Number
East Voe of Skellister, Nesting, Shetland 2n = 26, 28 (2 plants)
Scalloway, Shetland 2n = 24 (+2B)
Little Green Holm, Orkney 2n = 24 (+3B)
Muckle Green Holm, Orkney 2n = 24

Brock of Borwick, Orkney 2n = 24 (+2B)
Ramnageo, Orkney 2n = 24 (+1B)
Wifes Geo, Orkney 2n = 24

S. E. Swona, Orkney 2n = 24 (+2B)
Loch Sligachan, Skye 2n = 24

Loch Ainort, Skye 2n = 24(+2B)
Broadford Bay, Skye 2n = 24

Loch na Dal, Skye 2n = 24 (+2B)
Armadale Bay, Skye 2n = 24(+1B)
Barvas, Lewis 2n = 24 (+2B)
Garynahine, Lewis 2n = 24

Butt of Lewis, Lewis 2ii— 4
Balallan, Lewis 2n = 24 (+3B)
Husinish, Harris 2n = 24 (+2B)
Rudha Reamhur, Harris 2n = 24

Loch Scresort, Rhum 2n = 25 (1 plant)
Melvich Bay, Sutherland 2n = 24 (+ 1B)
Helmsdale, Sutherland 2n = 24
Plockton, Ross & Cromarty 2n = 24 (+3B)

Morar, Inverness 2n = 27 (1 plant)

396 SHORT NOTES

REFERENCES

CiapHaM, A. R. (1952), in CLAPHAM, A. R., TuTIN, T. G. & WARBURG, E. F. Flora of the
British Isles, p. 191. Cambridge. é

CLapPHaM, A. R. (1962), in CLAPHAM, A. R., TuTIn, T. G. & WARBURG, E. F. Flora of the
British Isles, 2nd ed., p. 151-152. Cambridge.

Cuater, A. O. & HEywoop, V. H. (1964), in TuTin, T. G., et alia, ed. Flora Europaea, vol. 1,
p. 314. Cambridge.

GaIRDNER, A. E. (1939), in MaupDeE, P. F. The Merton Catalogue. A list of the chromosome
numbers of species of British flowering plants. New Phytol., 38: 1-31.

I-53. BoGine

252/1. HiPPOPHAE RHAMNOIDES L.—On the Isle of Mull, Argyllshire.

Whilst on Mull with the botanical survey party from the British Museum (Natural
History) during September 1968, I had an opportunity to investigate carefully a
locality for Hippophaé rhamnoides, sea-buckthorn, discovered the previous year by
Mr Peter James (in Cannon & Bangerter 1968). Hippophaé is generally not considered
to be native on the west coast of Britain. The site lay along a small headland on the
north side of Camas Mor, near the grounds of Torosay Castle (GR 17/733.358). Over
a distance of 30 yards along the shore of this headland grew some twenty old, gnarled
and contorted trees of Hippophaé and at an estimate I would consider them to be
between 30 and 40 years old. Several had trunks at least six inches in diameter at
the base though all were much divided and branched a short way up from the ground.
Some of the largest branches (most usually the lowest) were dead or dying. Both male
and female trees were present, the females being in full fruit when I saw them on
17th September.

In spite of a thorough search only four seedling trees were found. These were growing
close to the outer trees of the colony on the south-west side, and were all between
12 and 18 inches high and probably no more than 3-5 years old.

No evidence of spread by vegetative means was observed amongst the rocks and
pebbles surrounding the site (there being no patches of sand present), nor in the
adjacent area of finer material, consisting of gravel and mud, nearer the shore. The
gravel and mud at this part of the shoreline was closely compacted and waterlogged
at every incoming tide. These factors in all probability precluded a successful growth
of a horizontal rooting system spreading outwards from the parent bushes. Nor was
there any sign of perennation on the landward side of the colony, the shade formed
by the woodland trees growing immediately behind being particularly dense.

The hooded crow (Corvus cornix) has, on occasions, been reported to feed on sea-
buckthorn fruits at localities elsewhere in Britain (Goddard 1949, Groves 1958,
Pearson & Rogers 1962). Although this crow was seen feeding elsewhere along the
coast of Mull, no evidence of droppings of the bird, or indeed of any other species,
that would indicate feeding on the sea-buckthorn fruits, was seen on the rocks beneath
the Hippophaé at this Torosay Castle site.

Sea-buckthorn at this locality is therefore almost certainly originally planted as is
the single tree of Rhamnus cathartica noticed growing nearby over a derelict stone
summer-house.

REFERENCES

CANNON, J. F. M. & BANGERTER, E. B. (1968). Plant records from Mull and the adjacent small
islands. Proc. bot. Soc. Br. Isl., 7: 365-372.

GODDARD, M. (1949). Shrubs of the sand hills. Countryside (New Ser.), 15: 77-79.

Groves, E. W. (1958). Hippophaé rhamnoides in the British Isles. Proc. bot. Soc. Br. Isl., 3:
1-21.

Pearson, M. C. & Rocers, J. A. (1962). Hippophaé rhamnoides L., in Biological Flora of the
British Isles. J. Ecol., 50: 501-513.

E. W. GROVES

SHORT NOTES 397

ECHINOPHORA SPINOSA L.—A member of our flora.

This Mediterranean Umbellifer was at one time included in British Floras even
though known to be extinct, but in recent Floras it has been excluded. Exclusion
of any species unsupported by valid evidence, especially with its specious appearance
of critical acumen is no less reprehensible than the indiscriminate inclusion of im-
probable records.

The justification for the omission of any mention of Echinophora spinosa was not
merely that it was extinct but it appeared not unreasonable to suppose that it was
here beyond its climatic range and, perhaps, a mere casual that neither persisted
nor reproduced. It therefore appeared desirable to ascertain by experiment the tolerance
of this species in our coastal climate. For this purpose ripe fruits were obtained from
the south of France and the seedlings derived from these were grown in a pocket of
sand and shingle in my garden on the coast at Felpham, Sussex. The plants were
quite unprotected and only two escaped depredation by birds. Echinophora spinosa
has now survived for five years and endured prolonged frost with winter temperatures
that have been below —9°C.

Its capacity for reproduction in our climatic conditions was apparently nil during
the first three years, although flowering occurred freely, since no seedlings were
produced. In the fifth season infructescences were developed. The produce of an entire
plant, all the fruits of which were dissected, was 114 mericarps that appeared to be
fertile. In a number of the fruits one of the pair of mericarps had obviously aborted
and in some the single fertile mericarp was exceptionally large.

The average weight, based on a sample of 96 mericarps, was 0-0148gm. This com-
pares favourably with the average weight for samples of mericarps harvested in the
south of France, which was 0:013gm. This lower average weight is perhaps correlated
with an appreciably larger output. The French mericarps showed 28-5°% germination,
beginning in February and ending in the middle of May.

One sample of 26 fruits from a plant in my garden was sown as soon as the in-
fructescences had withered in November. Germination began in April and continued
till the end of May. On the highly improbable assumption that all these fruits contained
two fertile mericarps the total seedlings would represent a germination of 32:5%.
Actually, in only a few instances did the seedlings appear in pairs so that the real
proportion might well have been about 60°%. These results warrant the belief that
reproduction on the south coast could have been adequate for the maintenance of
the species.

Of the records of Echinophora spinosa on our coasts, that from Lancashire we have
on the authority of Ray. Those from the Kentish localities we have on good authority.
The Dorset record has been called in question. Professor Good refers to the belief
that this was an error due to Crithmum maritimum being mistaken for Echinophora.
This appears highly improbable, quite apart from the fact that in genuine mistakes
the unfamiliar is normally assimilated to the familiar, not the reverse. No one who
knew Crithmum maritimum could mistake it for the other plant, since even in the
vegetative condition Echinophora has spiny, almost crisped leaves while those of
Crithmum have pointed tips with almost flat segments. (Figs. 1B and 1C). Furthermore
the Crithmum leaf emits the characteristic aromatic odour whilst that of Echinophora
has but a slight parsley-like scent. The two Dorset records would appear to be on the
authority of the Rev. A. Bloxham (Baxter, W. (1843) British Phaenogamous Botany,
vol. 6, p. 478). Bloxham was evidently a field botanist of competence to whom J. C.
Mansell-Pleydell, the author of the Flora of Dorsetshire (1874), pays tribute for
assistance with certain critical genera.

There are other southern species which are recognised members of the British flora
that have become extinct in a large proportion of their former stations. Otanthus
maritimus has been recorded from some twenty-three coastal areas from East Anglia
to the West of Ireland, but is now perhaps extinct from all but three of these (c 87%).

398 SHORT NOTES

According to the Atlas, Euphorbia peplis was formerly present in a similar number of
locations, though these were not quite so widespread, and it may now be extinct
from all but two (c 90%). Polygonum maritimum formerly occurred in some twelve
distinct areas though it is now probably present in not more than three (c 75°%).
These species have clearly all declined similarly, the prostrate and inconspicuous
annual spurge more severely than the two perennials. The extinction of Echinophora
spinosa is thus no evidence that it was in any different category to these other southern
types whilst its taller habit (up to half a metre) and stems that rather readily fracture
from the base, render it specially vulnerable to damage by pedestrians.

FicureE 1. A. Seedling of Echinophora spinosa. B. Lateral segment of adult leaf. C. Lateral
segment of Crithmum maritimum,. All material grown in Sussex.

SHORT NOTES 399

It is unlikely that any of these southern types were deliberately introduced since
they are not specially attractive nor accredited with medicinal or culinary merits,
although J. Hill (The Family Herbal (1722)) claims that the root of Echinophora is
comparable to the parsnip as a vegetable. It is not improbable that all these species
owed their presence on our coasts to accidental introduction by man, perhaps in
ballast from vessels returning relatively empty from the south of Europe for which,
in primitive times, soil from the foreshore would be the handiest source. This is mere
speculation, but the facts here recorded emphasise that there is no evidence to warrant
placing Echinophora in a separate category and any distinction between these southern
species is more likely to be one of degree rather than of kind.

E. J. SALISBURY

430/20. VERONICA HEDERIFOLIA L. sensu lato—In Wales.

The variable Veronica hederifolia has lately been divided into three species on cyto-
logical grounds by Fischer (1967); an English-language abstract appeared in Proc.
bot. Soc. Br. Isl., 7: 435 (1968). Two of these segregates occur in Wales—the pale-
lilac-flowered tetraploid (2n = 36) V. sublobata M. Fischer and the blue-flowered
hexaploid (2n = 54) V. hederifolia L. sensu stricto. The following are new descriptions
based on Welsh material observed in 1970.

V. sublobata M. Fischer. Plant less robust, greyish green with denser, shorter hairs.
Leaves with shallower more obtuse lobes. Stomata 26-35 wm, mean c 32 um. Pedicels
4—5 times the fruiting calyx. Calyx weakly accrescent. Corolla c 2:0 mm long, pale-lilac
to whitish. Anthers 0-6 mm, whitish, dehiscing before the flower opens. Pollen (fresh,
in water) (31-) c 33 (-35) pm.

The common segregate in Wales, occurring on rich humus in woods and on shady
roadsides and as a garden weed.

V. hederifolia L. sensu stricto. Plant more robust and fleshy, bright green with
sparser, longer hairs. Leaves with deeper more acute lobes. Stomata 35-48 »m, mean
c 42 wm. Pedicels 24-34 times the fruiting calyx. Calyx strongly accrescent. Corolla
2:5—3-0 mm long, blue. Anthers 1-0 mm, bright-blue, dehiscing after the flower opens.
Pollen (fresh, in water) (38-) c 40 (-42) um.

In Wales, more local and so far seen by the writer only as a garden weed.

Corolla size in both species becomes smaller towards the end of the flowering
period. Pollen from herbarium specimens, in water, gives measurements about 10%
less than fresh pollen in water.

An experiment carried out by the writer during 1970 supports the treatment of these
taxa as species. To try to induce a hybrid between them, 20 cross-pollinations were
made with V. sublobata as the intended Q parent and 18 in the opposite direction.
No capsules developed from these crossings although all the plants used fruited
normally from selfing. A hybrid might well be obtained if a large number of crossings
were made, but the negative result of this limited experiment seems proof of an effective
if not complete barrier against interbreeding between the two taxa; moreover a hybrid
would very probably be at least partially sterile.

Welsh material and records of the aggregate, particularly of V. hederifolia sensu
stricto, would be welcome for the compilation of distribution maps.

REFERENCE

FiscHER, M. (1967). Beitrage zur Cytotaxonomie der Veronica hederifolia-Gruppe (Scrophula-
riaceae). Ost. bot. Z., 114: 189-223.
P. M. BENOIT

457/1. PRUNELLA VULGARIS L.—Colour variants apparently introduced.

The distribution of the colour variants in this species has long gone largely unrecorded,
in the reasonable belief that, like so many trivial mutants, their incidence is purely

400 SHORT NOTES

random in character and unlikely to reveal any meaningful patterns. Recent fieldwork
in the Isle of Man suggests that this assumption is mistaken. Dr L. S. Garrad has
found that a variant with pink flowers characteristically occurs in leys, in which it has
been noted in several widely-separated parts of the island. In one such habitat, near
Castletown, a large population of the species is as much as about 80% pink-flowered.
Rather similarly, a variant with pale blue flowers occurs in plenty on garden lawns
in one village in the east of the island; it has not, however, been noticed anywhere
else, even singly. In both cases introduction with grass-seed seems to be indicated. Is
there, one wonders, a firm of seedsmen somewhere in the British Isles whose premises
are overrun with the products of one-time experimental work on the colour forms of
this species ?

A further point to which attention might usefully be drawn is that in some cases, at
least, abnormal corolla colour in this species seems to be associated with distinctiveness
in other characters. Var. pallida Gilmour, for instance, as originally described, has
besides pale bluish-mauve flowers, a smaller corolla and a low stature; it is also said
to be characteristic of dunes and dry grassland. N. Douglas Simpson (pers. comm.)
had in cultivation for many years a variant with pale pinkish-mauve flowers, viscous
foliage and yellow-green leaves. L. Cumming (Rep. botl Soc. Exch. Club Br. Isl., 6:
245 (1921)) likewise reported cultivating through seven generations a plant with
almost-white flowers which was distinctly more hoary-pubescent than the type.

D. E. ALLEN

582/1. ERIOCAULON AQUATICUM (Hill) Druce—On Ardnamurchan (Westerness, v.c. 97).

In 1967 Mr D. McClintock showed at the Annual Exhibition Meeting of the B.S.B.I.
a living specimen of Eriocaulon aquaticum (Hill) Druce (E. septangulare) found in the
inland hills of Ardnamurchan by Mr W. Dolling (Proc. bot. Soc. Br. Isl., 77:509 (1968)).
This is the first record of the plant on the Scottish mainland. Eriocaulon is recorded
in the Atlas from five 10 km grid-squares on the Scottish islands of Skye and Coll and
is frequent in much of western Ireland. The record was subsequently published as a
new vice-county record for Westerness, v.c. 97 (Proc. bot. Soc. Br. Isl., 7:566 (1969)),
but without locality.

In September 1970 the authors carried out a superficial survey of 12 scattered lochs
on Ardnamurchan. Eriocaulon was found to be very abundant and flowering freely in
Loch Caorach, GR 17/434.656, about 14 miles south-east of the Point of Ardnamurchan.
It appears to be absent from the neighbouring Loch Grigadale, Gr 17/431.669, and
the twin lochs at the head of Allt a’ Bhriaghlann, Gr 17/437.648, as well as the lochs
at GR 17/464.657 and 17/470.655 (Lochan na Crannaig) about three miles to the east.
Eriocaulon was found in a second locality at Lochan Dubh, Gr 17/490.706, about
one mile west of Faskadale on the north of the peninsula. Here it was locally abundant
in small patches growing in about 1 ft of water with Littorella. c

The loch at Ockle, Gr 17/558.705, and those in Glen Sordail, Lochan an Ime, GR
17/557.686, Lochan nan Dearcag and Lochan na Tuaidh, Gr 17/559.679, and unnamed
lochs at GR 17/556.680 and 17/560.689 were visited but Eriocaulon was not found.
Lochan na Creige Duibhe, Gr 17/640.667, south of Kentra Bay was found to now be
a reservoir with fluctuating water levels.

It would seem that Eriocaulon may be confined to the western third of the Ardna-
murchan peninsula but there are a large number of lochs on Ardnamurchan, which
to our knowledge remain to be investigated, both within the region where we have
found Eriocaulon and further east.

Specimens from the two localities are deposited in the herbarium at the British
Museum (Natural History) (BM), I.K. & L.F. Ferguson Nos. 2639 & 2667.

I.K. & L.F. FERGUSON

SHORT NOTES 40]

Eriocaulon was searched for independently in the following lochans in 1970 but
with no success:

south-east of Beinn nan Ord GR 17/44.64 Lochan an Dobhrain 17/47.70
Lochan Tom Mhic Iain 17/51.66 north of Beinn an Leathaid 17/51.68
lochans on Ben Laga 17/64.62

G. HALLIDAY

643/1b. DACTYLORHIZA FUCHSI subsp. OKELLYI (Druce) Sod6—Behaviour and charac-
ters in the Isle of Man.

Specimens of D. fuchsii with small and whitish flowers first attracted notice in the
Isle of Man in 1957. Their suspected identity with the predominantly Irish subsp.
okellyi was subsequently confirmed by several authorities who were sent fresh material
or else saw the plant in the field. Since then the taxon has been found, in very varying
quantities, in over twenty further localities scattered over a large part of the island.
Some of these are in areas where subsp. fuchsii is not known to occur. A number are
perhaps significantly in secondary habitats—notably quarries and sandpits—in
which subsp. fuchsii occurs, by comparison, much more seldom on the Island. In
several, solitary specimens only have been noted (whereas subsp. fuchsii almost
invariably occurs in groups), and several more have obvious hybrid swarms with either
subsp. fuchsii or D. maculata subsp. ericetorum.

This evidence, taken together, points to a taxon with an existence in the island quite
independent of subsp. fuchsii, compared with which it appears to be recent and in-
trusive. It is possible, indeed, that the sudden rush of discoveries made since 1957
reflects an actual spread and is not merely the product of greater taxonomic acumen.
In this connection the fact that the B.S.B.I. Field Meeting in late June, 1950, in which
V.S. Summerhayes took part, failed to produce a record of any such plant may perhaps
be significant. In the intervening period the myxomatosis epidemic has taken place
and, probably as a direct result of this, quite a number of Manx orchid species have
exhibited striking increases.

Most botanists acquainted with subsp. okellyi have seen it only in its Jocus classicus,
the Burren. To judge from the comments made, the situation there is abnormally
perplexing, with the result that many have come away unconvinced that this is a
maintainable taxonomic entity. In several of the Manx localities, however, exclusively
homogeneous populations occur and those who have seen these (for example, the
entire B.S.B.I. Field Meeting party in 1960) have had no difficulty in accepting their
distinctiveness. Several of the discoveries have in fact been made by persons with
little or no knowledge of the dactylorchids. The extent to which the taxon is involved
in hybridisation, and the relative commonness with which this occurs, is no more
than is to be expected in this genus and certainly no argument against taxonomic
recognition. If, as seems likely, this is a relatively recently evolved geographical race,
possessed of some selective advantage in an Atlantic climate, sufficient time may not
yet have elapsed for the development of more than a minimum of isolating barriers
—and these, by virtue of their fragility, may well have proved more than ordinarily
susceptible to the effects of human interference. There is some evidence that subsp.
okellyi prefers drier habitats than subsp. fuchsii, and its smaller, slighter habit seems
to point to an adaptation to shorter herbage. Odd, ‘weedy’ specimens of D. fuchsii
which occur here and there in Man on roadsides and waste ground are often suggestive
of hybrids with very probably subsp. okellyi in their make-up; and it could be that
the advent of this race has made possible an extension in the ecological range of the
species.

A further cause of scepticism about the validity of this taxon may well be the rather
inadequate descriptions of it that appear in the standard books. Accordingly it seems
worth drawing attention to the considerably larger number of characters that have
emerged from a study of the Manx populations. These may, of course, differ to some

402 SHORT NOTES

extent from the populations in Ireland and elsewhere, and it would be useful if the
following description could be checked against material throughout the full range of
the subspecies:

Plant typically small and slender, often no more than 7 cm. Leaves few, relatively
narrow, very faintly spotted, typically yellowish-green, the largest 7-11 cm in
length. Spike narrow and normally short (down to 3.5 cm). Flowers small (labellum
only 5-9 mm wide), cream-coloured, turning pinkish when dried. Outer lateral
perianth segments + equalling labellum, so that the overall profile of the flower
is square (in subsp. fuchsii considerably exceeding the labellum, so that the overall
profile is inversely triangular). Labellum with at most faint pinkish-purple striae,
the central loop markings running vertically (in subsp. fuchsii directed sideways);
middle lobe approximating the laterals in size and shape, triangular, crenate, with
somewhat involute margins (in subsp. fuchsii often considerably longer than the
laterals, triangular to spathulate, entire, not involute); lateral lobes conspicuously
truncate, shorter in relation to breadth than in subsp. fuchsii and with strongly
curved and rounded (instead of almost straight) outer margins.

The Irish populations are said to be fragrant (whereas subsp. fuchsii is almost
odourless), but whether this also holds true of those on the Isle of Man is not yet

certain.
D. E. ALLEN

698/1. CORYNEPHORUS CANESCENS (L.) BEAUV.—In West Suffolk.

This grass was found by the writer and M. Rutterford at Wangford Warren near
Brandon in the West Suffolk (v.c. 26) Breckland in January 1970. Later visits to the
site in June showed a colony of about 100 plants, confined to an area seven yards by
five on a sand bank in association with open colonisation by Carex arenaria and
Agrostis canina subsp. montana.

Although Hind’s Flora of Suffolk (1889) records ‘between Lakenheath and Wang-
ford, G. C. Druce and Bolton King: Lackford Heath, G. C. Druce 1883’, the former
locality is probably now occupied by the aerodrome, whilst it has not been rediscovered
on the heathland at Lackford which was converted to arable and new grazing some
years ago.

The Atlas of the British Flora (1962) shows thirteen post-1930 stations for this
grass, all in coastal areas, of which eleven are in Suffolk and Norfolk (v.c.’s 25, 27 and
28). Wangford Warren, which is leased by the Suffolk Trust for Nature Conservation,
lies on the eastern extremity of the West Suffolk fens and is 28 miles from the south-
eastern corner of the Wash. It is therefore possible to assume that the site was at one

time an area of maritime sand dunes.
P. J. O. TRIstT

Watsonia, 8, 403-411 (1971). 403

Plant Records

Records for publication must be submitted in the form shown below to the appropriate
vice-county Recorder (Watsonia, 8: 435-447 (1971)) and not to the Editors.

Records are arranged in the order given in the List of British Vascular Plants by J. E. Dandy
(1958) and in his subsequent revision (Watsonia, 7: 157-178 (1969)). With the exception of
collectors’ initials, herbarium abbreviations are those used in British Herbaria by D. H. Kent
(1958).

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.

1/3. LYCOPODIUM ANNOTINUM L. 104, North Ebudes: west branch of Allt Camus
na Geadaig, Scalpay, GR 18/58.32. Woodland. H. A. P. Ingram, 1969, field record.
2nd record.

4/8. EQUISETUM PRATENSE Ehrh. 108, W. Sutherland: Ben Hope, GR 29/47.49.
Grassy slopes on west side, 1500 ft. R. W. M. Corner, 1970, herb. R.W.M.C. 2nd
record.

21/3. DRYOPTERIS ABBREVIATA (DC.) Newm. *50, Denbigh: roadside wall,
Llanfihangel, GR 23/98.51. J. M. Brummitt, 1969, det. R. H. Roberts, NMW. (Nature
Wales, 12: 39 (1970)).

24/3. THELYPTERIS PHEGOPTERIS (L.) Slosson 80, Roxburgh: Sundhope New-
castleton, GR 35/51.98. Wooded banks of Lairds Linns. R. W. M. Corner, 1970, field
record. Ist post-1930 record. 82, Haddington: Pressmennan Woods, Stenton, GR
36/6.7. E. P. Beattie, 1969, E. 2nd record.

24/4. GYMNOCARPIUM DRYOPTERIS (L.) Newm. *15, E. Kent: under pines in
Lyminge Forest, GR 61/14.43. E. G. Philp, 1970, MINE.

28/1. BOTRYCHIUM LUNARIA (L.) Sw. 42, Brecon: 3 miles north of Llanbedr,
GR 32/21.25. Mountain ledge on Old Red Sandstone, 1750 ft. M. Massey & M.
Porter, 1970, field record. Ist post-1930 record.

37/1. TROLLIUS EUROPAEUS L. 83, Edinburgh: Heriot station, GR 36/4.5.
In marshy ground by the railway. E. P. Beattie, 1969, field record. Ist post-1930 record.

38/1. HELLEBORUS FOETIDUS L. +82, Haddington: Balgone Estate, North Berwick,
GR 36/5.8. E. P. Beattie, 1969, E. 2nd record.

46/10. RANUNCULUS AURICOMUS L. 84, Linlithgow: Broxburn, GR 36/09.69.
In wood below Almondell. E. P. Beattie, 1969, field record. 2nd record.

46/19. RANUNCULUS FLUITANS Lam. *42, Brecon: in R. Wye one mile east of
Llyswen, GR 32/15.38. M. Porter, 1970, NMW.
46/21. RANUNCULUS TRICHOPHYLLUS Chaix 73, Kirkcudbright: brackish pool,

Southwick Merse, GR 25/91.55. O. M. Stewart, 1968, det. W. Muirhead, E. 2nd
record.

46/23. RANUNCULUS BAUDOTII Godr. 73, Kirkcudbright: brackish pool, South-
wick Merse, GR 25/92.55. O. M. Stewart, 1970, E. 2nd record.

404 PLANT RECORDS

50/3. THALICTRUM MINUS L. 44, Carmarthen: Llyn y Fan Fach, Llandeusant,
GR 22/80.21. Old Red Sandstone cliffs, 1900 ft. D. Davies, 1970, NMW. 2nd record.

61/1. GLAUCIUM FLAVUM Crantz *82, Haddington: Aberlady Bay Nature
Reserve, GR 36/4.8. E. P. Beattie, 1968, field record.

65/4. CoRYDALIS LUTEA (L.) DC. 82, Haddington: on walls, Garvald, GR
36/5.7. E. P. Beattie, 1968, field record. 2nd record.

+69/3. RHYNCOSINAPIS CHEIRANTHOS (Vill.) Dandy 35, Monmouth: disused
railway, Mynydd Dimlaith, GR 31/15.90. G. Ellis & A. E. Wade, 1968, NMW. 2nd
record.

72/2. DIPLOTAXIS TENUIFOLIA (L.) DC. *+71, Isle of Man: derelict garden,
Kirkmichael, GR 24/3.9. M. Devereau & L. S. Garrad, 1968, DGS.

+80/2. CORONOPUS DIDYMUS (L.) Sm. *47, Montgomery: roadside, Erw Wen,
Welshpool, GR 33/23.07. D. Pugh, 1970, NMW.

+81/1. CARDARIA DRABA (L.) Desv. 45, Pembroke: waste ground, Broad Haven,
GR 12/86.13. C. C. Jones, 1970, NM'W. 2nd record.

+98/4. BARBAREA VERNA (Mill.) Aschers. 73, Kirkcudbright: north of Haugh
of Urr, GR 25/76.73. Shingle by R. Urr near Margley Farm. O. M. Stewart, 1970, det.
W. Muirhead, E. Ist post-1930 record.

113/1. VIOLA ODORATA L. *83, Edinburgh: Dreghorn Woods, GR 36/2.6. E. P.
Beattie, 1970, field record.

133/1. STELLARIA NEMORUM L. subsp. GLOCHIDOSPERMA Murb. *42, Brecon:
streambank, Llywel, GR 22/85.30. 750 ft. M. Porter & M. Massey, 1970, NMW;
streambank, Llangynidr, GR 32/14.19. M. Porter, 1970, NMW. ist and 2nd definite
records respectively for the species and subspecies.

140/1. MOEHRINGIA TRINERVIA (L.) Clairv. *71, Isle of Man: shady banks,
Ballakerka, Sulby, GR 24/3.9. M. Devereau, 1969, DGS. Ist definite record.

+149/2. MONTIA PERFOLIATA (Willd.) Howell 82, Haddington: dunes at Gosford,
GR 36/4.7. E. P. Beattie, 1968, field record. 2nd record.

4149/3. MONTIA SIBIRICA (L.) Howell 83, Edinburgh: Fountainhall, GR 36/41.46.
Abundant at Nether Shiels on the Lugate Water. E. P. Beattie, 1970, E. 2nd record.

4154/1. CHENOPODIUM BONUS-HENRICUS L. 72, Dumfries: 2 miles north of Ae
village, GR 25/98.92. Shingle by Water of Ae, 600 ft. O. M. Stewart, 1966, det. W.
Muirhead, E. Ist post-1930 record.

156/5. ATRIPLEX LACINIATA L. 73, Kirkcudbright: sandy shore, Southwick
Merse, GR 25/92.55. O. M. Stewart, 1970, det. M. McCallum Webster, field record.
Ist post-1930 record.

167/1. RADIOLA LINOIDES Roth 73, Kirkcudbright: stony track, Craigend, 3
miles north-west of New Abbey, GR 25/93.69. O. M. Stewart, 1969, det. W. Muirhead,
E. Ist post-1930 record.

4170/2. OXALIS CORNICULATA L. *73, Kirkcudbright: Shambellie Wood, New
Abbey, GR 25/96.66. Peaty path in woods, 100 ft. O. M. Stewart, 1967, EK.

+171/4. IMPATIENS GLANDULIFERA Royle *82, Haddington: Dirleton, GR
36/5.8. By path in Invereil Wood. E. P. Beattie, 1970, E. 83, Edinburgh: Warriston,
GR 36/2.7. Abundant along disused railway line. E. P. Beattie, 1970. 2nd record.

189/2. ONONIS SPINOSA L. 74, Wigtown: east of Isle of Whithorn, GR 25/48.36.
Just above the shore. H. K. C. Mair, 1968, field record. Ist post-1930 record.

PLANT RECORDS 405

+190/2. MEDICAGO SATIVA L. 74, Wigtown: 14 miles south of Glenluce, GR
25/19.55. H. K. C. Mair, 1969, field record. Ist post-1930 record.

4191/4. MELILOoTUS INDICA (L.) All. *35, Monmouth: Newport Docks, GR
31/32.84. A. E. Wade, 1968, NMW. (Nature Wales, 11:138 (1969)).

192/19. TRIFOLIUM FRAGIFERUM L. 82, Haddington: Longniddry dunes, GR
36/4.7. E. P. Beattie, 1969, field record. Only known existing locality.

4192/22. TRIFOLIUM AUREUM Poll. *83, Edinburgh: waste ground, Borthwick,
GR 36/3.6. E. P. Beattie, 1968, E. *84, Linlithgow: waste ground, Bo’ness, GR
26/98. E. P. Beattie, 1970, field record.

4203/1. CORONILLA VARIA L. *82, Haddington: Gullane, GR 36/4.8. Embank-
ment of disused railway. E. P. Beattie, 1968, E. Pencaitland, GR 36/4.6. Embankment
of disused railway line. E. P. Beattie, 1968, E. 2nd record. 83, Edinburgh: Edin-
burgh, GR 36/2.7. Embankment of disused railway. E. P. Beattie, 1967, E. 2nd record.
*86, Stirling: Grangemouth docks, GR 26/9.8. Long-established. E. P. Beattie, E.

206/2. VICIA TETRASPERMA (L.) Schreb. *44, Carmarthen: Cilycwm, Llandovery,
GR 22/75.39. Garden weed. I. M. Vaughan, 1970, NMW.

206/10. VICIA SYLVATICA L. (2, E. Cornwall: cliff at Morwenstow, GR 21/20.16
J. Paton, 1968, field record. Ist post-1930 record. *42, Brecon: Felindre, GR
32/19.35. By paths in coniferous wood, 750 ft. G. Hargest, 1970, NMW. Ist definite
record. 82, Haddington: in Humbie Wood, GR 36/4.6. E. P. Beattie, 1968, field
record. 2nd record.

2111/1. RUBUS CHAMAEMORUS L. *82, Haddington: Gifford, Lammermuir Hills,
GR 36/5.6. Scattered on the Hope Hills and Fasney Water. E. P. Beattie, 1970, E.

211/2. RUBUS SAXATILIS L. *43, Radnor: Radnor Forest, GR 32/18.62. On
rocks in Davy Morgan’s Dingle, 1500 ft. A. C. Powell, 1970, det. G. Ellis, NMIW.

211/9. RuBUsS CAEsIUS L. 73, Kirkcudbright: Port Ling, GR 25/88.53. Creeping
over rocks. O. M. Stewart, 1968, det. E. S. Edees, E. Ist post-1930 record.

211/11/18. RuUBUS LATIFOLIUS Bab. *73, Kirkcudbright: Rockcliffe, GR 25/
84.54. Path by the shore. O. M. Stewart, 1969, det. E. S. Edees, E.

211/11/52. RUBUS NEMORALIS P. J. Muell. 73, Kirkcudbright: Rockcliffe, GR
25/84.53, and Nether Clifton Merse, GR 25/91.56. Both O. M. Stewart, 1968, det.
E. S. Edees, E. Ist post-1930 records.

211/11/59. RUBUS LINDLEIANUS Lees *73, Kirkcudbright: Sandyhills, GR 25/
89.55, and New Abbey, GR 25/97.66. Both O. M. Stewart, 1969, det. E. S. Edees, E.
Ist and 2nd records respectively.

