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Watsonia, 16, 365-371 (1987) 365

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An introduction to the study of the British Hieracia, 1. History
and classification

P. D. SELL

Botany School, Downing Street, The University, Cambridge, CB2 3EA

ABSTRACT

A brief account is given of the historical background to the study of Hieracium L. in the British Isles. Pilosella
Hill is recognized as a distinct genus. The typification of and differences between Hieracium and Pilosella are
set out. The disparity between the Scandinavian and British schools of thought and those of central Europe as
regards the definition of species is discussed.

HISTORY

From the time of Linnaeus onwards the number of described species of Hieracium has grown with
ever-increasing rapidity. The first volume devoted to the genus in the British Isles was by
Backhouse junior (1856), which was based on the classification of Fries (1847-1848). It contains 33
species in eight sections. Eleven of the species were new and have been subsequently maintained
by the majority of authors. Eight of them belong to the series Alpina and series Subalpina, and the
account deals essentially only with Upper Teesdale and the Clova and Braemar districts of
Scotland. All the taxa are clearly defined and well represented by herbarium material (BM, CGE).
An account of Hieracium, based on the work of Backhouse, was also published by Babington
(1856) in the 4th edition of his Manual. The descriptions were probably drawn up by Babington
from a set of Hieracia given him by Backhouse and now in CGE. The Preface of both works is
dated 1st May 1856. I am indebted to William Stearn for the following information concerning the
problem of priority between the two works. Backhouse’s Monograph was reviewed in Gard.
Chron., 1856: 304 (3rd May 1856) and Ann. Mag. Nat. Hist. , Ser. 2, 17: 418 (May 1856), and was
received by the Linnean Society on 16th June 1856. Babington’s Manual, 4th ed., was reviewed in
Gard. Chron., 1856: 391 (7th June 1856) as “‘just appeared” and Ann. Mag. Nat. Hist., Ser. 2, 18:
163 (August 1856), and received by the British Museum on 11th June 1856. From this it is
reasonable to accept that Backhouse’s Monograph was published in (April—) May 1856 and
Babington’s Manual, 4th ed., in June 1856. This accords with Babington’s statement, presumably
inserted in the last proof, that he had availed himself of the British Hieracia “lately published by
my friend Mr James Backhouse, Jun.”’. A revised account of the genus, based on Backhouse’s
work, was published ten years later by Sowerby (1878). It adds one new species and four new
varieties.

From 1878 until the end of the century, no more advanced elaboration was shown in any British
Flora, and Hooker (1884) in fact reduced the number of species to ten.

Towards the end of the last century a number of amateur botanists became interested in
Hieracium. Foremost of these was F. J. Hanbury, who published a number of papers in the Journal
of Botany (London) (Hanbury 1888, 1889, 1892, 1893, 1894a,b) which culminated in 1894 with a
list of all the taxa known to him in the British flora, 104 species and 114 varieties. Hanbury himself
described as new 24 species and over 30 varieties. Although some of his descriptions were grossly
inadequate, his herbarium (BM) was one of the finest ever made, and there is no difficulty in
interpreting his taxa. Between 1889 and 1898 Hanbury started publishing An illustrated monograph
of the British Hieracia, with large plates reproduced from water-colour drawings by Miss Gulielma

366 P. D-ASELL

Lister. It was unfortunately never finished. The least satisfactory of Hanbury’s works is the account
with Miss R. F. Thompson of Hieracium in the 9th edition of Babington’s Manual of British Botany
(Hanbury & Thompson 1904), in which 97 species were described. Although the details of the
descriptions are perfectly true as regard to facts, they are too general and do not contrast the
important features and are utterly useless for the purpose of identification or classification.
Hanbury referred many of his specimens to the Scandinavian botanists C. J. Lindeberg and M.
Elfstrand for comment, and it is possibly this early contact that led British students of the genus to
follow the Scandinavian school of thought as opposed to that which developed in central Europe.

Simultaneous with the work of Hanbury was that of two brothers, the Revs E. F. and W. R.
Linton. Efficient field botanists, they published accounts of their new species and varieties which
were much more detailed and accurate than those of Hanbury. As well as papers in the Journal of
Botany (London) (E. F. Linton 1891, 1897, 1911; E. F. & W. R. Linton 1893; W. R. Linton 1890),
one of them brought out An account of the British Hieracia (W. R. Linton 1905). It contained 124
species, one subspecies and 135 varieties and forms. It was the most generally useful work until
Pugsley (1948). In total the brothers Linton described 16 new species and 37 new varieties. In 1896
they began the issue of a Set of British Hieracia which extended to 160 numbers by 1901, and an
additional fascicle of 25 numbers was sent out in 1906. The sets were filled partly by wild specimens
and partly by cultivated ones. This, together with the fact that some numbers were made up of
plants from different localities, means that the sets should be used with care, as the same number
sometimes contains more than one species.

Another connoisseur of hawkweeds was the Rev. A. Ley, who concentrated mainly on the
Welsh species. He published several papers describing new taxa (Ley 1895, 1898, 1899, 1900, 1901,
1907, 1909, 1910), the most important being on Brecon and West Yorkshire Hawkweeds (Ley
1909). W. H. Beeby collected and described some of the first new species in the series Alpestria
from Shetland (Beeby 1891a,b, 1908), and the Rev. W. H. Purchas named a new species from
Derbyshire and a new variety from the Wye Valley (Purchas 1895, 1899). Perhaps the most critical
of all British field botanists, the Rev. E. S. Marshall, described three new species and collected a
wealth of interesting material especially from Scotland (Marshall 1892a,b, 1894, 1913). The finds
made on these Scottish journeys are published, sometimes with F. J. Hanbury or W. A. Shoolbred,
in numerous papers in the Journal of Botany (London).

While all this activity was going on in the field, F. N. Williams was sitting in the British Museum
or Kew producing his Prodromus Florae Britannicae. His account of Hieracium (Williams 1902,
1903) contains 75 species and many varieties. The descriptions, though of considerable length,
show no salient or contrasting characters and the species are somewhat grotesquely arranged
within the sections. It is quite clear Williams knew little about the living plants. He was, however,
the first British botanist to find foliar glands in the series Alpina and to use the character of the
receptacle pits.

On the Continent of Europe, A. Jordan, C. Arvet-Touvet and H. Sudre had described many
new species from France, while C. J. Lindeberg, J. P. Norrlin, M. Elfstrand and H. Dahlstedt were
busy in Scandinavia. Although some of these authors started by giving their new taxa the rank of
subspecies, they usually finished up by accepting them as species. In central Europe, however, the
enormous works by Naegeli & Peter (1885-1886) and Zahn (1921-1923) were adopting a system of
species principales each with numerous subspecies which are taxonomically equivalent to the
species of British and Scandinavian botanists. The basic difference between the two schools of
thought, which still exists, is that the central European botanists believe they can tell how their
taxa originated and the British and Scandinavian botanists do not think this is possible. Zahn’s
(1921-23) monumental monograph in Engler’s Pflanzenreich describes 756 species principales in
four subgenera with a vast number of subspecies. The index includes approximately 18,000 names.
There are 47 sections, but a great many of the main taxa are regarded as species intermediae (each
with numerous subspecies) which are considered to be intermediate between sections. Although an
invaluable source of reference to the dedicated hieraciologist, it is useless for identification by any
normal procedure as the keys only cover typical members of the group, and species intermediae are
not included in the keys at all. Zahn’s knowledge of the British Hieracia was almost entirely based
on Linton’s Set of British Hieracia, for the contents of which he coined many new names.

Between the two World Wars very little was published on British Hieracia. H. H. Johnston and
G. C. Druce sent many specimens to H. Dahlstedt (then more interested in Taraxacum) and K. H.

HISTORY AND CLASSIFICATION OF HIERACIUM 367

Zahn (who had finished his monographic work). Numerous new subspecies and species were
described by them based on poor specimens from a flora they did not know well, and most of these
names have disappeared into synonymy. Two lists were produced in the 1920s, the first by Roffey
(1925) and the second by Druce (1928). Neither author knew much about Hieracium and both tried
to combine the work of British botanists with that of Zahn. Many new names were introduced to
the British literature.

This brings us to the fine monograph produced by Pugsley (1948). It provided detailed
descriptions of 260 species and brought together names used by all previous British authors as well
as linking them up with Continental works. Pugsley described 62 new species and 53 new varieties
and was responsible for ten new names and 36 new combinations.

Over the last 35 years, I have, in co-operation with Dr C. West, examined most of the specimens
cited by Pugsley and seen the majority of the species in the field, many in their type localities.
Some of Pugsley’s species have been reduced to synonymy and more new species have been
described. In Dandy (1958) we brought Pugsley’s species nomenclaturally up to date, and later
mapped all the species then known to us (Sell & West 1968). The new species and nomenclatural
corrections were published in separate papers in Watsonia (Sell & West 1955, 1962, 1965, 1967).

Work on the distribution of the Hieracia has continued, particularly by A. G. Kenneth and A.
McG. Stirling, who published an account of the hawkweeds of western Scotland (Kenneth &
Stirling 1970). M. McC. Webster included 88 species in her Flora of Moray, Nairn and East
Inverness (Webster 1978), and Dr West and I endeavoured to make a key to them. In 1981, a
Hieracium Study Group was started within the B.S.B.I., with 36 members. It has so far distributed
six series of notes. One of the most obvious results of recent work is that when areas such as Ross
and southern Scotland, -which have previously received little attention, are investigated, they
produce a number of new species, whereas more intensive investigation of Cardiganshire, v.c. 46,
by Chater (1984) has not produced any new species, but has added considerably to the number of
species known in the county. D. J. Tennant has devoted much time over the last 20 years working
on the species of the series Alpina. Not only has he seen all of them in the field, but he has
cultivated most of the species in his garden. The writing up of this group is in progress.

A word or two needs to be said about the genus in Ireland. Ireland has produced no
hieraciologist of its own. R. L. Praeger collected many specimens on his travels (DBN) and most of
the British botanists interested in Hieracium made at least one visit to Ireland. The material as a
whole is not very well preserved and good modern specimens are badly needed for a re-appraisal of
the Irish species. In particular, the rediscovery of H. hartii (F. J. Hanb.) P. D. Sell & C. West is
desirable. It is a species which, when originally described by Hanbury (1892), was said to be in
great profusion on Slieve League, Donegai.