211/11/113. RuBUs POLYANTHEMUS Lindeb. *73, Kirkcudbright: hedgerow,
Rockcliffe, GR 25/84.53. O. M. Stewart, 1969, det. E. S. Edees, E. *74, Wigtown:
2 miles north of Garlieston, GR 25/48.49. By woodland track near Innerwell. O. M.
Stewart, 1967, det. E. S. Edees, E.

211/11/129. RuBUs ULMIFOLIUS Schott 74, Wigtown: 2 miles north of Garlie,
ston, GR 25/48.49. Bank near Innerwell shore. O. M. Stewart, 1967, det. E. S. Edees-
E. 2nd or possibly Ist record.

211/11/165. RuBus vestTIrus Weihe & Nees. *74, Wigtown: 2 miles north of
Garlieston, GR 25/48.49. Near shore near Innerwell. O. M. Stewart, 1967, det. E. S.
Edees, E.

211/11/356. RUBUS DASYPHYLLUS (Rogers) Rogers 73, Kirkcudbright: thicket,
Rockcliffe, GR 25/84.54. O. M. Stewart, 1969, det. E. S. Edees, E. Probably Ist post-
1930 record.

406 PLANT RECORDS

218/2. AGRIMONIA PROCERA Wallr. *106, E. Ross: Craigdarroch Lodge, by Loch
Achilty, GR 28/4.5. Edge of oak wood by farm track, increasing. W. J. Sutherland,
1966, herb. U. K. Duncan.

222/1. SANGUISORBA OFFICINALIS L. 73, Kirkcudbright: Brighouse Bay, 2 miles
south of Borgue, GR 25/63.45. Short sandy grassland. H. K. C. Mair, 1968, field
record. Ist post-1930 record.

223/1. POTERIUM SANGUISORBA L. 73, Kirkcudbright: Kirkdale Bank, 5 miles
south-east of Creetown, GR 25/51.53. Meadowland by shore. H. K. C. Mair, 1966,
field record. Ist post-1930 record.

225/7. ROSA STYLOSA Desv. 44, Carmarthen: old hedge, St Anthony’s Well,
Llanstephan, GR 22/34.09. I. M. Vaughan, 1970, NMW. 2nd record and 1st authen-
ticated one.

~227/2. COTONEASTER SIMONSII Bak. *73, Kirkcudbright: cliffs between Douglas
Hall and Port Ling, GR 25/88.54. O. M. Stewart, 1970, det. W. Muirhead, E.

235/1. SEDUM ROSEA (L.) Scop. 44, Carmarthen: Llyn y Fan Fach, Llandeusant,
GR 22/8.2. 1000 ft. D. Davies, 1970, NMW. Ist record since 1908.

+235/3. SEDUM SPURIUM Bieb. 73, Kirkcudbright: rocks by R. Dee near Old
Tongland Bridge, GR 25/69.53. O. M. Stewart, 1970, det. W. Muirhead, E. 2nd record.

239/15. SAXIFRAGA HYPNOIDES L. *79, Selkirk: Buck Cleuch, GR 36/33.14.
Basic flush, 950 ft. R. W. M. Corner, 1970, field record.

7241/1. ToOLMIEA MENZIES (Pursh) Torr. & Gray *82, Haddington: Gifford,
GR 36/5.6. In deep shade in gorge of Newlands Burn. E. P. Beattie, 1970, E.

242/2. CHRYSOSPLENIUM ALTERNIFOLIUM L. 79, Selkirk: Banks of Ettrick, near
Thirlstane House, GR 36/28.15. Damp woodland, 750 ft. R. W. M. Corner, 1970,
herb. R.W.M.C. Ist post-1930 record.

247/1. DROSERA ROTUNDIFOLIA L. 84, Linlithgow: Raizie Moss, Blackridge,
near Armadale, GR 26/8.6. E. P. Beattie, 1970, E. 2nd record.

~254/13. EPILOBIUM NERTERIOIDES Cunn. *44, Carmarthen: Llyn y Fan Fach,
Llandeusant, GR 22/80.22. By track to lake, 2000 ft. D. Davies, 1970, NMW.

259/2. MYRIOPHYLLUM SPICATUM L. *48, Merioneth: duneland pool, Morfa
Harlech, GR 23/5.3. P. M. Benoit, 1970, NMW. Ist definite record.

261/1. HIPPURIS VULGARIS L. *84, Linlithgow: Easter Inch Moss, Blackburn,
GR 36/0.6. E. P. Beattie, 1968, field record. Pool in Tailend Moss, Bathgate, GR
36/0.6. E. P. Beattie, 1968, E. 2nd record.

294/2. PIMPINELLA MAJOR (L.) Huds. *83, Edinburgh: Warriston, GR 36/25.75.
Abundant on disused railway line. E. P. Beattie, 1970, E.

297/1. BERULA ERECTA (Huds.) Coville 73, Kirkcudbright: Nether Clifton,
GR 25/90.55. Wet rivulet in merse. O. M. Stewart, 1969, det. W. Muirhead, E. Ist
post-1930 record.

300/6. OENANTHE AQUATICA (L.) Poir. 47, Montgomery: by canal 14 miles south
of Arddleen, GR 33/25.13. V. J. Macnair, 1970, det. J. F. M. Cannon, BM. 2nd record.

+309/3. PEUCEDANUM OSTRUTHIUM (L.) Koch *74, Wigtown: 5 miles south of
Stranraer, GR 25/06.54. Edge of cultivated field by ditch, 350 ft. H. K. C. Mair,
1968, field record.

319/17. EUPHORBIA AMYGDALOIDES L. *+74, Wigtown: near Old House Point,
Cairnryan, GR 25/05.70. Waste ground near the shore. H. K. C. Mair, 1968, field
record.

PLANT RECORDS 407

7320/20. PoOLYGONUM SACHALINENSE F. Schmidt *73, Kirkcudbright: New
Abbey, GR 25/96.66. On bank of Pow Burn. O. M. Stewart, 1970, DFS.

$325/5. RUMEX ALPINUS L. *74, Wigtown: roadside, 2 miles west of Drummore,
GR 25/10.35. H. K. C. Mair, 1967, field record.

326/1. PARIETARIA JUDAICA L. 73, Kirkcudbright: Port Ling, GR 25/88.53.
Rocks by shore. O. M. Stewart, 1970, DFS. Ist post-1930 record.

343/1. SALIX PENTANDRA L. *42, Brecon: 2 miles north-west of Upper Chapel,
GR 22/99.43. Banks of moorland stream, 1150 ft. M. Porter, 1970, det. R. D. Meikle,
NMW.

343/6. SALIX PURPUREA L. *42, Brecon: 1 mile north-east of Glasbury, GR
32/18.40. By oxbow pool near R. Wye, 300 ft. M. Porter, 1970, det. R. D. Meikle,
NMW.

343/14. SALIX NIGRICANS Sm. 79, Selkirk: Buck Cleuch, GR 36/33.14. Steep,
flushed slope, 950 ft. R. W. M. Corner, 1969, det. R. D. Meikle, K. Ist post-1930
record. 80, Roxburgh: Lairds Linns, Sundhope, Newcastleton, GR 35/51.98.
Burn sides, 750 ft. R. W. M. Corner, 1970, det. R. D. Meikle, K. 1st post-1930 record.

358/1. VACCINIUM VITIS-IDAEA L. 44, Carmarthen: Llyn y Fan Fach, Llandeusant,
GR 22/80.21. Old Red Sandstone cliffs, 1900 ft. D. Davies, 1970, NMW. ist record
since 1908.

358/3. VACCINIUM ULIGINOSUM L. 69, Westmorland: Cliburn Moss, GR 35/57.25.
R. M. Burton, 1970, herb E. J. Clement. Ist post-1930 record.

358/4. WVACCINIUM OXYCOCCUS L. 43, Radnor: between Colva Hill and Caety
Traylow Hill, GR 32/19.55. On Sphagnum in peat pools, 1600 ft. A. C. Powell, 1970,
field record. 2nd record. 84, Linlithgow: Raizie Moss, Blackridge, near Armadale,
GR 26/8.6. E. P. Beattie, 1970, E. 2nd record.

359/1. PYROLA MINOR L. 83, Edinburgh: North Wood, New Park, GR 36/04.64.
E. P. Beattie, 1968, field record. 2nd record. *84, Linlithgow: Blackburn, GR
26/99.65. Birch wood by Easter Inch Moss. E. P. Beattie, 1968, field record. 108,
W. Sutherland: north-west slopes of Ben Hope, GR 29/47.50. R. W. M. Corner,
1970, field record. Ist post-1930 record.

+370/5. LYSIMACHIA PUNCTATA L. *82, Haddington: dunes at Bilsdean, GR

36/7.7. E. P. Beattie, 1957, E. *83, Edinburgh: North Wood, New Park,
Bellsquarry, GR 36/0.6. E. P. Beattie, 1967, E.
370/7. LYSIMACHIA THYRSIFLORA L. *83, Edinburgh: on the Union Canal, GR

26/2.7. E. P. Beattie, 1967, field record. Cobbinshaw Reservoir, GR 36/00.57. C.S.S.F.
Field Meeting 1968, field record. 2nd record. *86, Stirling: Union Canal between
Glen Village and the Avon viaduct, GR 26/9.7. E. P. Beattie, 1967, E.

371/1. TRIENTALIS EUROPAEA L. 83, Edinburgh: Torfichen Hill, Moorfoot Hills,
GR 36/3.5. E. P. Beattie, 1969, E. 2nd record.

+379/2. VINCA MAJOR L. 82, Haddington: roadside, Longniddry, GR 36/4.7.
E. P. Beattie, 1970, field record. 2nd record.

411/1. HyoscyAMUS NIGER L. 74, Wigtown: near shore, | mile south of Garlies-
ton, GR 25/48.45. H. K. C. Mair, 1968, field record. Ist post-1930 record.

7415/1. DATURA STRAMONIUM L. 47, Montgomery: 1 mile south of Trewern,
GR 33/27.10. C. T. Caswell, 1970, field record. 2nd record.

7416/3. VERBASCUM PHLOMOIDES L. *50, Denbigh: Pentrefoelas, GR 23/87.50.
On refuse tips. J. M. Harrop, 1968, det. I. K. Ferguson, K. (Nature Wales, 11:140
(1969)).

408 PLANT RECORDS

416/10. VERBASCUM VIRGATUM Stokes *+42, Brecon: roadside verge, Llanfrynach,
GR 32/08.26. M. Porter, 1970, NMW. *+71, Isle of Man: waste ground, Douglas,
GR 24/3.7. L. V. Crellin, 1968, det. D. E. Allen, DGS.

4425/1 x 2. MIMULUS GUTTATUS DC. x LUTEUS L. *69, Westmorland: Chapel
Beck, 14 miles south of Orton, GR 35/62.06. G. Halliday (72/70), 1970, det. R. H.
Roberts, LANC.

7428/1. ERINUS ALPINUS L. 80, Roxburgh: ruined walls of Jedburgh Abbey,
GR 36/65.20. J. H. Fremlin, 1970, field record. Ist post-1930 record.

4430/14. VERONICA PEREGRINA L. *6, N. Somerset: Bruton, GR 31/68.35.
Garden weed at Brulands. C. A. Howe, 1970, conf. E. B. Bangerter, BM.

440/3. OROBANCHE RAPUM-GENISTAE Thuill. 6, N. Somerset: Greyfield Wood,
Clutton, GR 31/64.58. On Cytisus scoparius in young conifer plantation. P. A.
Hickmott, 1970, det. R. G. B. Roe, field record. ist post-1930 record.

440/9. OROBANCHE PICRIDIS F. W. Schultz ex Koch *6, N. Somerset: Brean
Down, GR 31/28.58. South-facing cliffs. P. Macpherson, 1969, det. D. J. Hambler.

448/1. THYMUS PULEGIOIDES L. *42, Brecon: Clydach, GR 32/22.12. Limestone
grassland by wood, 700 ft. M. Porter, 1970, NMW.

459/6 x 7. STACHYS X AMBIGUA Sm. *106, E. Ross: north-west side of Loch
Garve near Garve, GR 28/4.6. Bank of stream. U.K. Duncan, 1970, herb. U.K.D.

465/1. GALEOPSIS ANGUSTIFOLIA Ehrh. ex Hoffm. *42, Brecon: 3 miles south-west
of Llywel, GR 22/84.25. Grass verge of lane, 1000 ft. M. Massey & M. Porter, 1970,
NMW.

465/4/2. GALEOPSIS BIFIDA Boenn. *44, Carmarthen: waste ground, Rhagwen
Point, Pendine, GR 22/22.07. I. M. Vaughan, 1969, NMW.
485/2. GALIUM BOREALE L. 80, Roxburgh: Girnwood Linn, 8 miles west of

Hawick, GR 36/36.11. Rocky sides of burn, 1000 ft. R. W. M. Corner, 1970, field
record. First post-1930 record.

502/2. BIDENS TRIPARTITA L. 74, Wigtown: by Soulseat Loch, 2 miles west of
Stranraer, GR 25/10.59. H. K. C. Mair, 1969, field record. Ist post-1930 record.

4503/1. GALINSOGA PARVIFLORA Cav. 83, Edinburgh: Edinburgh, GR 36/2.7.
In a nursery in the City. E. P. Beattie, 1969, E. 2nd record.

4506/4. SENECIO SQUALIDUS L. *82, Haddington: Longniddry, GR 36/4.7. On
disused railway line. E. P. Beattie, 1967, E. *86, Stirling: Grangemouth Docks,
GR 26/9.7. C. S. S. F. Field Meeting, 1969, field record.

506/4 x 7. SENECIO X LONDINENSIS Lousley *82, Haddington: Longniddry,
GR 36/4.7. On a disused railway line with both parents. E. P. Beattie, 1970, E.

506/5. SENECIO CAMBRENSIS Rosser 50, Denbigh: Colwyn Bay, GR 23/83.79.
Verge by A55. J. M. Brummitt, 1966, det. P. M. Benoit. A marked increase of range
from its origin in the east of the county. (Nature Wales, 11:32 (1968)).

506/8. SENECIO VULGARIS L. f. RADIATUS Hegi *71, Isle of Man: Douglas, GR
24/3.7. L. S. Garrad, 1967. *95, Moray: Elgin, GR 38/2.6. Garden weed. M.
McCallum Webster, 1969, E.

4506/9. SENECIO TANGUTICUS Maxim. *82, Haddington: bank of the Gifford
Water below the village, GR 36/5.6. E. P. Beattie, 1969, E.

4506/13. SENECIO FLUVIATILIS Wallr. *82, Haddington: roadside verge, Lennox-
love Mains, near Haddington and banks of Papana Water at Garvald. Both GR
36/5.7, E. P. Beattie, 1967, E. Ist and 2nd records respectively.

PLANT RECORDS 409

4509/4. PETASITES FRAGRANS (Vill.) C. Presl 74, Wigtown: roadside verge,
Cairnryan, GR 25/06.68. H. K. C. Mair, 1968, field record. 2nd record.

513/1. PULICARIA DYSENTERICA (L.) Bernh. 82, Haddington: Aberlady, GR
36/4.8. Banks of Marl Loch. E. P. Beattie, 1968, field record. 2nd record.

514/5. FILAGO MINIMA (Sm.) Pers. *35, Monmouth: colliery tips at Cross Keys,
GR 31/21.91, and New Tredegar, GR 32/15.06. A. E. Wade, 1968, NMW. (Nature
Wales, 11:138 (1969)).

515/1. GNAPHALIUM SYLVATICUM L. 71, Isle of Man: Point of Ayre, GR 25/4.0.
M. Devereau & L. S. Garrard, 1968, DGS. Ist record this century.

$51/2. PICRIS HIERACIOIDES L. *82, Haddington: near Longniddry, GR 36/4.7.
On disused railway line. E. P. Beattie, 1969, E.

555/1. MYCELIS MURALIS (L.) Dumort. 73, Kirkcudbright: near Kirkconnell,
14 miles north-east of New Abbey, GR 25/97.67. Steep grassy bank by road, 50 ft.
O. M. Stewart, 1970, det. H. Milne-Redhead, E. 2nd record.

+558/1/3. HIERACIUM SPELUNCARUM Arv.-Touv. *50, Denbigh: Colwyn Bay,
GR 23/83.79. J. M. Brummitt, 1969, det. P. D. Sell, CGE.

558/1/47. HiERACIUM SCHMIDTIL Tausch *50, Denbigh: limestone cliffs, Bryn
Euryn, Colwyn Bay, GR 23/83.79. J. M. Brummitt, 1968, det. P. D. Sell, CGE. (Nature
Wales, 11: 140 (1969)).

558/1/49. HIERACIUM LEYI F. J. Hanb. *50, Denbigh: Colwyn Bay, GR 23/83.79.
J. M. Brummitt, 1969, det. P. D. Sell, NMW. *73, Kirkcudbright: scree on Douglas
Hall Cliffs, GR 25/88.54. O. M. Stewart, 1969, det. P. D. Sell, CGE.

558/1/54. HIERACIUM VAGENSE (F. J. Hanb.) A. Ley *50, Denbigh: limestone
cliffs, Bryn Euryn, Colwyn Bay, GR 23/83.79. W. Elfyn Hughes, 1967, det. B. A.
Miles, herb. W.E.H. (Nature Wales, 11:140 (1969)).

558/1/73. HIERACIUM BRITANNICIFORME Pugsl. *50, Denbigh: limestone cliff,
Bryn Euryn, Colwyn Bay, GR 23/83.79. J. M. Brummitt, 1968, det. P. D. Sell, NMW.
(Nature Wales, 11: 140 (1969)).

558/1/98. HIERACIUM EXOTERICUM Jord. ex Bor. *50, Denbigh: Bodnant, GR
23/80.72. Under trees in park. J. M. Brummitt, 1968, det. P. D. Sell, NMW. (Nature
Wales, 11:140 (1969)).

558/1/102. H1ERACIUM SUBLEPISTOIDES (Zahn) Druce *50, Denbigh: roadside
bank, Colwyn Bay, GR 23/84.78. J. M. Brummitt, 1968, det. P. D. Sell, NMW. (Nature
Wales, 11:140 (1969)).

558/1/143. HTERACIUM CALEDONICUM F. J. Hanb. *73, Kirkcudbright: Port Ling,
GR 25/88.53. Rocks by the shore. O. M. Stewart, 1968, det. C. West, CGE.

558/1/155. HIERACIUM SUBMUTABILE (Zahn) Pugsl. 150, Denbigh: Llanfairtal-
haiarn, GR 23/92.70. J. M. Brummitt, 1969, det. P. D. Sell, CGE, NMW.

558/1/157. HYERACIUM DIAPHANOIDES Lindeb. *50, Denbigh: Maerdy, GR
23/99.44, and Llanrwst, GR 23/81.58. Both J. M. Brummitt, 1969, det. P. D. Sell,
NMW. ist and 2nd records.

558/1/199. H1ERACIUM SCABRISETUM (Zahn) Roffey 50, Denbigh: Cwm Penanner,
GR 23/89.47. J. M. Brummitt, 1969, det. P. D. Sell, CGE, NMW. Ist post-1930 record.

558/1/203. HimzRACIUM EBORACENSE Pugsl. *50, Denbigh: Llanfihangel, GR
23/98.50. J. M. Brummitt, 1969, det. P. D. Sell, CGE, NMW.

558/1/222. HIERACIUM SALTICOLA (Sudre) Sell & West *60, W. Lancs.: railway
embankment north of Carnforth, GR 34/50.72. G. Halliday (79/70), 1970, det. P. D.
Sell & C. West, LANC.

E

A10 PLANT RECORDS

558/1/228. HIERACIUM ORNATILORUM Sell & West *60, W. Lancs.: shady
verge, Dalton, Burton-in-Kendal, GR 34/5.7. G. Halliday (92/70), 1970, det. P. D. Sell
& C. West, LANC. Ist record for the species for 30 years; previously only known
from Yorkshire.

561/1. BALDELLIA RANUNCULOIDES (L.) Parl. *42, Brecon: bog, 2 miles west of
Libanus, GR 22/9.2. 1050 ft. M. Porter, 1970, NMW.

562/1. LURONIUM NATANS (L.) Raf. 47, Montgomery: canal, 12 miles north-east
of Guilsfield, GR 33/23.13. V. J. Macnair, 1970, det. F. H. Perring, field record.
2nd record (lst extinct).

7571/1. LAGAROSIPHON MAJOR (Ridl.) Moss *35, Monmouth: quarry pool at
Gelligroes, GR 31/17.94. A. E. Wade, 1968, NMW. (Nature Wales, 11:138 (1969)).

576/2. ZOSTERA ANGUSTIFOLIA (Hornem.) Reichb. *73, Kirkcudbright: Auchen-
cairn Bay, GR 25/82.50. O. M. Stewart, 1970, E.
577/6 x 8. POTAMOGETON X NITENS Weber H15, SE. Galway: river south-west

of Kiltartan crossroads and north of Gort, GR 93/4.8. G. Halliday (137/70) & D. A.
Webb, 1970, det. J. E. Dandy, BM, LANC, TCD. Ist post-1930 record.

577/7. POTAMOGETON ALPINUS Balb. *45, Pembroke: 1:5 km east of Ludchurch,
GR 22/15.10. D. A. Cadbury, 1966, det. J. E. Dandy, BM, NMW.
577/13. POTAMOGETON PUSILLUS L. *44, Carmarthen: pond, Moat Llandyry,

near Kidwelly, GR 22/43.05. I. M. Vaughan, 1970, det. B. Seddon, NMW. _—«x4H1I5, SE,
Galway: Coole Lough, near Gort, GR 93/4.7. G. Halliday (133/70) & D. A. Webb,
1970, det, J. E. Dandy, BM, LANC. 2nd record.

577/20. POTAMOGETON FILIFORMIS Pers. H16, W. Galway: White Isle, Lough
Mask, GR 94/1.3. Shallow sandy bay on east side. G. Halliday (121/70) & D. A.
Webb, 1970, conf. J. E. Dandy, BM, LANC, TCD. Ist post-1930 record and confirma-
tion of the only v.c. record.

580/1. ZANNICHELLIA PALUSTRIS L. *73, Kirkcudbright: Carlingwark Loch,
Castle Douglas, GR 25/76.61. E. I. Biggar, 1969, conf. H. Milne-Redhead, field record.

584/1. NARTHECIUM OSSIFRAGUM (L.) Huds. 83, Edinburgh: marshes by Cob-
binshaw Reservoir, GR 36/01.57. E. P. Beattie, 1968, field record. 2nd record. *84,
Linlithgow: Tailend Moss, Bathgate. E. P. Beattie, 1968, E. Raizie Moss, Blackridge,
GR 26/87.66. E. P. Beattie, 1970, field record. 2nd record.

597/1. GAGEA LUTEA (L.) Ker-Gawl. 83, Edinburgh: Millholm Ford, near
Dalhousie, Newtongrange, GR 36/3.6. Woodland bank on R. South Esk. E. B.
Basden, 1967, field record. 2nd record.

4605/2. JUNCUS TENUIS Willd. *35, Monmouth: colliery tip at Cross Keys,
GR 31/21.91. A. E. Wade, 1968, NMW. (Nature Wales, 11: 138 (1969)). 36,
Hereford: above R. Wye, GR 32/54.15. Grassy cart track. L. E. Whitehead, 1967,
field record. 2nd record; not recorded since 1889.

7607/7. ALLIUM CARINATUM L. 72, Dumfries: by R. Nith, Dumfries, GR 25/
96.76. O. M. Stewart, 1970, DFS. 2nd record.

7607/11. ALLIUM PARADOXUM (Bieb.) G. Don *73, Kirkcudbright: woodland
by R. Urr, Corbieter, GR 25/79.66. E. I. Biggar, 1969, conf. H. Milne-Redhead, field
record.

1620/1. CROCOSMIA X CROCOSMIFLORA (Lemoine) N. E. Br. *35, Monmouth:
roadside near Gelligroes, GR 31/18.95. A. E. Wade, 1968, NMW. (Nature Wales, 11:
138 (1969)).

625/4. EPIPACTIS LEPTOCHILA (Godfery) Godfery *36, Hereford: Great Doward,
GR 32/55.16. Side of track in open woodland. E. Watkins, 1970, det. D. P. Young,
field record.

PLANT RECORDS 411

629/1. NEOTTIA NIDUS-AVIS (L.) Rich. 2, E. Cornwall: beechwood, Looe, GR
20/24.53. L. J. Margetts, 1969, field record. Ist post-1930 record.

633/1. CORALLORHIZA TRIFIDA Chatel. *79, Selkirk: Loch of the Lowes, GR
36/23. 19. Willow carr at south end, in two areas, 800 ft. R. W. M. Corner, 1970, field
record.

635/1. COELOGLOSSUM VIRIDE (L.) Hartm. 82, Haddington: Dirleton, GR
36/5.8. In short grass on golf course south oH Yellow Craig. E. P. Beattie, ‘for many
years’, field record. 2nd record.

638/1. PLATANTHERA CHLORANTHA (Custer) Reichb. 83, Edinburgh: Gladhouse
Reservoir, GR 36/30.53. E. P. Beattie, 1968, field record. 2nd record. 84, Linlith-
gow: Blackburn, GR 26/96. Pasture bordering Easter Inch Moss. E. P. Beattie, 1968,
field record. Only known existing locality.

643/3. DACTYLORHIZA INCARNATA (L.) So6 47, Montgomery: marsh, # mile
south-west of Llanfihangel, GR 33/08.16. H. Webster, 1970, field record. 2nd record.

7646/1. ACORUS CALAMUS L. 72, Dumfries: by R. Nith, Dumfries, GR 25/96.76.
O. M. Stewart, 1970, field record. Ist post-1930 record.

649/1. ARUM MACULATUM L. 82, Haddington: Gifford, by Gifford Water,
GR 36/5.6. E. P. Beattie, ‘for many years’, E. 2nd record.

650/2. LEMNA TRISULCA L. *79, Selkirk: Kingside Loch, near Tushielaw, GR
36/34.13. 1150 ft. R. W. M. Corner, 1970, herb. R.W.M.

655/8. SCIRPUS LACUSTRIS L. *83, Edinburgh: Cobbinshaw Reservoir, GR
36/0.5. E. P. Beattie, 1967, E. *84, Linlithgow: pond on Tailend Moss, GR
36/0.6. E. P. Beattie, 1967, E.

663/7b. CAREX LEPIDOCARPA Tausch subsp. scoTicA E. W. Davies *71, Isle of
Man: Strandhall, GR 24/2.6. In alluvial marsh. D. E. Allen, M. Devereau & L. S.
Garrad, 1969, det. A. C. Jermy, DGS. The first definite record for this subspecies.

663/20. CAREX RIPARIA Curt. 82, Haddington: Danskine Loch, Gifford, GR
36/5.6. E. P. Beattie, 1967, field record. 2nd record.

673/2. PUCCINELLIA DISTANS (L.) Parl. *46, Cardigan: sea-wall north-west of
Tre’r-ddol, Dovey estuary, GR 22/64.93. G. Halliday (200/70), 1970, LANC, NMW.
675/1. NARDURUS MARITIMUS (L.) Murb. *33, E. Gloucs.: Lower Harford,

Naunton, GR 42/1.2. On ballast of disused railway line. B. Nash, 1968, conf. E. Philp,
field record.

676/11. POA ANGUSTIFOLIA L. *35, Monmouth: Newport Docks, GR 31/32.84.
A. E. Wade, 1968, NMW. (Nature Wales, 11:138 (1969)).

681/2. MELICA NUTANS L. 80, Roxburgh: Black Burn, 2 miles north-west of
Newcastleton, GR 35/46.89. Rock ledge by waterfall, 650 ft. R. W. M. Corner, 1970,
field record. Ist post-1930 record.

687/1. HORDEUM SECALINUM Schreb. *+71, Isle of Man: near Ronaldsway, GR
24/2.6. In ley beside Silverburn. L. S. Garrad, 1968, field record.

412 Watsonia, 8, 412-422 (1971).

Book Reviews

The Oxford Book of Wild Flowers. Pocket Edition. Edited by S. Ary and M. Gregory.
Pp. vii + 222 with 580 illustrations by B. E. Nicholson. Oxford University Press,
London. 1970. Price £1.25.

This is not another book of coloured pictures of British wild plants to add to the
growing number of such productions but ‘a small replica of the library edition’ of
a work first published in 1960 and twice reprinted, with corrections, in 1962 and 1965.
That it lives up to its latest title by fitting neatly into a jacket pocket is its first recom-
mendation—indeed it could be carried by hand into the field without being an en-
cumbrance. In order to achieve this reduction in size the illustrations have, of course,
become smaller and the print of the text has just about reached the limit for ease in
reading. Two chapters, on ‘Flower Families’ and ‘Ecology of Plants,’ have been sac-
rificed, no doubt to allow for a proportionate decrease in thickness; their loss, though
a pity, should not detract from the declared ‘first purpose’ of the book—‘to enable
the user to identify the wild flowers of the British countryside’.

The items retained are: an Introduction, telling the user how to set about his purpose
and instilling a sense of discretion in picking specimens; a Glossary, limited to one
page, explaining the few technical terms used; Illustrations in colour of some 550
species and in black and white of 30 more, the latter mainly of trees and shrubs, where
shape of leaf, flower and fruit rather than colour is significant; a page of text opposite
each plate (which includes from 5 to 8 figures), giving a simple description with brief
habitat notes and times of flowering of each species, aided by black and white drawings
of particular morphological details; 4 pages of “Tree Shapes’, summer and winter,
of 14 common trees; a page devoted to ‘Naming Plants’ and just over a page to ‘classi-
fying Plants’, wherein some explanation of the use of Latin names and the basic
principles of classification are given with, wisely, no attempt to delve too deeply into
the intricacies of taxonomy; a 7-page index to scientific and English names to end the
volume.

Also, several species closely allied to those illustrated are described in the text,
thus enabling about twice the number to be covered. The plates are primarily arranged
by flower colour but the family to which each species belongs is always stated. The
English name is the first detail given, followed by the scientific name, which is taken
from the Flora of the British Isles by Clapham, Tutin & Warburg (1952). Omitted
entirely are grasses and sedges (pocket-size works on these groups, usually left until
later by the beginner, are now-a-days available). Critical genera such as Rubus, Rosa,
Euphrasia, Taraxacum and Hieracium are sufficiently dealt with as aggregate species
or by major groups.

The Introduction makes it clear that the main ‘user’ envisaged is the one ‘with little
or no botanical training’, so the value of the book is mainly to be assessed by the
adequacy of its figures plus descriptions to guide the beginner to a correct identification
of his plant. Perfection in colour reproduction is notoriously difficult and prohibitively
expensive to attain, so it must not be looked for in such a modestly priced work as
the present one. Some figures are of high standard, the majority are from just to
completely adequate and a few are scarcely or not at all adequate. Figure 3 on p. 113,
for example, would not assist many learners to identify the common milk-wort;
whereas the ‘berried’ species of Ericaceae on p. 121 are clear and distinctive, not enough
distinction is evident between the heath species on p. 119. Now and again inaccuracies
occur, such as depicting Veronica officinalis on p. 175 with an apparent terminal
spike-like inflorescence instead of with axillary racemes. On the same page, fig. 1

BOOK REVIEWS 413

does not represent Veronica persica as a characteristically creeping, prostrate plant,
nor does the text opposite indicate this; it is indeed an example of the tendency to
formalise and to miss the essential habit of many species. Such faults as these, however,
are not so serious as to impair the usefulness of the book as a whole; tested on a
beginner it gave the correct answer for a half-dozen plants in a very reasonable time.
It has no pretensions to being ‘a must’ for the expert or a necessity for the knowledge-
able, but its very handy size, general adequacy and moderate price recommend it to
the beginner and to those seeking a suitable gift for their young relatives.

E. B. BANGERTER

Phytochemical Phylogeny. Edited by J. B. Harborne. Pp. xiv + 335. Academic Press,
London. 1970. Price £5.50.

Phylogenetic speculation is a pursuit which none will begrudge the taxonomist who
wishes to indulge in it as a relief from, and a reward for, the rigorous discipline of his
professional way of life. Others also are lured by its attraction, and among these now
appear the chemists engaged in various aspects of the study of plant life and plant
products.

The book under review appears almost simultaneously with another important
publication: the Symposium on Major Evolutionary Events and the Geological
Record of Plants (Biol. Rev., 45:317—454 (1970)). For the palaeobotanical material
the sources must overlap and the conclusions reached must necessarily in large measure
agree. Both rest heavily on the palynological evidence, and it is this, both palynologi-
cally and phytochemically, which would seem to hold great promise for future develop-
ments in knowledge of the early history of present-day plants. One chapter, that by
G. Shaw on ‘Sporopollenin’, deals specifically with the contribution from this quarter,
sporopollenin itself being a collective term for the nitrogen-free, cellulose-free resistant
wall material of pollen grains.

As regards the phylogeny of the higher plants, phytochemistry is tending to work
backwards from the situation as it exists at present. How does evolution seem to be
working here and now? Proteins, fats, nucleic acids, enzymes, and all manner of
secondary constituents are dealt with in detail in this connection. The situation in the
primitive vascular plant is surmised, and the likely course of evolution therefrom
traced. For the higher plants there is no one chapter devoted to a comprehensive survey
—the phytochemical material at present to hand will scarcely allow of this; but for
the origin of plants as such the first chapter, by P. Echlin, does succeed in drawing
together numerous threads and reaching positive conclusions.

This book, therefore, is only a scratching of the surface of the subject claimed in its
title. There is no doubt that chemistry will have a massive contribution to make, all
in good time, to the phylogeny of plants, the more effectively as it becomes organised
under some such heading as ‘Chemical Taxonomy’ or ‘Biochemical Systematics’. The
immediate need is for an interpretation of morphology in terms of chemistry, and this
is dramatically growing in the current expansion of knowledge of plant hormones
and the way in which they determine and affect the structure of the plant and its
organs. It is here that the so-called ‘secondary’ plant constituents become of such
importance. They are the substances that differ as between plants, and it is from them
that the linkages are being forged with the morphological characters on which system-
atics is based. What can already be achieved in this direction is well illustrated by the
chapter by J. McClure on the secondary constituents of aquatic angiosperms; but at
a more detailed level those by B. L. Turner on molecular approaches to population
problems at the infraspecific level and T. J. Mabry on sesquiterpene lactones in
Ambrosia are equally significant. These chapters are selected as dealing more especially
with higher plants, but the fungi and unicellular organisms are not omitted from

consideration.
E. C. BATE-SMITH

414 BOOK REVIEWS

Principles and Methods of Plant Biosystematics. O. T. Solbrig. Pp. xiii + 226 + 70
figures. The Macmillan Company, London. 1970. Price £5.

This attractively produced introductory textbook of plant biosystematics is clearly
aimed at the American college student who has studied elementary botany or biology.
It is not assumed that the reader has had a course in genetics. The first hundred pages
are about the principles of biosystematics. The seven chapters discuss the process
(sic) of speciation and the forces that control it. Chapter headings read: Synthetic
Theory of Evolution, Patterns of Phenetic Variability, Breeding Systems, Speciation,
Hybridization, The Species Problem and Classification. In the middle of the book there
is a complete change of approach. Part II, again of some hundred pages, reports on
special techniques for the study of species, with chapters on Genetics, Cytology,
Chemistry, Mathematics and Statistics.

There are several excellent chapters in the book, those on plant variation and
breeding systems being very useful. The section on techniques leaves the reader in no
doubt as to the importance of the different approaches to plant variation. Nevertheless,
despite good material, this remains a disappointing book. It may convince the reader
that modern techniques are used by students of evolution, but the basic construction
of the book, in two more or less self-contained areas, means that there is serious frag-
mentation of content. To give but one example, polyploidy is discussed most fully
and from first principles in Chapter 9; but much of this material is needed at an earlier
point, for instance in Chapter 7. A student with little or no genetics might find the
going hard.

Throughout the book the author has chosen up-to-date examples to illustrate his
theme. One is disappointed, however, by his parochial attitude. Almost all the examples
are from the work of United States biosystematists. This approach gives a student no
notion of the excellent studies carried out in Poland, Canada, Japan, etc.

A number of omissions and errors have been noted. The text contains practically no
mention of apomixis. It is unfortunate that an error in the formula for variance in
Chapter 11 has escaped correction in proof-reading and there is no definition of OTU’s
in the section on numerical taxonomy or in the glossary, which makes Fig. 11-3
inexplicable for the beginner. The first name given to a plant is not necessarily the
valid name. Mendel did not propose the principles of inheritance. The section on
Linnaeus’s view on classification is very misleading. The glossary defines analysis of
variance but nowhere is this analysis discussed in the text. Also certain controversial
areas of the subject are not presented in sufficient balance; for example Thoday and
Boam’s experiments on disruptive selection in Drosophila have now been repeated and
extended by many geneticists, often with quite different results (Thoday & Gibson,
American Naturalist, 104: 219-30 (1970), give a full bibliography of the researches in
this area).

D. BRIGGS

Trees and Shrubs—Their Identification in Summer or Winter. C. T. Prime and R. J.
Peacock. 6th edition (revised and enlarged by C. T. Prime). Pp. 139. Heffer & Sons
Ltd, Cambridge. 1970. Price 65p.

This attractive new paperback is a substantially enlarged and revised sixth edition of
the authors’ booklet of 1935 How to identify Trees and Shrubs from Leaves and Twigs
in Summer or Winter. The short introductory chapter introduces the reader to the
trees and shrubs of our country, their biology, important diagnostic features and their
place in the landscape of Britain. Then follow two keys, one for use in summer and
the other in winter, with a short introduction on ‘How to use the Keys’—which the
author has rightly labelled ‘Important’. This is a feature far too rarely provided by
the authors of keys intended for use by non-specialists, and, when provided, usually
neglected by those casually wishing to name plants. The keys have obviously been

BOOK REVIEWS 415

compiled with great care and it is clear that serious thought has been given to the
selection of the simplest channels through which the user can be led to a correct
identification. The keys contain a number of trichotomous leads but, while most
systematists are of the opinion that keys should be uniformly dichotomous on grounds
of simplicity, it must be admitted that these particular examples seem to be without
any undesirable complexities. The reviewer’s son (aged 13) experienced no problems
in using the keys under trial conditions. Then follow concise descriptions of about a
hundred species, including all the common natives, with a selection of the more frequent
and interesting introductions. For each species a paragraph after the introduction
mentions points of historical interest, horticultural aspects, related species, economic
uses and autecological attributes. As always in Dr Prime’s work, biological information
is prominent, and it is clear that the book stems from much time spent observing the
living plants and is in no way a library or herbarium compilation. The work concludes
with twelve photographs of barks and buds in close-up and twenty pages of drawings
by R. J. Jones of leaves, twigs and silhouettes. The leaves and twigs are excellent,
being drawn with great directness and simplicity, capturing the essential features with
a minimum of superfluous detail. The silhouettes are much less successful and lack
the authority and conviction of the other drawings. As in most books the reviewer
can find minor points for comment. Thus, under Gooseberry we find the rather odd
statement ‘A native of Central and Southern Europe, probably wild only in the North
of England’; likewise the Giant Sequoia of California (here referred to by the unusual
vernacular name Mammoth Tree) is treated as a species of Sequoia along with its
relative the Redwood. Specialists today seem to be agreed that it should be accorded
recognition as the monotypic genus Sequoiadendron, a procedure which incidentally
allows the retention of the familiar epithet giganteum. Although this book was origin-
ally produced for teaching purposes and has already passed through five editions, it
is clear that in its new form it has evolved into a work of more general utility. At 65p
it is not unduly expensive by modern standards and can be confidently recommended
to the beginner who wants to learn to identify our trees and shrubs in a thoroughly
scientific manner and not merely by facile (and error-prone) picture-matching. More
experienced users of this book will find many new and interesting pieces of information,
which are likely to stimulate even the most jaded of botanical palates.

J. F. M. CANNON

Cellular Differentiation in Plants and Other Essays. C. W. Wardlaw. Pp. 160. Manchester
University Press, Manchester; Barnes & Noble Inc., New York. 1970. Price £2.

Definitive treatment for the professional and simplified texts for the beginner are
the necessary partners in the advancement of learning. Informal discourse, on the
other hand, is a beneficial luxury. It enables us to read why the writer, admitting that
actions and opinions are not exclusively determined by ‘the facts’, thinks as he does.
In this vein, Professor Wardlaw has written these essays, mentioning his regret that
the senior botanists of his youth did not similarly express themselves.

The good-natured banter of the first essay, ‘On writing botanical essays’, serves to
assure us that our author is still his old self and we embark on the main business
with ‘Aspects of cellular differentiation in plants’. The design is historical as it is
part of Wardlaw’s intention to show how problems of differentiation have been
handled by workers of different interests and periods. Considerable attention is given
to the impact of molecular biology. This and the following essay, ‘Reconciliations
among apex lovers’, will certainly help Honours and research students, and also
teachers, to find their bearings. Discussion predominates and factual material is
present in just the right amount. These essays are ideal for tutorials.

The notions of preformation and epigenesis enter into the treatment of these themes
and in the next essay, ‘Enigmas of Epigenesis’, Wardlaw gives them further attention.

416 BOOK REVIEWS

He does not, however, consider the philosophical analysis of the concepts as, for
instance, Woodger has done.

In the last essay, Wardlaw looks back on the recent phase of experimental mor-
phology and reflects on the probability that many of the exhibits should be put in the
rather negative category of disorganizations. Optimism reasserting itself, he is soon
surveying the possibilities of neo-organizations which ‘transcend the parental genetic
limits’.

Austerity is not a part of the style of these essays and some readers may crave for
it by the end. The embellishment is, however, an expression (an epigenesis?) of the
author’s natural (preformed?) gaiety. It is worth much to have a discussion of con-
flicting attitudes without too firm a set of jaw.

F. Cusick

Flora of Norfolk. C. P. Petch and E. L. Swann. Pp. 288 + 115 coloured illustrations.
Jarrold & Sons, Ltd, Norwich. 1968. Price £2.25.

The Flora of Norfolk was published to coincide with the Centenary of the Norfolk
and Norwich Naturalists’ Society. Its somewhat lavish production is a gesture by the
publishers to Norfolk natural history. In order to attract buyers it has 26 pages of
colour photographs, some bad, some good, and a few very good indeed; all are of
flower portraits and not always typical of the Norfolk flora. I would like to have seen
some general pictures of Norfolk habitats and well-known botanical sites besides these
plant portraits.

There are three preliminary chapters: those by Swann on the history of botany
in Norfolk and by G. P. Larwood on the geology and fossil floras are very concise,
but that of Petch on the plant communities leaves something to be desired. It is doubtful
if the terms ‘fen’ and ‘bog’ can be defined without recourse to the plant communities
that are associated with them. Certainly the type of ‘bog’ or mire seen in several
Norfolk sites, but especially at Buxton Heath, should have been mentioned. Con-
siderable work has been done on the hydrosere of the broadland marshes by Dr Joyce
Lambert; but her classic papers are not mentioned and the east river valley associations
are dismissed in two lines. “True fen’, it is said, is characterised by having Cladium
mariscus dominant; this is a marshman’s definition rather than that of an ecologist.
There might well have been mention here of Nature Reserves in Norfolk, as the county
has the oldest, if not the most active, Naturalists’ Trust and has been subjected to
some intense study by the Nature Conservancy.

The bulk of the Flora deals with the distribution of some 28 taxa of Characeae,
361 of Bryophyta and 1563 of vascular plants. The problem for the compilers has
been, of course, the old one—what to leave out rather than what to put in. I feel the
result has not done justice to the careful and detailed notes that I know at least one
of the authors has on his files. Both have known and worked in the county for some
40 years. Tight budgets and perhaps a rush to produce the book for the Centenary
may be the cause of the scanty, often variable notes on each species. There is a regret-
table although understandable bias for records from Vice-County 28; both authors
have lived or based themselves in the west of the county and Norfolk is well over 80
miles across. Nicholson’s Flora (1914) was biased in the opposite direction; but,
because of the changing landscape, pollution of the Broads and suchlike, the balance
is not redressed. However, if past and present records had been amalgamated, trends
and distribution would have been shown, or at least the need for confirmation would
have been exposed. An example of this is seen under Potamogeton coloratus, where
Petch and Swan give no records for v.c.27, whilst Nicholson gives 6 from the Broads.
First or other historical records are given on some occasions and not on others; and
some obvious ones have been missed out, e.g. Carex divulsa, first recorded and des-
cribed as a new species by Stokes from v.c.28. The typeface is clear, but lack of varia-

BOOK REVIEWS 417

tion in size or the use of bold face makes finding a particular genus or species very
difficult.

Probably the most irksome aspect of the Flora is the lack of any decent map giving
even the rudiments of the National Grid. Throughout the text reference is made to
the 10km squares (by a two-figure reference which, however, is not ambiguous). But
unless the reader knows the county well he cannot form a picture of the distribution
of its flora without recourse to at least the } in. O.S. map. Other maps, or at least one
showing soils, would have been useful.

Most botanists will, by now, have bought this book; but to those who are still
undecided I can recommend it as well worth the price. Like most (but fortunately
not all) County Floras, it is not a book for the young or uninitiated—in spite of the
pictures.

A. C. JERMY

Grass: a story of Frankenwald. Edward Roux. Pp. xx + 212 with 18 plates + 44 text
figures. Oxford University Press, Cape Town. 1969. Price £4.

Primarily this is a lively record of co-operative human endeavour: a history of the
early years of research at the biological field station of the University of the Wit-
watersrand. It describes the experiments and discusses the conclusions of the scientists
who have worked there since 1931.

The research reported mainly concerns one problem: what are the factors which
determine the course of secondary succession in the High Veld? For anyone with only
a theoretical knowledge of ecology, here is an exciting account of the practice. The
chapters deal piecemeal with the complex environment without losing sight of its
wholeness. In the process, a number of traditional axioms of pasture management
in Africa are found to be surprisingly inadequate. The proper use of the world’s
resources is the real subject of this book.

J. LEwIs

Millefiori della Valle d’ Aosta. Degiovanni Luciana. Pp. 285 + 125 plates in colour.
SGS, Torino. 1969. Price 2,500L.

This book provides brief descriptions, frequencies, localities and medical uses for
about a thousand of the more interesting plants of the Val d’Aosta. In spite of short-
comings, it is to be recommended as an inexpensive guide to a rich botanical area
for which information is not otherwise readily available.

J. E. LOUSLEY

Index to European Taxonomic Literature for 1966. R. K. Brummitt and I. K. Ferguson,
Pp. 245. Regnum Vegetabile. Vol. 53. International Bureau for Plant Taxonomy and
Nomenclature, Utrecht. 1968. Price £1.25 (I.A.P.T. members), £2.90 (others).

Index to European Taxonomic Literature for 1967. R. K. Brummitt and I. K. Ferguson.
Pp. 202. Regnum Vegetabile. Vol. 61. International Bureau for Plant Taxonomy and
Nomenclature, Utrecht. 1969. Price £1.65 (I.A.P.T. members), £3.40 (others).

As the number of botanical publications in the field of taxonomy and related discip-
lines is increasing every year, it has become more and more difficult for individual
botanists to keep up to date with recent discoveries, and they have to spend much
valuable time searching through the literature relevant to their studies. Since the
work on the Flora Europaea was initiated this problem has become more acute,
especially to the contributors of that Flora. The existing reference and abstract periodi-
cals Fortschritte der Botanik (covering various fields of botany) and Excerpta Botanica

418 BOOK REVIEWS

(taxonomy and chorology), which are devoted to all plant groups from different
parts of the world, were too incomplete with regard to the European taxonomic
literature on vascular plants, especially from East Europe. This gap was filled when the
International Bureau for Plant Taxonomy and Nomenclature began to publish
annually an Index to European taxonomic literature in the Regnum Vegetabile.

The first issue of this Index (for 1965) was compiled by R. K. Brummitt and pub-
lished in 1966 as Regnum Vegetabile, vol. 45. The next two are mentioned above, and
the fourth (for 1968) is in the press for spring 1971.

The Index covers firstly the whole of Europe (as defined in Flora Europaea, vol. 1,
p. xvi (1964)), as well as neighbouring areas, such as Madeira and the Canary Islands
in the West, N. Africa in the south, and Cyprus, Asiatic Turkey, Syria, Lebanon,
Israel, Jordan and the Caucasus in the East.

The compilers have examined the literature received at Kew, the British Museum
(Natural History), the Department of Forestry of the University of Oxford, the
Department of Agriculture of the University of Cambridge and the Ministry of
Agriculture (London).

The Jndex is easy to use. All items are treated uniformly and organised in alpha-
betical order. The author’s name is given first, then the title of the publication, followed
by a reference to the periodical or publisher if it is a book. Next is given the year of
publication followed by an indication of whether the summary is in a language
other than that of the publication itself. The title is given in its original form (Cyrillic
scripts are transliterated), followed by whatever language translation of it the author
has given in the publication. Useful information regarding new taxa described is also
included and an indication given (by the use of heavy type) that these have been
validly published.

The Index begins with some smaller chapters covering topics of general interest,
such as biography, bibliography, botanical institutions and gardens, phytogeography,
mapping and chromosome surveys. These are followed by a chapter on the Floras,
grouped according to the countries. The main bulk of the Jndex consists of references
to papers and books dealing with the taxonomy of vascular plants arranged systemati-
cally.

The Jndex for 1966 contains more than 3100 references and includes about 1600
new names of various taxa, while that for 1967 has about 3500 references and more
than 1500 new names.

These references are supplemented by an appendix, consisting of a list of the full
titles of the periodicals that are abbreviated in the main part of the Index. In the
issue for 1966 the compilers have cited 712 periodicals and a further 196 are given
in the issue for 1967.

The references are compiled by the authors and grouped with great skill and care,
so that this Index is of immense value, not only for students of the European Flora,
but also for all botanists who are studying the taxonomy, ecology and distribution
of any taxon of vascular plants.

The whole /ndex is well prepared and nicely produced. The compilers and publishers
should be congratulated on such an important contribution to the study of the systema-
tics of European vascular plants.

A. MELDERIS

Modern Methods in Plant Taxonomy. Edited by V. H. Heywood. Pp. xv + 312.
Published for the Botanical Society of the British Isles and the Linnean Society of
London by Academic Press, London and New York. 1968. Price £4.20.

A detached observer might find it remarkable that plant taxonomy apparently provides,
simultaneously, a stereotype of all that is drab, outmoded and dispensible in botany,
and an ever-rolling bandwaggon upon which new developments in the plant sciences
(and outside them) seem perennially ready to climb. This implies no disrespect or

BOOK REVIEWS 419

ingratitude to those who have enriched systematic botany with their contributions
in the past, or are so enriching it now, but it epitomises a situation in which any
survey of the present state of the subject is likely to be of more than passing interest.

The present volume contains the text of the contributions to the conference organised
by the B.S.B.I. in association with the Linnean Society, held in Liverpool in September
1967. The occasion was an impressive one. The speakers were well chosen to cover a
broad field, and included some very eminent names. As a result, the book contains a
well-balanced and fascinating cross-section of plant systematics in the 1960s. The
decision not to print the discussion that followed the papers was undoubtedly right;
one’s great regret (obviously shared with the editor) is that this has deprived us of the
contribution to the conference made by Professor Takhtajan.

After a general introduction by the editor, the papers are presented in four groups:
‘the continuing role of the herbarium in modern taxonomic research’, ‘the role of
experimental data’, “biochemistry, computers and taxonomy’ and ‘geography and
ecology’. It is eminently right that the book should begin with A. Cronquist and
J. P. M. Brenan’s outlines of the functions of major herbaria. These, after all, are the
places where most of the real and useful work of systematic botany is done. J. McNeill
contributes a valuable and instructive discussion of the functions and problems of
regional and local herbaria; it is a little sadly that one wonders whether the Liverpool
University herbarium will always exist in as favourable a milieu as it did in 1967. C. R.
Metcalfe gives a down-to-earth appraisal of the value of systematic plant anatomy,
touching briefiy on the comparable (if currently more fashionable) field of chemo-
taxonomy. The second group of papers embrace the field which has become known as
“experimental taxonomy’; much work in this category is concerned with micro-
evolution rather than with systematics as such. D. M. Moore’s paper on the karyotype
occupies an interesting borderline position; the chromosomes may be regarded simply
as providing additional characters to supplement those drawn from other sources,
or their behaviour may be studied to draw directly phylogenetic conclusions. O. T.
Solbrig gives a useful discussion of the relation between morphological and genetic
discontinuity. C. D. K. Cook analyses three cases of heterophylly in aquatic plants,
ranging from a situation with a strong element of heteroblastic development in Synnema
triflorum to the autoregulatory mechanism, switched by photoperiod, of Ranunculus
aquatilis, and discusses the taxonomic consequences of these situations and of plasticity
in general. W. H. Wagner’s paper on interspecific hybrids includes a consideration
of the evolutionary effects of hybridisation, and of the practical taxonomic treatment
of hybrids. The third group of papers brings us back to pure taxonomy, allied with
new techniques. D. E. Fairbrother’s paper will be valued as a condensed introduction
to the techniques and literature of chemotaxonomy and systematic serology. J. Cullen
discusses, critically but not antagonistically, some of the problems that are likely to
arise in the practical application of numerical methods. M. B. Dale gives a brief outline
of the sequence of procedures in a numerical taxonomic investigation, with a note
on list-structures as a possible means of representing data. M. P. Johnson & R. W.
Holm compare numerical classifications of Sarcostemma obtained using correlation
coefficients and taxonomic distance with floral and vegetative characters and conclude
(not surprisingly) that the non-specificity hypothesis holds only in part, and that
detailed analysis of character-sets may be of great interest in evolutionary and ecological
studies. W. T. Stearn briefly discusses principal component and cluster analyses of
data from Columnea and Alloplectus in relation to a taxonomic revision of the Jamaican
species of these genera. The final group of papers includes a short general consideration
of adaptive relationships by D. A. Wilkins, a stimulating paper on ecogeographical
relationships (and a good deal else besides) in relation to taxonomy by F. J. F. Fisher,
and a substantial review of the evolution of patterns of infraspecific differentiation by
F. Ehrendorfer.

Modern Methods in Plant Taxonomy partakes of the usual strengths and weaknesses
of symposium volumes. Each contributor can write with authority about his own

420 BOOK REVIEWS

corner of the field, but the general effect is inevitably somewhat disjointed and the
reader is left to make his own broad synthesis. What, in fact, is the field of plant tax-
onomy ? Clearly it embraces and must always be centred around the practical systematic
work that produces the Floras and monographs that we notice chiefly when they are
missing. A number of topics considered in this book, such as chemotaxonomy and
numerical classificatory techniques (which may well be seen as the major fields of
advance of the 1960s), are extensions of, or ancillary to, traditional taxonomic methods.
Systematists have no doubt always been interested in the origin of the entities they
classify, and since the time of Darwin phylogenetic speculation has seemed a natural
and legitimate part of a systematist’s work; since the rise of genecology the interest
has shifted largely (and profitably) from a macro-evolutionary to a micro-evolutionary
level. Sometimes the phylogenetic tail has tried to wag the taxonomic dog, but on the
whole the contributions to this book maintain a more critical and level-headed per-
spective than could have been expected a few decades ago. Phylogenetic conclusions
are still in essence descriptive and demand their own explanation and synthesis, and
this is perhaps why I, at least, found the final chapters of this book the most stimulating.
It is in this region that the boundaries of systematics are most diffuse, and the con-
nections with neighbouring disciplines richest. There is no hard-and-fast line between
experimental taxonomy and population genetics. We cannot understand the effect
of genes and of natural selection without considering the limitations on the possible
imposed by pathways of development. The evolution and functioning of genetic
systems are very closely linked with patterns of taxonomic differentiation on the one
hand and ecosystemic relationships on the other. If present-day ecosystems are built
up of (to borrow a phrase from Margalef (1968)) ‘prefabricated parts’ called individuals
and species, we can surely only understand these parts by considering their function
in the overall design, the production-line on which they are made, the transport systems
by which they reach their destinations, and the mechanisms by which they are fitted
into their places alongside the other components of the ecosystem. Phytogeography
and dispersal have also been traditional interests of systematists. Perhaps, as the
recent book by MacArthur & Wilson (1967) suggests, their problems are neither as
isolated from the field we are considering nor as intractable as they have sometimes
appeared. However, the next great synthesis of the living world will no doubt have
to wait for a latter-day Darwin. Meanwhile, we can be grateful to the organisers of
the conference, the contributors and the editor for giving us as good a book as this.
Probably the honours botany student or the incipient postgraduate systematist will
gain the most from it, but parts at least will be of interest to the keen amateur, and
there will surely be few professionals who will not learn from it something new about
how the other half of the taxonomic world lives. The book is well produced, and the
price is not exorbitant by present-day standards.

REFERENCES
MacArTHur, R. H. & WILSON, E. O. (1967). The theory of island biogeography. Princeton.

MARGALEF, R. (1968). Perspectives in ecological theory. Chicago.
M. C. F. PROCTOR

Viruses in plant hosts. Form, Distribution and Pathologic Effects. K. Esau. Pp. vill + 225
University of Wisconsin Press, Madison, Milwaukee and London. 1968. Price $10.00"

This book is based on the subjects covered in a series of three lectures given by Pro-
fessor Katherine Esau at the second J. C. Walker Conference held in Madison, Wis-
consin in 1968. These conferences deal with restricted subject-matter discussed in
greater depth than is usually possible, and the subject in 1968 was the morphological
and anatomical aspects of plant disease.

Professor Esau largely restricts her subject to two plant viruses, beet yellows and
tobacco mosaic, which she and her colleagues have investigated over a number of

BOOK REVIEWS 421

years. Other viruses are discussed, but in less detail, and for this reason the title of
the book is perhaps somewhat misleading.

By its comprehensive coverage of the development of virus-related protein and its
relationship with cell components and cell type, the transport of virus, and the effects
of infection on the cell, this book neatly collates the observations on beet yellows and
tobacco mosaic. In addition, there are 137 superb electron micrographs which illustrate
the ultrastructure of infected cells, and are of exceptionally high quality.

S. M. Ross

Evolution and Phylogeny of Flowering Plants. John Hutchinson. Pp. xxv + 717 with
517 text figures. Academic Press, London and New York. 1969. Price £8.50.

The subtitle to this book (Dicotyledons: Fact and Theory) gives a clue to its contents.
In it, Dr Hutchinson has found room to discuss (and often illustrate) many of the
most extraordinary and interesting genera of Dicotyledons that he was unable to treat
in sufficient detail in his previous works on the families and genera of Flowering
Plants. Here one finds reasons for many of the decisions about family relationships
that are incorporated in his well-known classification (but not, it must be added, for
the basic division of the Dicotyledons into Lignosae (mainly woody) and Herbaceae
(mainly herbaceous)). Each Dicotyledon family in that classification is described
informally and its relationships discussed. A more detailed description of the type
genus is included, often with a citation of its type-species ; and references are also made
to other genera that have especially remarkable characters or some economic impor-
tance. To complete the factual side, Dr Hutchinson has included a large number of
line drawings and some maps, many of them depicting the above-mentioned type
species and mostly from his own pen. These in themselves make it a fascinating book
to browse through.

Fascinating ? Yes, but also exasperating. The text is an amalgam of factual exposition
and opinion with, every now and then, a ‘hit’ at some person, persons or viewpoint
with which the author disagrees. This provides entertaining reading; but, for the
reviewer at least, it raised doubts as well as smiles. Not all the facts are facts. To take
one example, if Xylopia is ‘the only truly natural genus common to the tropics of
both hemispheres’ (p. 22), then what about the genus Annona (Custard Apples), which
is native to both Africa and America? Likewise, not all the theories are in accord with
the most recent research. Thus the Cactales continue to repose among the ‘mainly
woody’ Lignosae, quite divorced from the Portulacaceae and other Centrospermous
families, although evidence continues to accumulate from various sources, chemical
as well as morphological, that these groups are closely related. Again, the Eucry-
phiaceae are still included in the Guttiferales, although in the reviewer’s opinion it is
quite anomalous in that order and appears to be much more at home near the
Cunoniaceae, where Bausch placed it in 1938 (Bausch 1938).

Despite the inaccuracies scattered through the text, however, the reader of this
distillation of a lifetime’s experience of Dicotyledon taxonomy will obtain a clear
picture of the characters of the various Hutchinsonian orders and their suggested
inter-relationships, as well as much information not otherwise easily accessible.
Whether the student mentioned so often in the text will be able to afford the book is
another matter. It is well produced, with remarkably few printing errors. Anyone
interested in the inter-relationships of flowering plant families—or even those merely
fascinated by the variety of form that they exhibit—should make a point of dipping
into this book. They may well decide that, even at £8.50, it is very good value.

REFERENCE
Bauscu, J. (1938). A revision of the Eucryphiaceae. Bull. Misc. Inf. R.B.G. Kew, 1938: 317-349.

N. K. B. ROBSON

422 BOOK REVIEWS

Numerical Taxonomy. Edited by A. J. Cole. Pp. xv + 324. Academic Press, London.
1969. Price £3.

This is a symposium volume, consisting of papers presented at a colloquium on
numerical taxonomy held at St Andrews University in September 1968. The papers
cover a very wide range of subjects in the field, from the more abstract mathematical
aspects to its application in fields as diverse as business enterprises, social geography
and botanical taxonomy.

There are 18 papers in all, arranged quite arbitrarily on alphabetical sequence of
authors’ names. This is slightly disturbing at first; but since there are probably as
many ways of classifying the papers as there are methods which they enumerate, it
is as reasonable a system as any other. Of these papers seven are either on botanical
topics or by botanists and two are geological; zoology, geography, economics, bio-
chemistry and microbiology are represented by one each, while the remaining four
are by theoreticians with computing backgrounds. All the authors are convinced
that numerical taxonomy is a Good Thing. There are therefore no papers reiterating
what it is and why it should be used. Discussion in general in many of the papers
revolves round comparisons of different methods.

Goronzy (on business enterprises), McNeill et alia (on Caucalid Umbellifers) and
Wilmot and Grimshaw (social geography) restrict their techniques to cluster analysis,
while Boyce and Ivimey-Cook (on anthropoids and the genus Ononis respectively)
use both cluster analysis and Principal Components-type techniques. Among the
papers concerned with applications of numerical methods is one by Parker-Rhodes
and Jackson, who give an ecological/fungal classification using a programme originally
devised for linguistic work. This scheme is unusual in that it allows overlapping
classes to be formed.

Two papers deal with the comparison of classifications. Sneath’s is a useful review
article, dealing with the comparisons in qualitative rather than quantitative terms.
Jackson’s is a theoretical paper dealing with the comparison of class'fications, either
with themselves or with the original data, and concluding with a graphical technique
of potential use in determining the ‘goodness’ of a classification.

The volume is produced by offset from typewritten copy and is thus slightly tiring
to read, though the type-face is slightly shaded and therefore easier than some. Mis-
prints are refreshingly few. Each paper is followed by a transcript of the ensuing
discussion. These proved helpful in clearing up difficulties encountered in the texts
of some of the papers. An appendix gives specifications or sources of the programmes
used by the contributors.

This book is not for the beginner, but as a source book of ideas and improvements
for someone already involved in numerical taxonomy, it should prove reasonable
value for money—the price is not exorbitant for this day and age.

D. B. WILLIAMS

Watsonia, 8, 423-424 (1971). 423

Obituary
BEVERLEY ALAN MILES
(1937-1970)

Beverley Alan Miles died suddenly in his sleep on 26th January, 1970 at the early age
of 32. He will be sadly missed by many friends, and especially those interested in Rubi.

He was born at Porthcawl in Glamorgan on 17th November, 1937 and educated at
Bridgend Elementary School and Bridgend Grammar School for Boys. He was
awarded a State Scholarship and went up to St John’s College, Cambridge in October
1956, receiving first class Honours in Part I of the Natural Sciences Tripos in 1958.
In 1958-59 he took the course for Part II of the Natural Sciences Tripos in Psychology,
without however sitting the examination. He also took a course in Danish, passing
the Oral examination. Miles received his B.A. in 1959 and his M.A. in 1963.

In October 1959 Miles took up the position of Biology Master at St Mary’s Grammar
School for Boys at Sidcup, being in charge of that subject until his resignation in
December 1969. He was respected by masters and pupils alike for his scholarship,
and his capacity for informal discussion and keen sense of humour were much
appreciated. Although he suffered from a painful form of bronchial trouble, especially
during the winter months, which often made teaching laborious and difficult, he rarely
complained and was always cheerful. The respect of his senior pupils was shown by
the large number of sixth-formers present at his funeral.

Miles was appointed Curator of the South London Botanical Institute on May
30th, 1969, but did not take up residence until a month before his death. It was his
intention to spend his whole time on botanical research.

His interest in plants started at the very early age of 7 or 8 years, when he went on
country walks with his parents and collected flowers to take home and press in a book.
By the time he had reached the age of 14 he had built up a collection of botanical
books, and all his spare time and holidays were spent collecting plants, his parents
taking him to many parts of the British Isles. My first acquaintance with him was when
I received, on the 10th August 1954, a letter asking me if I would check the identifica-
tion of 20 specimens of Hieracium. To my surprise 18 of the 20 specimens were correctly
named, and (what I did not know at the time) by a self-taught 17-year-old. On his
arrival at Cambridge in 1956 he paid many visits to the Herbarium, and towards the
end of his stay approached me for a suggestion as to a critical genus on which to work.
I suggested Taraxacum or Rubus, and after one summer’s work on Taraxacum he
finally turned to Rubus, and chose as his College Prize W. C. R. Watson’s Rubi of
Great Britain and Ireland. We drew up together a programme on which he could work,
based on the similar problems and experiences I had had with Hieracium. It was
particularly unfortunate, in view of his early death, that I impressed upon him the
necessity of working for about 10 years before he made any major publication. He
was preparing his first papers at the time of his death.

Miles’ main contribution to the study of British plants lies in the work he had done
on Rubus in the Cambridge Herbarium (CGE). From his days at Cambridge onwards
he was allowed completely free access to the Rubus herbarium and, when he moved to
Sidcup, was given permission to take as much material as he liked to his home for study.
The Rubus collection at Cambridge now consists of 21,795 sheets and contains the
original herbaria of J. Lindley, C. C. Babington, E. S. Marshall, W. H. Mills and
J. E. Woodhead, as well as a large proportion of that of W. C. R. Watson, and many
sheets of W: A. Leighton, T. B. Salter, F. A. Lees, E.F. and W. R. Linton, W. M.
Rogers and W. C. Barton. During the 10 years Miles worked on Rubus he examined
and annotated the major part of this material. The Watson herbarium in 1959 was
unmounted and labelled only with copious abbreviated notes scribbled on the margins
of newspapers. One of Miles’ first tasks was to mount this material himself and

424 OBITUARY

decipher the labels. The Babington Collection of Rubi was sent on permanent loan
to the British Museum in 1934, but was never incorporated into their collection. In
view of Miles’ work on the Cambridge collection, permission was obtained to return
the Babington collection. Miles personally carried out this transference, collecting
it from the British Museum, annotating it and returning it in sections to Cambridge.
While annotating the Cambridge herbarium he selected many lectotypes, a task not
attempted by Watson in his revision of the British Rubi.