CLASSIFICATION

When preparing the accounts of Hieracium for Flora Europaea (Sell & West 1976) and Flora of
Turkey (Sell & West 1975a), Dr West and I had to consider the classification of the genus
throughout its range and to decide which of the two schools of thought to adopt as regards the
definition of species.

Zahn (1921-1923) has four subgenera. One of these, Pilosella Hill, we had treated as a separate
genus in Sell & West (1968), and continued to do so in the Flora of Turkey (Sell & West 1975a).
For Flora Europaea, the editors, Tutin et al. (1976), after consultation with regional advisers,
decided against this treatment, preferring continuity with the treatment of Zahn. Our separation of
Pilosella from Hieracium was based on the characters in the following descriptions.

HIERACIUM L., Sp. Pl., 799 (1753); Gen. Pl., 5th ed., 350 (1754).
Lectotype: H. murorum L. (vide N. L. Britton & A. Brown, Ill. Fl. N. United States, 2nd ed., 3:
328 (1913)).

Perennial herbs, the descending or oblique rootstock with thick fibres, not stoloniferous. Stems
usually solitary, sometimes few. Basal leaves, when present, in a rosette, sometimes withered at
anthesis; cauline leaves none to numerous; all entire to deeply dentate, at least the basal usually

368 P. D. SELE

distinctly petiolate. Margins of receptacle pits more or less dentate, sometimes with more or less
long fimbriae. Ligules yellow, sometimes dingy yellow, rarely greenish. Achenes 3-5 mm, 10-
ribbed, ribs apically confluent in an obscure ring; pappus hairs in two series, unequal. About
10,000 species, probably mainly apogamous, described from Europe, N.W. Africa, N. and W. Asia
and N. America.

Hieracium murorum was first described by Linnaeus (1753, p. 802). His diagnosis, Hieracium
caule ramoso, foliis radicalibus ovatis dentatis; caulino minori, is taken from the Hortus
Cliffortianus no. 6 on page 388 without change. There is a sheet in the Hortus Cliffortianus (BM)
labelled ‘‘Hieracium macrocaulon, hirsutum, folio rotundiore” and in an unknown hand ‘“‘murorum
6’’. The sheet contains a stem with a bract and an inflorescence of 20 capitula, a second stem with
four capitula and one large, ovate, nearly entire leaf, and two unattached basal leaves which are
large, ovate and nearly entire. The two stems belong to different species. The one with 20 capitula
certainly belongs to the aggregate species Hieracium murorum as understood by most Continental
botanists, but it is too inadequate to identify as to segregate species. The second stem I cannot
place at all. It does not belong to the aggregate species Hieracium murorum. I would not like to say
to which stem, if either, the two basal leaves belong. I have selected the stem with 20 capitula as
the lectotype of Hieracium murorum L. It will enable the Continental botanists to continue its
usage in the aggregate sense. As the type species of the genus it does not disagree with Linnaeus’
generic diagnosis. A second sheet in the Hortus Cliffortianus labelled ‘‘Hieracium murorum
laciniatum minus, pilosum” is referable to Hieracium murorum f of Linnaeus (1753, p. 803).

PILOSELLA Hill, Brit. Herb., 441 (1756).
Holotype: Pilosella major repens hirsuta Bauhin = P. officinarum C. H. & F. W. Schultz
(Hieracium pilosella L.).

Perennial herbs, with horizontal or oblique rhizomes and persistent rosettes of leaves from the
axils of which prostrate leafy, or underground scaly stolons are often developed. Stems (1—) few to
numerous. Leaves entire or slightly denticulate, never distinctly petiolate, often all basal; cauline
leaves when present usually small. Margins of receptacle pits shortly dentate. Ligules yellow, outer
sometimes with a red stripe on outer face, rarely reddish. Achenes up to 2:5 mm, 10-ribbed, each
rib projecting to form a crenulate truncate apex; pappus hairs in 1 series with a few shorter than the
rest. 18 species, probably mostly sexual, in Eurasia and N. W. Africa. Hybrids between most
species that grow together have been recorded.

The two remaining subgenera of Zahn’s monograph are mainly American, with isolated species
in S. Africa and southern Europe. Jeffrey (1966) concluded that the S. African species of subgenus
Stenotheca Fries, H. capense L., should really belong to the genus Tolpis Adanson, where it was
placed by Schultz-Bip. in 1861; and in Flora Europaea (Sell & West 1976), H. staticifolia All. of
southern Europe was also included in Tolpis as Schultz-Bip. had earlier concluded. Whether Tolpis
is the correct place for these species or whether they should be in a separate genus (Chlorocrepis
Griseb. is available), West and I were not certain, but we considered they should not be included in
Hieracium.

The American species of the subgenera Stenotheca and Mandonia Arvet-Touvet bear no
resemblance in general facies to the majority of species in Hieracium proper, but they fit in much
better with the facies of species included in Crepis. If the generic descriptions in Zahn’s
monographic treatment of Hieracium and Babcock’s (1947) description in his monographic
treatment of Crepis L. are compared it will be seen there are no characters which will absolutely
distinguish the two genera. In Europe and western Asia, the main area of diversity of the two
genera, they can be readily separated by the arrangement of the involucral bracts and their general
facies. In Hieracium the involucral bracts are in a graduated series, in Crepis there is an inner row
of long bracts and an outer row of lax, short bracts giving the appearance of a cup and saucer. In all
Hieracium species the pappus is discoloured and rather stiff, and in Crepis it is mainly white and
soft, but in some species (including C. paludosa) it is like Hieracium. In Hieracium the apex of the
achene is always broad and truncate, while in Crepis it is often narrowed and sometimes beaked.
The American species of Hieracium of the subgenera Stenotheca and Mandonia have the general

HISTORY AND CLASSIFICATION OF HIERACIUM 369

facies and involucral bracts of Crepis and the pappus of Hieracium. The achenes are sometimes
narrowed or shortly beaked at the apex. The basic chromosome number of Hieracium is 9, a
number not recorded in Crepis, although the basic number of that genus varies from 3 to 13. Those
American species of Hieracium subgenus Stenotheca whose chromosomes have been counted have
a base number of 9. Nevertheless, on morphological grounds they are best placed in Crepis, which
includes great morphological variation and has by some authors been split into several genera. This
leaves Hieracium more clearly defined, and including only the subgenus Eu-Hieracium of the
classification of Zahn.

The only entity below the rank of genus that has any useful meaning is that which the
Scandinavian and British botanists call a species and the central European botanists call a
subspecies. They are not subspecies in the normal sense of the word (that is, either geographically
or ecologically discrete taxa) as several can grow intermingled, nor is the near relationship of all
the subspecies included by Zahn under a species principalis certain. What is known about the
subspecies of Zahn’s species principales suggests that most are apogamous triploids and tetraploids
which almost certainly have a hybrid origin, as do those subspecies within species intermediae
considered by him to be intermediate between sections. The present known diploids are few
(Merxmiiller 1975) and most are in southern Europe. Even if all the chomosome numbers of the
taxa were known, it is doubtful if a workable phylogenetic classification could be suggested as some
at least of the ancestral species will have died out. As it is, the greater part of any suggested
phylogenetic relationship in Hieracium is pure guesswork and it is better to group the taxa
according to their morphological affinities. When several taxa grow together in one locality, all the
specimens can be put into these taxa: there are no intermediates. It is therefore better to give all
such taxa binomials and regard them as species. Any grouping of the species results in
intermediates between groups in many directions. It is possible to make major groups into which
large numbers of species fall absolutely and many others fall on a majority of characters. As these
are mainly a collection of apogamous taxa which may or may not have any phylogenetic
relationship it is best to give them the lowest possible rank, that of series. An added incentive to
use this rank is that the epithets used for these groups by Fries (1847-1848) and most later authors
can in the main be retained.

In Flora Europaea (Sell & West 1976), we divided the account of subgenus Hieracium into 38 of
these series, but we did not give them names in deference to our central European colleagues who
still wished to use Zahn’s classification. It was not practical to include all the vast number of species
that had been described and some grouping was therefore necessary. The most sensible thing to do
was to use most of Zahn’s species principales and intermediae as groups and arrange them within
our series. It sometimes proved difficult to correlate our classification with that of Zahn, as we had
some groups which were different from those of Zahn and we transferred some segregates to other
groups. These groups are in no sense comparable with ordinary sexual species. We did, however,
try to find the correct name for each group (contrary to the statement by Jeffrey (1978, p. 499)) for
those who wished to call them species. To use ‘group’ and not ‘sensu lato’ with an authority was the
decision of the editors, not the authors. Those who wish to use Zahn’s complete classification will
have to make many hundreds, if not thousands, of new combinations at all ranks to bring it up to
date. Dr West and I pointed out some of the problems involved (Sell & West 1975b).

ACKNOWLEDGMENTS
Although it has fallen to my lot to write this paper, it is in fact the result of 30 years of joint work

with the late Dr Cyril West. I am indebted to A. O. Chater, D. J. Tennant and S. M. Walters for
reading through the paper and making useful comments.

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supplement to the atlas of the British flora, pp. 75-134. London.

HISTORY AND CLASSIFICATION OF HIERACIUM S71

SELL, P. D. & WEsT, C. (1975a). Hieracium L. and Pilosella Hill, in Davis, P. H., ed. Flora of Turkey, 5:
696-763. Edinburgh.

SELL, P. D. & WEsT, C. (1975b). Hieracium L., in HEywoop, V. H., ed. Flora Europaea. Notulae systematicae
ad floram Europaeam spectantes. No. 19. Bot. J. Linn. Soc., 71: 256-267.

SELL, P. D. & West, C. (1976). Hieracium L., in Tutin, T. G. et al., eds. Flora Europaea, 4: 358-410.
Cambridge.

SoweErsy, J. (1878). English Botany, 3rd ed., vol. 5. London.

TutTin, T. G. et al. (1976). Flora Europaea, 4. Cambridge.