His nearness to the large London herbaria and libraries allowed him to check on
other types and also to examine the prolific literature on the subject. He worked
meticulously through all the British literature, extracting every name ever used in
Rubus, and made attempts to check the original place of publication. In the course
of his checking of the literature and the selection of types he found many earlier
combinations not in the present checklist of British Rubi, and realised that many
name changes had to take place. The only one of these published was the typification
of Rubus echinatus Lindley and its synonymy (Watsonia, 6: 295 (1957)). Fortunately
the remainder of this work is set out in his manuscripts. He also examined specimens
in other herbaria and for private individuals, records of these specimens being found
in his notebooks.

As well as working in the herbarium he travelled with his close friend, Roy Jones,
to many parts of the British Isles, building up a large collection of his own, and in
particular making gatherings from many type localities. His herbarium contains 4,560
sheets of Rubus, beautifully pressed, copiously annotated and mounted to the best
advantage. After an excursion to the Cornish peninsula he added his opinion to
that of the late W. H. Mills in thinking that Watson was wrong in not recognising as
distinct the many species described by F. Rilstone from that area.

His interest in Rubus was not confined to the British species. He had looked at
Rubus from many parts of the world and, just before his death, had started work on
the account of Rubus for the Flora of Turkey. R. L. Amor, working on Rubus in
Australia, sent Miles several boxes of specimens of European brambles introduced
into Australia. These Miles had named, and there is a nearly complete manuscript
giving an account of them, which it is hoped can be published in the near future in
conjunction with Amor.

Miles’ early interest in Hieracium, particularly the Section Alpina, was continued
throughout his life. From 1965 onwards he spent several weeks each summer camping
in the Scottish mountains so that he could make a particular study of this Section.
His herbarium contains 778 specimens of Hieracium, including some very fine samples
of Alpina and representing most described British species of the genus.

His herbarium, consisting of 7068 sheets in all, contains an excellent series of
Sorbus and many sheets from his native county, Glamorgan. Miles left no will. He
always gave his friends to understand that he wished his herbarium to go to Cambridge,
but on accepting the Curatorship of the South London Botanical Institute had
announced his intention of adding his collection to theirs. The Council of the Institute,
however, decided they would be happy for his herbarium and manuscripts to go to

Cambridge, in accordance with the wishes of his father.
P. D. SELL

I should like to pay tribute to the friendliness and generosity of Beverley Miles.
I spent a few days at his house three years ago and he gave me free access to his her-
barium and was extremely helpful and kind. He lived at the heart of W. C. R. Watson’s
countryside and was in a good position to study Watson’s species as living bushes.
The greatest value of Miles’ herbarium is that it contains reliable specimens of nearly
all the brambles recognised by Watson. If the student armed with Watson’s Handbook
wants to know what Watson meant by a name, he can usually find the answer in
Miles’ herbarium. The next step is to test Watson’s identifications and correct them
where they are wrong. Miles had only just begun to do this and there is a long way

to go.
E. S. EDEES

Watsonia, 8, 425-433 (1971). 425

Reports

CONFERENCE REPORT
HEDGES AND LOCAL HISTORY

Some four years ago the Meetings Committee decided to extend the scope of the
Society’s regular annual programme by holding special meetings from time to time
jointly with other national bodies. The aim of this was twofold: to provide members
with a greater variety of indoor meetings, especially during the largely uncatered-for
winter months (without, if possible, duplicating programmes already arranged under
other auspices); and to extend the Society’s range of contacts with bodies sharing
at least a part of its particular field of interest. The first experimental venture of this
kind was a joint evening meeting in February 1967 with the History of Pharmacy
Section of the Pharmaceutical Society of Great Britain. This was sufficiently successful
to encourage a search for other appropriate topics, of fairly general interest—ideally
to both bodies in equal measure—that would lend themselves to a symposium-type
discussion of either an evening or a day in duration.

Accordingly, the news that Dr M. D. Hooper, of Monks Wood Experimental
Station, had developed a method of dating hedges by means of a simple count of
the number of shrub species contained in them could hardly have come at a better
moment. Here, clearly, was a topic which could not merely serve to bring the field
botanists and the local historians together (as it later transpired, apparently for the
very first time ever), but which was sufficiently important in itself to warrant a meeting
specially for the purpose at this juncture. An approach was thereupon made to the
Standing Conference for Local History, who duly agreed to act as co-sponsors of a
one-day conference on the subject. And at the instance of Dr Hooper, and by kind
permission of the Director, Monks Wood was decided on as the venue.

Some 90 people—almost exactly half botanists, half local historians—in the event
attended, on Saturday, 14 June 1969, filling the lecture-hall at Monks Wood to more
than capacity.

Dr K. MELLANBY opened by welcoming those present to the Station and read out
a message which the Society had received from its Patron, expressing her keen personal
interest in the subject of the conference.

The following five talks were then given :—

Professor W. G. Hoskins (University of Leicester): Hedges and the Local
Historian.

Dr M. D. Hooper (Nature Conservancy, Monks Wood): Plants and Hedges.
D. E. ALLEN: Bramble-dating: a Promising Approach.

R. H. Richens (Commonwealth Bureau of Plant Breeding and Genetics): Elms
as Historical Indicators.

Professor A. D. BRADSHAW (University of Liverpool): Hedges and Hawthorn
Hybridisation.

Professor A. M. Everitt (University of Leicester) contributed the final summing
up.

A half-hour following the break for lunch was given over to an inspection of the
tall hedge fronting the Station, which documentary evidence indicates was planted
by William the Conqueror’s niece. This provided all those attending with an oppor-
tunity to try out for themselves the ‘Hooper Method’ of dating.

Throughout the day the community of interest between field botanists and local

*E

A426 REPORTS

historians was much in evidence, and it came as no surprise that at the close of the
conference there was a general call for another joint meeting on the topic reasonably
soon. This, it was suggested, might perhaps be focussed on the results of some of the
more intensive, localised surveys of hedges which it was confidently expected that
teams of local botanists and historians would now be undertaking in collaboration—
and which it had been a particular aim of this conference to promote.
The full proceedings are to be published as a booklet by the Standing Conference
for Local History. A short account appeared in Nature, Lond., 223: 18 (1969).
D. E. ALLEN

PIEE DEMME E RUNG Sa o7.0

SWITZERLAND AND ITALY

BOURG ST PIERRE. AUGUST

A party of four attended the third B.S.B.I. meeting to be based on Bourg St Pierre.
Botanising began on the Furka Pass where two stops were made during the long
drive from Zurich. Below the Tiefenbach hotel were rich slopes where we examined
such typical plants as Campanula barbata, Rhododendron ferrugineum, Tofieldia
calyculata and Trifolium montanum, and were delighted to find Polygala chamaebuxus.
Pedicularis tuberosa was the yellow-flowered member of this genus on the acidic soils
of this region. At the top of the pass, Soldanella alpina, Alchemilla pentaphylla and
Cardamine resedifolia marked areas where the snow had only recently melted.

The first two days at Bourg St Pierre were spent in splendid weather mostly in the
vicinity of the village. Two excursions were made in the Valsorey where several un-
common British plants were frequent, notably Phyteuma spicatum, Peucedanum
ostruthium and Polystichum lonchitis, and Lilium martagon under a clear blue sky
kept the photographers busy. During the ensuing thunderstorm which enlivened the
evening, the smaller Caryophyllaceae and Euphrasia spp. were identified.

An afternoon at the Great St Bernard Col produced four buttercups: Ranunculus
aconitifolius, R. pyrenaeus, R. acris subsp. steveni and R. glacialis in both white and
pink forms, the latter being noticeably more vigorous. Gagea fistulosa also occurred
above the lake. Across the border into Italy Doronicum grandiflorum occurred in
abundance on an area of block scree, not far from a small patch of Senecio doronicum,
a far less spectacular plant. On an area of scree six dwarf willows and Adenostyles
leucophylla were among plants identified, while a stream-side produced Pedicularis
rostrato-spicata, the most handsome of the seven members of this genus seen during
the week.

After the earlier excellence of the weather, it was disappointing to wake up the next
day to find the village shrouded in cloud, and armed with pullovers and anoraks we
hurried south through the St Bernard tunnel to the relative warmth of Italy.

On a south-facing slope in the Val di Gran San Bernardino lists made were at first
surprisingly familiar to some of the party: Artemisia campestris, Silene otites, Phleum
phleoides, Medicago falcata and M. x varia with Herniaria glabra and Alyssum
alyssiodes on bare patches. However the occurrence of Melica ciliata and Bupleurum
ranunculoides helped us to become re-orientated. A short stop was made above the Val
d’Aosta to admire Eryngium campestre, and the rest of the day was spent in the Cogne
Valley near the Gran Paradiso National Park. Here the flora had a much more southern
appearance with Juniperus sabina, Ononis natrix and Astragalus exscapus on the hill-
sides, and areas of Myricaria germanica on the shingle ridges by the river.

Monday began with a quick exploration of the sadly overgrown alpine garden of
La Linnaea, belonging to the University of Geneva. Only the stronger growing plants

REPORTS 427

have survived, and beautifully inscribed labels are all that remain of the less invasive
plants. From this rather melancholy spot we drove down the valley to Vichéres and
then worked our way up the Combe de I|’A. Below the tree-line the streamside path
was blocked by a tangled mass of spruce trees brought down by a recent avalanche, one
of many which occurred in this area of the Alps during the spring. Above it the water
flowed beneath deep snow beds, an indication of both the severity of the previous
winter and the cool summer. In the spruce forest the ground flora was dominated by
Melampyrum sylvaticum and Luzula nivea with occasional plants of Epipactis atro-
rubens, Ajuga pyramidalis and Cicerbita alpina. Above the spruce was a small wood
of Alnus viridis with Adenostyles alliariae beneath.

The stream which flows down the valley follows a fault line, one which must have
caused considerable displacement, as the western side is of steep limestone, in contrast
to the darker, acidic rocks opposite. Once across on to the limestone, the flora changed
equally abruptly, and an area of scree produced Coronilla vaginalis, Kernera saxatilis,
Polygala alpestris, Helianthemum alpestre and Pedicularis adscendens, while the lime-
flushed ground by the stream had Viola rupestris, Veronica aphylla, Draba aizoides
and Pinguicula alpina amongst its many plants.

The weather on the last day, during which we returned to Ziirich, was more like a
caricature of an English November, and crossing the high pass at ‘snail’s pace’ in
thick mist meant that a brief stop to examine Digitalis grandiflora and Campanula
thyrsoides was all the botanising possible.

Although no completely exhaustive list was kept during the week, over five hundred
plants were identified, and a wide range of alpine and sub-alpine habitats examined.

G. TUCK

ENGLAND

CONSERVATION OF CHALK GRASSLAND IN BEDFORDSHIRE. 15TH—17TH MAY

The object of this field meeting, which was attended by 34 members and visitors, was
to study problems of the conservation of chalk downland in various stages of its
development. Extremes were seen in two small nature reserves. The first, visited on
the Saturday morning, was Knocking Hoe National Nature Reserve, claimed as virgin
downland and of great interest for its assemblage of rare plant species. As it is now
undergrazed, experiments are in progress to estimate the value of mowing as an alter-
native to grazing. The second, visited on the Sunday morning, was Totternhoe Knolls
Local Nature Reserve, a medieval castle site with spoil heaps of previous workings
for Totternhoe Stone. Here the turf is at most six centuries in its development and
until management was undertaken by the Bedfordshire and Huntingdonshire
Naturalists’ Trust it had become overgrown with scrub. Sheep will be re-introduced,
which may maintain the population of the Man and Musk Orchids but destroy butter-
flies in their pupal stage. To remedy this, grazing will be at different times of the year
in fenced-off sections. At a third site, Barton Hills, visited on Saturday afternoon,
there is a greater expanse of scenically attractive downland which had until about twelve
years ago suffered badly from scrub invasion. Part was ploughed for a ley on which
sheep were put and at a later stage allowed to graze on the downland from which scrub
had been cleared. Much of the site is now overgrazed, to the benefit of the flowering
plants but to the detriment of the insect life, which is the subject of continuous study
both on the downland and in enclosures. Attention was drawn from time to time in
these visits to the need to have a clear objective in conservation.

In the passage to and from these sites, a chalk stream, a beech plantation and arable
fields were studied in less detail. The meeting ended with a visit to beech woods near
Tring which are probably the eastern limit of natural woodland in the Chilterns.

J. G. Dony & T. C. E. WELLS

ap

428 REPORTS

GUERNSEY. 29TH MAY—/TH JUNE

The exigencies of term-time meant that only about 25 people took part in the follow-up
of the previous year’s visit to Guernsey, and as a result, under 60°% of the Island’s
squares were listed. But this perhaps made the haul all the more remarkable. The
gathering again had local support. H.E. The Governor regretted he was away and
could not attend, but invited a party into the grounds of Government House, an area
hitherto unvisited, where 140 plants were listed. There was a special service at St
James the Less, with the Lesson read by the Leader; a joint meeting with the Société
Guernesiaise on the first Friday evening, at which plants were discussed and trans-
parencies shown; and several Guernsey people participated in the field for some or
all of the time.

What was found? Nothing dramatically new, but plenty of value in filling gaps and
getting greater accuracy. For example, Aphanes arvensis had hitherto always proved
to be A. microcarpa; the typical subspecies of Pilosella officinarum was at last detected
in several places; and ‘Juncus bulbosus’, not noted for nearly half a century, when
found at Saints Bay proved to be J. kochii. Better, new indeed to the Channel Islands,
were two fine tufts of Fumaria purpurea on soil dumped, from where? Everyone
seemed delighted at the perennial Amaranthus deflexus in good quantity at the Vale
Power Station, Sampsons, which is therefore no longer only a Jersey plant. The same
is true of Ranunculus ficaria subsp. ficariiformis at Caledonia; and of Potentilla
anglica, at last. Montia perfoliata is almost certainly new to Guernsey. There was a
1929, unvouched, record at La Favorita, in St Martins, but only M. sibirica, which
was noted nowhere in the Island till 1968, is now there (where another new weed was
Meconopsis cambrica). M. perfoliata was in fact in at least three different spots in
the north of the island, one of them, again, on dumped soil, which begs the question.
The Carpobrotus with small pink flowers, which has been erroneously called C.
acinaciformis (but may prove to be C. glaucescens), was found on the north coast.
Previously it was known only in Alderney and Herm. Ribes uva-crispa was spotted
in a hedge, also new to the Island; and half-a-dozen or so casuals and throw-outs
were noted for the first time, plants such as Hypericum calycinum, Lupinus polyphyllus,
Atriplex hortensis, Spiraea japonica, Chrysanthemum maximum and Polygonatum x
hybridum.

No less, or even more, satisfactory, were the plants refound after many years,
many decades even. Carex laevigata, of which there were only two, 75-and 80-year-old,
records was found well away from its previous, long-lost, locality; and who would have
guessed that the next day it would also be found in two more places. A chance deviation
at last located Sambucus ebulus, still in the area where it was known in the 1830’s.
Peplis portula and Apium inundatum were half a century ago in a pool near Fort Doyle,
where there is now no pool. Both were found in a small pond hidden among gorse on
the opposite side of L’Ancresse Common. Silene alba x dioica, also collected, had
only one earlier definite record, in 1907.

But most satisfactory of all was the Gaudinia story. This grass is the subject of a
separate note, so suffice it here that specimens were produced from the area of its
only previous, 1928, record. Wholesale search the next day failed to locate the plants.
But later a patch 20 yards across was found two miles away in native turf on L’Ancresse
Common.

To be fair, several yearned-for plants were not refound. Gaudinia came to light
when Ononis reclinata was being re-re-searched for (experience in Alderney suggests
it was having a year off, but it does seem to be rarer). None of the three old localities
for Trifolium strictum seemed still to hold it, while surely Druce’s 1906 specimen of
Hornungia must be mis-labelled. Erodium glutinosum remains unconfirmed—all the
plants like it were E. cicutarium subsp. dunense; Myosurus defeated much search,
although its locality still seemed suitable.

Nor was Geranium pyrenaicum to be seen in the south-east; but it was in the north-

REPORTS 429

west, near to where G. cf. submolle was recorded in 1968, which itself was found in a
brand new locality, on a hedge bank 1,000 yards to the north. Stellaria graminea
was yet again searched for in vain in the south-east, but was discovered 3 mile to the
west, also in a new place. Sanguisorba minor subsp. minor, long known in the north-
west, was seen in the north-east and, surprise, surprise, on the cliffs on the south-west.

Allium roseum was gay and increasing in its 1957 locality at Rousse and also found
in two new places. A. neapolitanum proved not to be rare, nor Nothoscordum inodorum,
in sweet fragrance at the foot of at least half a dozen walls. The very rare and elusive
Anthyllis vulneraria was seen in situ, where it proved to be of the same subspecies,
corbieri, as in Sark and different from those in Jersey and Alderney. Ranunculus
parviflorus, Sisymbrium orientale, Trifolium squamosum, Vicia bithynica, V. lutea and
Lactuca serriola were all in new places. On one of the two available dumps in the
island were bushes of Centaurea diluta up to 4 ft high. Two good colour forms seen
were Hyacinth Blue (H.C.C.40) Polygala vulgaris and Rose Madder (H.C.C.23/2)
Silene gallica.

Useful work was done recording fresh places for such plants as Trifolium occidentale,
Spergularia bocconei, Asparagus officinalis subsp. prostratus and Gladiolus byzantinus;
and the oh, so slow spread of Senecio squalidus and Veronica filiformis was noted.
Lavatera cretica showed itself widespread and often plentiful. Orchis laxiflora was
mapped in some detail. It is no longer in some places where it used to grow, one at
least of which had been ploughed, for potatoes; and it was said to be a bad year for
orchids anyway. But even so, it was found in over half a dozen different places, with
perhaps 2,500 plants in all—over 2,200 of them in one district of St Peter-in-the-Wood,
beautifully mapped by Mrs Burridge. The aibino re-appeared. But O. morio could not
be refound, nor its hybrid, O. x alata.

There was much else. In all, perhaps 9,000 records were entered in the cards, helping
in the knowledge of distribution; but some task to assess and enter! With so much
useful and enjoyable work, and so much obviously still awaiting finding in these 24
square miles, is it surprising that the visit is to be repeated again in 1971, from 9th-17th
July ?

Finally, thanks are due to at least 16 specialists, so far, who have confirmed or
made all the important critical examinations.

D. McCLINTOcK

WESTMORLAND. 11TH—-12TH JULY

This field meeting was arranged to satisfy a number of members who had had their
botanical appetites for v.c. 69 whetted during the A.G.M. excursion in May 1969.

On the Saturday morning we visited Cunswick Scar, near Kendal, a continuation
northward of the better-known Scout Scar. We started well by finding Stachys x
ambigua and Hieracium strumosum in the car park. The latter is here near its northern
limit as also is Hippocrepis comosa, which was growing in profusion along the scar.
The Sorbi (S. lancastriensis/S. rupicola) growing on the face of the scar provoked much
discussion; Carex ornithopoda was found in the shallow limestone grassland. We
descended to Cunswick Tarn to admire Cladium mariscus. After lunch some drove
to Sunbiggin Tarn and climbed up to the limestone pavement to the north, finding
Dryopteris villarii and Thelypteris robertiana. The rest of the party looked for alpines
near the road at the head of Longsleddale; as is so characteristic of the Lake District,
Alchemilla alpina and Cryptogramma crispa were growing in profusion.

The more energetic of the party ascended the fells above Kentmere on the Sunday.
The weather was superb and there were sufficient of the Lake District rarities to keep
the alpine enthusiasts happy: Epilobium alsinifolium, Myosotis brevifolia, Hieracium
anglicum, H. leyi, Juncus triglumis and Salix herbacea.

The second group visited Roudsea Wood, by the Leven estuary, and Roanhead
dunes, by the Duddon. These had also been visited the previous year. At Roudsea

430 REPORTS

the Nature Conservancy wardens described the woodland management policy and
discussed the problem of saline flooding from the estuary at high tide. The dunes were
disappointing as a result of the June drought. Pyrola rotundifolia subsp. maritima
and Epipactis palustris were still fairly abundant but E. dunense was virtually absent.

G. HALLIDAY & L. T. HYDE

EAST SUFFOLK. 3RD—5TH JULY

The purpose of this meeting was to collect records in underworked areas of East
Suffolk for the Nature Conservancy, under the leadership of Mr P. Nicholson, former
deputy regional officer, East Anglia.

Thirteen members met at the ‘Swan Hotel’, Southwold, on Friday evening, when
Mr Nicholson outlined the geology of the Sandlings area of Suffolk. Colour trans-
parencies were used to illustrate local problems of conservation.

On Saturday, members were divided into four groups. One group visited Benacre
Broad where Lathyrus japonicus was noted. Among other places visited were Iken
heath and marsh, and Fen Wood. Each group visited one or two hornbeam coppices
and notes on these were compared in the evening. It was agreed that the ground flora
was altogether very uniform and sparse, owing to the poor drainage and the density
of the canopy. An exposure value of 54 was recorded in one wood. An area near one
coppice had been cleared and replanted with poplar, and had become recolonised
with a large number of marsh and heathland species. Sorbus torminalis was found, as
well as Allium ursinum and Sedum telephium. Three members were fortunate enough
to find Trifolium ochroleucon, en route.

On Sunday, members collected records for the Suffolk Naturalists’ Trust at their
newly acquired reserve at Hollesley heath. The party then drove to Redgrave fen.
Utricularia vulgaris, Rumex palustris and Epipactis palustris were among the many
interesting plants listed. The lateness of the flowering of Gymnadenia conopsea vat.
densiflora compared with its dry chalkland counterpart, was remarked upon. It was
interesting too, to be able to compare the calcicole and calcifuge floras, respectively,
of the fen and an adjoining area of wet, acid heath. A few members went on to another
fen to try and refind Liparis loeselii but were disappointed to discover that the fen
had been drained.

Other plants of interest found during the meeting were: in woods Asplenium
adiantum-nigrum, Crassula tillaea and Iris foetidissima; in a salt-marsh Sonchus palustris
and on Southwold shore Lupinus arboreus.

H. M. PRoOcTOR

WALES

KIDWELLY, CARMARTHENSHIRE. 13TH JUNE

18 members and guests attended this meeting. The vegetation of the disused ports—
our intended objective—was desiccated by drought, wherefore the meeting was switched
to Towyn Burrows. Under Corsican Pine in the Forest a patch of Pyrola rotundifolia,
forward in bud, measured some 8 ft by 10 ft, whilst Ophioglossum vulgatum and
Eleocharis quinqueflora had enough moisture in the rides for survival. On the open
dunes, though many Dactylorhiza species were in abundance, the Orchis morio of a
month earlier had gone to ground completely as also had Centaurium pulchellum and
C. littorale. It was too early for Gentianellas or for Liparis loeselii. Calystegia soldanella
was much reduced but great mounds of Lathyrus sylvestris were unharmed by the
drought.

A pool which a month earlier had held Ranunculus tripartitus had disappeared
without trace save for one disconsolate frog.

At nearby Pinged Halt Linaria supina had seeded freely despite British Rail’s dose
of weedkiller. Thence to Moat Llandyry where the party was entertained to a mag-

REPORTS 431

nificent tea by the kindness of Mr and Mrs Tallowin. We then toured their interesting
wet woodlands (Galanthus nivalis and Impatiens glandulifera extensively naturalised)
and took Ranunculus trichophyllus and Potamogeton pusillus from their pond. A
torrential thunderstorm made a sudden end to the meeting.

Mrs H. R. H. VAUGHAN

SHROPSHIRE UNION CANAL (MONTGOMERY BRANCH). 14TH JULY

A party of 11 met to record plants of the S.S.S.I. sector between Tan House bridge
and Burgedin Lock and of the Guilsfield Arm. A stop was made where road and canal
run parallel to see Ranunculus circinatus in flower and look for Luronium natans
previously located here but not refound.

In the S.S.S.I. sector the banks were partly choked with Glyceria maxima despite
which plants such as Cardamine amara, Oenanthe fistulosa, O. crocata, Rumex hydro-
lapathum, Scrophularia aquatica, Butomus umbellatus, Sparganium erectum, S. emersum,
Acorus calamus, Berula erecta and Mimulus guttatus were well represented. Sedges
included Carex pseudocyperus, C. riparia, C. acutiformis, C. paniculata, and C.
muricata. Hypericum tetrapterum here supplants the locally more common H. mac-
ulatum.

Eleven species of Potamogeton have been recorded but owing to the heavy growth
of Ceratophyllum only four were collected on this occasion: Potamogeton natans, P.
compressus, P. obtusifolius and P. friesii. Three species of Lemna were found, Lemna
minor, L. polyrhiza and L. trisulca, and in an adjacent marsh Oenanthe aquatica grew
with Stellaria palustris.

In the Guilsfield Arm Luronium natans was found but not Dactylorhiza praetermissa,
present here in 1969. About 154 plants were recorded in the S.S.S.I. and 56 along the
Guilsfield Arm but time prevented a full coverage of the latter, which urgently needs
clearance of overgrown alders.

Mrs H. R. H. VAUGHAN

NATIONAL MUSEUM OF WALES, CARDIFF. 5TH SEPTEMBER

A small group of members was conducted round the ‘Man and the Countryside’
exhibition, the Museum’s main contribution to European Conservation Year 1970,
occupying the whole of the Main Hall and parts of the Botany and Zoology Galleries
and the Library, from Ist July to 16th September. The exhibition illustrated many
aspects of man’s impact upon the animals, plants and scenery of the countryside from
prehistoric times to the present day. It included exhibition material provided by Welsh
Region members and alluded to the work of the B.S.B.I. in recording and mapping
the flora of the British Isles.

‘Man and Nature’, a photographic exhibition prepared jointly by the Nature
Conservancy and Kodak Ltd, supplemented ‘Man and the Countryside’.

The party visited the Welsh National Herbarium, where members had an oppor-
tunity to see the extent of the collections and to examine records and specimens.
For the duration of ‘Man and the Countryside’, part of the Herbarium was used as a
reference library, where visitors could read books and numerous pamphlets on topics
relating to conservation and the countryside.

S. G. HARRISON

SCOTLAND

ISLAND OF BUTE. 23RD—25TH MAY

This meeting was organised at the request of members of the Buteshire Natural
History Society, and the Andersonian Naturalists of Glasgow were invited to attend.

432 REPORTS

A party of 14, including the vice-county recorder, travelled to the island, and was
joined by 3 local ladies, members of the Buteshire Society.

On 23rd May the party travelled to St Ninian’s Bay; where two groups were formed
to cover as much as possible of the shore. The area yielded over 160 species, including
Asplenium marinum, Ranunculus bulbosus, Cochlearia danica, Hypericum elodes,
Carum verticillatum, Oenanthe lachenalii, Ligusticum scoticum, Samolus valerandi,
Mertensia maritima and Listera cordata.

A small wooded area south-east of Loch Quien proved to have a very limited
ground flora.

In the evening, at the invitation of members of the local society, the visitors assem-
bled at the Museum, Rothesay. Here they were welcomed by Mr Harrison, president
of the society, and shown round the premises, and later Miss Marshall and Mr Stephen,
two society members, showed slides.

The following day opened with a visit to deciduous woodland west of Kames Bay,
where nearly 90 records were made; the most noteworthy were Phyllitis scolopendrium,
Thelypteris phegopteris, Hypericum androsaemum, Quercus petraea, Betonica officinalis,
Listera ovata, Dactylorhiza purpurella, Poa nemoralis and Milium effusum. The steep
wooded slopes above the Ettrick Burn in Glen More added Polystichum aculeatum,
Hypericum tetrapterum and Melica uniflora to the day’s total.

In the afternoon the south end of the island was visited. Parietaria diffusa was noted
on the wall of Kingarth Church, and the rocky coast of Dunagoil Bay yielded Poly-
podium interjectum, Ligusticum scoticum, Allium vineale, A. ursinum, Orchis mascula
and Carex otrubae. At the little un-named loch east of St Blane’s Hill Nuphar lutea,
Typha latifolia and Carex paniculata were recorded.

On the final day short visits were made to Loch Ascog, Kirk Dam and Greenan
Loch. A small colony of Ceterach officinarum was found near Kirk Dam, while at
Greenan Loch Nymphaea alba, Nuphar lutea, Apium inundatum and Potamogeton
alpinus were noted. At Balnakailly Wood, in the north end of the island, young shoots
of Carex laevigata were showing, and several interesting woodland ferns were seen;
these included Hymenophyllum tunbrigense, Dryopteris aemula, Thelypteris phegopteris
and 7. dryopteris.

R. MACKECHNIE

GLENDARUEL, ARGYLL. 14TH JUNE

The object of the excursion, at which 8 members and friends attended, was to examine
the flora of the serpentine outcrop in Glendaruel, and in particular to try to confirm
a suspected occurrence of Orchis morio. Unfortunately the Orchis turned out to be
O. mascula. Other species seen were Cardaminopsis petraea, which is fairly common
on all the serpentine cliffs, and Dryas octopetala, which is present in small quantity
and confined to a single crag. .

H. A. MCALLISTER

WEST LOTHIAN MOSSES. 11TH JULY

Three West Lothian mosses were visited. In the morning two Edinburgh members
visited Raizie Moss which is two miles west of Blackridge. It is a very fine raised bog
with an abundance of Oxycoccus palustris, Drosera rotundifolia and Narthecium
ossifragum.

In the afternoon when the party was joined by two members from Glasgow, Easter
Inch Moss, near Blackburn, was explored. This moss had previously been worked
for its peat by a Dutch company and now West Lothian County Council are draining
it with a view to converting the area into recreation grounds. It was fascinating to
examine the plant remains in the successive vegetation layers which the 12 feet deep

REPORTS 433

drains had exposed. The County Council have, however, given an area to the Lothians
Branch of the Scottish Wildlife Trust as a reserve. This area comprises a birch wood
in which Pyrola minor was abundant; a tract of moor on which Drosera rotundifolia
was plentiful; three ponds and adjacent marshes in which Potamogeton natans, Hip-
puris vulgaris, Sparganium erectum, Potentilla palustris, Myriophyllum alterniflorum,
Carex rostrata, C. hirta, C. nigra and C. curta occurred among many other plants;
and a pasture where Platanthera chlorantha was one of the most abundant species—
some 300 plants. Salix pentandra and S. repens were also present.

Tailend Moss is two miles south-east of Bathgate. This is also a fine raised bog.
There are also two ponds and a marsh. In the larger of the ponds fine stands of Scirpus
lacustris and Typha latifolia were noted. Potamogeton natans, Sparganium erectum,
Hippuris vulgaris, Potentilla palustris and Salix pentandra were also recorded. On the
moss Oxycoccus palustris was found in several places together with Narthecium
ossifragum but Drosera rotundifolia was very rare.

E. P. BEATTIE

FORT WILLIAM, INVERNESS-SHIRE. 11TH-18TH JULY

The meeting started on Saturday, 11th July, at Banavie Cross where 10 persons
assembled, of whom only the leader and one other was a member of the B.S.B.I. or
B.S.E. The party repaired to Glen Loy and in showery weather recorded the 5 x 5km
square 27/18 SW. No plants of great interest were seen but there was an interesting
contrast between the south-facing and north-facing sides of the aqueduct carrying
the Caledonian Canal across the Loy River.

The following day was wet but four keen botanists (three of whom were society
members) submitted to a soaking in Glen Righ.

On Monday, 13th July, a party of 8 set off for Glen Roy where 27/38 NW and
27/39 SW were explored.

Tuesday, 14th July, was hot and sunny but with a strong wind and mist on the hills.
A party of 8 explored the ‘Dark Mile’ of Achnacarry and the shores of Loch Lochy,
all in 27/18 NE. Trientalis europaea and Goodyera repens were two plants seen.

On Wednesday, 15th July, a small party of 4 visited Blarmafoldach, where a seam
of limestone in 27/06 NE gave a variety of plants including Listera ovata, but Ranun-
culus bulbosus, said to be near here could not be found.

Thursday, 16th July, promised to be the best day of this stormy week, and for an
hour or two Ben Nevis was clear. An ascent into Coire Leis was made by a party of
5, where Luzula arcuata, Cerastium cerastoides and others were recorded for 27/17 SE.

On Friday, 17th July, a party of 8 botanized around the head of Loch Shiel whenever
it was reasonable to get out of the cars. As the rain increased an early return was made.

Saturday, 18th July, was again inclement and abortive attempts to botanize were
quickly abandoned.

Generally the weather set the limits to the work accomplished in the week, but an
appreciation of the interest and energy of the Natural History Society at Fort William
should be placed on record. It was also interesting to meet a conservation-minded
landowner who was reluctant to allow publication of plant records found on his land.

A. A. SLACK

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Watsonia, 8, 435-447 (1971). 435

The Vice-County Guide
compiled by
F. H. PERRING

INTRODUCTION

This Guide is essentially a revised compilation of the following papers previously
published by the Society: ‘Local Organisation’ (Anon 1967), ‘A Guide for Local
Flora Writers’ (Perring 1964) and ‘Local Floras currently available for sale’ (Perring
1969). The contents are arranged in vice-county order and include within each the
name of the Recorder and, when they are kept, the location of the card index and
vouchers for that vice-county. Also included is the name of any Flora which can be
bought, with the address to write to and the cost (including postage), and a note of
any work known to be in progress towards the preparation of a Flora or Check-list.
To avoid unnecessary repetition, if the most recent Flora for a county is however no
longer available for sale other than second hand, it has been omitted. These may be
found in ‘A Guide for Local Flora Writers’.

Members of the Society are invited to offer help to any Flora writer whose county
they may be interested in. In any case ali records thought suitable for publication in
Plant Records in Watsonia must be sent first to the Recorder. Those which meet the
following criteria are likely to be accepted.

1. First vice-county records.

2. Second vice-county records, except for Hieracium, Rubus and Taraxacum.

3. First record in a vice-county since 1930.

4. First record, or the first since 1930, for the major islands in vice-counties
102-104 and 110-112.

5. Any record not included above which nevertheless extends the range of the
species by over 50 km.

Records for species which are not native in a particular area will only be published if
the plants have established themselves some distance from gardens and similar sources
of origin.