WessTER, M.McC. (1978). Flora of Moray, Nairn & east Inverness. Aberdeen.

WILLIAMS, F. N. (1902, 1903). Hieracium in Prodromus florae Britannicae. Brentford.

Zaun, K. H. (1921-1923). Compositae—Hieracium, in ENGLER, H. G. A., ed. Das Pflanzenreich, TV. 280.

Leipzig.
(Accepted October 1986)

i)

CAL om

bi

Watsonia, 16, 373-382 (1987) 373

A biometric study of the Arenaria ciliata L. complex
(Caryophyllaceae)

M. B. WYSE JACKSON and J. A. N. PARNELL

School of Botany, Trinity College, Dublin 2, Ireland

ABSTRACT

25 morphological characters were recorded for 178 herbarium specimens belonging to the Arenaria ciliata L.
complex. Multivariate analysis suggested that they be referred to three species, A. ciliata L., A. norvegica
Gunnerus and A. gothica Fries. A. moehringioides J. Murr was included as a subspecies of A. gothica, A.
gothica subsp. moehringioides (J. Murr) M. B. Wyse Jackson & J. Parnell, comb. nov. Plants of A. gothica
subsp. gothica from Sweden and Switzerland were found to be morphologically distinct and were treated as
varieties i.e. A. gothica subsp. gothica var. gothica for the Swedish plants and A. gothica subsp. gothica var.
fugax (Gay ex Gren.) M. B. Wyse Jackson & J. Parnell, comb. nov. for the Swiss plants.

Four of the five subspecies of A. ciliata recognized by Jalas & Suominen (1983) were maintained, as was the
variety A. ciliata subsp. ciliata var. hibernica (Ostenf. & O. C. Dahl) G. C. Druce. Multivariate analysis
confirmed the inclusion of A. ciliata subsp. bernensis in A. ciliata sensu stricto and not A. gothica subsp.
moehringioides as had been previously suggested.

A. ciliata subsp. polycarpoides (Rouy & Fouc.) Br.-Bl. was seen to intergrade with A. gothica subsp.
moehringioides and does not warrant separate status.

INTRODUCTION

Arenaria L. is a medium sized genus of about 160 species which occurs mainly in arctic and
temperate regions of the northern hemisphere. It is usually placed in the sub-family Alsinoideae of
the Caryophyllaceae. This sub-family is characterized by its opposite leaves, absence of stipules,
usually well-developed petals and free or only partly fused sepals. In the most recent
comprehensive treatment, McNeill (1962) split Arenaria into ten subgenera, the most widespread
of which is subgenus Arenaria. This contains eleven sections including the pan-arctic section
Rariflora Williams, the type species of which is Arenaria ciliata L.

A. ciliata sensu lato has a confused taxonomic history. The very wide variation both in
morphology and in the level of ploidy together with geographic isolation has led to the evolution of
a number of closely related taxa which together form the so-called A. ciliata complex. The complex
has been taken to include as many as seven and as few as two species. However as a basis for this
study we follow Hess, Landolt & Hirzel (1967) and take it to include the four species A. ciliata
sensu stricto, A. moehringioides Murr (= A. multicaulis L.), A. norvegica Gunnerus and A.
gothica Fries.

A. ciliata s.s. occurs through the Alps, the Tatra (as subsp. tenella (Kit.)Br.-Bl.) in the
Carpathians, the northern Apennines and with single stations in the Dinaric (western-central
Yugoslavia) and northern Albanian Alps. North of the Tatra it occurs only in north-western
Ireland (subsp. hibernica Ostenf. & O. C. Dahl) and, as subsp. pseudofrigida Ostenf. & O. C.
Dahl, in Finland, Norway (extreme north), Spitzbergen, the Russian arctic and eastern Greenland.
A. moehringioides has a more western distribution than A. ciliata s.s., occurring in northern Spain,
the Pyrenees, Jura, and Alps to Vorarlberg and northern Italy. A. norvegica is a northern species
occurring throughout Iceland and scattered through the mountains of Norway and northern
Sweden. It also occurs in Shetland, Scotland and western Ireland. The Irish station is the
southernmost for the species (53°N), but it has not been seen there for 25 years. A. norvegica is
also found in western Yorkshire in England as subsp. anglica Halliday. A. gothica is a species with

374 M. B. WYSE JACKSON AND J. A. N. PARNELL

a very disjunct distribution. It occurs in southern mainland Sweden and on the island of Gotland
and also in the Swiss Jura. All members of the complex are mountain plants, except for the lowland
A. gothica, which mostly grow in open habitats on base-rich soils.

A. ciliata s.s. is a low-growing herbaceous chamaephyte which characteristically has small (<10
mm long), entire, non-fleshy, oblanceolate or spathulate leaves which are ciliate at least at the
base. The flowers are usually solitary or paired (rarely in threes), with white petals that are 1-5-2
times as long as the sepals.

Despite the splitting of the A. ciliata complex, A. ciliata s.s. remains a very polymorphic species.
Thus it in turn has been divided into a number of geographically disjunct subspecies. Jalas &
Suominen (1983), for example, recognize the following: A. ciliata subsp. ciliata, subsp. bernensis
Favarger, subsp. hibernica Ostenf. & O. C. Dahl, subsp. pseudofrigida Ostenf. & O. C. Dahl and
subsp. tenella (Kit.) Br.-Bl.

The first two subspecies, A. ciliata subsp. bernensis and subsp. ciliata, are in fact the only two
subspecies which have partially overlapping ranges, both occurring in the Bernese Alps in
Switzerland. Favarger (1960, 1963) separated the former from subsp. ciliata on morphological,
cytological and ecological differences. However the occurrence of some intermediate populations
has led Jalas & Suominen (1983) to question their separate status.

The other subspecies of A. ciliata s.s. have also posed taxonomic problems. Halliday (1960a)
recognized that subsp. pseudofrigida and subsp. tenella are morphologically very similar. They also
have the same chromosome number of 2n = 40 (Horn 1948; Favarger 1963). In their turn, other
workers have noted that these two subspecies are similar to subsp. ciliata and subsp. hibernica (see
Clapham 1962; Chater & Halliday 1964).

It is clear that although the A. ciliata complex has been split into a number of taxa the
distinctions between them are still somewhat unclear. Our aim was to try to clarify the variation
pattern of the complex in general, whilst concentrating especially on four subspecies of A. ciliata
s.S., i.e. subsp. bernensis, subsp. ciliata, subsp. hibernica and subsp. pseudofrigida.

Despite the obviously complex nature of the variation pattern shown by the A. ciliata complex,
all previous accounts of it have relied on orthodox taxonomic techniques. We felt that a new
approach was needed. The approach we chose involved the use of multivariate statistics to analyze
measurements made on herbarium specimens.

MATERIALS AND METHODS

178 pressed specimens of the above four species were identified using the appropriate regional
Floras and scored for 25 morphological characters (Table 1, Fig. 1). The characters chosen were a
combination of those used in appropriate local Floras or descriptions of the taxa, together with
characters which we felt would give optimal species or subspecies separation.

Some characters measured were found not to be as useful as initially hoped. Characters 19-25
(see Table 1) are qualitative and can thus only be used in a multivariate analysis in averaged or
percentage form to show general tendencies in the data. Character 15 (seed length) could not be
recorded for all plants examined and so is only used when average values are being considered.
The ratios (Characters 16-18) were found, like Characters 19—25, to be useful only in expressing
the broadest variation patterns in the data. The only specimen of A. ciliata subsp. tenella that we
saw was too young to be of use in this study. Measurements of A. norvegica subsp. anglica were
made on a single cultivated specimen in the Trinity College Botanic Garden (Dublin). This taxon
was included only in the analysis presented in Fig. 4.

The data were standardised for use in a principal components analysis (PCA) program
(BMDP4M). Bivariate scatter diagrams showing the morphological relationships of the plants
scored are shown in Figs. 2, 3, 4 & 5. These scatters form the basis for all further discussion of the
A. ciliata complex in this paper.

RESULTS AND DISCUSSION

Not all of the characters measured were equally useful at all stages of the analysis. Thus, where
very general trends in the data were being considered and where the data were or could be

STUDY OF ARENARIA CILIATA 375

TABLE 1. MORPHOLOGICAL CHARACTERS RECORDED FOR MULTIVARIATE ANALYSES
The figures in brackets refer to the number of measurements made for each character on each specimen. See
Fig. 1 for illustrations of some of the characters.

. Petal length (3)

. Sepal length (3)

. Leaf length (3)

. Leaf width (3)

Flower diameter (3)

. Pedicel length (3)

No. of flowers per cyme (3)

. Maximum length of flowering shoot (1)
. No. of flowering shoots (1)

10. No. of sterile shoots (1)

11. No. of styles per flower (3)

12. No. of capsule teeth (3)

13. Length of leaf ciliation (3)

14. Length of sepal ciliation (3)

15. Seed length (5)

16. Petal length/Sepal length

17. Leaf length/Leaf width

18. Pedicel length/Sepal length

19. Plants perennial or annual

20. Lateral nerves of leaf conspicuous or not (3)
21. Lateral nerves of sepals conspicuous or not (3)
22. Backs of sepals pubescent or not (3)
23. Leaf midrib distinct or not (3)

24. Stems pubescent or not (3)

25. Pedicels pubescent or not (3)

WOIDNAARWNE

IS

Figure 1. Some of the characters used in the multivariate analyses. Top row |.-r.: Petal, Sepal, Leaf. Bottom
TOW l.-r.: Seed, Inflorescence of A. ciliata s.s.

376 M. B. WYSE JACKSON AND J. A. N. PARNELL

©) oF

14

PE2

=-2 -1

0
PC
FicureE 2. Principal Components Analysis of individual specimens (excluding A. norvegica subsp. anglica) in
the A. ciliata complex. Components 1 (PC1) and 2 (PC2) are illustrated. PC1 accounts for 26-2% of the
variation and PC2 for 16-5%. © = A. ciliata subsp. polycarpoides, O = A. ciliata subsp. hibernica, @ = A.
ciliata subsp. pseudofrigida, @ = A. ciliata subsp. ciliata, VV = A. ciliata subsp. bernensis, A = A. norvegica
subsp. norvegica, ® = A. norvegica subsp. anglica (only on Fig. 4), 0 = A. moehringioides, W = A. gothica
(Switzerland), A = A. gothica (Sweden)

considered as percentage or mean data, all 25 characters were used in the analysis (Fig. 4). Where
this was not the case, only the unmeaned data for characters 1-14 were used (Figs. 2, 3 & 5).