When submitting records please check that all the data published in Plant Records
are included.

Whenever possible, and when it does not conflict with the Society’s Code of Conduct,
voucher specimens should be sent to the Recorder to support records for publication.
Any other records, particularly if they appear to be new to 10 km squares when com-
pared to the Atlas of the British Flora (Perring & Walters 1962), should be sent,
preferably on Record Cards, either to the Recorder, or to the Biological Records
Centre, Monks Wood Experimental Station, Abbots Ripton, Huntingdon, from where
supplies of cards can always be obtained.

The Recorders are always willing to answer queries from members about the flora
of their areas, though this does not include supplying information about localities for
rare plants. Many of them maintain a card index which is generally available to mem-
bers for consultation.

The compiler accepts full responsibility for any inaccuracies in this Guide and
wishes to thank all Recorders who have helped by supplying information about their
counties, and Mr B. W. Ribbons, who compiled the data for Scotland, though he suspects
that some Recorders are being modest about their achievements. As it is intended to
republish the Guide periodically, additions or corrections will always be welcome.

436 F. H. PERRING
REFERENCES

ANON. (1967). Local Organisation. Proc. bot. Soc. Br. Isl., 6: 427-432.

PERRING, F. H. (1964). A Guide for Local Flora Writers. Proc. bot. Soc. Br. Isl., 5: 283-302.

PERRING, F. H. (1969). Local Floras currently available for sale. Prec. bot. Soc. Br. Isl., 7: 615—
617.

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

THE GUIDE

The addresses of the Recorders will be found in the List of Members and Subscribers
published periodically by the Society. The following abbreviations refer to institutions
which are mentioned more than twice in the text (herbaria in bold type).

ABD = Dept. of Botany, St Machar Drive, Aberdeen, AB9 2UD.
BEL = Ulster Museum, Stranmillis Road, Belfast, BT9 SAB.
BM = Dept. of Botany, British Museum (Natural History), Cromwell Road,
London, S.W.7.
CGE = Botany School, Downing Street, Cambridge.
DBN = National Museum (Botany Dept.), Botanic Gardens, Glasnevin, Co. Dublin.
EK = The Herbarium, Royal Botanic Garden, Inverleith Row, Edinburgh, 3.

NMW = Dept. of Botany, National Museum of Wales, Cardiff, CF1 3NP.

OXF = Botany School, South Parks Road, Oxford.

TCD = School of Botany, Trinity College, Dublin, 2.
UNPS = Periodicals Room, University Library, Queen Victoria Road, Newcastle-

upon-Tyne, NEI 7RU.

The names of the vice-counties (excluding Ireland) have been taken from J. E. Dandy’s
(1969) Watsonian Vice-Counties of Great Britain.

ENGLAND

S, CHANNEL ISLES. Jersey: Mrs F. Le Sueur. Index: home. Vouchers: The Museum,
Société Jersiaise, 9 Pier Road, Jersey.
Other Islands: D. McClintock. Index (Guernsey): N. Jee, La Houguette, Castel,
Guernsey. Vouchers: Guille-Allés Library, St Peter Port, Guernsey.
FLORAS: A Check List of the Flowering Plants and Ferns Wild in Alderney and its
Off-Islets. F. Le Sueur & D. McClintock. 1964. £1:10. Wild Flowers of Herm.
F. Le Sueur & D. McClintock. 1962. 10p. A Check List of Flowering Plants and
Ferns Wild on Sark and its Off-Islets. F. Le Sueur & D. McClintock. 1963. 224p.
All obtainable from Guille-Allés Library, St Peter Port, Guernsey.
PRESENT WORK: Mrs Le Sueur and Mr McClintock are collecting data for a Flora
onal x 1 km basis.

1, WEST CORNWALL. L. J. Margetts. Index and Vouchers: home.
FLORAS: A Natural History of Porthleven. H. B. Sargent. 1961. A revised and en-
larged 2nd edition in typescript has been lodged with Truro Museum. Wild Flowers
in Cornwall and the Isles of Scilly. J. A. Paton. 1968. £1-:35. D. Bradford Barton,
Ltd, Truro Bookshop, 18 Frances Street, Truro. (Paperback, 65p).
PRESENT WORK: Dr D. E. Coombe and Dr L. C. Frost are mapping the distribution
of the plants of the Lizard Peninsula on a 1 x 1 km basis.

1b, ISLES OF SCILLY. J. E. Lousley. Manuscript and Vouchers: home.
FLORA: Flowering Plants and Ferns in the Isles of Scilly. J. E. Lousley. Reprinted
with additions, 1970. 20p. The Museum, St Mary’s, Isles of Scilly. See also under 1.
PRESENT WORK: Mr Lousley intends to publish a full Flora to replace the above
Check-list by early 1972.

10,
12,

118},

14,

15,
16,

THE VICE-COUNTY GUIDE 437

EAST CORNWALL. Mrs E. L. Almond. Index and Vouchers: home.
FLORA: see under |.

SOUTH DEVON. &

NORTH DEVON. Miss M. A. Turner. Index (3 only): home. Vouchers: Dept. of
Biological Sciences, Hatherly Laboratories, Exeter.
PRESENT WORK: a project has been initiated by the Botanical Section of the
Devonshire Association and the Dept. of Biological Sciences to collect data for
a new Flora on a tetrad basis.

SOUTH SOMERSET. A. D. Hallam. Index: home. Vouchers: Somerset County
Museum, Taunton Castle, Taunton.

NORTH SOMERSET. Capt. R. G. B. Roe. Index: home. Vouchers: Somerset County
Museum. An index of the flora of the Bristol region is maintained by Professor
A. J. Willis at the Dept. of Botany, The University, Sheffield, S10 2TN.

PRESENT WORK: Capt. Roe is collecting data for a new Flora on a tetrad basis.

NORTH WILTS. L. F. Stearn. Index: home. Vouchers: J. G. Grose.
PRESENT WORK: material is being assembled by Miss W. M. Stevenson for the
publication of a supplement to the Flora of Wiltshire (J. G. Grose 1956). Wiltshire
Plant Notes are published annually in the Wiltshire Archaeological Magazine.

SOUTH WILTS. Miss A. M. Hutchinson. Index: home.
PRESENT WORK: see under 7.

porseT. Dr H. J. M. Bowen. Jndex: home. Vouchers: Dorset County Museum,
Dorchester.
FLORA: A Geographical Handbook of the Dorset Flora. R. d’O. Good. 1948. £1-50.
Addenda: No. 1. 1955. 15p. No. 2. 1961. 10p. Dorset County Museum, Dor-
chester. Prof. Good’s unpublished maps of the distribution of Dorset plants are
kept by the Nature Conservancy at Furzebrook, Wareham.

ISLE OF WIGHT, 11, SOUTH HANTS. &

NORTH HANTS. Mrs I. S. Yule. Index and Vouchers: home.

PRESENT WORK: the collection of data for a new Flora on a tetrad basis has now
been completed. The Flora will be published by Mrs Yule in collaboration with the
Hampshire and Isle of Wight Naturalists’ Trust.

WEST SUSSEX. O. Buckle. Index: I. M. Frenguelli and P. C. Hall, homes.

FLORA: A. H. Wolley-Dod. 1937. Reprint 1970. £5-:25. John Roberts Bookshop,
43 Triangle West, Bristol, BS8 IES.

PRESENT WORK: Mrs M. Briggs is secretary of a committee sponsored by the
Sussex Naturalists’ Trust which is collecting data for a new Flora on a tetrad basis.

EAST SUSSEX. P. C. Hall. Index: home. Vouchers: BM.
FLORA and PRESENT WORK: See under 13.

EAST KENT &

WEST KENT. E. Philp, Maidstone Corporation Museum. Index and Vouchers:
Maidstone Museum.

FLORA: (London area only) A Handlist of the Plants of the London Area. D. H.
Kent & J. E. Lousley. 1951-7. 574p. Mrs L. M. P. Small, 13 Woodfield Crescent,
London, W.5S.

PRESENT WORK: Dr F. Rose has completed the collection of data for a new Flora;
Mr Philp, with the Kent Field Club, is organising the gathering of records for an
Atlas of the distribution of Kent plants on a tetrad basis; the Botanical Section
of the Thanet Natural History Society has prepared data for a Flora of the Isle of
Thanet on a tetrad basis; the Botanical Section of the London Natural History
Society is collecting data on a tetrad basis for that part of v.c. 16 which lies
within 20 miles of St Paul’s Cathedral.

438 F. H. PERRING

17, suRREY. Dr D. P. Young. Index: home. Vouchers: home and South London
Botanical Institute, 323 Upper Norwood Road, London, S.E.24.
FLORA: (London area only) see under 16.
PRESENT WORK: collection of data for a new Flora on a tetrad basis has been
completed by the Surrey Flora Committee of which the editor and recorder is
Dr Young. The work should be published early in the 1970's.

18, SOUTH ESSEX &

19, NORTH ESSEX. S. T. Jermyn. Index and Vouchers: home.
FLORA: (London area only) see under 16.
PRESENT WORK: collection of data for a new Flora on a 10 x 10 km and parish
basis has been completed by Mr Jermyn. The work should be published early in
the 1970's.

20, HERTS. Dr J. G. Dony. Index: home. Vouchers: Hitchin Museum & Art Gallery,
Paynes Park, Hitchin.
FLORA: J. G. Dony, 1967. £2-10. Hitchin Museum.
PRESENT WORK: Dr Dony maintains an annotated copy of the Flora with amenda-
tions and additions. Yearly report in Transactions of the Hertfordshire Natural
History Society.

21, MIDDLESEX. D. H. Kent. Vouchers: BM.
FLORA: (London area only) see under 16.
PRESENT WORK: Mr Kent has prepared an MS. which will be published by the Ray
Society during 1971.

22, BERKS. Dr H. J. M. Bowen. Index: home. Vouchers: Reading Museum, Blagrave
Street, Reading and OXF.
FLORA: H. J. M. Bowen. 1968. £2:25. 20 Winchester Road, Oxford.

23, OXFORD. A. R. Perry. Index and Vouchers: OXF.
PRESENT WORK: a group including Mr A. R. Perry and Dr S. R. J. Woodell is
collecting data for a new Flora on a tetrad basis.

24, pucks. P. R. Knipe. Index: home.
FLORA: (London area only) see under 16.
PRESENT WORK: Mr Knipe is co-ordinating the collection of data for a new Flora
on a tetrad basis.

25, EAST SUFFOLK, &

26, WEST SUFFOLK. F. W. Simpson. Jndex and Vouchers: County Borough Museum,
High Street, Ipswich.
PRESENT WORK: Mr Simpson, assisted by Mr W. E. Lewis, is preparing the MS.
of a new Flora of the county. Mr P. J. O. Trist is chairman of a committee which is
collecting data for a Flora of Breckland on a1 x 1 km basis.

27, EAST NORFOLK. E. A. Ellis. Index: home. Vouchers: home and Castle Museum,
Norwich.
FLORA: Flora of Norfolk. C. P. Petch & E. L. Swann. 1968. £2-25. Jarrold & Sons
Ltd, Norwich.

28, WEST NORFOLK. E. L. Swann. Index: home. Vouchers: home and Castle Museum,
Norwich.
FLORA: ‘West Norfolk Plants Today’. C.P.Petch & E.L. Swann. 1962. Supplement
Proc. bot. Soc. Br. Isl., 4. 60p. E. W. Classey Ltd, 353 Hanworth Road, Hampton,
Middlesex. See also under 27.
PRESENT WORK: Mr P. J. O. Trist is chairman of a committee which is collecting
data for a Flora of Breckland on a1 x I km basis.

29, CAMBRIDGE. Dr F. H. Perring. Index: home and CGE. Vouchers: CGE.

FLORA: F. H. Perring et alia. 1964. £1-874. Cambridge University Press, 200
Euston Road, London, N.W.1.

30,

31,

32,

33,

34,

36,
37,

38,

39,

40,

53;
34,

THE VICE-COUNTY GUIDE 439

PRESENT WORK : a group led by Dr Perring and Dr S. M. Walters will begin collecting
data on a tetrad basis in 1971. An annotated copy of the Flora is maintained by
Mr P. D. Sell. Yearly report in Nature in Cambridgeshire.

BEDFORD. Dr J. G. Dony. Index: home. Vouchers: Luton Museum, Wardown
Park, Luton.

FLORAS: J. G. Dony. 1953. £2:10. Luton Museum. Additional Notes on the Flora
of Bedfordshire, Proc. bot. Soc. Brit. Isl., 7: 523-535 (1969).

PRESENT WORK: Dr Dony is co-ordinating the collection of data on a tetrad basis.
Yearly report in Bedfordshire Naturalist.

HUNTS. T. C. E. Wells. Index: home. Vouchers: Monks Wood Experimental
Station, Abbots Ripton, Huntingdon.

FLORAS: J. L. Gilbert. 1965. 374p. Peterborough Museum.

Flora of Oundle (15 mile radius of Oundle). J. Rees. 1969. 574 p. Biology Dept.,
Oundle School, Peterborough.

PRESENT WORK: Mr Wells is collecting data for a new Flora on a tetrad basis.

NORTHAMPTON. Mrs G. M. Gent. Index: home. Vouchers: Kettering Natural
History Society’s Clubrooms, 183a Rockingham Road, Kettering.

FLORA: (15 mile radius of Oundle) see under 31.

PRESENT WORK: Mrs Gent is collecting data for a new Flora on a 5 x 5 km basis.
Mrs P. Rixon is collecting data for the Soke of Peterborough on a tetrad basis.

EAST GLOUCESTER. Mrs S. C. Holland. Index and Vouchers: North Gloucestershire
Naturalists’ Society.

PRESENT WORK: Mrs S. C. Holland and the North Gloucestershire Naturalists’
Society are collecting data on a tetrad basis.

WEST GLOUCESTER. West of R. Severn: Mrs S. C. Holland. Index and Vouchers:
see under 33.

East of R. Severn: Prof. A. J. Willis. Index: Dept. of Botany, The University,
Sheffield, S10 2TN. Vouchers: Dept. of Botany, The University, Bristol, 8.

HEREFORD. Mrs L. E. Whitehead. Index: home.
PRESENT WORK: Mrs Whitehead has completed the MS. for a new Flora.

WORCESTER. F. Fincher. Index and Vouchers: home.
PRESENT WORK: Mr Fincher is collecting information for a new Check-list.

WARWICK. Dr S. M. Coles. Index and Vouchers: Dept. of Botany, The University,
Birmingham, B15 2TT.

FLORA: Check lists of the Vascular Plants and Bryophytes of Warwickshire. R. C.
Readett, J. G. Hawkes, D. A. Cadbury. 1965. 75p. Birmingham Natural History
Society, The Midland Institute, Birmingham, 1.

PRESENT WORK: a new Flora, prepared by Mr Readett, Prof. Hawkes and Miss
Cadbury will be published by Academic Press in 1971.

STAFFORD. E. S. Edees. Index and Vouchers: home.

PRESENT WORK: since 1941 Mr Edees has been collecting data for a new Flora on
a tetrad basis. A MS. has been completed which he hopes to publish early in the
1970’s.

SALOP. C. A. Sinker. Index: Preston Montford Field Centre, Montford Bridge,
Shrewsbury, SY4 1DX.

FLORAS: Handlist of the Shropshire Flora. L. C. Lloyd and E. M. Rutter. 1957.
50p. Transactions of the Caradoc and Severn Valley Field Club, 16, 1961-67 (numer-
ous additions). 50p. Both obtainable from Mr W. T. Lewis, Berries Mount,
Hazler Road, Church Stretton.

SOUTH LINCOLN. &
NORTH LINCOLN. Miss E. J. Gibbons. Index and Vouchers: home.

440 F. H. PERRING

FLORA: Checklist of Lincolnshire Plants. Rev. E. A. Woodruffe-Peacock. 1909.
With additional map showing Botanical Divisions. £1-00 Public Library, Lincoln.
PRESENT WORK: Miss Gibbons has been collecting data for a Flora which will be
published early in the 1970’s.

55, LEICESTER. Rev. A. L. Primavesi. Index: Ratcliffe College, Syston, Leicester.
Vouchers: City Museum and Art Gallery, New Walk, Leicester.
PRESENT WORK: a Leicestershire Flora Committee has begun collecting data on a
tetrad basis. The chairman is Mr I. M. Evans of the Museum, and the secretary is
the Rev. A. L. Primavesi.

55b, RUTLAND. K. G. Messenger. Index: home. Vouchers: Uppingham School.
FLORA: (15 mile radius of Oundle) see under 31.
PRESENT WORK: Mr J. H. Chandler and Mr Messenger have been collecting data
since 1953 for a Flora on a tetrad basis which will be published by the City
Museum and Art Gallery, New Walk, Leicester in 1971. £2-00.

56, NoTTs. R.C. L. Howitt.
FLORA: R. C. L. & B. Howitt. 1963. £1:50. R. C. L. Howitt, Farndon, Newark.

57, DERBY. Miss K. M. Hollick. Index and Vouchers: The Museum and Art Gallery,
The Strand, Derby.
FLORAS: A. R. Clapham, ed. 1969. £2:25. The Museum and Art Gallery, Derby.
Natural History of Derbyshire (with a section on plants). 1971. 30p. The Museum
and Art Gallery, Derby.

58, CHESTER. A. Newton. Index: home. Vouchers: home or Grosvenor Museum,

Chester.
FLORA: A. Newton. 1971. £2:95. The General Secretary, Cheshire Community
Council, Watergate House, Watergate Street, Chester, CH1 2LW.

59, SOUTH LANCASTER. Miss V. Gordon. Index: home. Vouchers: City of Liverpool
Museums, William Brown Street, Liverpool, L3 8EN.
FLORA: Travis’s Flora of South Lancashire. J. P. Savidge et alia. 1963. £1:65. Miss
V. Gordon, 23 Alder Grove, Waterloo, Liverpool, 22.

60, WEST LANCASTER. E. F. Greenwood. Index and Vouchers: City of Liverpool
Museums, William Brown Street, Liverpool, L3 8EN. -
PRESENT WORK: Mr Greenwood is co-ordinating the collection of data for a new
Flora on a tetrad basis.

61, SOUTH-EAST YORK. Miss F. E. Crackles. Index: home. Vouchers: home and Dept.
of Botany, The University, Hull.
PRESENT WORK: Miss Crackles is collecting data for a new Flora of the East Riding
on a tetrad basis.

62, NORTH-EAST YORK. Miss C. M. Rob. Index: home.

63, SOUTH-WEST YORK. &

64, MID-wEST yorK. Dr W. A. Sledge. Vouchers: Dept. of Botany, The University,

Leeds.
PRESENT WORK: Mr D. R. Grant and Mr T. Schofield are collecting data for a

Check-list of the West Riding.
65, NORTH-WEST YORK. Miss C. M. Rob. Index: home.

66, DURHAM. Rev. G. G. Graham. Index: home. Vouchers: home and, eventually,
Museum and Art Gallery, Sunderland.
PRESENT WORK: Mr Graham is co-ordinating the collection of data for a new
Flora on a random | x | km basis.

67, NORTHUMBERLAND SOUTH &
68, CHEVIOTLAND. Dr G. A. Swan. Index and Vouchers: home.

69,

70,

TL

35,

41,

42,

45,

46,

47

w

48,

49

N)

50

»

THE VICE-COUNTY GUIDE 44]

PRESENT WORK: Dr and Mrs Swan have made an exhaustive survey of the county
and are preparing maps of some species on a 5 x 5 km basis.

WESTMORLAND (& N. LANCASHIRE). Mrs F. L. Woodman. Index: home. Vouchers:
Brathay Hall Field Study Centre, Ambleside, Westmorland.

CUMBERLAND. Miss C. W. Muirhead.
PRESENT WORK: Dr D. A. Ratcliffe is collecting data for a new Flora.

ISLE OF MAN. D. E. Allen. Register: home. Vouchers: The Manx Museum,
Douglas.

FLORA: The Flowering Plants and Ferns of the Isle of Man. D. E. Allen. 1970. 30p.
Manx Museum, Douglas.

PRESENT WoRK: Mr Allen has a larger Flora of the island in an advanced state of
preparation.

WALES

MONMOUTH. R. G. Ellis, National Museum of Wales, Cardiff. Index and Vouchers:

NMW.
FLORA: A. E. Wade. Published by the National Museum in 1971. £2-40.

GLAMORGAN. R.G.Ellis, National Museum of Wales, Cardiff. Index and Vouchers:
NMW.

PRESENT WORK: a Flora committee under Mr R. G. Ellis and Dr Q. O. N. Kay
has begun collecting data ona 5 x 5 km basis for a new Flora under the auspices
of the Cardiff Naturalists’ Society.

BRECON. M. Porter. Index: home. Vouchers: NMW.
PRESENT WORK: Mr Porter began collecting records on a 5 x 5 km basis for the
County Naturalists’ Trust in 1968, and intends to prepare a Flora.

RADNOR. Miss A. C. Powell. Index: home. Vouchers: home and NMW.
PRESENT WORK: Dr J. P. Savidge intends to publish a Check-list in about 1972.

CARMARTHEN. Mrs H. R. H. Vaughan. Index: home. Vouchers: NMW.

FLORA: A list of the Flowering Plants and Ferns of Carmarthenshire. R. F. May.
1968. 55p. West Wales Naturalists’ Trust, 4 Victoria Place, Haverfordwest,
Pembrokeshire.

PEMBROKE. T. A. W. Davis. Index: home. Vouchers: NMW.

FLORA: Plants of Pembrokeshire. T. A. W. Davis. 1970. 55p. West Wales Naturalists’
Trust, 4 Victoria Place, Haverfordwest.

PRESENT WORK: Mr Davis is collecting data for a full Flora ona 5 x 5 km basis.

CARDIGAN. Dr J. P. Savidge.

FLORA: A Supplement to the Flowering Plants and Ferns of Cardiganshire. A. E.
Wade. 1952. 20p. University of Wales Press, Cathays Park, Cardiff.

PRESENT WORK: Dr Savidge intends to publish a Check-list in about 1972.

MONTGOMERY. Miss V. J. MacNair. Index and Vouchers: home.
PRESENT WORK: Dr J. P. Savidge intends to publish a Check-list in about 1972.

MERIONETH. P. M. Benoit. Index: home. Vouchers: NMW.

PRESENT WORK: Mr Benoit hopes to produce a new Flora in due course.
CAERNARVON. M. Morris.

PRESENT WORK: the collection of data for a new Flora (with Anglesey) on a tetrad
basis began in the early 1960’s, at the Dept. of Botany, University College of
N. Wales, Bangor.

DENBIGH. J. M. Brummitt. Index: home. Vouchers: NMW.

442
51,

52,

72,
74,

i)

~)

76,

Ta

~

78

w

71D)
80,

81,

82,
84,

85,

86,

87
89

7

F. H. PERRING

FLINT. G. Wynne. Index: home. Vouchers: Cartrefle College of Education, Wrex-
ham, Denbighshire.

ANGLESEY. R. H. Roberts. Index: home. Vouchers: NMW.
PRESENT WORK: see under 49.

SCOTLAND

DUMERIES., 73, KIRKCUDBRIGHT. &
WIGTOWN. Dr H. Milne-Redhead. Index: home. Vouchers: home and Dumfries
Borough Museum, The Observatory, Corberry Hill, Dumfries, CGE and E.

AYR. R. Mackechnie.
FLORA: The Flora of the Clyde Area. J. R. Lee. 1933. 574p. John Smith & Son
(Glasgow) Ltd, 57-61 St Vincent Street, Glasgow, C.2.

RENFREW. R. Mackechnie.

FLORA: see under 75.

PRESENT WORK: Mr B. W. Ribbons has been co-ordinating the collection of data
for a Check-list which should be published early in the 1970’s.

LANARK. R. Mackechnie.
FLORA: see under 75.

PEEBLES. D. J. McCosh, 13 Cottesmore Gardens, London, W.8. J/ndex: home.
Vouchers: E.

PRESENT WORK: Mr McCosh is collecting data ona 5 x 5 km basis and intends to
publish a Check-list.

SELKIRK. &

ROXBURGH. Dr R. W. M. Corner, 36 Wordsworth Street, Penrith, Cumberland.
Index and Vouchers: home.

PRESENT WORK: Dr Corner has begun to collect data for a Flora.

BERWICK. A. G. Long. Index: home. Vouchers: Hancock Museum, Newcastle-
upon-Tyne, NE2 4PT.

PRESENT WORK: Mr Long is co-ordinating the collection of data for a Flora which
will include Bryophytes.

HADDINGTON, 83, EDINBURGH. &

LINLITHGOW. Miss E. P. Beattie. Index: home. Vouchers: E.

FLORA: A Field Club Flora of the Lothians. 1. H. Martin. 1934. 25p. Miss N.
Henderson, 11 St Fillans Terrace, Edinburgh, 10.

PRESENT WORK: Miss Beattie is collecting data for a new Flora of the Lothians
ona5 x 5 km basis.

FIFE. G.H. Ballantyne. Index: home. Vouchers: E.

FLORA: A List of the Flowering Plants and Ferns recorded from Fife and Kinross.
W. Young. 1936. 50p. A. D. Lilly, Charenton, Cliff Road, Hythe, Kent. Checklist
of Flowering Plants of Kirkcaldy and District. G. H. Ballantyne. 1970. 424p.
PRESENT WORK: Mr Ballantyne is collecting data foranew Floraona5S x Skmbasis.

STIRLING. B. W. Ribbons.
FLORA: The Flora of the Clyde Area (see under 75) covers a small part of the west

of the county.
WEST PERTH., 88, MID PERTH. &
EAST PERTH. A. W. Robson. Jndex and Vouchers: City Art Gallery and Museum,

Perth. (Enquiries through Mr Robson.)
PRESENT WORK: Mr Robson is co-ordinating the collection of data for a Flora ona

5 x 5 km basis.

90,

91,

92;
93,

94,
95,

96,

96b

97,

98,

99,

100,

101,

102,

103,

THE VICE-COUNTY GUIDE 443

FORFAR. Miss U. K. Duncan. Jndex and Vouchers: home.
PRESENT WORK: no decision has yet been made as to the preparation of a Flora
based on an MS. produced many years ago by Mr and Mrs R. H. Corstorphine.

KINCARDINE. Mrs A. H. Sommerville. Index and Vouchers: ABD.
PRESENT WORK: Mrs Sommerville is collecting data with a view to publishing a
Flora.

SOUTH ABERDEEN. &
NORTH ABERDEEN. Professor C. H.Gimingham. Vouchers: ABD.

BANFF. Miss M. McCallum Webster. Vouchers: ABD and E.

ELGIN. Miss M. McCallum Webster. Index: home. Vouchers: ABD and E.

FLORA: A Check List of the Flora of the Culbin State Forest. M. McCallum Web-
ster. 1968. 30p. Miss M. McCallum Webster, Rose Cottage, Dyke, By Forres,
Moray.

PRESENT WORK: Miss McCallum Webster is collecting data for a Flora (with Nairn).

EASTERNESS. Position vacant. Index: Miss M. McCallum Webster, home.
Vouchers: E.

PRESENT WORK: the Committee for the Study of the Scottish Flora is co-ordinating
the collection of data for a Flora of Easterness and Westerness, excluding the por-
tion in Argyllshire, on a 5 x 5 km basis. Miss McCallum Webster has already
covered most of Easterness in detail.

NAIRN. Miss M. McCallum Webster. /ndex: home. Vouchers: ABD and E.
PRESENT WORK: see under 95.

WESTERNESS. A. A. Slack.
PRESENT WORK: see under 96 and 98.

MAIN ARGYLL. Mrs E. J. Montgomery.
PRESENT WORK: Prof. MacLeay has been collecting data with the intention of pub-
lishing a Flora of the whole of the political county of Argyllshire.

DUNBARTON. A. McG. Stirling. Index: home. Vouchers: E.

FLORA: Flora of Helensburgh. R. R. Mill. 1967. 324p. Macneur & Bryden Ltd,
Helensburgh. See also under 75.

PRESENT WORK: Mr Stirling is collecting data for a Flora ona5 x 5 km basis.

CLYDE ISLES. Mrs A. H. Sommerville. Index and Vouchers: ABD.

FLORA: see under 75.

PRESENT WORK: Mrs Sommerville is collecting data with a view to publishing a
Flora.

KINTYRE. A. G. Kenneth. Vouchers: CGE and E.

FLORA: ‘Flora of Danna’. A. G. Kenneth. 1964. In Trans. Proc. bot. Soc. Edinb.,
39 (5). £1.00. A. D. Lilly, Charenton, Cliff Road, Hythe, Kent.

PRESENT WORK: Mr Kenneth, in collaboration with Miss M. H. Cunningham, is
preparing a Check-list. See also under 98.

SOUTH EBUDES. Miss C. W. Muirhead. Index and Vouchers: E.

FLORAS: ‘Flora of Islay and Jura.’ J. K. Morton. 1959. Supplement Proc. bot. Soc.
Br. Isl., 3. 35p. E. W. Classey Ltd, 353 Hanworth Road, Hampton, Middlesex. “The
Flora of Easdale and the Garvellochs.’ C. W. Muirhead. 1962. In Trans. Proc. bot.
Soc. Edinb., 39 (3). £1. A. D. Lilly, Charenton, Cliff Road, Hythe, Kent.

PRESENT WORK: Miss Muirhead is preparing a Flora of Colonsay and Oronsay
which will include Bryophytes and Marine Algae. See also under 98.

MID EBUDES. Mull and adjacent small islands: J. F. M. Cannon. Index and
Vouchers: BM.
Coll and Tiree: Miss C. W. Muirhead. Vouchers: E.

444 F. H. PERRING

FLORAS: ‘Flora of the Isles of Coll, Tiree and Gunna.’ J. W. Heslop Harrison et alia.
1941. In Proc. Univ. Durham Phil. Soc., 10 (4). 25p. UNPS.

PRESENT WORK: the Dept. of Botany, British Museum (Natural History) is collect-
ing data for a Flora of Mull which will include all groups of plants. See also under 98.

104, NORTH EBUDES (except Rhum): Mrs C. W. Murray. Index and Vouchers: home.
Rhum: Dr W. J. Eggeling.
FLORAS: ‘List of the Plants of Rhum.’ W. J. Eggeling. 1965. In Trans. Proc. bot.
Soc. Edinb., 40 (1). £1-00. A. D. Lilly, Charenton, Cliff Road, Hythe, Kent. ‘The
natural history of the Isle of Raasay and of the adjoining islands of South Rona,
Scalpay, Fladday and Longay.’ J. W. Heslop Harrison. 1937. In Proc. Univ.
Durham Phil. Soc., 9 (5). 25p. UNPS. ‘The Flora of the Islands of Rhum, Eigg,
Canna, Sanday, Muck, Eilean nan Each, Hyskeir, Soay and Pabbay.’ J. W. Heslop
Harrison. 1939. In Proc. Univ. Durham Phil. Soc., 10 (2). 25p. UNPS.
PRESENT WORK: Mrs Murray is collecting data for a Flora of Skye and Raasay ona
10 x 10 km basis.

105, weEsT Ross. D. M. Henderson. Vouchers: E.
FLORA: ‘Observations on the flora around Reiff, Rhu More, Wester Ross, 1953.’
D. Morgan. 1956. In Proc. Univ. Durham Phil. Soc., 12 (13). 124p. UNPS.

106, EAST ROSS. Miss U. K. Duncan. Jndex and Vouchers: home and E.
PRESENT WORK: Miss Duncan is collecting data on a 10 x 10 km basis and hopes
to publish a Flora by the end of 1972.

107, EAST SUTHERLAND &

108, WEST SUTHERLAND. J. Anthony, 120 Trinity Road, Edinburgh, 5. Index: home.
Vouchers: E.
PRESENT WORK: Mr Anthony has been collecting data for a Flora for over 10 years
ona1l0O x 10kmand parish basis. It should be published in 1971.

109, CAITHNESS. J. M. Gunn.

110, OUTER HEBRIDES. Miss M. S. Campbell. Index: home. Vouchers: BM.
FLORAS: The Flora of Uig. M.S. Campbell. 1945. 55p. T. Buncle & Co. Ltd,
Market Place, Arbroath. The following from Proc. Univ. Durham Phil. Soc. are
available at UNPS: ‘Noteworthy plants from North Uist, Baleshare, Monach
Islands, Harris, Taransay, Mingulay and Berneray.’” W. A. Clark. 1939. 10 (2).
25p. ‘Noteworthy plants from Great and Little Bernera (Lewis), Pabbay and
Bernara (Harris), and the Uig District of Lewis.’ W. A. Clark & J. Heslop Harrison.
1940. 10 (3). 25p. ‘A preliminary Flora of the Outer Hebrides.’ J. W. Heslop Harri-
son. 1941. 10 (4). 25p. ‘The vascular plants of Stuley Island, the Isles of Grimsay and
Ronay.’ J. W. & J. Heslop Harrison. 1950. 10 (7). 25p. “Observations on the Flora
of the Isle of Lewis, Isle of Harris and the Shiant Isles in 1952.’ J. W. Heslop Harri-
son. 1951. 11 (6). 124p. ‘Botanical investigations in the Isles of Lewis, Harris,
Taransay, Coppay and Shillay in 1953.’ J. W. Heslop Harrison. 1954. 11 (12). 124p.
‘Botanizing in the Outer Hebrides in 1955 and 1956.’ J. W. Heslop Harrison. 1956.
12 (14). 1254p.
PRESENT WORK: Miss Campbell is collecting data for a Flora.

111, ORKNEY. Miss E. R. Bullard. Index: home. Vouchers: E.
FLORAS: a Check-list prepared by Miss Bullard is being published in parts as a
Supplement to the Orkney Field Club Bulletin.
PRESENT WORK: Miss Bullard is collecting data for an Ecological Flora.

112, ZETLAND. W. Scott. Index: home. Vouchers: home, OXF and Zetland County
Museum, Lerwick.
FLORA: A Check-list of the Flowering Plants and Ferns of the Shetland Islands.
R. C. Palmer & W. Scott. 1969. 42p. W. Scott, Easterhoull, Scalloway.
PRESENT WORK: Mr Palmer and Mr Scott are collecting data for a new Flora ona
parish and 10 x 10 km basis which will be completed about 1975.

THE VICE-COUNTY GUIDE 445

IRELAND

H1, SOUTH KERRY &

H2, NORTH KERRY. Prof. D. A. Webb.
PRESENT WORK: Mr B. Goggin and Mr M. Long are collecting data for a supple-
ment to the Flora.

H3, WEST CORK, H4, MID CORK &
HS5 EAST cork. Miss M. P. S. Scannell. Vouchers: DBN.

H6, WATERFORD. DrI.K. Ferguson. Index: Royal Botanic Gardens, Kew. Vouchers:
BM, DBN or TCD.
FLORA: Notes on the Flora of Co. Waterford. I. K. Ferguson. 1968. Irish Nat. J.,
16: 176-177. 5p. I. K. Ferguson, Royal Botanic Gardens, Kew.

H7, SOUTH TIPPERARY. Prof. D. A. Webb.
H8, LIMERICK. Prof. D. A. Webb.

H9, CLARE. Prof. D. A. Webb. Vouchers: TCD.
PRESENT WORK: see under H16.

H10, NORTH TIPPERARY. R. McMullen. Index: St Columba’s College, Rathfarnham,
Dublin, 14. Vouchers: DBN.
PRESENT WORK: Mr McMullen is collecting data for a Flora on a 10 x 10 km basis.

H11, KILKENNY. Prof. D. A. Webb.

H12, wexrorD. Dr A. M. O’Sullivan. Index and Vouchers: Agricultural Institute,
Johnstown Castle, Wexford.
PRESENT WORK: Dr O’Sullivan is collecting data on the botanical composition of
the grasslands and heaths.

H13, CARLow. Miss E. M. Booth. Index: home. Vouchers: DBN.
PRESENT WORK: Miss Booth has collected data which would be the basis for a Check-
list of the flora. Dr A. M. O’Sullivan has carried out a survey of the grasslands,
heaths and bogs, which is to be published.

H14, Ler. Prof. D. A. Webb.

H15, SOUTH-EAST GALWAY. Dr M.E. Mitchell. Vouchers: Dept. of Botany, University
College, Galway.
PRESENT WORK: see under H16.

H16, west GALWAY. Prof. D. A. Webb. Index and Vouchers: TCD.
PRESENT WORK: the whole of H16, parts of H15 and H17, and the north of H9
will be covered by a Flora of the Galway Region being prepared by Prof. Webb for
publication about 1973.

H17, NORTH-EAST GALWAY. Dr M.E. Mitchell. Vouchers: Dept. of Botany, University
College, Galway.
PRESENT WORK: see under H16.

H18, oFFALY. Dr J. G. D. Lamb. Index: The Agricultural Institute, Horticulture &
Forestry Division, Kinsealy, Malahide Road, Dublin, 5.

H19, KiLpare. D. Doogue. Index: 12 Glasilawn Road, Dublin, 11. Vouchers: DBN.

H20, wickKLow. H. J. Hudson.
FLORA: J. P. Brunker. 1951. £1:35. Enquires about purchase to H. J. Hudson.

H21, pusBLIN. H. J. Hudson.
FLORA: A Supplement to Colgan’s Flora of the County Dublin. 1961. 374p. Govern-
ment Publications Sales Office, G.P.O. Arcade, Dublin.

446 F. H. PERRING

H22, MEATH. D. Synnott. Index and Vouchers: DBN.
PRESENT WORK: Mr Synnott, in collaboration with the Dublin Naturalists’ Field
Club, is collecting data for a Flora which it is hoped will be published in the 1970s.

H23, WEST MEATH &
H24, LONGFORD. Prof. D. A. Webb.

H25, RoSCOMMON. Dr M. E. Mitchell. Index and Vouchers: Dept. of Botany, Univer-
sity College, Galway.
PRESENT WORK: Dr Mitchell is co-ordinating the collection of data for a Flora.
H26, EASTMAYO &
H27, WEST MAYO. Position vacant.

H28, sLico &
H29, LEITRIM. Prof. D. A. Webb.

H30, CAVAN. R. C. Faris.

H31, LouTH. D. Synnott. Index and Vouchers: DBN.
FLORA: Wild Flowers of County Louth. D. Synnott. 1970. 10p. Clarks (Ireland)
Ltd, Shoemakers, Dundalk.

H32, MONAGHAN. Fr J. J. Moore. Index: DBN.
PRESENT WORK: Miss M. P. H. Kertland and Mr D. Synnott have begun collecting
data for a Flora.

H33, FERMANAGH. R.D. Meikle. Index and Vouchers: Royal Botanic Gardens, Kew.
PRESENT WORK: for over 20 years Mr Meikle has been collecting data for a Flora;
a MS. has already been prepared.

H34, EAST DONEGAL. Miss M. N. Hamilton.
H35, WEST DONEGAL. Prof. D. A. Webb.

H36, TYRONE. Miss M. P. H. Kertland. Vouchers: BEL.
FLORA: a MS. of work started by the Tyrone Field Club about 1920 was added to
and typed in 1956. It can be consulted at the Science Library, Queen’s University,
Belfast, 9.

H37, ARMAGH. Miss Dawson, The County Library, The Mall, Armagh. Index: Science
Library, Queen’s University, Belfast, 9. Vouchers: BEL.
FLORA: A preliminary report on the survey of County Armagh Flora; 1965-1967.
274p. The Librarian, Science Library, Queen’s University, Belfast, 9.
PRESENT WORK: the Botanical Section of the Belfast Naturalists’ Field Club is
organizing the collection of data for a Floraona5 x 5 km basis.

H38, Down. P. Hackney. Index and Vouchers: BEL.
FLORA: A Flora of the North-east of Ireland. Ed. 2. R. L. Praeger & W. R. Megaw.
1938. £1-10. The Hon. Secretary, Belfast Naturalists’ Field Club, Old Museum, 7
College Square North, Belfast, 1.
PRESENT WORK: the Botanical Section of the Belfast Naturalists’ Field Club is pre-
paring a supplement to the Flora as a first step towards a new edition in 1988.

H39, ANTRIM. Miss G. E. Drennan. Index: home. Vouchers: BEL.
FLORA and PRESENT WORK: see under H38.

H40, perry. C. R. Nodder, Coastguard Cottage, Ballintoy, Ballycastle, Co. Antrim.
FLORA: Preliminary Survey of the Magilligan Area. Route Field Club. 1967. S5Op.
J. Donaldson, Black Rock Cottage, Runkerry, Bushmills, Co. Antrim. See also
under H38.
PRESENT WORK: see under H38.

THE VICE-COUNTY GUIDE 447

PRIORITIES FOR WORK ON LOCAL (COUNTY) FLORAS

Using the above Guide as a basis, an attempt has been made to place each vice-county
into one of five arbitrary categories from the highest to the lowest priority for the publica-
tion of a new Flora or Supplement. The categories are as follows:

I
Il

Ill
IV
Vv

No Flora has ever been published.

The Flora published is either incomplete, lacking localities and/or some species,
or was published before 1850.

The last Flora (or Supplement) was published between 1850 and 1900.

The last Flora (or Supplement) was published between 1900 and 1950.

The last Flora (or Supplement) has been published since 1950.

A vice-county has been placed in a priority category one higher if its last Flora (or
Supplement) was deficient in some other way. The numbers of vice-counties in which
active work towards a Flora, Supplement or Check-list is in progress are shown in

bold.

I

II

Ul

IV

42, 43, 47, 79, 86, 96, 96b, 97, 98, 101, 106, 107, 108, H6, 7, 10, 11, 12, 13, 14,
18, 19, 22, 23, 24, 30, 31, 32, 36

53, 54, 76, 77, 78, 80, 81, 90, 91, 92, 99, 100, 103, 104, 109, H8, 9, 15, 16, 17, 25,
DO 28, 29, 33

15, 16, 18, 19, 21, 25, 26, 36, 39, 49, 50, 51, 52, 58, 66, 67, 68, 69b, 70, 72, 73,
74, 75, 87, 88, 89, 110, H3, 4, 5, 34, 35

S, 1, 2, 3, 4, 5, 6, 9, 10, 11, 12, 13, 14, 17, 23, 24, 31, 32, 33, 34, 37, 41, 55, 60,
61, 62, 63, 64, 65, 69, 82, 83, 84, 85, 93, 94, 95, 102, 105, 111, H1, 2, 37, 38,
39, 40

1b, 7, 8, 20, 22, 27, 28, 29, 30, 35, 38, 40, 44, 45, 46, 48, 56, 57, 59, 71, 112, H20,
jell

Irish Naturalists’ Journal

A MAGAZINE OF IRISH NATURAL HISTORY

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LOCAL FLORAS

Edited by P. J. Wanstall

The purpose of this volume is to make available to a wider public the papers
read at the Conference held by the Botanical Society of the British Isles in 1961.
The contributors include D. E. Allen, Dr H. J. M. Bowen, Dr J. G. Dony,
J. S. L. Gilmour, Prof J. G. Hawkes, D. H. Kent, J. E. Lousley, D. McClintock,
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FLORA OF A CHANGING
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The Report of the 1969 Conference which reviewed factors responsible for
changes in the British flora, particularly since the end of the Second World War,
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Topics covered included the effects on the flora of climatic changes, atmos-
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Edited by E. Milne-Redhead

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INDEX
Compiled by R. K. Brummitt

Abbreviated Titles of Biological Journals
(compiled by P. C. Williams), review-
ed by R. K. Brummitt, 63-64

Abutilon theophrasti (in Middx), 187

Acaena inermis (in Mull), 149

Aceras anthropophorum (in Belgium—N.
France), 313

Adenolinum, 208; tommasinii, 212

Adiantum (in Guernsey), 335

Aerodynamics of winged seeds, 334

Aethusa cynapium Gn Mull), 150

Agropyron pungens xX repens (on R.
Stour), 185

Agrostemma githago (in Mull), 148

Agrostis semiverticillata (in Middx), 187

Alchemilla alpina Gn Mull), 146; vestita
(on Perthshire shingle beds), 189

Alice Holt (Field Meetings 1969), 186

Alien Euphorbias in the British Isles (by
A. R. Smith), 94

Allen, D. E—Conference Report: Hedges
and Local History, 425-426

Allen, D. E.—Dactylorhiza fuchsii subsp.
okellyi (Druce) So6—behaviour and
characters in the Isle of Man, 401-
402

Allen, D. E.—Prunella vulgaris L.—colour
variants apparently introduced, 399-

Allen, D. E.—The Victorian Fern Craze, a
history of pteridomania, reviewed by
J. W. Dyce, 314-315

Allen, D. E.—(Goint author with J. G.
Dony)—Report of the Council for
1968, 75-78

Allium neapolitanum (in Guernsey), 429;
paradoxum (Gloucestershire popula-
tions), 379-384 ; reseum (in Guernsey),
429; vineale (Qn Mull), 146, 152

Alopecurus bulbosus (on R. Stour), 185

Amaranthus blitoides (in Surrey), 187;
caudatus (in Surrey), 187; deflexus (in
Guernsey), 428; hybridus x _ retro-
flexus (in Middx), 187; standleyanus
Gn Middx), 187

Ambrosia artemisiifolia (in Middx), 187

Amelanchier alnifolia, 157; arborea, 157,
159, 161; asiatica, 157; canadensis,

156, 157, 159-162; confusa, 155-157,
159; florida, 160; grandiflora, 155,
156; humilis, 155; intermedia, 156,
159 lacvisee ISO ST. w t59=1oo-
lamarckii, 155-162; lamarckii var.
scharnkeana, 157; ovalis, 155, 156;
spicata, 155-157; stolonifera, 155

Ammocalamagrostis baltica (in Norfolk),
185; baltica (in Sutherland), 193

Anacamptis pyramidalis (in Tiree), 194;
pyramidalis (pollination of), 337

Anemone nemorosa ‘Vestal’ in Guernsey),
183-184

Angus, A.—Monotropa hypopitys L.—
new to Fife (v.c. 86), 163

Annual General Meeting, May 3rd, 1969,
81-82; May 9th, 1970, 329-331

Anogramma leptophylla (from Guernsey),
90

Anthyllis vulneraria (on tip in S. Lancs),
301; vulneraria subsp. corbieri (in
Guernsey), 429

Aosta (Flora reviewed), 417

Apera spica-venti (in Surrey), 187

Aphanes arvensis (in Guernsey), 428.

Arabis hirsuta Gn Mull), 148; hirsuta (on
Raasay), 190

Arenaria balearica (on Raasay), 190;
ciliata (seed characters), 333; lepto-
clados, 333; norvegica (seed charac-
ters), 333; norvegica subsp. norvegica
(in Shetland), 193; serpyllifolia, 333

Argyll (Field Meetings 1970), 432

Armeria maritima, 273-274

Arum maculatum (in Mull), 152

Arundinaria anceps (in Guernsey), 184

Asimov, 1.—Photosynthesis, reviewed by
M. C. F. Proctor, 322

Asparagus officinalis subsp. prostratus (in
Guernsey), 429

Asplenium septentrionale (in Carmarthen),
314: trichomanes (in Shetland), 192;
viride (on Iona), 147

x Asplenophyllitis microdon, 184; micro-
don (in Guernsey), 89-90

Astrantia major (in Montgom.), 187

Athyrium flexile, 68

Atriplex hortensis (in Guernsey), 428

451

452

Australian orchids, 94

Avena fatua, 274

Balance sheet, 31 December 1968, 79; 31
December 1969, 328

Ball, P. W. & Brown, K. G.—A bio-
systematic and ecological study of
Salicornia in the Dee estuary, 27-40

Ballantyne, G. H.—Salix and Epilobium
species and hybrids from lowland
Fife (v.c. 85), 332-333

Balquhidder (Field Meetings 1969), 191

Bangerter, E. B.—review of The Oxford
Book of Wild Flowers. Pocket Edition,
412-413.
Bangerter, E. B. & Ferguson, L. F.—
Some often confused plants, 333
Bangerter, E. B.—(joint author with J.
F. M. Cannon)—Plant records from
Mull and the adjacent small islands,
2, 145-153

Barbarea intermedia (in Mull), 147

Barling, D. M.—Studies on Gloucester-
shire populations of Allium para-
doxum (Bieb.) G. Don, 379-384

Barratt, T. B.—Fungi of Northampton-
shire woodlands, 84

Barton Hills—Field Meetings 1970, 427

Bate-Smith, E. C.—review of J. B.
Harborne, ed., Phytochemical phylo-
geny, 413

Beattie, E. P.—Committee for the Study of
the Scottish Flora, fourteenth annual
report to 31 December 1968, 83-84.

Beattie, E. P.—Field Meetings 1969:
Pease Dean, Berwickshire, 8 June,
189

Beattie, E. P.—Field Meetings 1969:
Skinflats, Stirlingshire, 31 August,
194

Beattie, E. P.—Field Meetings 1970:
West Lothian mosses, 11th July, 432—
433

Bedfordshire (Field Meetings 1970), 427

Bell, F. G.—Stereoscan studies on seed
morphology, 333-334

Belvoir (Field Meetings 1969), 185

Benoit, P. M.—Festuca altissima in
Merioneth, 84

Benoit, P. M.—Veronica hederifolia L.
sensu lato—(in Wales,) 399

Biosystematic and ecological study of
Salicornia in the Dee estuary (by
P. W. Ball & K. G. Brown), 27-40

Biscutella neustriaca (in Belgium—N.
France), 313

Blackburn, K. B.—(obituary by D. H.
Valentine), 69-70

Book reviews, 63-68, 168-180, 313-322,
412-422

Botanical keys generated by computer (by
R. J. Pankhurst), 357-368

INDEX TO WATSONIA VOLUME 8

Botrychium lunaria (new to Handa Is.),
193

Bottle, R. T. & Wyatt, H. V.—The Use of
Biological Literature, reviewed by
P. I. Edwards, 64-65

Bourg St Pierre (Field Meetings 1970),
426-427

Bowen, H. J. M.—review of J. B.
Harborne & T. Swain, eds., Perspec-
tives in Phytochemistry, 168

Boyle, P. J—Chromosomes of Spartina
pectinata Link in Ireland, 85

Brambles (six new species), 369-377

Brassica cheiranthos, 293; napus (in Mull),
146

Briggs, D.—review of O. T. Solbrig,
Principles and Methods of Plant
Biosystematics, 414

Briggs, D. & Walters, M.—Plant Variation
and Evolution, reviewed by D.
Wilkins, 179-180

Bromus carinatus (in Middx), 187; inermis,

0

Bromus pumpelianus—an introduced
species at Walthamstow, S. Essex,
v.c. 18 (by A. Melderis), 300

Bromus sterilis (in Mull), 146; thominii Gn
Mull), 153

Brookes, B. S.—Field Meetings 1969:
Handa Island and Durness, Suther-
land, 2-3 August, 193-194

Brown, K. G.—(joint author with P.
W. Ball)—A biosystematic and ecol-
ogical study of Salicornia in the
Dee estuary, 27-40

Brummitt, R. K.—review of P. C.
Williams, Abbreviated titles of Biologi-
cal Journals, 63—64

Brummitt, R. K. & Ferguson, I. K.—
Index to European Taxonomic Litera-
ture for 1966 and 1967, reviewed by
A. Melderis, 417-418

B.S.B.JI. Abstracts, 1

B.S.B.I. Membership (by Mrs J. G.

Dony), 86

B.S.B.1. Symphytum survey (by F. H.
Perring), 91

Bussey, Mrs Winifred—(watercolour

drawings by), 334
Bute (Field Meetings 1970), 431-432
Byrne Hill, Ayrshire (Field Meetings
1969), 189
Cakile maritima (on tip in S. Lancs), 301
Caladenia (in Australia), 94
Calamagrostis canescens (in Norfolk),
185; epigejos (in Norfolk), 185
Calystegia soldanella (in Carmarthen-
shire), 430
Camddwr Valley, Cardiganshire (Field
Meetings 1969), 188
Camelina sativa (in Middx), 186

INDEX TO WATSONIA VOLUME 8

Campanula latifolia (in Mull), 151;
rotundifolia, 83

Cannabis sativa (in Middx), 186

Cannon, J. F. M.—review of C. Jeffrey,
An Introduction to Plant Taxonomy,
168-169

Cannon, J. F. M.—review of C. T. Prime
& R. J. Peacock, Trees and Shrubs—
Their Identification in Summer or
Winter, ed. 6, 414-415

Cannon, J. F. M. & Bangerter, E. B.—
Plant records from Mull and the
adjacent small islands, 2, 145-153

Cardamine flexuosa, 45; hirsuta, 45;
impatiens, 45; matthiola, 45

Cardamine—pollen (by R. P. Ellis &

B. M. G. Jones), 45

ras Ai pratensis, 45; raphanifolia, 45

Cardamine raphanifolia in East Glouces-
tershire (by Mrs R. Dudley-Smith),
86

Cardamine resedifolia, 45

Cardaminopsis petraea (in Mull), 146

Cardiff, National Museum of Wales
(Field Meetings 1970), 431

Carduus acanthoides (in Mull), 146, 152

Carex appropinquata (in Belgium—N.
France), 313

Carex aquatilis in the central and eastern
Highlands (by J. S. Faulkner), 87

Carex bigelowii, 87; bigelowii (in Mull),
146; diandra (in Belgium—N.
France), 313; flava (in Ireland), 91;
laevigata (in Guernsey), 428 ; laevigata
(on Raasay), 190; Jasiocarpa (in
Belgium—N. France), 313; Jepido-
carpa (hybridises with C. flava), 91;
limosa (on Raasay), 190, 191;
maritima (in Shetland), 192; nigra,
87; paucifiora (on Raasay), 190-191;
vesicaria (in Mull), 153

Carmarthenshire (Field Meetings 1970),
430-431; (plant list reviewed), 313-
314

Carpobrotus acinaciformis auct. (in
Guernsey), 428

Carthamus tinctorius (in Middx), 186

Catabrosa aquatica (in Shetland), 192;
aquatica (in Sutherland), 193

Cellular Differentiation in Plants and other
Essays (by C. W. Wardlaw), reviewed
by F. Cusick, 415-416

Centaurea diluta (in (Guernsey), 429;
diluta (in Middx), 186

Centranthus ruber (in Mull), 146

Cerastium arcticum subsp. edmondstonii,

3

Ceratophyllum submersum var. haynaldi-
anum (in Cambridgeshire), 93; sub-
mersum var. submersum (in Cam-
bridgeshire), 93

Px

453

Chandler, J. H. & Gilbert, J. L._—Yellow-
berried Crataegus monogyna, 85
Channel Isles (Geranium submolle in ),47
Charnwood Forest (Field Meetings 1969),

185
Chater, A. O.—review of R. F. May, A
list of the Flowering Plants and Ferns
of Carmarthenshire, 313-314
Chenopodium x bontei, 336; carinatum,
335; cristatum, 335-336; ficifolium (in
Middx), 187; glaucum (in Surrey),
187; opulifolium (in Middx), 187;
probstii (in Middx), 186
Chenopodium pumilio—a Kentish obit-
uary? (by D. McClintock), 335-336
Cherleria sedoides (in Mull), 146
Children’s Centre Club, British Museum
(Natural History)—A study of the
aerodynamics of winged seeds, 334
Chromosomes of Spartina pectinata Link
in Ireland (by P. J. Boyle), 85
Chrysanthemum maximum (in Guernsey),
428
Chrysanthemum uliginosum (in Surrey),
187
Cicendia filiformis (in Pembs.), 188
Cicerbita macrophylla (in Mull), 152
Citrullus lanatus (in Middx), 187
Cladium mariscus (on Raasay), 190
Clapham, A. R., ed.—Flora of Derbyshire,
reviewed by D. L. Hawksworth, 171-—
174
Clematis balearica (in Guernsey), 184;
vitalba (in Mull), 147
Clement, E. J. & Ryves, T. B.—Field
Meetings 1969: alien hunt about
London, 4-5 October, 186-187
Cochlearia arctica, 395; groenlandica, 395;
oblongifolia, 395; officinalis, 395
Cochlearia scotica Druce—does it exist in
northern Scotland? (by J. J. B. Gill),
395-396
Coeloglossum viride (on Raasay), 190
Cole, A. J.,—Numerical Taxonomy, re-
viewed by D. B. Williams, 422
Coles, S. M.—The Ranunculus acris L.
complex in Europe, 237-261
Committee for the Study of the Scottish
Flora, fourteenth Annual Report to
31 December 1968 (by E. P. Beattie),
83-84; fifteenth Annual Report to
31 December 1969 (by B. W. Rib-
bons), 331-332; (Report of Council
1968), 77-78; (Report of Council
1969), 326
Computer-generated keys (by R. J.
Pankhurst), 336-337; 357-368
Conference Report: Hedges and Local
History (by D. E. Allen), 425-426
Conservation Committee (Report 1968),
77; (Report 1969), 325

454

Conservation in Britain, 195-203

Conservation of chalk grassland in Bed-
fordshire (Field Meetings 1970), 427

Corallorhiza (in N. Lancs.), 184

Coriandrum sativum (in Middx), 187

Coronilla varia (in Stirlingshire), 194

Cors Goch, Anglesey (Field Meetings
1969), 189

Corynephorus canescens (in Norfolk), 185

Corynephorus canescens (L.) Beauv.—in
West Suffolk (by P. J. O. Trist), 402

Cotoneaster bullata (an Miull), 149;
frigidus (an Mull), 149; microphyllus
(in Mull), 149; simonsii (Gn Mull),
149

Coverlon (for flower mounting), 337

Cowslips, 289-291

Crabbe, J. A., Jermy, A. C. & Walker S.—
The distribution of Dryopteris assimi-
lis S. Walker in Britain, 3-13

Crassula aquatica (new to Scotland), 331

Crassula aquatica (L.) Schonl.—new to
Scotland (by A. J. Souter), 294

Crataegus monogyna (yellow-berried), 85;
racemosa, 156

Crisp, P. & Jones, B. M. G.—Introgressive
hybridisation between British annual
Senecio species, 85

Crisp, P. & Jones, B. M. G.—Senecio
squalidus L., S. vulgaris L. and S.
cambrensis Rosser, 47-48

Cronquist, A.—The Evolution and Classi-
fication of Flowering Plants, reviewed
by N. K. B. Robson, 177-179

Cucumis melo (in Middx), 186

Cucubita pepo (in Middx), 186

Cusick, F.—review of C. W. Wardlaw,
Cellular Differentiation in Plants and
other Essays, 415-416

Cutler, D. F.—review of E. G. Cutter,
Plant Anatomy: Experiment and
Interpretation. Part I. Cells and
Tissues, 169

Cutter, E. G.—Plant Anatomy: Experi-
ment and Interpretation. Part I.
Cells and Tissues, reviewed by D. F.
Cutler, 169

Cypripedium calceolus (conservation of),
195, 201

Cyrtomium falcatum (in Guernsey), 183

Cystopteris fragilis (in Cambridgeshire),
93; fragilis (on Foula, Zetland), 301

Cytisus scoparius (with particular reference
to prostrate forms), 345-356

Cytisus scoparius subsp. maritimus, 345-—
356

Cytotaxonomic study of Lamiastrum
galeobdolon (L.) Ehrend. & Polats-
chek in Britain (by S. Wegmiiller),
277-288

Dactylis maritima, 164

INDEX TO WATSONIA VOLUME 8

Dactylorhiza maculata subsp. ericetorum,

Dactylorhiza fuchsii subsp. okellyi (Druce)
Soo—behaviour and characters in
the Isle of Man (by D. E. Allen),
401-402

Danthonia, 298

Davis, P. H. & Cullen, J.—The Identifica-
tion of Flowering Plant Families,
reviewed by J. H. Thomas, 68

Davis, T. A. W.—Field Meetings 1969:
Pembrokeshire, 16-17 August, 188

Davis, T. A. W.—Morphological develop-
ment of Spartina ‘X’, 85-86

Derbyshire (review of Flora), 171-174

Development and Rules Committee
(Report 1968), 76; (Report 1969), 325

Dicentra formosa (in Mull), 147

Digitaria ciliaris (in Middx), 187; ischae-
mum (in Surrey), 187; sanguinalis (in
Middx), 186-7

Diphasium alpinum (not in Mull), 147

Diplotaxis muralis (on tip in S. Lancs.),
301

Distribution of Dryopteris assimilis S.
Walker in Britain (by J. A. Crabbe,
A. C. Jermy & S. Walker), 3-13

Distribution of Juniperus communis n
southern England (by L. K. Ward),
94

Dony, J. G.—The journals of the Botani-
cal Society of the British Isles, 1-2

Dony, J. G. & Allen, D. E.—Report of
the Council for 1968, 75-78

Dony, J. G. & Wells, T. C. E.—Field
Meetings 1970: Conservation of
chalk grassland in Bedfordshire, 427

Dony, Mrs J. G.—B.S.B.I. membership,
86

Dore, W. G.—Sieglingia decumbens (L.)
Bernh.—pulvini of palea, 297-299

Draba incana (on Raasay), 190; incana (on
Tiree), 194

Drosera intermedia (in Pembs.), 188

Dryas octopetala (in Sutherland), 194

Dryopteris aemula, 4, 12; assimilis, 3-13;
carthusiana, 3-7, 12; carthusiana (in
Norfolk), 185; cristata (in Norfolk),
185; dilatata complex, 3-13; dilatata
var. collina, 7; dilatata var. pseudo-
spinulosa, 7; lanceolatocristata, 3; x
pseudoabbreviata (on Mull), 334

Dudley-Smith, Mrs R.—Cardamine raph-
anifolia in East Gloucestershire, 86

Duncan, U. K.—Lathyrus maritimus
Bigel. subsp. acutifolius (Bab.) Peder-
sen in eastern Scotland, 87

Duncan, U. K.—Senecio x _ ostenfeldii
Druce in Scotland, 87

Duncan, U. K.—Ten species of Taraxacum
from Scotland, 334

INDEX TO WATSONIA VOLUME 8

Durness, Sutherland (Field Meetings
1969), 193-194

Dyce, J. W.—review of D. E., Allen, The
Victorian Fern Craze, a history of
pteridomania, 314-315

Echinophora spinosa L.—a member of our
flora (by E. J. Salisbury), 397— 399

Edees, E. S.—Obituary: Beverley Alan
Miles, 424

Edmundson, T.—Persistence of Silene
maritima and other plants onan inland
tip in South Lancashire, v.c. 59, 301

Edwards, P. I.—review of R. T. Bottle &
H. V. Wyatt, The Use of Biological
Literature, 64-65

Elatine hexandra, 45—46; hydropiper, 83

Elatine hydropiper—new to Scotland (by
E. T. Idle, J. Mitchell & A. McG.
Stirling), 45-46

Eleocharis acicularis (in Shetland), 192

Eleogiton fluitans (in Shetland), 193

Ellis, R. P. & Jones, B. M. G.—Cardamine

—pollen, 45
Epilobium (computer-generated keys to
species), 363-366; species and

hybrids, 331; species and hybrids in
v.c. 85, 332-333

Epilobium alsinifolium (in Mull), 146;
nerterioides (in Carmarthenshire),
314; nerterioides (new to Guernsey),

Epipactis atrorubens (on Raasay), 190;
atrorubens (in Sutherland), 194;
dunense, 430; palustris, 430

Equisetum x litorale (in Anglesey), 189;
palustre var. polystachyum, 68; tel-
mateia (in Guernsey), 335; variegatum
(in N. Lancs.), 184

Erica tetralix (on Norfolk dunes), 185

Eriocaulon aquaticum (Hill) Druce—on
Ardnamurchan (Westerness, v.c. 97)
(by I. K. & L. F. Ferguson and G.
Halliday), 400-401

Eriophorum _ gracile
France), 313

Erodium cicutarium subsp. cicutarium (on
tip in S. Lancs.), 301; glutinosum (not
in Guernsey), 428; reichardii, 295

Eryngium campestre (in Guernsey), 89

Esau, K.—Viruses in Plant Hosts, re-
viewed by S. M. Robb, 420-421

Euphorbia ceratocarpa, 94; corallioides,
94; esula subsp. esula, 94; esula
subsp. tommasiniana x subsp. esula,
94; oblongata, 94; x pseudovirgata,
94; uralensis, 94; virgata, 94

Euphrasia arctica, 42; borealis, 41-44;
borealis var. speciosa, 41, 44; borealis
var. zetlandica, 41, 44

Euphrasia brevipila and E. borealis in the
British Isles (by P. F. Yeo), 41-44

Gn Belgium—N.