GENERAL PATTERN OF VARIATION

Fig. 2 shows the first two Components of the PCA of all 178 individuals in the A. ciliata complex.
Aberrant individuals, usually of exceptional size, lie in the top right of the scatter (numbered 1-5)
and are not further discussed. Fig. 3 shows Components 1 and 3 of the PCA of all 178 individuals in
the complex. Based on these results there is little evidence for strong distinctions between any of

STUDY OF ARENARIA CILIATA 377

3 O

=v, -1 O 1 2 3

PCi

Ficure 3. Principal Components Analysis of individual specimens (excluding A. norvegica subsp. anglica) in
the A. ciliata complex. Components 1 (PC1) and 3 (PC3) are illustrated. See Fig. 2 for explanation of symbols.
PCi accounts for 26:2% of the variation and PC3 for 13-0%.

the species or subspecies. This is somewhat surprising as the characters chosen are those that are
usually used to distinguish taxa in the complex, indicating that the distinctions between them are
not as clearcut as is generally held. However three loose groupings (emphasized in Fig. 2 by the
envelopes surrounding them) can be recognized.

Group (A) towards the bottom left corner of Fig. 2 contains virtually all plants assignable to A.
norvegica. However, though A. norvegica is usually recognized as a separate species, it is apparent
from this scatter that it shows some morphological overlap with A. ciliata subsp. pseudofrigida.
This overlap is also seen in Fig. 3. These two taxa do, however, mostly lie in different areas of this
scatter and are, on the whole, quite distinct.

Group (B) to the right of Group (A) in Fig. 2 is made up almost entirely of plants assignable to
A. ciliata s.s. This group is discussed in more detail below.

Group (C) in the upper middle part of Fig. 2 contains plants of A. moehringioides and A.

378 M. B. WYSE JACKSON AND J. A. N. PARNELL

0 1 2 3
PC1
Figure 4. Principal Components Analysis of meaned data for all taxa in the A. ciliata complex. Components 1

(PC1) and 2 (PC2) are illustrated. See Fig. 2 for explanation of the symbols. PC1 accounts for 28-2% of the
variation and PC2 for 20-0%.

gothica. Three points are evident in relation to the scatter of this group. Firstly, it seems impossible
to effectively separate A. moehringioides and A. gothica. This is supported by Fig. 3 in which A.
gothica from both Sweden and Switzerland grade into A. moehringioides. Secondly, A. ciliata
subsp. polycarpoides (Rouy & Fouc.) Br.-Bl. intergrades with typical A. moehringioides. This is
also seen in Fig. 3 though not as clearly as in Fig. 2. The degree of overlap is large considering the
small sample size. We feel that the weight of evidence is against giving A. ciliata subsp.
polycarpoides separate status in A. moehringioides. Thirdly, it has been suggested by Rouy &
Foucaud (1896), Williams (1898) and Chater & Halliday (1964) that Swiss and Swedish A. gothica
are significantly different. Chater & Halliday (1964), whose observations were based on cultivated
plants, describe Swiss A. gothica as being less ciliate and with larger cymes than Swedish plants of
the same species. Our analysis supports such a difference. In Fig. 2 only a few plants of Swiss A.
gothica show morphological overlap with Swedish A. gothica. In Fig. 3 Swiss A. gothica is in fact
almost completely separated from Swedish A. gothica.

STUDY OF ARENARIA CILIATA 379

B
a
B
[| VV VV,
1 Vv
Vv @ Vv Vv Vv
| B | Vv VVvV
Vv
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ae a B e ev 4
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0 es @ 0e6ee
@ Vv OO
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exe O
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eal
Ficure 5. Principal Components Analysis of individual specimens of A. ciliata s.s. (Group 2 of Fig. 4).

Components 1 (PC1) and 2 (PC2) are illustrated. See Fig. 2 for explanation of the symbols. PC1 accounts for
30:7% of the variation and PC2 for 18-3%.

In order to emphasize the differences between the groups (A to C) in Fig. 2, the data for each
taxon were meaned and a PCA and a Cluster Analysis (BMDP2M) performed.

Fig. 4 shows the first two Components of the PCA of the meaned data for all taxa measured. A
comparison of Figs. 2 & 4 indicates that Fig. 4 emphasizes the differences that occur between and
within groups. We use it to illustrate five points. Firstly, there is evidently a clear difference
between Group 1 (Fig. 4) and the rest of the complex. So although individuals of A. norvegica
show some morphological overlap with A. ciliata subsp. pseudofrigida, on average, these two taxa
are distinct. Secondly, within A. norvegica s.1., the rare taxon subsp. anglica is distinguishable from
subsp. norvegica. Thirdly, A. ciliata s.s. (Group 2 of Fig. 4) can be seen to contain four elements
that show a gradation from subsp. pseudofrigida through subsp. ciliata and subsp. hibernica to
subsp. bernensis. This trend represents a morphological shift from small plants with weakly ciliate
leaves, short petals and small seeds to larger plants with large, strongly ciliate leaves, longer petals

380 M. B. WYSE JACKSON AND J. A. N. PARNELL

and larger seeds. Closer inspection of Fig. 2 also reveals the trend, though naturally it is less clear.
Fourthly, there is a very clear separation between A. ciliata subsp. bernensis and A.
moehringioides. This does not support the suggestion of Welten & Sutter (1982) that the former
should be included in the latter. This separation can also be clearly seen in Fig. 2. Finally, A.
gothica and A. moehringioides (Group 3 of Fig. 4) are clearly separated from the rest of the
complex; though again these two species are not clearly separable from each other. Component 3
in the PCA of the average values accounts for 16.8% of the variation but does not significantly alter
the groupings seen in Fig. 4.

In summary, Figs. 2, 3 & 4 show there to be three principal groups present in the A. ciliata
complex. The first group contains A. norvegica s.1., which may, on average, be distinguished by its
weak leaf and sepal ciliation, fleshy leaves, sparse pedicel pubescence and more or less equally
long petals and sepals. The second group contains A. ciliata s.s. which may, on average, be
distinguished by a combination of its large flowers (9-0—14-5(—15) mm in diameter), long petals
((5:0—)6-0—8-0 mm), few flowers per cyme (1—2(-—3)), usually strongly ciliate leaves, pubescent
pedicels and petals 1-5—2-0 times as long as the sepals. The third group contains A. moehringioides
and A. gothica which may, on average, be distinguished by their small flowers (5-5—8-0(—9) mm in
diameter), short petals (3-5—6-0 mm), greater number of flowers per cyme ((2—)3—-6(—9)) and
leaves c. 2-2-5 times as long as broad. The morphological similarity of these two species is
emphasized by the fact that the most striking difference between them (Character 19, Table 1) did
not effectively separate them in these analyses. Within this group, A. moehringioides, A. gothica
and A. ciliata subsp. polycarpoides intergrade and do not warrant separate specific status. These
analyses do however support separation of Swiss and Swedish A. gothica. The groupings seen in
the PCA scatters were confirmed by the Cluster Analysis.

VARIATION OF A. CILIATA S.S.

Fig. 5 shows the first two Components of a PCA of individuals of A. ciliata s.s. This analysis
provides a clearer resolution of the relationships of the subspecies within A. ciliata s.s. A. ciliata
subsp. ciliata occupies the centre of the scatter and around it are arranged the other subspecies.
Subsp. bernensis lies towards the top right hand corner of the scatter and is clearly separable from
its nearest neighbours, subsp. ciliata and subsp. hibernica. A. ciliata subsp. pseudofrigida is also
well separated from the rest of the subspecies, lying as it does on the left-hand side of the scatter
and showing little overlap with any of the other taxa. Such clear separation however cannot be said
to exist between subsp. hibernica and subsp. ciliata — there is in fact a significant overlap between
them. Certainly these two subspecies are morphologically very similar. This scatter also shows that
subsp. hibernica can be distinguished from subsp. pseudofrigida. We found that subsp.
pseudofrigida has in comparison with subsp. hibernica smaller flowers (c. 9-12 mm in diameter cf.
10-13 mm) and shorter, narrower leaves (3-5 mm X 1-1-5 mm cf. 3-5—6(—7) mm X 1-5-2 mm)
which are never (cf. almost always) ciliate to the tip. They are, however, cytologically and
chemically very similar. For the Irish plant we obtained a chromosome count identical to Horn’s
(1948) count of 2n = 40 for subsp. pseudofrigida. (Halliday’s (1960a) count for subsp. hibernica is
reported as 2n = c.40 in Clapham (1962).) The results of a chemical analysis that we performed on
these two subspecies indicated that they also have an almost identical pattern of leaf flavonoids.
However this scatter suggests that these two subspecies are morphologically less closely related to
each other than they are to subsp. ciliata. Component 3 accounts for 14-0% of the variation but
does not significantly alter the groupings seen in Fig. 5.

Halliday (pers. comm.) has indicated the close relationship of A. ciliata subsp. tenella to subsp.
hibernica. However, both Kitaibel (1814) and Graebner (1919) describe the former taxon as having
leaves ciliate at the base. Based on these descriptions it seems that subsp. tenella is probably less
closely related to subsp. hibernica, which has leaves almost always ciliate to the tip, than it is to
subsp. pseudofrigida which has leaves ciliate at the base.

CONCLUSION
Multivariate analysis of the A. ciliata complex has facilitated the clarification of some of the

relationships of the taxa normally included in it. Thus three groups can clearly be recognized: A.
norvegica s.l., A. ciliata s.s. and A. moehringioides/A. gothica.

STUDY OF ARENARIA CILIATA 381

A. norvegica s.l. has proved to be closely related to, though distinct from, A. ciliata s.s. Two
infra-specific taxa corresponding to the two hitherto recognized subspecies of A. norvegica have
been distinguished in this analysis.