455

Euphrasia confusa, 44; nemorosa, 42;
rostkoviana f. borealis, 43; stricta, 42,
43

Evolution and Classification of Flowering
Plants (by A. Cronquist), reviewed
by N. K. B. Robson, 177-179

Evolution and Phylogeny of Flowering
Plants (by J. Hutchinson), reviewed
by N. K. B. Robson, 421

Exaculum pusillum (in Guernsey), 89

Exhibition Meeting, 1968, 84-95; 1969,
332-338

Exotic Amelanchier species naturalised in
Europe and their occurrence in Great
Britain (by F. G. Schroeder), 155-162

Fahn, A.—Plant Anatomy, reviewed by
V. H. Heywood, 317-318

Farnpflanzen Zentraleuropas (by K. & H.
Rasbach & O. Wilmanns), reviewed
by F. H. Perring, 318

Faulkner, J. S.—Carex aquatilis in the
central and eastern Highlands, 87

Ferguson, I. K.—Annual General Meet-
ing 1970, 329-331

Ferguson, I. K.—Field Meetings, 1969:
Conference Excursion, 21st Septem-
ber, 186

Ferguson, I. K.—Some interesting Co.
Waterford plants, 334—335

Ferguson, I. K.—South-east Region
Meeting 1970, 331

Ferguson, I. K. & Ferguson, L. F.—
Eriocaulon aquaticum (Hill) Druce—
on Ardnamurchan (Westerness, v.c.
97), 400

Ferguson, I. K.—(joint author with R. K.
Brummitt)—Jndex to European Taxo-
nomic Literature for 1966 and 1967,
reviewed by A. Melderis, 417-418

Ferguson, I. K.—(joint author with E.
Milne-Redhead)—Report of the
Council for 1969, 323-327

Ferguson, L. F.—Naufraga balearica Con-
stance & Cannon—refound, 294-295

Ferguson, L. F.—(joint author with I.
K. Ferguson)—Eriocaulon aquaticum
(Hill) Druce—on Ardnamurchan
(Westerness, v.c. 97), 400

Ferguson, L. F.—(joint exhibitor with
E. B. Bangerter)—Some often con-
fused plants, 333

Festuca altissima in Merioneth (by P. M.
Benoit), 84

Festuca (distinction from Nardurus and
Vulpia), 333

Ficus carica (in Guernsey), 184

Field Meetings 1969, 183-194; 1970, 426—
433

Field Natural History—a Guide to Ecology
(by A. Leutscher), reviewed by E. W.
Groves, 169-170

456

Flatford Mill, Essex (Field Meetings
1969), 185

Flora Europaea, Volume 2, reviewed by F.
Markgraf, 66

Flora of Derbyshire (ed. A. R. Clapham),
reviewed by D. L. Hawksworth, 171-
174

Flora of Norfolk (by C. P. Petch & E. L.
Swann), reviewed by A. C. Jermy,
416-417

Flowering Plants, Origin and Dispersal (by
A. Takhtajan), reviewed by N. K. B.
Robson, 315-316

Flowers of Europe (by O. Polunin),
reviewed by R. D. Meikle, 321-322

Flora of Guernsey (exhibit by D.
McClintock), 89-90

Floral aberrations in Oxalis acetosella L.
in Epping Forest (by R. J. Giles), 88

Flore de la Belgique, du Nord de la France
et des régions voisines (ed. W.
Mullenders), reviewed by P. A. Stott,
S518}

Fort William, Inverness-shire (Field Meet-
ings 1970), 433

Foula, Zetland, 301

Frequencies of pin and thrum plants in a
wild population of the Cowslip,
Primula veris L. (by D. R. Lees), 289-
291

Friends of John Gerard (1545-1612),
Surgeon and Botanist (by R.
Jeffers), reviewed by J. E. Lousley,
65-66

Fumaria bastardii (in Mull), 146; purpurea
(in Guernsey), 428

Fungi of Northamptonshire woodlands
(by T. B. Barratt), 84

Gagea lutea (in Derbyshire), 173

Galeobdolon luteum (see Lamiastrum
galeobdolon), 277; montanum, 278

Galeopsis tetrahit (in Guernsey), 335

Galinsoga ciliata, 186, 187; parviflora, 186,
187

Galium palustre, 333; uliginosum, 333;
uliginosum (in Mull), 151; uliginosum
(in Pembs.), 188

Gardiner, J. C.—Teesdale Defence Com-
mittee Report, 329

Gardiner, J. C.—Treasurer’s Report, 1969
79-80, 1970 328

Gastrosophy, 320

Gaudinia (in Guernsey), 428

Gentianella amarella subsp. septentrionalis
(in Shetland), 192

Geranium retrorsum, 47; c.f. robertianum
(octoploid in Guernsey), 90; c.f.
submolle (in Guernsey), 89

Geranium submolle Steud. (or G. core-core
Steud.)—in the Channel Isles (by D.
McClintock), 47

INDEX TO WATSONIA VOLUME 8

Gerard, J. (1545-1612), 65-66

Gilbert, J. L.—Scabiosa columbaria show-
ing “‘bird’s nest”’ proliferation, 87-88

Gilbert, J. L.—QGoint exhibitor with J. H.
Chandler)—Yellow-berried Cratae-
gus monogyna, 85

Gilbert-Carter, H. (obituaries by H.
Godwin and T. G. Tutin), 71-74

Giles, R. J.—Floral aberrations in Oxalis
acetosella L. in Epping Forest, 88

Gill, J. J. B—Cochlearia scotica Druce—
does it exist in northern Scotland ?,
395-396

Gill, J. J. B. & Walker, S.—Studies on
Cytisus scoparius (L.) Link with
particular reference to the prostrate
forms, 345-356

Ginkgo (in Harrow), 88

Gladiolus byzantinus, 88; byzantinus (in
Guernsey), 429; communis, 88; illyri-
cus, 88; italicus, 88

Glamorgan (Field Meetings 1969), 187-—
188

Glendaruel, Argyll (Field Meetings 1970),
432

Gloucestershire (Allium paradoxum popu-
lations), 379-384

Glyceria borreri, 17; pseudodistans, 17

Globularia vulgaris (in Belgium—N.
France), 313

Glycine max (in Middx), 187

Gnaphalium uliginosum (in Tiree), 194

Godwin, H.—Obituary: Humphrey Gil-
bert-Carter (1884-1969), 71-72

Goulimis, C. N.—Wild Flowers of Greece,
reviewed by O. Polunin, 316-317

Grant, K. A. & Grant, V.—Humming
Birds and their Flowers, reviewed by
L. van der Pijl, 67

Grass: a Story of Frankenwald (by E.
Roux), reviewed by J. Lewis, 417

Gray, A. J.—Morecambe Bay: northern
and southern elements in the flora,
335

Greenwood, E. F.—North-West Region
Meeting 1969, 82

Groves, E. W.—Hippophae rhamnoides
L.—on the Isle of Mull, Argyllshire,
396

Groves, E. W.—review of A. Leutscher,
Field Natural History—a Guide to
Ecology, 169-170

Grubb, P. J.—review of M. Husova,
Synékologische Studie der Waldgesell-
schaften auf | Amphibolitgestein.
Vegetace CSSR A3, 170-171

Guernsey, 89-90; (Field Meetings 1969),
183-184; (Field Meetings 1970), 428-
429

Guernsey flora in
McClintock), 335

1969 (by D.

INDEX TO WATSONIA VOLUME 8

Guide to the Naming of Plants: with
special reference to Heathers (by D.
McClintock), reviewed by R. Lan-
caster, 319

Gymnadenia conopsea (in Raasay), 190;
conopsea var. densiflora (in E.
Suffolk), 430

Gymnocarpium dryopteris (in Shetland),
193; dryopteris (on Raasay), 190

Hall, P. C—Field Meetings 1969:
Midhurst, Sussex, 24-26 May, 184-
185

Halliday, G.—Eriocaulon aquaticum (Hill)
Druce—on Ardnamurchan (Wester-
ness, v.c. 97), 401

Halliday, G.—Field Meetings 1969:
Roanhead and Roudsea, North
Lancashire, 4 May, 184

Halliday, G. & Hyde, L. T.—Field

Meetings 1970: Westmorland, 429-
430

Hamilton, A. P.—A study of some western
Eurasian Gladioli, 88

Hammarbya paludosa (in Pembs), 188;
paludosa (in Tiree), 194

Handa Island, Sutherland (Field Meetings
1969), 193

Harborne, J. B., ed.,—Phytochemical
Phylogeny, reviewed by E. C. Bate-
Smith, 413

Harborne, J. B. & Swain, T., eds.,
Perspectives in Phytochemistry, re-
viewed by H. J. M. Bowen, 168

Harrison, S. G., ed.—Welsh Ferns, Club-
mosses, Quillworts and Horsetails (by
Fiydes cH. As & Wade, A: E.),
reviewed by W. A. Sledge, 67-68

Harrison S. G.—Field Meetings 1970:
National Museum of Wales, Cardiff,
431

Harrison, S. G., Masefield, G. B. &
Wallis, M.—The Oxford Book of Food
Plants, reviewed by E. Launert, 319-
320

Harrow County School for Boys—a tree
survey in the London Borough of
Harrow, 88

Hartman, F. K. & Jahn G., Waldgesell-
schaften des mitteleuropdischen Gebir-
gsraumes nordlich der Alpen, reviewed
by M. C. F. Proctor, 177

Hawksworth, D. L.—review of A. R.
Clapham, ed., Flora of Derbyshire,
171-174

Hawksworth, D. L.—Two additions to
the flora of Foula, Zetland, v.c. 112,
301

Hawkweeds of western Scotland, 97-120

Heathers (guide to naming of), 319

Hedges and Local History (by D. E.
Allen), 425-426

457

Heiser, C. B.—WNightshades. The Para-
doxical Plants, reviewed by D.
McClintock, 320-321

Helianthus annuus (in Mull), 152

Heracleum mantegazzianum (in Mull), 150

Heslop-Harrison, J. W. (1881-1967)
ena by J. A. Richardson), 181-

Hesperolinon, 232

Heterostyly in Linum, 208-209, 231; in
Primula veris, 289-291

Heywood, V. H., ed..— Modern Methods
in Plant Taxonomy, reviewed by
M. C. F. Proctor, 418-420

Heywood, V. H.—review of A. Fahn,
Plant Anatomy, 317-318

Hieracium aggregatum, 104, 105; alpinum,
101, 107, 111, 112, 114; ampliatum,
100-106, 108-112, 114; anfractiforme,
104, 106, 110, 111; anglicum, 101-
106, 108, 110-114; angustisquamum,
102, 109, 113; argenteum, 101-103,
107, 108, 111-114; attenuatifolium,
192; australius, 193; breadalbanense,
106, 107; britanniciforme, 101, 108;
caesiomurorum, 98, 102, 104, 106-111,
113; caledonicum, 98, 100-103, 106—
114, 192; calenduliflorum, 104; callis-
tophyllum, 104-106; camptopetalum,
111-113; carpathicum, 48; centri-
petale, 104, 111, 112; cheriense, 103;
chloranthum, 100-111, 114; clovense,
105; coniops, 103; cravoniense, 98, 99,
102, 105, 106, 108, 111; cuspidens,
104, 105; cymbifolium, 103, 110, 111;
dasythrix, 102-108, 111-114; dewarii,
104, 107; diaphanoides, 99, 107, 111;
diaphanum, 99; dicella, 101-103, 108,
114; dilectum, 192; dipteroides, 100,
104, 111; dissimile, 108; dovrense;
112, 113, 336; drummondii, 103, 108,
109; duriceps, 98, 100-102, 106-108,
110-114; ebudicum, 102, 106-108;
euprepes, 99, 101-105, 108, 109, 111,
112; eximium, 104, 105, 107, 111, 112,
114; exotericum complex, 114; floc-
culosum, 108; fratrum, 103; glanduli-
dens, 104, 107, 111-113; globosifio-
rum, 111-114; gothicoides, 109, 193;
gracilifolium, 104-106, 111; grandidens,
105, 107, 114; gratum, 193; hanburyi,
111; hastiforme, 105; hebridense, 102,
103, 108, 111; hethlandiae, 192;
holosericeum, 99, 101, 104, 107, 111,
113; iricum, 107, 109, 110, 112-114;
jovimontis, 101, 102, 113, 115;
langwellense, 103-105, 108, 110, 114;
lasiophyllum, 101; latobrigorum, 100,
101, 103, 106-108, 110, 111, 113, 114;
latobrigorum f. angustifolium, 106;
leyi, 98, 99; lingulatum, 104-106, 110-

458

112, 114; lintonianum, 104, 105;
lissolepium, 100, 101; magniceps, 49;
marginatum, 111, 112; maritimum,
98, 103, 106-108, 110; marshallii, 101,
104, 105, 110; memorabile, 104;
mirandum, 48; nigrisquamum, 104—
106, 111, 336; nitidum, 104, 111-113,
115; orimeles, 99, 107, 109, 110, 192;
oxyodus, 104-106, 111, 112; pauculi-
dens, 112; perpropinquum, 99, 101,
107, 109, 111, 115; petrocharis, 101-
104, 108; pictorum, 101, 102, 104, 105,
LOT PAAOS MOMSASS 13362 pill-
gerum, 104, 105, 107, 108, 110, 111;
pollinarioides, 111-113, 336; polli-
narium, 113; pilosella, 115 (see also
Pilosella); praetermissum, 112, 113;
praethulense, 192; pseudanglicoides,
104, 105, 112; pseudanglicum, 101,
103-106, 111, 114; pseudostenstroemii,
102; pugsleyi, 192; reticulatum, 105,
107, 108, 110, 114; rhomboides, 101,
1029110 %rivale, 107, 708, 1145 112:
rubiginosum, 99-113; sanguineum,
102; sarcophylloides, 102, 103, 108,
110-113; saxorum, 106, 109; schmidtii,
99, 100, 106; scoticum, 103, 109, 113,
192; senescens, 101, 104-106, 109,
111, 112, 114; shoolbredii, 101-105,
107-114, 336; sinuans, 104, 111;
solum, 107, 108, 113, 336; sommer-
feltii, 101, 105, 109, 114; sparsifolium,
99, 101, 104, 105, 112-114, 192, 193;
stewartii, 101; strictiforme, 101-105,
108, 110, 113, 114; strumosum, 429;
subcrocatum, 101, 103, 109, 114;
subhirtum, 99, 101-112, 114; sub-
planifolium, 99, 108, 109; subrude, 99,
100, 108, 112, 113; subtenue, 111-
113; subumbellatiforme, 98, 104, 114;
tenuifrons, 104, 105; triviale, 102;
uiginskyense, 102, 103, 114; uistense,
104, 107, 108, 111; wisticola, 102, 103,
108; umbellatum, 99, 100, 108, 109,
115; vagense, 99; variifolium, 113;
vennicontium, 102, 104-106, 108;
vinicaule, 192; vulgatum, 98, 101-104,
106-114; zetlandicum, 192

Hieracium—two Hieracia sect. Alpestria
from the British mainland (by J. N.
Mills & J. R. J. Mills), 48-49

Hippocrepis comosa (in Westmorland),
429

Hippophae rhamnoides L.—on the Isle of
Mull, Argyllshire (by E. W. Groves),
396

History of British Vegetation (by Winifred
Pennington), reviewed by J. R.
Matthews, 175-177

Holcus lanatus (in mixed population), 185;
mollis (in mixed population), 185

INDEX TO WATSONIA VOLUME 8

Husova, M.—Synékologische Studie der
Waldgesellschaften auf Amphibolit-
gestein. Vegetace CSSR A3, reviewed
by P. J. Grubb, 170-171

Humming Birds and their Flowers (by K. A.
Grant & V. Grant), reviewed by L.
van der Pijl, 67

Hutchinson, J.—Evolution and Phylogeny
of Flowering Plants, reviewed by
N. K. B. Robson, 421

Hybrids in Juncus section Genuini (by
CHA Stace)1338

Hyde, H A. & Wade, A. E.— Welsh Ferns,
Clubmosses, Quillworts and Horse-
tails, ed. 5 (by S. <G. *Haerisen):
reviewed by W. A. Sledge, 67-68

Hyde, L. T. (Goint author with G.
Halliday)—Field Meetings 1970:
Westmorland, 429-430

Hymenophyllum tunbrigense, 83; tunbri-
gense (on Raasay), 191; wilsonii, 83;
wilsonii (on Raasay), 190; wilsonii Gn
Shetland), 192

Hypericum calycinum (in Guernsey), 428;
calycinum (in Iona), 148

Hypericum canadense _.—in England (by
J. E. Lousley), 293

Hypericum canadense L.—in the British
Isles (by R. D. Meikle), 90

Hypericum montanum (in v.c. 85), 331;
undulatum (in Pembs.), 188

Identification of Flowering Plant Families
(by P: H. Davis’ & 2 32¢Guilen):
reviewed by J. H. Thomas, 68

Idle, E. T., Mitchell, J. & Stirling, A.
McG.—Elatine hydropiper—new to
Scotland, 45-46

Index Kewensis Plantarum Phaneroga-
marum, Supplementum XIV, reviewed
by N. K. B. Robson, 322

Index to European Taxonomic Literature
for 1966 and 1967 (by R. K. Brummitt
& I. K. Ferguson), reviewed by A.
Melderis, 417-418

Introduction to Plant Taxonomy (by C.
Jeffrey), reviewed by J. F. M. Cannon,
168-169

Introgressive hybridisation between
British annual Senecio species (exhibit
by P. Crisp & B. M. G. Jones), 85

Inula helenium (in Mull), 152; helenium
(on Raasay), 190

Inverness-shire (Field Meetings 1970),
433

Ipheion uniflorum (new to Guernsey),
3

Ireland (new vascular plants), 91

Tris pseudacorus (pollination of), 337

Irish Regional Committee (Report 1968),
78; (Report 1969), 326

Irish vice-counties (by D. A Webb), 51

INDEX TO WATSONIA VOLUME 8

“Is your plate camera really necessary ?”
(by M. C. F. Proctor), 92

Isle of Man (Dactylorhiza fuchsii subsp.
Okellyi in), 401-402

Isoetes histrix (from Guernsey),
lacustris (in Sutherland), 193

Jeffers, R. H.—The Friends of John
Gerard (1545-1612), Surgeon and
Botanist, reviewed by J. E. Lousley,
65-66

Jeffrey, C.—An Introduction to Plant
Taxonomy, reviewed by J. F. M.
Cannon, 168-169

Jermy, A. C.—review of C. P. Petch &
E. L. Swann, Flora of Norfolk, 416—
417

Jermy, A. C.—(joint author with J. A.
Crabbe and S. Walker)—The Distri-
bution of Dryopteris assimilis S.
Walker in Britain, 3-13

Jermy, A. C. & Martin, B. S.—A note
on the preparation of material for
the scanning electron microscope,
14-15

Jones, B. M. G.—Field Meetings 1969:
Flatford Mill, Essex, 28 June, 185

Jones, B. M. G.—(joint exhibitor with P.
Crisp)—Introgressive hybridisation
between British annual Senecio
species, 85

Jones, B. M. G.—(joint author with P.
Crisp)—Senecio squalidus L., S.
vulgaris L. and S. cambrensis Rosser,
47-48

Jones, B. M. G.—(joint author with R. P.
Ellis)—Cardamine—pollen, 45

Jones, B. M. G. & Newton, L. E.—The
status of Puccinellia pseudodistans
(Crép.) Jansen & Wachter in Great
Britain, 17—26

Jones, H.—(joint exhibitor with B. Batts
and C. A. Stace)—Nardurus, Vulpia
or Festuca, 333

Journals of the Botanical Society of the
British Isles (by J. G. Dony), 1-2

Juncus acutus (in Carmarthenshire), 314;
arcticus (hybridisation), 338; balticus
(hybridisation), 338; balticus (in
Sutherland), 194; balticus (in Tiree),
194; bulbosus, 89; bulbosus sensu lato
(in Pembs.), 188; compressus (in
Ireland), 91; conglomeratus x effusus
(in Pembs.), 188; conglomeratus
(hybridisation), 338; effusus x balti-
cus, 338; effusus x inflexus, 338;
filiformis (hybridisation), 338; in-
flexus ~x_ balticus, 338; jacquinii
(hybridisation), 338; kochii, 89;
triglumis (in Mull), 146

Junior Activities Sub-Committee (Report
1968), 76; (Report 1969), 324

90;

459

Juniperus communis (in Southern
England), 94

Kenneth, A. G. & Stirling, A. McG.—
Notes on the Hawkweeds (Hieracium
sensu lato) of western Scotland, 97-
120

Kent, D. H., 1

Kent, D. H —Spartina glabra Muhl.—in
the British Isles, 163-164

Keys (by computer), 357-368

Kidwelly, Carmarthenshire (Field Meet-
ings 1970), 430-431

Kiernan, J. A.— Rhynchosinapis —the
Worcestershire records, 293

Knocking Hoe National Nature Reserve
(Field Meetings 1970), 427

Kochia scoparia (in Middx), 186

Kovanda, M.— Observations on Phyteuma
tenerum R. Schulz in England, 385—
389

Lamiastrum galeobdolon, 277-288 ; galeob-
dolon (pollination of), 337; galeob-
dolon subsp. flavidum, 277, 285-287;
galeobdolon subsp. montanum, 277—
286, 288

Lamium galeobdolon (see Lamiastrum
galeobdolon), 277, 278

Lancaster, R.—review of D. McClintock,
Guide to the Naming of Plants: with
special Reference to Heathers, 319

Lastrea dilatata var. alpina, 3, 7; var.
fraseri, 7; var. micromera, 7; var.
ordeanae, 7; var. pseudo-spinulosa, 7

Lathraea squamaria (in Montgom.), 187

Lathyrus maritimus Bigel. subsp. acuti-
folius (Bab.) Pedersen in eastern
Scotland (by U. K. Duncan), 87

Lathyrus sylvestris (in Mull), 149

Launert, E.—review of S. G. Harrison,
G. B. Masefield & M. Wallis, The
Oxford Book of Food Plants, 319-
320

Laurentia tenella, 295

Lawrence, W. J. C.—Plant Breeding,
reviewed by C. J. Marchant, 174

Lees, D. R.—Frequencies of pin and
thrum plants in a wild population of
the Cowslip, Primula veris L., 289-
291

Leicestershire (Field Meetings 1969), 185-
186; (Senecio vernalis in), 90

Lepidium campestre, 333; heterophyllum,

Leutscher, A.—Field Natural History—a
Guide to Ecology, reviewed by E. W.
Groves, 169-170

Lewis, J.—review of E. Roux, Grass: a
Story of Frankenwald, 417

Ligusticum scoticum (in Shetland), 193

Limerick (grasslands of), 321

Linaria dalmatica (on tip in S. Lancs.), 301

460

Linaria supina (in Carmarthenshire), 430

Linum alpinum, 205, 211, 214, 216, 218,
220, 227; alpinum subsp. anglicum,
208; alpinum subsp. montanum, 208;
anglicum, 208, 210; austriacum, 205,
206, 208, 213, 216, 220, 223-232, 234;
austriacum subsp. collinum, 206, 208,
211, 212, 215-220, 223, 225-231, 234;
austriacum subsp. squamulosum, 208 ;
bavaricum, 207, 208, 210, 213; bienne,
206; collinum, 205, 212; darmstadi-
num, 207, 210, 213; euxinum, 213;
extraaxillare, 205, 211, 216; julicum,
207, 211, 214, 227; laeve, 209; leonii,
206, 208-210, 212, 218, 219, 222-232,
234; lewisii, 205; limanense, 207, 212,
217; loreyi, 207, 212, 219; marschal-
lianum, 213; montanum, 205, 211,
214, 216; perenne 205-211 213-216,
220-232, 235; perenne x alpinum,
220; perenne subsp. alpinum, 206,
210, 211, 213-216, 218, 220, 222-226,
228-232, 235; perenne subsp. angli-
cum, 206, 208, 210, 213, 220-223,
225-230, 232, 235; perenne subsp.
extraaxillare, 206, 208, 210 211, 215,
246; 220, 2223-225 L227 evesOwet 235:
perenne subsp. montanum, 206, 208,
QUT 215;5 216) 220. 223-225; 228) 229.
232, 235; perenne subsp. petryi, 208;
petryi, 207, 212, 219; provinciale, 207,
208, 212, 217; punctatum, 206, 207,
210, 212, 218; pycnophyllum, 207,
212, 218; pycnophyllum subsp. kurdi-
cum, 218; ruscinonense, 207, 212, 217,
218; saxicola, 207, 212, 217; subbi-
florum, 209; tommasinii, 207, 212,
217, 218; trinervium, 207, 212, 217,
223

Liriodendron (in Harrow), 88

List of the Flowering Plants and Ferns of
Carmarthenshire, (by R. F. May),
reviewed by A. O. Chater, 313-314

Listera cordata (on Raasay), 190; ovata
(on Raasay), 190; ovata (pollination
Of); 337

Lolium multiflorum Lam. x L. rigidum
Gaud.—at Blackmoor, N. Hants,
v.c. 12 (by A. Melderis), 299-300

Lolium rigidum, 299

Lolium species and hybrids in Britain (by
A. Melderis), 336

Long Mountain (Field Meetings 1969), 187

Lonicera periclymenum (in Shetland), 192,
193; periclymenum (pollination of),
337,

Lothian, West (Field Meetings 1970),
432-433

Lotus pedunculatus (in Shetland), 192

Lousley, J. E-—Hypericum canadense L.—
in England, 293

INDEX TO WATSONIA VOLUME 8

Lousley, J. E——review of D. Luciana,
Millefiori della Valle d’ Aosta, 417

Lousley, J. E.—review of R. H. Jeffers,
The Friends of John Gerard (1545-
1612), Surgeon and Botanist, 65-66

Lowland Grasslands of Co. Limerick, (by
A. M. O’Sullivan), reviewed by R. D.
Meikle, 321

Luciana, D.—Millefiori della Valle d’ Aosta,
reviewed by J. E. Lousley, 417

Lunaria annua (new to Guernsey), 335

Lupinus arboreus (in E. Suffolk), 430;
nootkatensis (colour forms in Perth-
shire), 189; polyphyllus (in Guernsey),
428

Luzula spicata (in Mull), 146

Lycium barbarum (in Iona), 150

Lycopersicon esculentum (in Mull), 150

Lycopodium alpinum (not in Mull), 146,
147; alpinum (on Raasay), 191;
annotinum (in Mull), 147; annotinum
(on Raasay), 191; clavatum (in Mull),
146; inundatum (in Hants.), 186;
inundatum (in Pembs.), 188

Lysimachia nummularifolium (in Tiree),
194; punctata (in Mull), 150; punc-
tata, 333; vulgaris, 333; vulgaris (in
Mull), 150

McAllister, H. A.—Field Meetings 1969:
Byrne Hill, Ayrshire, 15 June, 189

McAllister, H. A.—Field Meetings 1970:
Glendaruel, 432

McClintock, D.—Chenopodium pumilio—
a Kentish obituary ?, 335-336

McClintock, D.—Field Meetings 1969:
Guernsey, 11-20 April, 183-184

McClintock, D.—Field Meetings 1970:
Guernsey, 428-429

McClintock, D.—Geranium submolle
Steud. (or G. core-core Steud.)—in
the Channel Isles, 47

McClintock, D.—Guide to the Naming of
Plants: with special reference to
heathers, reviewed by R. Lancaster,
319

McClintock, D.—review of C. B. Heiser,
Nightshades: the Paradoxical Plants,
320-321

McClintock, D.—The flora of Guernsey
(exhibit), 89-90

McClintock, D.—The Guernsey flora in
1969, 335

Mackechnie, R.—Field Meetings 1970:
Island of Bute, 431-432

Maianthemum bifolium (L.) Schmidt—its
status in the British Isles (by E. L.
Swann), 295-297

Manchester Poplar (by C. A. Stace), 391-
393

Marchant, C. J.—review of W. J. C.
Lawrence, Plant Breeding, 174

INDEX TO WATSONIA VOLUME 8

Markgraf, F.—review of Flora Europaea,
Volume 2, 66

Martin, B. S.—(joint author with A. C.
Jermy)—A note on the preparation
of material for the scanning electron
microscope, 14-15

Masefield, G. B.—(joint author with S. G.

Harrison and M. Wallis)—The
Oxford Book of Food Plants, reviewed
by E. Launert, 319-320

Matthews, J. R.—review of W. Penning-
ton, The History of British Vegetation,
175-177

May, R. F.—A List of the Flowering
Plants and Ferns of Carmarthenshire,
reviewed by A. O. Chater, 313-314

Meconopsis cambrica (in Guernsey), 428

Meetings Committee (Report 1968), 76;
(Report 1969), 324

Meikle, R. D.—AHypericum canadense L.
in the British Isles, 90

Meikle, R. D.—review of A. M. O’Sul-
livan, The Lowland Grasslands of
Co. Limerick, 321

Meikle, R. D.—review of O. Polunin,
Flowers of Europe, 321-322

Melderis, A.—Bromus pumpelianus—an
introduced species, at Walthamstow,
S. Essex, v.c. 18, 300

Melderis, A.—Lolium multiflorum Lam.
x L. rigidum Gaud.—at Blackmoor,
N. Hants, v.c. 12, 299-300

Melderis, A.—review of R. K. Brummitt
& I. K. Ferguson, Index to European
Taxonomic Literature for 1966 and
1967, 417-418

Melderis, A.—Species of Lolium and their
hybrids in Britain, 336

Membership of B.S.B.I., 86

Mentha x niliaca (in Mull), 151; spicata
(in Mull), 151; x_ verticillata (in
Mull), 151; x villosa (in Mull), 151

Mercurialis annua (in Carmarthenshire),
314

Merioneth (Festuca altissima in), 84
Mertensia maritima (in Mull), 150

Messenger, K. G.—Senecio vernalis
Waldst. & Kit. in Leicestershire
(v.c. 55), 90

Metasequoia (in Harrow), 88
Microsorium diversifolium (in Guernsey),

Middlesex (Field Meetings 1969), 186-187

Midhurst, Sussex (Field Meetings 1969),
184-185

Miles, Beverley Alan (obituaries by P. D.
Sell and E. S. Edees), 423-424

Milium scabrum (from Guernsey), 90

Millefiori della Valle d’Aosta, (by D.
pean): reviewed by J. E. Lousley,

17

461

Mills, J. N. & Mills, J. R. J.—Hieracium
—two Hieracia sect. Alpestria from
the British mainland, 48—49

Mills, J. R. J—(joint author with J. N.
Mills)—Hieracium—two — Hieracia
sect. Alpestria from the British
Mainland, 48-49

Milne-Redhead, E.—Presidential Address
1970: Botanical conservation in
iat past, present and future, 195-
20

Milne-Redhead, E. & Ferguson, I. K.—
Report of the Council for 1969, 323-
327

Mimulus cupreus x guttatus (in Shetland),
192; guttatus <x cupreus (in Mull),
151; moschatus (in Mull), 151

Minuartia verna (in Ayrshire), 189; verna
(in Mull), 148

Mitchell, J—(joint author with E. T. Idle
and A. McG. Stirling)—Elatine
hydropiper—new to Scotland, 45-46

Modern Methods in Plant Taxonomy (ed.
V. H. Heywood), reviewed by
M. C. F. Proctor, 418-420

Monks Wood National Nature Reserve,
90-91

Monotropa hypopitys L.—new to Fife
(v.c. 86) (by A. Angus), 163

Montgomery Branch of Shropshire Union
Canal (Field Meetings 1970), 431

Montgomeryshire (Field Meetings 1969),
187

Montia perfoliata (in Guernsey), 428;
sibirica (in Guernsey), 183, 428
Morecambe Bay—northern and southern
elements in the flora (by A. J. Gray),
335

Morphological development of Spartina
‘X’ (by T. A. W. Davis), 85-86

Mull, 89, 145-153, 334; (Hippophae
rhamnoides on), 396

Mullenders, W. ed.,—Flore de la Belgique,
du Nord de la France et des régions
voisines, reviewed by P. A. Stott, 313

Murray, Mrs C. W.—Field Meetings
1969: Isle of Raasay, Inverness-
shire, 28th June—Sth July, 190-191

Myosotis sylvatica (in Guernsey), 184

Myosurus (? not in Guernsey), 428

Mpyrica gale (in Pembs.), 188

Mpriophyllum alternifiorum (in Guernsey),
335; alterniflorum (in Pembs.), 188;
spicatum (in Shetland), 193

Naias marina (in Norfolk), 185

Nardurus, Vulpia or Festuca (by B. Batts,
H. Jones & C. A. Stace), 333

‘Nardus stricta (on Norfolk dunes), 185

National Museum of Wales, Cardiff
(Field Meetings 1970), 431

National Woodland Survey, 90

462

Naufraga balearica Constance & Cannon
—refound (by L. F. Ferguson), 294—
295

Neottia nidus-avis (on Raasay), 190, 191

Newton, A.—Six brambles (Rubi) from
the north Midlands, 369-377

Newton, L. E.—(Goint author with
B. M. G. Jones)—The Status of
Puccinellia pseudodistans (Crép.)
Jansen & Wachter in Great Britain,
17-26

Nicholson, B. E.—(illustrations by), 319

Nightshades: the Paradoxical Plants (by
C. B. Heiser), reviewed by D.
McClintock, 320-321

Norfolk, East (Field Meetings 1969),
185

Norfolk (Flora reviewed), 416-417

North Hants (Lolium hybrid in), 299

North-West Region Meeting 1969 (by
E. F. Greenwood), 82

Note on the preparation of material for
the scanning electron microscope (by
A. C. Jermy & B. S. Martin), 14-15

Notes on the Hawkweeds (Hieracium
sensu lato) of western Scotland (by
A. G. Kenneth & A. McG. Stirling),
97-120

Nothoscordum inodorum (in Guernsey),
429

Numerical Taxonomy (ed. A. J. Cole),
reviewed by D. B. Williams, 422

Nuphar lutea (on Raasay), 191

Nymphaea alba (in Sutherland), 193

Obituaries, 69-74, 181-182, 423-424

Observations on Phyteuma tenerum R.
Schulz in England (by M. Kovanda),
385-389

Ockendon, D. J.—Taxonomy of the
Linum perenne group in Europe, 205—
235

Ockendon, D. J. & Walters, S. M.—
Studies in Potentilla anserina L., 135-
144

Oenothera biennis (on tip in S. Lancs), 301

Olearia forsteri (in Guernsey), 183

Ononis reclinata (in Channel Is.), 428

Ononis repens (on tip in S. Lancs.), 301

Ophioglossum lusitanicum (from Guern-
sey), 90

Orchis x alata (? not in Guernsey), 429;
laxiflora (in Guernsey), 429; morio (?
not in Guernsey), 429; simia, 95

Orobanche alba (on Raasay), 190

Orthilia secunda (in Mull), 150

Osmunda regalis (in Leics.), 186; regalis
(in Norfolk), 185

O’Sullivan, A. M.—The Lowland Grass-
lands of Co. Limerick, reviewed by
R. D. Meikle, 321

Oxalis acetosella (floral aberrations), 88

INDEX TO WATSONIA VOLUME 8

Oxford Book of Food Plants (by S. G.
Harrison, G. B. Masefield and M.
Wallis), reviewed by E. Launert, 319-
320

Oxford Book of Wild Flowers: Pocket
Edition (ed. S. Ary & M. Gregory),
reviewed by E. B. Bangerter, 412-413

Oxyria digyna (in Mull), 146

Paish, D. M.—Field Meetings 1969: Long
Mountain, Montgomeryshire, 15th
June, 187

Panel of referees and specialists, 339-344

Panicum capillare (in Middx), 187

Pankhurst, R. J.—Botanical keys genera-

ted by computer, 357-368

Pankhurst, R. J.—Computer generated
keys, 336-337

Papaver argemone, 263, 266, 267, 270,
274-275; dubium, 263, 266-271, 273-
275; dubium (in Mull), 146, 147;
hybridum, 263, 266, 267, 270, 275;
lecoqii, 263, 266-270, 273-275; nudi-
caule, 271; rhoeas, 263-275; rhoeas
(in Mull), 147; rhoeas x dubium,
271; rhoeas f. strigosum, 271; somni-
ferum (in Mull), 147; strigosum, 271

Peacock, R. J.—(joint author with C. T.
Prime)—Trees and Shrubs—their
Identification in Summer or Winter,
ed. 6, reviewed by J. F. M. Cannon,
414-415.

Pedicularis sylvatica subsp. hibernica (in
Mull), 151

Pembrokeshire (Field Meetings 1969), 188

Pennington, W.—The History of British
Vegetation, reviewed by J. R.
Matthews, 175-177

Peplis portula (in Guernsey), 428

Perring, F. H.—B.S.B.[. Symphytum
survey, 91

Perring, F. H.—review of K. & H.
Rasbach & D. Wilmanns, Die
Farnpflanzen Zentraleuropas, 318

Perring, F. H.—The Vice-County Guide,
435-447

Perring, F. H.—Vascular plants new to
Ireland, 91

Persistence of Silene maritima and other
plants on an inland tip in South
Lancashire, v.c. 59 (by T. Edmund-
son), 301

Perspectives in Phytochemistry (ed. J. B.
Harborne, & T. Swain), reviewed by
H. J. M. Bowen, 168

Perthshire (recording in), (Field
Meetings 1969), 189-190

Petasites albus (in Mull), 152; fragrans (in
Mull), 152

Petch, C. P. & Swann, E. L.—Flora of
Norfolk, reviewed by A. C. Jermy,
416-417

93;

INDEX TO WATSONIA VOLUME 8

Peucedanum palustre (in Norfolk), 185

Phleum arenarium (in Mull), 153

Photomicrographs of a section of Orchis
simia root (by H. M. Wilks), 95

Photosynthesis (by I. Asimov), reviewed
by M. C. F. Proctor, 322

Phragmites australis (in Shetland), 193

Physalis peruviana (in Middx), 186

Phyteuma orbiculare complex, 385-388;
tenerum (in England), 385-389

Phytochemical Phylogeny (ed. J. B.
Harborne), reviewed by E. C. Bate-
Smith, 413

Pickering, E. & Stace, C. A.—Variation
in the Vicia sativa aggregate, 92

Pilosella aurantiaca, 83; aurantiaca subsp.
aurantiaca, 117; aurantiaca subsp.
brunneocrocea, 117; caespitosa subsp.
colliniformis, 118; flagellaris subsp.
bicapitata, 118; flagellaris subsp. bi-
capitata (in Shetland), 193; flagellaris
subsp. /flagellaris, 118; Jlactucella
subsp. helveola, 118; lactucella subsp.
lactucella, 118; officinarum, 83, 115-—
117, 428; officinarum subsp. concin-
nata, 115, 116, 117; officinarum subsp.
nigrescens, 116, 117; officinarum
subsp. tricholepia, 116, 117; offici-
narum subsp. trichoscapa, 115, 117;
officinarum subsp. unnamed, 116,
117; peleteriana subsp. peleteriana,
117; peleteriana subsp. tenuiscapa,
117; praealta subsp. arvorum, 118;
praealta subsp. praealta, 118; praealta
subsp. spraguei, 118; x rubrum, 83

Pinguicula lusitanica (in Pembs.), 188

Plant Anatomy (by A. Fahn), reviewed by
V. H. Heywood, 317-318

Plant Anatomy: Experiment and Inter-
pretation, Part I, Cells and Tissues (by
E. G. Cutter), reviewed by D. F.
Cutler, 169

Plant Breeding (by W. J. C. Lawrence),
reviewed by C. J. Marchant, 174

Plant records, 53-61, 165-167, 302-312,
403-411

Plant records from Mull and the adjacent
small islands, 2 (by J. F. M. Cannon
& E. B. Bangerter), 145-153

Plant variation and Evolution (by D.
Briggs & S. M. Walters), reviewed by
D. A. Wilkins, 179-180

Platanthera bifolia (in Tiree), 194

Pease Dean, Berwickshire (Field Meetings
1969), 189

Poa bulbosa (in Guernsey), 183; compressa
Gin Mull), 153; infirma (from
Guernsey), 90

Pollichia montanum, 278; vulgaris, 278

Pollination photographs (by M. C. F.
Proctor), 337

463

Polunin, O.—Flowers of Europe, reviewed
by R. D. Meikle, 321-322

Polunin, O.—review of C. N. Goulimis,
Wild Flowers of Greece, 316-317

Polygonatum x hybridum (in Guernsey),
428

Polygonum arenarium (in Middx), 187;
baldschuanicum (in Mull), 150; cam-
panulatum (in Mull), 150; cuspidatum
Gn Mull), 150; oxyspermum subsp.
raii (in Shetland), 193; patulum, 187;
pensylvanicum (in Middx), 187; poly-
stachyum (in Mull), 150; pulchellum,
187; sachalinense (in Mull), 150;
viviparum (on Raasay), 190

Polypodium australe, 12\1—134; interjectume
121-128, 130-133; vulgare, 121-123,
125. 11ZouelsO~ 13k 31336 woulgar:
subsp. prionodes, 121; vulgare subsp.
serratum, 121; vulgare subsp. serru-
latum, 121; vulgare var. serratum,
121

Polypogon monspeliensis (in Middx), 187

Poplar (Manchester), 391-393

Populus x canadensis, 391, 392; x
canadensis var. serotina, 391; del-
toides, 391; ‘eugenei’, 392; nigra, 391,
392; nigra subsp. betulifolia, 391;
nigra var. betulifolia, 391, 392;
nigra var. italica, 392; tremula (in
Shetland), 192

Portulaca oleracea (in Middx), 187

Potamogeton alpinus (in Mull), 152;
berchtoldii (in Mull), 152; coloratus
(in Tiree), 194; filiformis (in Shetland),
192; filiformis (in Tiree), 194; grami-
neus (in Shetland), 193; /ucens (not in
Shetland), 193; x nitens (in Tiree),
194; pectinatus (in Tiree), 194;
perfoliatus (in Tiree), 194; rutilus (in
Shetland), 193; rutilus (in Tiree), 194;
x sparganifolius (in Tiree), 194; x
suecicus (in Tiree), 194; x zizii (new
to Shetland), 193

Potentilla anglica (in Guernsey), 428;
anserina, 135-144; anserina subsp.
egedii, 140, 144; norvegica (on tip in
S. Lancs), 301; tabernaemontani,
143

Poterium sanguisorba (in Perthshire), 190

Presidential Address 1970 (by E. Milne-
Redhead), 195-203

Primavesi, A. L.—Field Meetings 1969:
Belvoir and Charnwood Forest,
Leicestershire, 1-3 August, 185-186

Prime, C. T. & Peacock, R. J.—Trees and
Shrubs—their Identification in
Summer or Winter, ed. 6, reviewed by
J. F. M. Cannon, 414-415

Primula scotica (in Sutherland),
veris, 289-291

194;

464

Principles and Methods of Plant Bio-
systematics (by O. T. Solbrig),
reviewed by D. Briggs, 414

Proceedings of the Botanical Society of the
British Isles, 1-2

Proctor, H. M.—Field Meetings 1970:
East Suffolk, 430

Proctor, H. M.—The Upper Teesdale
Flora in June, 92

Proctor, M. C. F.—“‘Is your plate camera
really necessary ?’’, 92

Proctor, M. C. F.—Pollination photo-
graphs, 337

Proctor, M. C. F.—review of F. K.
Hartman & G. Jahn, Waldgesellschaf-
ten des mitteleuropdischen Gebirgs-
raumes nordlich der Alpen, 177

Proctor, M. C. F.—review of I. Asimov,
Photosynthesis, 322

Proctor, M. C. F.—review of V. H.
Heywood, ed., Modern Methods in
Plant Taxonomy, 418-420

Proposed list of British hybrids (by C. A.
Stace), 94

Prunella vulgaris W.—colour variants
apparently introduced (by D _ E.
Allen), 399-400

Prunella vulgaris var. pallida, 400

Prunus cerasifera (in Guernsey),
padus (on Raasay), 191

Psoralea americana (in Surrey), 187

Pterostylis (in Australia), 94

Publications Committee (Report 1968),
76-77; (Report 1969), 325

Publications Fund transactions in 1968,
80; transactions in 1969, 329

Puccinellia capillaris (in Shetland), 193;
distans, 17-26; fasciculata, 17-26;
fasciculata (on R. Stour), 185;
fasciculata f. pseudodistans comb.
nov., 26; maritima, 21; pseudodistans,
17-26; rupestris, 24

Pyrola minor (on Raasay), 191; rotundi-
folia (in Carmarthenshire), 430;
rotundifolia subsp. maritima (in N.
Lancs.), 184; rotundifolia subsp.
maritima (in Westmorland), 430;
rotundifolia subsp. rotundifolia (on
Norfolk dunes), 185; secunda (on
Raasay), 190, 191

Raasay (Field Meetings 1969), 190-191

Ranunculus acris L. complex in Europe
(by S. M. Coles), 237-261

Ranunculus acris f. multifidus, 240; acris f.
villosus, 240; acris subsp. borealis,
237, 241, 246, 248, 249, 251, 252, 257,
260; acris subsp. friesanus, 237-239,
241-244, 246-253, 255, 257, 261;
acris subsp. granatensis, 237, 239;
acris subsp. strigulosus, 237, 239;
acris var. atlanticus, 239; acris var.

SSE

INDEX TO WATSONIA VOLUME 8

multifidus, 242; acris var. pumilus,
~ 237, 240, 243, 246-249, 251, 257, 260;

acris var. pyrenaeus Coles, 237, 238,
241, 242, 246, 247, 249, 257, 261;
acris var. steveni, 242; acris var.
villosus (Drabble) Coles, 237, 238,
240, 243, 247, 249, 257, 259; atlanti-
cus, 239; borealis, 241; boreanus, 240;
csatoi, 239; ficaria subsp. ficariiformis
(in Guernsey), 428; friesanus, 241;
granatensis, 237-240, 242-243, 245,
247, 249-252, 255, 256, 258; maloco-
phyllus, 239; nemorivagus, 241; serbi-
cus, 237; steveni, 241; strigulosus,
237-243, 245, 247, 249-252, 255, 256,
258; tomophyllus, 240; trichophyllus
subsp. trichophyllus (in Shetland), 193

Raphanus raphanistrum (in Mull), 146;
sativus (in Mull), 147

Rasbach, K. & H. & Wilmanns, O.—Die
Farnpflanzen Zentraleuropas, re-
viewed by F. H. Perring, 318

Ravensthorpe, Western Australia (orchids
of), 94

Recording in Perthshire (by A. W.
Robson), 93

Records Committee (Report 1968), 77;
(Report 1969), 326

Referees, 339-344

Report of the Council for 1968 (by J. G.
Dony & D. E. Allen), 75—78; for 1969
(by E. Milne-Redhead & I. K.
Ferguson), 323-327

Revision of some of the taxonomic
characters of Polypodium australe Fée
(by R. H. Roberts), 121-134

Rhamnus catharticus (in Mull), 148

Rhinanthus serotinus subsp. apterus (in
Shetland), 192

Rhynchosinapis cheiranthos, 293 ; monensis,
293

Rhynchosinapis—The Worcestershire re-
cords (by J. A. Kiernan), 293

Ribbons, B. W.—Committee for the Study
of the Scottish Flora, fifteenth annual
report to 31 December 1969, 331-
332

Ribes uva-crispa (in Guernsey), 428

Richards, A. J.—Taraxacum in Upper
Teesdale, 337

Richards, A. J.—Taraxacum section Ery-
throsperma in the British Isles, 93

Richardson, J. A.—Obituary: John Wil-
liam Heslop-Harrison (1881-1967),

181-182

Roanhead (Field Meetings 1969), 184

Robb, S. M.—review of K. Esau, Viruses
in Plant Hosts, 420-421

Roberts, R. H.—A revision of some of the
taxonomic characters of Polypodium
australe Fée, 121-134

INDEX TO WATSONIA VOLUME 8

Robson, A. W.—Field Meetings 1969:
The shingle beds of the Tummel and
Tay, Perthshire, 21-22 June, 189-190

Robson, A. W.—Recording in Perthshire,
93

Robson, N. K. B.—book notices, 174-175

Robson, N. K. B.—review of A.
Cronquist, The Evolution and Classi-
fication of Flowering Plants, 177-179

Robson, N. K. B.—review of A.
Takhtajan, Flowering Plants, Origin
and Dispersal, 315-316

Robson, N. K. B.—review of Index
Kewensis Plantarum Phanerogama-
rum, Supplementum XIV, 322

Robson, N. K. B.—review of J.
Hutchinson, Evolution and Phylogeny
of Flowering Plants, 421

Rogers, S.—Studies on British Poppies, 4.
Some aspects of variability in the
British species of Papaver and their
relation to breeding mechanisms and
ecology, 263-276

Romulea columnae (albino, in Guernsey),
183; rosea (in Guernsey), 183; rosea
var. australis (in Guernsey), 335

Rorippa x sterilis (in Pembs.), 188

Rosa canina (in Shetland), 192

Roudsea (Field Meetings 1969), 184

Roux, E.—Grass: a story of Frankenwald,
reviewed by J. Lewis, 417

Rubus (work by B. A. Miles), 423-424;
accrescens A. Newton, 369; adenan-
thoides A. Newton, 374; adenanthus,
375; arcticus (in Mull), 149; cinerosus,
372; disjunctus, 377; distractiformis
A. Newton, 375; distractus, 377;
echinatus, 375; flexuosus, 373; goa-
ronii vat. foliolatus, 370; larktonensis,
370; menkei, 377; mucronatus, 372;
mucronifer, 372; muenteri var. robii,
370; porphyrocaulis A. Newton, 372;
pseudadenanthus, 375; robii (W. C. R.
Wats.) A. Newton, 370; sprengelii,
370; warrenii, 377; wirralensis A.
Newton, 371

Ruppia cirrhosa (in Mull), 152; maritima
(in Mull), 152

Ryves, T. B. (with E. J. Clement)—Field
Meetings 1969: alien hunt about
London, 4-5 October, 186-187

Sagina maritima (in Mull), 148

Salicornia dolichostachya, 27-40; euro-
paea, 27-40; europaea (new to
Guernsey), 335

Salicornia in Dee estuary, 27-40

Salisbury, E. J.—Echinophora spinosa L.
—a member of our flora, 397-399

Salix and Epilobium species and hybrids
from lowland Fife (v.c. 85) (by G. H.
Ballantyne), 332-333

465

Salix atrocinerea (on Raasay), 191;
aurita X cinerea (in Guernsey), 184;
basfordiana (in Guernsey), 184; calo-
dendron (in Guernsey), 335; daph-
noides (in Guernsey), 184; fragilis (in
Guernsey), 184; herbacea (in Mull),
146; herbacea (on Raasay), 191;
hybrids, 331; phylicifolia (in Perth-
shire), 191

Salvia glutinosa (pollination of), 337;
refiexa (in Middx), 187

Sambucus ebulus (in Guernsey), 428;
racemosa (in Mull), 151

Sanguisorba minor subsp. minor (in
Guernsey), 429

Saponaria officinalis (on tip in S. Lancs.),
301

Sarothamnus scoparius subsp. prostratus,
3

Sasa veitchii (in Guernsey), 184

Saussurea alpina (in Mull), 146

Saxifraga hypnoides (in Miull),
stellaris (in Mull), 146

Scabiosa columbaria showing ““bird’s nest”’
proliferation (by J. L. Gilbert), 87

Scanning electron microscope, 4, 14-15

Schroeder, F.-G.—Exotic Amelanchier
species naturalised in Europe and
their occurrence in Great Britain,
155-162

Scilla verna (in Sutherland), 194

Scirpus cernuus, 333; setaceus, 333

Scott, W.—Field Meetings 1969: Shet-
land, 1-8 August, 191-193

Sedum stoloniferum (in Guernsey), 183;
telephium (in Mull), 146

Selaginella selaginoides (in Cardigan-
shire), 188

Selby, H. E. W.—An experiment in
flower mounting, 337-338

Sell, P. D.—Obituary: Beverley Alan
Miles, 423-424

Sell, P. D. & Walters, S. M.—Two species
new to Cambridgeshire, 93

Senecio cambrensis, 47-48; x londinensis,
48, 85; x ostenfeldii (in Mull), 151

Senecio x ostenfeldii Druce in Scotland
(by U. K. Duncan), 87

Senecio smithii (in Shetland), 193; squali-
dus, 85; squalidus (in Guernsey), 429;
squalidus (in Stirlingshire), 194

Senecio squalidus L., S. vulgaris L. and
S. cambrensis Rosser (by P. Crisp &
B. M. G. Jones), 47-48

Senecio vernalis Waldst. & Kit. in
Leicestershire (v.c. 55) (by K. G.
Messenger), 90

Senecio viscosus, 48, 85; vulgaris, 47-48,
85; vulgaris var. denticulatus, 47;
vulgaris var. hibernicus, 85

Sequoia sempervirens (in Montgom.), 187

146;

466

Setaria lutescens (in Middx), 186

Shetland (Field Meetings 1969), 191-
193

Shingle beds of Tummel and Tay (Field
Meetings 1969), 189-190

Shropshire Union Canal (Field Meetings
1970), 431

Sieglingia decumbens (L.) Bernh.—pulvini
of palea (by W. G. Dore), 297-299

Sieglingia decumbens (on Norfolk dunes),
185

Silene acaulis (in Sutherland), 194; alba
x dioica, 428; maritima (on tip in S.
Lancs.), 301; vulgaris (in Mull), 148

Silphium perfoliatum (in Surrey), 187

Sisymbrium columnae, 293; irio (in
Middx), 187; Joesellii Gn Middx),
186; officinale (in Shetland), 192

Six brambles (Rubi) from the north
Midlands (by A. Newton), 369-377

Skinflats, Stirlingshire (Field Meetings
1969), 194

Slack, A. A —Field Meetings 1970: Fort
William, Inverness-shire, 433

Sledge, W. A.—review of H. A. Hyde &
A. E. Wade, Welsh Ferns, Clubmosses,
Quillworts and Horsetails, ed. 5 by
S. G. Harrison, 67-68

Slides of orchids from Ravensthorpe,
Western Australia (by L. E. White-
head), 94

Smith, A. R.—Some alien Euphorbias in
the British Isles, 94

Solanaceae (book review), 320-321

Solbrig, O. T.—Principles and Methods of
Plant Biosystematics, reviewed by D.
Briggs, 414

Some often confused plants (by E. B.
Bangerter & L. F. Ferguson), 333

Some Scottish hawkweeds (by A. McG.
Stirling), 336

Sonchus palustris (in Norfolk), 185

Sorbus aria (in Mull), 149; aucuparia (in
Shetland), 192; intermedia (in Mull),
149: rupicola (on Raasay), 190;
rupicola/lancastriense (in Westmor-
land), 429

Sorghum halepense (in Middx), 187

Souter, A J.—Crassula aquatica (L.)
Schonl.—new to Scotland, 294

South Essex (Bromus pumpelianus in), 300

South Lancashire (Leigh tip), 301

Spartina alterniflora var. glabra, 163, 164

Spartina glabra Muhl.—in the British
Isles (by D. H. Kent), 163-164

Spartina maritima subsp. glabra, 164;
pectinata, 85; stricta var. glabra, 164,
x townsendii (on R. Stour), 185; ‘X’,
85—86

Species of Lolium and their hybrids in
Britain (by A. Melderis), 336

INDEX TO WATSONIA VOLUME 8

Spergularia bocconei (in Guernsey), 429;
rubra (in Carmarthen), 314

Spiraea japonica (in Guernsey), 428

Stace, C. A.—A proposed list of British
hybrids, 94

Stace, C. A.—Hybrids in Juncus section
Genuini, 338

Stace, C. A.—The Manchester Poplar,
391-393

Stace, C. A.—(joint exhibitor with B.
Batts and H. Jones)—Nardurus,
Vulpia or Festuca, 333

Stace, C. A.—(joint exhibitor with E.
Pickering)—Variation in the Vicia
sativa aggregate, 92

Stachys sylvatica (pollination of), 337

Status of Puccinellia pseudodistans (Crép.
Jansen & Wachter in Great Britain
(by B. M. G. Jones & L. E. Newton),
17-26

Stearn, W. T. ed., C. N. Goulimis, Wild
Flowers of Greece, reviewed by O.
Polunin, 316-317

Stellaria graminea (in Shetland), 192

Stellaria (seed characters), 334

Stereoscan, 15

Steroscan studies on seed morphology (by
F. G. Bell), 333-334

Still more about Mull (by Dept. Bot.,
British Museum (Nat. Hist.)), 89

Stirling, A. McG.—Field Meetings 1969:
Braes of Balquhidder, Perthshire, 12
July, 191

Stirling, A. McG.—Some Scottish hawk-
weeds, 336

Stirling, A. McG.—(joint author with
A. G. Kenneth)—Notes on the
Hawkweeds (Hieracium sensu lato) of
western Scotland, 97-120

Stirling, A. McG.—(joint author with
E. T. Idle and J. Mitchell)—Elatine
hydropiper—new to Scotland, 45-—
46

Stott, P. A.—review of W. Mullenders,
ed.,—Flore de la Belgique, du Nord
de la France et des régions voisines,
313

Studies in Potentilla anserina L. (by D. J.
Ockendon & S. M. Walters), 135-144

Studies on British Poppies, 4. Some
aspects of variability in the British
species of Papaver and their relation
to breeding mechanisms and ecology
(by S. Rogers), 263-276

Studies on Cytisus scoparius (L.) Link
with particular reference to the
prostrate forms (by J. J. B. Gill & S.
Walker), 345-356

Studies on Gloucestershire populations of
Allium paradoxum (Bieb). G. Don
(by D. M. Barling), 379-384

INDEX TO WATSONIA VOLUME 8

Study of some western Eurasian Gladioli
(by A. P. Hamilton), 88

Subularia aquatica (in Shetland), 193

Succisa pratensis (in Sutherland), 193

Suffolk, East (Field Meetings 1970),
430

Swann, E. L.—Maianthemum bifolium (L.)
Schmidt—its status in the British
Isles, 295-297

Swann, E. L.—(joint author with C. P.
Petch), Flora of Norfolk, reviewed by
A. C. Jermy, 416-417

Symphytum asperum, 91; officinale var.
ochroleucum, 91; officinale var. pur-
pureum, 91; x uplandicum, 91

Synokologische Studie der Waldgesell-
schaften auf Amphibolitgestein. Vege-
tace CSSR A3, (by M. Husova),
reviewed by P. J. Grubb, 170-171

Takhtajan, A.—Flowering Plants, Origin
and Dispersal, reviewed by N. K. B.
Robson, 315-316

Taraxacum acutum (new species), 93;
argutum, 93; austriacum (new to
British Isles), 93; brachyglossum, 184;
brachyglossum (in Teesdale), 337;
brachyglossum (in v.c. 90), 334;
brachyglossum (new to British Isles),
93; canulum, 93; commixtum, 93;
cyanolepis (in v.c. 106), 334; degelii,
93; disseminatum, 93; drucei (in V.c.
90), 334; dunense (new to British
Isles), 93 ; euryphyllum, 184; faeroense,
184; faeroense (in Teesdale), 337;
faeroense (in v.c. 106), 334; fulvi-
forme, 93; fulvum, 93; fulvum (in V.c.
90), 334; glauciniforme, 93; gotlandi-
cum (in v.c. 90), 334; gotlandicum
(new to British Isles), 93; hispanicum
(new to British Isles), 93

Taraxacum in Upper Teesdale (by A. J.
Richards), 337

Taraxacum lacistophyllum, 93; lacisto-
phyllum (in v.c. 106), 334; laetiforme,
93; laetum, 93; landmarkii (in Tees-
dale), 337; maculigerum (in Teesdale),
337; oxoniense, 93; placidum (new
species), 93; platyglossum (in v.c. 90),
334; praestans (in Teesdale), 337;
proximiforme (new to British Isles),
93; proximum, 93; proximum (in
Teesdale), 337; pseudolacistophyllum,
93 ; pseudolarssonii (in Teesdale), 337;
pseudonordstedtii (in. Teesdale), 337;
retzii (new to British Isles), 93;
rubicundum, 93

Taraxacum section Erythrosperma in the
British Isles (by A. J. Richards), 93

Taraxacum silesiacum, 93; simile, 93;
spectabile (in Teesdale), 337; specta-
bile (in v.c. 90), 334; unguilobum (in

467

Teesdale), 337; unguilobum (in v.c.
88), 334

Taxodium (in Harrow), 88

Taxonomy of the Linum perenne group in
BuEOE (by D. J. Ockendon), 205-
235

Teesdale Defence Committee Report, 80,

Teesdale (in June), 92; (Taraxacum in),

Teesdalia nudicaulis (in Mull), 147

Tellima grandiflora (in Guernsey), 183

Ten species of Taraxacum from Scotland
(by U. K. Duncan), 334

Thalictrum alpinum (in Mull), 146;
alpinum (on Raasay), 190, 191; minus
subsp. majus (in Perthshire), 190

Thomas, J. H.—review of P. H. Davis &
J. Cullen, The Identification of
Flowering Plant Families, 68

Tiree (Field Meetings 1968), 194

Tolmiea menziesii (in Mull), 149; menziesii
(in Perthshire), 189

Totternhoe Knolls Local Nature Reserve
(Field Meetings 1970), 427

Towyn Burrows, Glamorgan (Field Meet-
ings 1969), 187-188

Towyn Burrows (Field Meetings 1970),
430-431

Trachelium caeruleum (in Guernsey), 335

Trachyspermum ammi (in Middx), 187

Trachystemon orientale (in Guernsey), 183

Treasurer’s Report (by J. C. Gardiner),
79-80; 328-329

Tree survey in the London Borough of
Harrow (by Harrow County School
for Boys), 88

Trees and Shrubs—their Identification in
Summer or Winter, ed. 6, (by C. T.
Prime & R. J. Peacock), reviewed by
J. F. M. Cannon, 414-415

Trifolium hybridum (on Foula, Zetland),
301; hybridum Gin Mull), 148 ; occiden-
tale (in Guernsey), 90, 429; squamo-
sum (in Guernsey), 429; strictum (not
in Guernsey), 428

Trist, P. J. O.—Corynephorus canescens
(L.) Beauv.—in West Suffolk, 402

Tuck, G.—Field Meetings 1969: East
Norfolk, 26-27 July, 185

Tuck, G.—Field Meetings 1970: Bourg
St. Pierre, 426—427

Tutin, T. G.—Obituary: Humphrey
Gilbert-Carter (1884-1969), 72-74

Two additions to the flora of Foula,
Zetland, v.c. 112 (by D. L. Hawks-
worth), 301

Two species new to Cambridgeshire (by
P. D. Sell & S. M. Walters), 93

Typha angustifolia (on Raasay), 191

Umbilicus rupestris (in Leics.), 186

468

Upper Teesdale flora in June (by H. M.
Proctor), 92

Use of Biological Literature (by R. T.
Bottle & H. V. Wyatt), reviewed by
P. I. Edwards, 64-65

Utricularia intermedia (on Raasay), 190;
minor (on Raasay), 190; vulgaris (in
Norfolk), 185

Vaccinium oxycoccus (in Mull), 150

Valentine, D. H.—Obituary: Kathleen
Beever Blackburn, 69-70

Valerianella locusta subsp. dunensis (in N.
Lancs.), 184

van der Pijl, L.—review of K. A. Grant &
V. Grant, Humming Birds and their
Flowers, 67

Variation in the Vicia sativa aggregate (by
E. Pickering & C. A. Stace), 92

Vascular plants new to Ireland (by F. H.
Perring), 91

Vaughan, Mrs H. R. H.—Field Meetings
1969: Camddwr Valley, Cardigan-
shire, 30 August, 188

Vaughan, Mrs H. R. H.—Field Meetings
1969: Cors Goch, Anglesey, 28
September, 189

Vaughan, Mrs H. R. H.— Field Meetings
1969: Towyn Burrows, Glamorgan,
5 July, 187-188

Vaughan, Mrs H. R. H. — Field Meetings
1970: Kidwelly, Carmarthenshire,
430-431

Vaughan, Mrs H. R. H.—Field Meetings
1970: Shropshire Union Canal
(Montgomery Branch), 431

Veronica agrestis, 333; catenata (in
Guernsey), 335; chamaedrys, 333;
filiformis (in Guernsey), 429

Veronica hederifolia L. sensu lato—in
Wales (by P. M. Benoit), 399

Veronica montana, 333; polita, 333;
sublobata, 399; sublobata (in Guern-
sey), 184

Viburnum opulus (on Raasay), 190,
191

Vice-County Guide (compiled by F.
Perring), 435-447

Victorian Fern Craze, a History of
Pteridomania (by D. E. Allen), re-
viewed by J. W. Dyce, 314-315

Vicia lathyroides, 183; hirsuta (in Mull),
149; sativa aggregate, 92

Viola lutea (not in Mull), 148; tricolor (in
N. Lancs.), 184; ‘tricolor subsp.
curtisii (in Mull), 148

Viruses in Plant. Hosts (by K. Esau),
reviewed by S. M. Robb, 420-421

Vulpia (distinction from Nardurus and
Festuca), 333

Waldgesellschaften des mitteleuropdischen
Gebirgsraumes nérdlich der Alpen (by

INDEX TO WATSONIA VOLUME 8

F. K. Hartman & G. Jahn), reviewed
by M. C. F. Proctor, 177

Walker, S.—(joint author with J. A.
Crabbe and A. C. Jermy), The
distribution of Dryopteris assimilis S.
Walker in Britain, 3-13

Walker, S.—(joint author with J. J. B.
Gill)—Studies on Cyfisus scoparius
(L.) Link with particular reference to
the prostrate forms, 345-356

Wallace, E. C. (compiler)—Plant records,
53-61

Wallis, M.—(joint author with S. G.
Harrison and G. B. Masefield), The
Oxford Book of Food Plants, re-
viewed by E. Launert, 319-320

Walters, S. M.—(joint author with D. J.
Ockendon)—Studies in Potentilla
anserina L., 135-144

Walters, S. M.—({joint author with D.
Briggs), Plant Variation and Evolution,
reviewed by D. A. Wilkins, 179-180

Walters, S. M.—(joint exhibitor with
P. D. Sell), Two species new to
Cambridgeshire, 93

Warburg, E. F., 1

Warburg Memorial Fund, 78, 326

Ward, L. K.—The distribution of Juni-
perus communis in southern England,
94

Wardlaw, C. W.—Cellular Differentiation
in Plants and Other Essays, reviewed
by F. Cusick, 415-416

Watercolour drawings of British plants by
the late Mrs Winifred Bussey (by G.
Bussey), 334

Waterford (interesting plants of), 334-335

Watson, H. C., 1

Watsonia, 1-2

Webb, D. A.—The Irish vice-counties, 51

Weemiuller, S—A cytotaxonomic study
of Lamiastrum galeobdolon (L.)
Ehrend. & Polatschek in Britain,
277-288

Wells, T. C. E.—(Goint author with J. G.
Dony)—Field Meetings 1970: Con-
servation of chalk grassland in
Bedfordshire, 427

Welsh Ferns, Clubmosses, Quillworts and
Horsetails (by H. A. Hyde & A. E.
Wade, ed. 5 by S. G. Harrison),
reviewed by W. A. Sledge, 67-68

Welsh Regional Committee (Report 1968),
78; (Report 1969), 326

West Lothian mosses (Field Meetings
1970), 432-433

Westmorland (Field Meetings 1970), 429-—
430

Whitehead, L. E.—Slides of orchids from
Ravensthorpe, Western Australia,
94

INDEX TO WATSONIA VOLUME 8

Wild Flowers of Greece (by C. N.
Goulimis, ed. W. T. Stearn), re-
viewed by O. Polunin, 316-317

Wilkins, D. A.—review of D. Briggs &
S. M. Walters, Plant Variation and
Evolution, 179-180

Wilks, H. M.—Photomicrographs of a

~ section of Orchis simia root, 95

Williams, D. B.—review of A. J. Cole,
Numerical Taxonomy, 422

Williams, P. C.—Abbreviated Titles of
Biological Journals, reviewed by R. K.
Brummitt, 63-64

Wilmanns, D.—(joint author with K. & H.
Rasbach), Die Farnpflanzen Zentral-

469

europas, reviewed by F. H. Perring, 318
Woodland survey, 90
Worcestershire (Rhynchosinapis in), 293
Xanthium strumarium (in Surrey), 187
Year Book, |
Yellow Archangel, 277-288
Yellow-berried Crataegus monogyna (by
J. H. Chandler & J. L. Gilbert), 85
Yeo, P. F.—Euphrasia brevipila and E.
borealis in the British Isles, 41-44
Yet more about Mull, 334
Zannichellia palustris, 193
Zannichellia palustris (in v.c. 72), 331
Zetland, 301
Zostera angustifolia Gn Mull), 152

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INSTRUCTIONS TO CONTRIBUTORS

PAPERS and short notes concerning the systematics and distribution of British and European vascular plants
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Names of genera and species should be underlined, but any other typographical indications should be
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Papers and short notes should be sent to Dr G. Halliday, Dept. of Biological Sciences, The University,
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vice-county recorders.

Watsonia

July 1971 Volume eight Part four

Contents

GILL, J.J. B. and WALKER, S. Studies on Cytisus scoparius (L.) Link

with particular reference to the prostrate forms
PANKHUuRST, R. J. Botanical keys generated by computer
Newron, A. Six brambles (Rubi) from the north midlands

BARLING, D. M. Studies on Gloucestershire populations of A//ium

paradoxum (Bieb.) G. Don

KOvANDA, M. Observations on Phyteuma tenerum R. Schulz in

England
Stace, C. A. The Manchester Poplar
SHORT NOTES
PLANT RECORDS
Book REVIEWS ...
OBITUARY
REPORTS
Conference Report
Field Meetings, 1970

THE VICE-COUNTY GUIDE compiled by F. H. PERRING

345-356

357-368

369-377

379-384

385-389

391-393

395-402

403-411

412-422

423-424

425-426

426-433

435-447

Published and sold by the Botanical Society of the British Isles,
c/o Department of Botany, British Museum (Natural History),
London, S.W.7.

Watsonia, 8 (4). Price Two pounds

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