Within the A. moehringioides/A. gothica group it has proved impossible to separate A. gothica
from A. moehringioides. However, the differences in life-history, chromosome number, habit and
geographical distribution of the two taxa warrant their separation at subspecific level. (A. gothica is
an annual or biennial with 2n = 100 and is taller and more erect than A. moehringioides. A.
moehringioides on the other hand is a perennial with 2n = 40.) This solution is of the same type as
that proposed by Halliday (1960b) who included the short-lived A. gothica auct. angl. in A.
norvegica as subsp. anglica. Additionally our analysis failed to effectively separate A. ciliata subsp.
polycarpoides from A. moehringioides and we think that these two taxa should be considered
synonymous. Finally, A. gothica has proved to be variable, and the Swiss plants warrant separation
from the Swedish plants at the varietal level. Swiss A. gothica tends to be taller, to have more
flowers per cyme, less ciliate leaves and slightly larger seeds than Swedish A. gothica. The former
also rarely (cf. commonly) has cilia on the back of the sepals.

Within A. ciliata s.s. there is a general trend from subsp. bernensis (with parts generally large) to
subsp. pseudofrigida (with parts generally small). This latter subspecies has proved to be less
closely related to subsp. hibernica than had been previously thought. This work has also shown that
subsp. hibernica intergrades with typical subsp. ciliata and does not warrant taxonomic recognition
at subspecific level.

LIST OF TAXA AND SELECTED SYNONYMY
1. ARENARIA NORVEGICA Gunnetus, Fl. Norvegica, 2: 144 (1772).

a. Subsp. NORVEGICA
A. ciliata subsp. norvegica (Gunnerus) Fries, Novitiae Florae Suecicae, 2nd ed., 2: 34 (1839).

b. Subsp. ANGLICA Halliday in Watsonia, 4: 209 (1960).

2. ARENARIA CILIATA L., Sp. Pl., 425 (1753).

a. Subsp. CILIATA

i. Var. CILIATA

ii. Var. HIBERNICA (Ostenf. & O. C. Dahl) G. C. Druce in Rep. botl Soc. Exch. Club Br. Isl. , 5:
279 (1919).

Arenaria ciliata L. subsp. hibernica Ostenf. & O. C. Dahl, in Nyt Mag. Naturvid., 55: 216 (1917).

b. Subsp. BERNENSIS Favarger in Ber. schweiz. bot. Ges., 73: 176 (1963).

c. Subsp. PPEUDOFRIGIDA Ostenf. & O. C. Dahl in Nyt Mag. Naturvid., 55: 217 (1917).

A. pseudofrigida (Ostenf. & O. C. Dahl) Juz. ex Schischkin & Knorring in Komarov, Fl. USSR, 6:
537 (1936).

d. Subsp. TENELLA (Kit.) Br.-Bl., Sched. Flor. Raet. exsicc., 10: 279 (1927).

A. multicaulis var. tatrensis Zapal., Consp. Fl. Galic., 3: 42 (1911).

A. ciliata subsp. tatrensis (Zapal.) Favarger in Ber. schweiz. bot. Ges., 73: 174 (1963).

3. ARENARIA GOTHICA Fries, Novitiae Florae Suecicae, 2nd ed., 2: 33 (1839).

a. Subsp. GOTHICA

i. Var. GOTHICA

382 M. B. WYSE JACKSON AND J. A. N. PARNELL

A. ciliata subsp. gothica (Fries) Hartman, Svensk Norsk Excurs.-Fl., 63 (1846).
A. ciliata ‘‘forme”’ A. gothica « gothica (Fries) Rouy & Fouc., pro parte, Fl. Fr., 3: 248 (1896).

ii. Var. FUGAX (Gay ex Gren.) M. B. Wyse Jackson & J. Parnell, comb. nov.

Basionym: Arenaria ciliata B fugax Gay ex Gren. in Grenier & Godron, Fl. Fr., 1: 259 (1848).
A. ciliata c. laxior Gremli, Excurs.-Fl. Schweiz, 3rd ed. 103 (1878).

A. ciliata “‘forme” A. Gothica B jurana Genty ex Rouy & Fouc., Fl. Fr., 3: 248 (1896).

b. Subsp. MOEHRINGIOIDES (J. Murr) M. B. Wyse Jackson & J. Parnell, comb. nov.
Basionym: Arenaria moehringioides J. Murr, Allg. bot. Z. 12: 176 (1906).

A..-ciliata L. var. B, Sp. Pl. 425 (1753).

A. multicaulis L., Syst. Nat. 10th ed. 2:1034 (1759), pro parte, nom. illeg.

A. ciliata “forme” A. polycarpoides Rouy & Fouc., Fl. Fr., 3: 247 (1896).

A. ciliata subsp. polycarpoides (Rouy & Fouc.) Br.-Bl., Sched. Flor. Raet. exsicc., 10: 279 (1927).
A. ciliata subsp. moehringioides (J. Murr) Br.-Bl., Sched. Flor. Raet. exsicc., 10: 279 (1927).
A. ciliata subsp. multicaulis (L.) O. Schwarz, Mitt. thiiring. bot. Ges., 1: 98 (1949), non Arcangeli.

ACKNOWLEDGMENTS

We are grateful to the curators of those herbaria in which we examined specimens: TCD, K and
especially to those that kindly lent specimens: BERN, GB, H, MAF, O, S, TRN, UPS, Z, ZT. M. B.
W. J. was in receipt of a financial assistance grant from Trinity College, Dublin, when he carried
out the biometric work during the period October 1983 to April 1984.

REFERENCES

CuatTerR, A. O. & HALLipAy, G. (1964). Arenaria L., in TuTIn, T. G., et al., eds. Flora Europaea, 1: 116-123.
Cambridge.

CLAPHAM, A. R. (1962). Caryophyllaceae and Arenaria L., in CLAPHAM, A. R., TuTin, T. G. & WARBURG, E.
F. Flora of the British Isles, 2nd. ed., pp. 211-215, 254-257. Cambridge.

FavARGER, C. (1960). Recherches cytotaxonomiques sur les populations alpines d’Arenaria ciliata L. (sens.
lat.). Ber. schweiz. bot. Ges., 70: 126-140.

FAVARGER, C. (1963). Nouvelles recherches sur les populations alpine et carpathiques d’Arenaria ciliata L. sens.
lat. Ber. schweiz. bot. Ges., 73: 161-178.

GRAEBNER, P. (1919). in ASCHERSON, P. & GRAEBNER, P. Synopsis der Mitteleuropdischen Flora, 5(1): 508.
Leipzig.

Hatupay, G. (1960a). Taxonomic and ecological studies in the Arenaria ciliata and Minuartia verna
complexes. Ph.D. thesis, Cambridge University.

Hacuipay, G. (1960b). The identity of Arenaria gothica auct. angl. Watsonia, 4: 207-210.

Hess, H. E., LANDOLT, E. & HIRZEL, R. (1967). Flora der Schweiz, 1: 822. Basle.

Horn, K. (1948). in Love, A. & L6vE, D. Chromosome numbers of Northern plant species. Rit LandbDeild.
Ser. B no. 3: 50.

JALAS, J. & SUOMINEN, J., eds. (1983). Atlas Florae Europaeae, 6: 20-22. Helsinki.

KITAIBEL, P. (1814). in ScHULTES, J. A. Osterreichs Flora 2nd. ed., 1: 662. Wien.

MCNEILL, J. (1962). Taxonomic studies in the Alsinoideae: 1. Generic and infra-generic groups. Notes R. bot.
Gdn Edinb. , 24: 79-155.

Rouy, G. & Foucaup, J. (1896). Flore de France, 3: 246-248. Rochefort and Asniéres.

WELTEN, M. & Sutter, R. (1982). Verbreitungsatlas der Farn — und Blutenpflanzen der Schweiz, 1. Basle.

WILLIAMS, F. N. (1898). A revision of the genus Arenaria. Bot. J. Linn. Soc., 33: 326-437.

(Accepted July 1986)

Watsonia, 16, 383-388 (1987) 383

Notes on a hybrid spearwort, Ranunculus flammula L.
x R. reptans L.

R. J. GORNALL

Botany Department, The University, Leicester, LE1 7RH

ABSTRACT

Hybrid Creeping Spearworts (Ranunculus flammula L. x R. reptans L.) are given the binomial Ranunculus x
levenensis Druce ex Gornall, hybr. nov. The character most useful in distinguishing the hybrid from its parents
is the relative length of the achene beak. In the British Isles, hybrids occur sporadically on a few northern lake
shores; they also occur in southern Scandinavia, northern-central Germany and the Alps. Their presence in the
British Isles is due to hybridization between native R. flammula and plants of R. reptans that have probably
been dispersed to these islands by waterfowl migrating from Iceland, Scandinavia and the northern part of the
U.S.S.R.

INTRODUCTION

Padmore (1957) and Candlish [née Padmore] (1975) concluded that the Creeping Spearworts in the
British Isles are hybrid derivatives of Ranunculus flammula L. and R. reptans L. (Ranunculaceae).
This hybrid hypothesis, which can be traced to Druce (1913), has received substantial experimental
support (Gibbs & Gornall 1976). Since most British collections are of the hybrid, rather than of R.
reptans, it seems desirable to validate the binomial bestowed on them as a nomen nudum by Druce
(1929a,b), so as to provide them with a convenient name.

DESCRIPTION

Ranunculus X levenensis Druce ex Gornall, hybr. nov.
TYPUS: Damp, sandy, and gravelly margin of Loch Leven, Kinross, Aug. 1912, G. C. Druce
(Holotypus: OXF! Isotypi: BM! E! OXF! STA! and probably elsewhere). Specimens of artificially
synthesized hybrids have been placed in LTR and STA.

Hybrida inter Ranunculus flammula L. et R. reptans L. Caulis procumbens, 0.50—2.75 mm diam.,
ad nodos radicans, internodiis saepe arcuatis. Folia basalia plerumque manifesta petiolata, lamina
5—20(—30) mm longa x 1—4(—10) mm lata, lineares vel spathulata usque anguste ovata. Flores
1—2(-6) in inflorescentia, 6—12(—15) mm diam., petalis obovatis, 1.2—3.0(—4.5) mm latis ad
latissimam partem. Achenia ellipsoidea vel ovata, (1.0—)1.2—1.5(—1.7) mm longa (rostro excluso),
in rostrum plus minusve abrupte angustata, (0.15—)0.2—0.4 mm longum, longitudine achenii rostro
c.5-plo longiore. Chromosomatum numerus 2n=32.

Hybrid between Ranunculus flammula L. and R. reptans L. Stems procumbent, 0.50—2.75 mm in
diameter, rooting at the nodes, the internodes often arcuate. Basal leaves usually with a distinct
petiole, the blades 5—20(—30) mm long x 1—4(—10) mm wide, linear or spathulate to narrowly
ovate. Flowers 1—2(—6) per inflorescence, 6—12(—15) mm in diameter, the petals obovate,
1.2—3.0(—4.5) mm wide at the widest point. Achenes ellipsoid to ovate, (1.0—)1.2—1.5(-1.7) mm
long (excluding beak), narrowed more or less abruptly to a beak, (0.15—)0.2—0.4 mm long, the
ratio of beak length/achene length being about 1/5. Chromosome number 2n=32.

384 R. J. GORNALL

TABLE 1. CHARACTERS USED TO SEPARATE RANUNCULUSXLEVENENSIS FROM_ ITS
PARENTS

Values quoted are means, standard errors (in brackets) and sample sizes. All measurements have been made

on wild-collected, herbarium material. Equivalent data on cultivated plants can be found in Gibbs & Gornall

(1976).
R. flammula
Character erect procumbent R. Xlevenensis R. reptans
Stem diameter, mm 3°35(0714) 1.9 (0.08) 1.2 (0.06) 0.9 (0.03)
109 58 81 87
Width of largest 10.2 (0.59) 4.8 (0.47) 1.7 (0.10) 0.9 (0.04)
basal leaf, mm 100 56 79 87
Diameter of largest 13,5 :(0:27) 10.4 (0.38) 9:2:(0:22) 8.8 (0.16)
flower, mm 103 36 64 73
Width of largest 4.6 (0.14) 3:3\(047) 2.5 (0.09) 2.1 (0.06)
petal, mm 103 36 64 73
Achene beak length/ 11.1 (0.67) 11.1 (0:55) 18.7 (0.52) 26.8 (0.84)
achene length, % 28 45 48 16
Habit in cultivation erect erect procumbent procumbent

The hybrid can usually be distinguished from its parents by a combination of the characters shown
in Table 1. Unfortunately, most of these characters are highly plastic, and sometimes it can be
particularly difficult to tell the hybrid from either the procumbent plastodeme of R. flammula (var.
tenuifolius Wallr.) without a cultivation experiment, or from pure R. reptans, although Table 1
shows that the achene beak length/achene length ratio is reasonably diagnostic in both cases.
Extreme difficulties arise in situations where the hybrid is a product of backcrossing to one parent
or where it is a parental-type segregate. In such cases there is no easy answer except arbitrarily to
treat all variants that fall within the established morphological ranges of the parents as pure

TABLE 2. LOCALITIES FOR RANUNCULUSXLEVENENSIS IN THE BRITISH ISLES
Asterisked (*) localities are major arrival centres for migrating waterfowl. Subsidiary centres are indented and
are within c.25 km of the ‘parent’ centre.

LOCH AWE, Main Argyll, v.c. 98: GR 27/1.2 (LTR).
*LOCH OF STRATHBEG, N. Aberdeen, v.c. 93: GR 48/0.5 (E).
*LOCH LEVEN, Fife, v.c. 85: GR 36/1.9, 37/1.0 (BM, E, STA).
Lindores Loch, Fife, v.c. 85: GR 37/2.1 (STA)?.
Luthrie Water Works, Fife, v.c. 85: GR 37/3.1 (STA)?.
*SOLWAY FIRTH
Loch Ken, Kirkcudbrights., v.c. 73: GR 25/7.6 (E)
Loch Whinyeon, Kirkcudbrights., v.c. 73: GR 25/6.6 (E) (Stewart 1975).
Lochmaben, Dumfriess., v.c. 72: GR 35/0.8 (Scott-Elliot 1896)?.
ULLSWATER, Cumberland, v.c. 70: near Pooley Bridge, GR 35/4.2 (BM); Aira Beck, GR 35/3.1 (BM)?.
Westmorland, v.c. 69: Glencoyne Bridge, GR 35/3.1 (RNG); near Patterdale, GR 35/3.1 (LTR); Pooley Bridge
to Sandwick, GR 35/4.2, (BM)?.
CONISTON WATER, Westmorland, v.c. 69: south shore, GR 34/3.9 (LIV).
WINDERMERE, Westmorland, v.c. 69: Thompson’s Holm Island, GR 34/3.9 (LIV)?.
*LOUGH NEAGH area
Lough Fea, Co. Londonderry, v.c. H40, GR 23/7.8 (herb. Wareham River Lab.).

Intermediate specimens of uncertain identity (usually without fruits) have been collected from the following

localities:

Cumberland, v.c. 70: Bassenthwaite Lake (BM)*, Derwentwater (TCD)?.

Westmorland, v.c. 69: Windermere, western shore, near ferry, GR 34/3.9 (BM, E)*, River Leven, GR 34/3.8
(KDL)* (Martindale 1886).

“19th century collections only; no 20th century material seen.

NOTES ON A HYBRID SPEARWORT 385

40°

Figure 1. The distributions of Ranunculus flammula (..... ), R. reptans ( ) and R. X levenensis
(@) in Europe. Ranges of the parents are taken from Meusel et al. (1965). Localities for hybrids are based on
specimens housed at BM, E and LTR.

species, even if hybrids occur in the same population. The high levels of pollen sterility in natural
hybrids that were reported by Padmore (1957) and Candlish (1975) were possibly due to chance
environmental effects; Gibbs & Gornall (1976) found that hybrids, both synthetic and natural,
form 16 bivalents at meiosis, show at least 88% pollen stainability, and set a full complement of
fruit on cross-pollination.

DISTRIBUTION
The distributions of R. xX Jevenensis and its parents in Europe are shown in Fig. 1. Except in the

British Isles and northern-central Germany, it can be seen that R. X lJevenensis occurs where the
ranges of its parents overlap, viz. in southern Scandinavia and the Alps.

386 R. J. GORNALL

Intermediate specimens from Iceland (BM, STA) need further investigation, as R. flammula is
not known to be present in the Icelandic flora (Léve 1983). With this in mind, the possible
occurrence of the North American R. reptans var. ovalis Torrey & A. Gray in northern Europe
should be considered. Such plants are very similar to R. X_ Jevenensis in gross
morphology.

The distribution of the hybrid in the British Isles is documented in Table 2. The earliest
reference to R. reptans in the British Isles was by J. Hope (Balfour 1901) who noted its occurrence
at Loch Leven in 1764, and there is a fine illustration of the plant from that locality on the
frontispiece of Flora Scotica (Lightfoot 1777) — labelled R. reptans but clearly referable to R. x
levenensis. Padmore (1957) believed there to be no pure R. reptans left in Britain, but as I
shall suggest later, I think it is likely to occur, at least intermittently. If, however, R. reptans is not
at present a regular member of our native flora, this raises the question of how the hybrid arose.

ORIGIN OF HYBRIDS IN THE BRITISH ISLES

There are three possible modes of origin for R. X Jevenensis in the British Isles: a)
hybridization between native populations of R. reptans and R. flammula; b) introduction of foreign
R. reptans and subsequent hybridization with native R. flammula; or c) dispersal of hybrid plants
into these islands.

Padmore (1957) favoured hypothesis (a), believing that it was ‘‘probable that R. reptans was
present in these islands in the Late Glacial and that relict populations survived at Loch Leven and
Ullswater and became modified at a later date by hybridization’’, leaving no genetically pure R.
reptans in the British Isles. However, two pieces of evidence argue against this interpretation.
Firstly, there is no unequivocal fossil evidence for the existence of R. reptans in the British Isles
during the Late Glacial, unlike R. flammula which has a good fossil record dating from as far back
as the Anglian stage (Godwin 1975). Secondly, both Loch Leven and Ullswater were fully covered
by ice sheets during the last glaciation (Godwin 1975) and therefore these stations cannot be
considered as refugia, although the species may have advanced there from the south, following the
retreating ice.

The alternative hypotheses, (b) and (c), involve long-distance dispersal (Gibbs & Gornall 1976).
Creeping Spearworts occur in a few, scattered localities, mainly in the north-east and points due
south; they are strikingly absent from the shores of lochs in the northern and western Highlands,
and from most of Ireland. In the case of the Irish and Scottish localities at least, the sites of R. x
levenensis/R. reptans correspond to, or are very close to, major arrival centres for waterfowl
migrating from Iceland, Scandinavia and the northern part of the U.S.S.R. (Table 2, Fig. 1). The
birds arrive in north-eastern, rather than north-western, Scotland, and progress southwards
through the central lowlands to the Solway Firth and suitable points beyond (Thom 1969; Boyd &
Ogilvie 1969; M. Owen, pers. comm. 1985). This provides good, albeit circumstantial, evidence in
support of the case for long-distance dispersal. The presence of R. X levenensis at the
Cumbrian stations is a little harder to explain since, at least at present, the English Lakes are not
important haunts of migratory waterfowl. However, they are likely to receive visits from stray
birds roosting in the Solway Firth area; and in the past, they may have played host in the autumn to
migrant Whooper Swans from Iceland (M. Owen, pers. comm. 1985).

Because Loch Leven has a well-studied waterfowl population, it is appropriate to use it as an
example to elaborate on some additional points relating to long-distance dispersal. The loch is a
major arrival point in Scotland for waterfowl and, at the peak of the autumn migration, it holds
nearly 20,000 birds (Thom 1969; Allison & Newton 1974). R. reptans could be imported from
different countries, corresponding to the departure points of the birds (Table 3). R. reptans is in
fruit in September—October, and this fact, together with the data on numbers, arrival times and
food preferences of the birds, suggests that the principal carriers of the species in recent years are
likely to be Pink-footed Geese, Mallard and Wigeon (Table 3). The bulk of the Greylag Geese
arrive during December, too late to have fed recently on achenes of R. reptans, which by then
would have fallen from the plants.

According to Jones & Gillmor (1955), Pink-footed Geese leave central Iceland, where they feed
on graminoids and seed-heads (Gardarsson 1972), in huge skeins as soon as the first snows fall.

NOTES ON A HYBRID SPEARWORT 387

TABLE 3. RECENT AUTUMN POPULATIONS OF THE MAIN, MIGRANT VEGETARIAN
WATERFOWL AT LOCH LEVEN, FIFE
Compiled from Allison & Newton (1974) and Thom (1969).

Date of first Date of

Species Origin*® Peak nos. or main influx peak nos.

Pink-footed Goose Is Gd 10,000-—12,500 Sept.—Oct. Sept.—Oct.

Greylag Goose Is 4000-5 ,000 ? Dec.—Feb.

Mallard Is No Su 2,200-3,500 Aug. Late Sept.
Ge Be Ho

Wigeon Is mainly 1,000-2,000 Late Sept. Jan.
No Su Fe Rs

Pochard Da Europe 500-2 ,500 Aug. Oct.
Rs (B)

Tufted Duck Is No Su Da up to 4,000 Mar.—Apr. Late Sept.
Ge Ga Hb

Teal Is No Su Po 300-—1,200 Aug. Sept.—Oct.
Ho Be Ga Rs

Goldeneye No Su (north) up to 1,000 Sept.—Nov. Mar.—Apr.

Shoveler Hs Ho Be Ge up to 650 Aug. Late Sept.
No Su Da

Whooper Swan i up to 430 Oct. Nov.

Goosander No Su Da up to 150 Oct. Feb.—Mar.

Pintail Mainly Is 50 Late Aug. Sept.—Oct.

* Territory abbreviations as in Tutin et al. (1964-1980); Gd=Greenland.

Some birds arrive in Britain the same day, others the day after departure; if ingested, spearwort
achenes are therefore likely to be still inside the alimentary tracts of the birds on arrival at Loch
Leven (M. Owen, pers. comm.). Vlaming & Proctor (1968) have provided evidence that some
seeds can remain in a Mallard’s gut for up to three days and be viable on excretion. At Loch Leven,
Newton & Campbell (1970) observed that several thousand Pink-footed Geese arrive within two
days of the first influx, and assemble on a mud bank in a few centimetres of water off the northern
shore of St Serf’s Island. Initially they drink and sleep on this mud bank, sometimes wading into
deeper water, splashing and preening for long periods. It is during this initial period that any
achenes present are likely to be excreted from the gut and/or washed from the body into the Loch.
Within a day or so, many of the geese proceed southwards to the Solway Firth and beyond.

The distribution of Creeping Spearworts in the British Isles is therefore likely to depend on the
arrival of a bird, carrying ripe fruits, at a place with a suitable habitat. To some extent then,
occurrences will be a matter of serendipity. However, where major migration centres have suitable
habitats, then R. reptans and/or its hybrid with R. flammula can be expected to occur in greater
numbers. This is the case in Scotland, where Loch Leven and the Loch of Strathbeg are
strongholds. Neighbouring freshwater bodies can also be expected to support the plants, although
in smaller numbers and possibly on a less permanent basis. Similar long-distance dispersal may be
involved in the origin of the hybrids in northern-central Germany.

Since the potential for long-distance dispersal has existed for about 10,000 years, i.e. the length
of time that the waterfowl migration routes have been in existence (M. Owen, pers. comm. 1985),
the introduction of spearworts is likely to be a repeated event. Given the northern origins of most
of the migrating waterfowl, I suspect that pure R. reptans is imported, rather than the hybrid,
which is both rarer and more southerly in its distribution. Indeed, a few collections referable to R.
reptans have been made from Loch Leven (e.g. Salmon, 8th September 1896) and Ullswater (e.g.
Bickham, 6th August 1917) (both BM, E). It is therefore my opinion that R. reptans is an
intermittent member of the British flora and that, on arrival, it quickly interbreeds with R.
flammula, or existing hybrids, to produce progeny which, on the loch shore, frequently closely
resemble R. reptans.

However, since R. reptans is perennial, stoloniferous and partially self-fertile (Gibbs & Gornall
1976), there ought to be a population build-up.of the species as well as of the hybrid, but this does

388 R. J. GORNALL

not seem to happen. Possibly one explanation is that the pure species is at a selective disadvantage
compared with the hybrid when growing on the shores of lochs with fluctuating water levels. There
is experimental evidence that the leaves of the hybrids are phenotypically more plastic than those
of R. reptans (Gibbs & Gornall 1976), and that this plasticity in leaf shape is an advantage in a
seasonally variable habitat (Cook & Johnson 1968).

ACKNOWLEDGMENTS

I should like to thank G. Halliday, A. McG. Stirling, D. F. Westlake and the curators of BM, E,
LIV, OXF, RNG, STA and TCD for loaning or allowing me access to herbarium specimens. I
should also like to thank M. Owen, Assistant Director of Research at the Wildfowl Trust, for
information concerning the migratory habits of various ducks and geese. |

REFERENCES

ALLISON, A. & NeEwron, I. (1974). Waterfowl at Loch Leven, Kinross. Proc. R. Soc. Edinb., B, 74: 365-381.

Ba.rour, I. B. (1901). 18th century records of Scottish plants. Ann. Scot. nat. Hist., 1901: 37-48.

Boyp, H. & Oaitvie, M. A. (1969). Changes in the British-wintering population of the Pink-footed Goose
from 1950 to 1965. Wildfowl, 20: 33-46.

CANDLISH, P. A. (1975). Ranunculus L. subg. Ranunculus, in Stace, C. A., ed. Hybridization and the flora of
the British Isles, pp. 124-125. London. |

Cook, S. A. & JoHNson, M. P. (1968). Adaptation to heterogeneous environments, 1. Variation in
heterophylly in Ranunculus flammula L. Evolution, 22: 496-516.

Druce, G. C. (1913). Ranunculus reptans L. Rep. botl Soc. Exch. Club Br. Isl., 3: 225.

Druce, G. C. (1929a). Notes on the second edition of the “‘British Plant List’. Rep. botl Soc. Exch. Club Br.
Isl. , 8: 867-877.

Druce, G. C. (1929b). British plant list (Edition II). Additions and corrections. Rep. botl Soc. Exch. Club Br.
Isl., 8: 878-883.

Garparsson, A. (1972). Research in Thjorsaver, C. Iceland, and a preliminary account of the summer food of
the Pink-footed Goose. /bis, 114: 581.

Gipss, P. E. & GornattL, R. J. (1976). A biosystematic study of the Creeping Spearworts at Loch Leven,
Kinross. New Phytol., 77: 777-785.

Gopwin, H. (1975). History of the British flora, 2nd ed. Cambridge.

Jones, N. G. B. & Git_mor, R. (1955). Observations on gathering and departure of Pink-footed Geese at
Asgard in central Iceland. Ann. Rep. Wildfowl Trust, 1953-1954, 7: 153-169.

Licutroot, J. (1977). Flora Scotica. Edinburgh.

Love, A. (1983). The flora of Iceland. Reykjavik.

MARTINDALE, G. E. (1886). Ranunculus? Ann. Rep. Watson bot. Exch. Club, 2: 4.

MEusEL, H., JAGER, E. & WEINERT, E. (1965). Vergleichende Chorologie der zentraleuropaischen Flora.
Karten. Jena.

Newton, I. & CAMPBELL, C. R. G. (1970). Goose studies at Loch Leven in 1967/8. Scott. Birds, 6: 5-18.

Papmorg, P. A. (1957). The varieties of Ranunculus flammula L. and the status of R. scoticus E. S. Marshall
and of R. reptans L. Watsonia, 4: 19-27.

Scott-E ior, G. F. (1896). The flora of Dumfriesshire.

STEWART, O. M. (1975). Plant record in Watsonia, 10: 421.

Tuom, V. H. (1969). Wintering duck in Scotland, 1962-68. Scott. Birds, 5: 417-466.

TuTINn, T. G. et al. (1964-1980). Flora Europaea, vols. 1-5. Cambridge.

VLAMING, V. DE & Procror, V. W. (1968). Dispersal of aquatic organisms: viability of seeds recovered from
the droppings of captive Killdeer and Mallard ducks. Amer. J. Bot., 55: 20-26.

(Accepted June 1986)

Watsonia, 16, 389-396 (1987) 389

The genus Barbarea R.Br. (Cruciferae) in Britain and Ireland

T. C. G. RICH

Biological Records Centre, Monks Wood Experimental Station, Abbots Ripton,
Huntingdon, PE17 2LS

ABSTRACT

An account of the genus Barbarea R.Br. (Cruciferae) in Britain and Ireland is presented. A general description
of the genus, and the four species that occur in these islands, Barbarea stricta Andrz., B. vulgaris R.Br., B.
intermedia Boreau and B. verna (Miller) Asch., is given, together with a key and notes on their identification
and distribution.

INTRODUCTION

The genus Barbarea R.Br. (Cruciferae) contains some 16 species, 13 of which occur in Europe.
Four of these have been recorded from Britain and Ireland: Barbarea stricta Andrz., B. vulgaris
R.Br., B. intermedia Boreau and B. verna (Miller) Asch. However, as the species are superficially
similar and the published keys and descriptions available often unreliable, they have been
frequently confused. During the last five years I have studied the genus in cultivation, field and
herbarium, and the purpose of this paper is to clarify the taxonomy of these four species and to
draw attention to characters by which they may be distinguished.

Barbarea is a well defined, natural genus and is usually placed in the tribe Arabideae with
Arabis, Cardamine, Cardaminopsis and Rorippa. Barbarea may be defined in the British Isles as
follows:

Biennial or short lived perennial (rarely annual) herbs, glabrous or sparsely pubescent below
with simple hairs. Rosette leaves pinnate. Upper cauline leaves amplexicaul, sessile, entire to
pinnatifid. Sepals erect, the inner slightly saccate at the base. Petals yellow, indistinctly clawed.
Stigma entire to slightly emarginate. Style persistent in fruit, sessile or forming a short, sterile
beak. Fruit a terete to four-angled, bivalvate, dehiscent siliqua, the valves with a strong median
vein and weaker reticulate lateral veins. Seeds in one row in each loculus.

CYTOLOGY

There are no reported chromosomal data from native British material. Counts from European or
cultivated material suggest that most of the species are diploid with a chromosome number of
2n=16 (Manton 1932), although B. stricta has counts of 2n=14—-18 (Manton 1932; Tischler 1934)
and B. vulgaris of 2n=14-16 and 2n=18 (Tischler 1934; Dvorak et al. 1981).

REPRODUCTIVE BIOLOGY

All four species begin flowering in spring or early summer, continuing under favourable conditions
into July. Barbarea intermedia and B. verna are usually the first to come into flower in early April.
Plants may flower in the autumn if they have been damaged earlier in the year. The pollination of
B. vulgaris has been described in detail by Proctor & Yeo (1973) and the general mechanisms are
probably essentially the same for the other species. Insects, mainly bees and flies, are attracted to
the yellow inflorescences to collect nectar and/or pollen and effect self- or cross- pollination in the
process. If that fails, or in dull weather, the two shorter outer stamens bend in towards the stigma
and self-pollination occurs. Abundant seed is usually set in all species, although occasional plants
of B. vulgaris with completely or partially aborted fruits have been found. It is unknown whether
these plants are self-sterile. There are no specialized seed dispersal mechanisms.

390 TC. G: RICH

Vegetative reproduction by cauline rosettes has been reported in B. vulgaris (MacDonald &
Cavers 1974; Rich 1984).

HYBRIDS

B. stricta x B. vulgaris (B. X schulzeana Hausskn.) and B. vulgaris x B. intermedia (B. xX
gradlii J. Murr.) have been recorded from Europe (Hess, Landolt & Hirzel 1970; Lange
1937), but not from Britain or Ireland. They are apparently highly sterile and of rare occurrence.
No hybrids with other genera have been recorded.

The hybrid B. intermedia x B. verna was synthesized in 1961 by P. M. Benoit from parents of
Welsh origin (specimen in K). The resulting plants had leaves intermediate between the two
parents and 60% sterile pollen. Fruit formation appeared to be normal, and the siliquae
resembled those of B. intermedia in size, although it is not known if the seeds were viable.

IDENTIFICATION

The main characters used to separate the four species are as follows:

(a) Rosette leaves. The rosette leaves are very variable in outline and cannot usually be relied
upon for identification, despite their use by Clapham (1962).

(b) Stem leaves. Failure to examine the uppermost leaves, which become inflorescence bracts if
the lateral racemes develop, is the most common reason for mis-identification of Barbarea
specimens. For instance, in B. vulgaris the leaves show a gradual transition from the pinnate
rosette leaves to the nearly entire uppermost leaves, which are not pinnatifid, although they may
be lobed. If the middle stem leaves are examined, this may prompt a mis-identification to B.
intermedia as these leaves are often pinnate. Part of the problem also lies in the terms used to
describe the leaves. The uppermost leaves of B. intermedia and B. verna should strictly be
described as pinnatifid rather than pinnate, as the lateral lobes do not quite reach the midrib.
The uppermost leaves of B. stricta are usually entire, but those of B. vulgaris may be entire or
have one pair of lateral lobes at the base of the leaflet, whose divisions do not reach as close to
the midrib as in B. intermedia and B. verna.

With practice, these leaves can provide the most useful characters, and typical uppermost stem
leaves from the four species are shown in Fig. 1. The stem leaves of both B. stricta and B.
vulgaris are diagnostic, but it is not possible to separate B. verna and B. intermedia using these
alone.

(c) Flower buds. B. stricta can usually be distinguished from the other species by the sparsely
hairy tips of the sepals, most easily seen in silhouette on the flower buds. However, these hairs
are often lost in herbarium specimens with age, and very rarely B. vulgaris may have a few hairs
on the sepals.

(d) Flower colour. There is much variability in flower colour and it cannot be used to separate
the species reliably. However, B. intermedia does often seem to have paler flowers than the other
species, and B. stricta has more uniformly deeper yellow petals.

(e) Length of petals and sepals. The ratio of sepal:petal length has been used to distinguish the
species (e.g. Wiggington & Graham 1981), but is highly variable and is not reliable. A much
better character is the length of the fresh petals, most easily measured by dissecting flowers and
sticking the petals from at least five mature flowers onto sticky tape.

(f) Fruit length, measured from the base of the valve to the tip of the persistent style. This is a
very useful character for distinguishing B. verna from the other species, as specimens usually
have at least some fruits more than 4 cm long, which is beyond the range of the other species.
Short-fruited variants of B. vulgaris also occur.

(g) Persistent style length, measured from the apex of the valves to the tip of the style. This
character is very useful for distinguishing B. vulgaris in Britain and Ireland, but is not so reliable
for European material.

(h) Seed size. The mean of at least ten seeds from different siliquae should be recorded.

BARBAREA IN BRITAIN AND IRELAND 391

H2O e

49 920

FicureE 1. Uppermost stem leaves of Barbarea species. A—B, B. stricta. C-E, B. vulgaris. F—H, B. intermedia.
I-J, B. verna.

SPECIES DESCRIPTIONS

KEY TO SPECIES

1

D
3)
3)

Uppermost cauline leaves and bracts simple, toothed or shallowly lobed,
sometimes with 1 or 2 pairs of linear, lateral lobes, the terminal lobe
broad, ovate to obovate; mean seed sizet1.6 mm ............c ccc eee e scene ee 2.

Uppermost cauline leaves and bracts pinnatifid with 2 or more pairs of
lateral lobes, the terminal lobe narrow, oblong to oblanceolate; mean

SEC MESIZe mie IMM te ace ec Gia sia eite eee ea lee Seb tnca | Meoninac 5 Sec eal Me cids he ck dards 3
Flower buds at least sparsely pubescent at tips; persistent style of fruit

Ase) (G23 MATIN eee teense cals scam nae en Acgebe doetuuia sitet. cs vodnad Sa WSeacaines 1. B. stricta
Flower buds glabrous; persistent style of fruit 2.0—-3.5 mm ...................... 2. B. vulgaris
Fresh petals 4.0—5.6 mm; fruits 1.5—3.1(—3.6) CM ........... ccc ccc eee se eee ee ees 3. B. intermedia
Fresh petals (5.6—)6.0—8.4 mm; fruits (2.8—)3.5-—7.1 cM ........ ee eee eee e eee 4. B. verna

The following descriptions have been prepared from British and Irish specimens: European
material is usually more variable. Synonyms in use in the British and Irish literature are given. No
infraspecific taxa merit recognition within the British Isles, with the possible exception of variants
of B. vulgaris.

1. B. STRICTA Andrz. in Besser, Enum. Pl. Volhyniae, 72 (1822).

Common names: Upright Wintercress, Small-flowered Wintercress

392 PVC. Ge RICH

Glabrescent biennial or short-lived perennial herb up to 100 cm. Basal leaves with a large, ovate-
oblong terminal leaflet and (0O—)1—3(—4) pairs of much smaller lateral leaflets. Stem leaves smaller,
sessile and entire, the uppermost broad, ovate to obovate, coarsely but shallowly sinuately lobed.
All foliage yellow-green. Inflorescence branches straight and erect, with dense racemes at anthesis.
Buds at least sparsely hairy at tips. Petals 3.5—6.0 mm, suberect, deep yellow all over including the
claw. Fruits on appressed (rarely more spreading) pedicels 3—7 mm, the siliqua 1.3—2.8(—3.5) cm,
usually straight, with a stout persistent style 0.5—1.6(—2.3) mm. Seeds (1.0—)1.1—1.6(-1.7) mm,
mean 1.4 mm. Flowering May to September.

B. stricta is closely related to B. vulgaris, and also to B. orthoceras Ledeb. of North America and
eastern Asia. Sprague & Hutchinson (1908) give an excellent account of its distinction from B.
vulgaris in the British Isles. It is a very local, lowland plant of canal, stream and river banks, quiet
backwaters and also rarely as a casual on drier ground. It occurs in similar habitats to B. vulgaris
but is perhaps more restricted to damper sites next to slow-moving water. It is widely distributed in
England and eastern Wales, but is very rare in Scotland, and absent from Ireland.

Jackson (1908) described its history and distribution in Britain. The species was first reported in
1843 by W. Borrer from Yorkshire (v.c. 63) and Northants. (v.c. 32), but its status in the British
Isles is uncertain. On the basis of the criteria of Webb (1985), it is impossible to come to any firm
conclusion as to whether the plant is native or introduced. There are no reliable fossil records
(Godwin 1975) since it is not possible to distinguish seeds of B. stricta from those of B. vulgaris.
The relatively late first record could be due to the plant being overlooked as B. vulgaris or due to
its recent arrival. In this respect, records from the River Thames are of interest, as although the
species has been persistent on the banks near Kew since at least 1871, neither Brown (1812) nor
Brewer (1863) mentioned the plant. This suggests that the Thames localities may be recent.
Jackson (1908) pointed out that its habitats on the Continent are much the same as some of those in
England, and its European distribution is consistent with it being a native plant. It is known to have
become naturalized elsewhere (ignoring for the moment localities within Britain), but although it
has reached New Zealand, it has yet to be recorded from Ireland. The species shows no obvious
geographical pattern of variation. It reproduces by seed and, although fluctuating in abundance
from year to year (e.g. Lousley 1976) in a similar manner to other biennials, it is remarkably
persistent in some localities. Seeds of other Barbarea species may be introduced with grain (c.f. B.
verna) and in some localities B. stricta is believed to have been brought in by barges (Gibbons
1975), presumably with ballast. Thus in the absence of any further information, the plant is best
regarded as ‘probably native’.

The world distribution of B. stricta is the western Mediterranean and central Europe to northern
Scandinavia, eastwards to the Russian provinces of Orenburg and Perm, and southwards to Bosnia
and Bulgaria (Sprague & Hutchinson 1908). It has been introduced to North America, New
Zealand and probably Japan.

2. B. VULGARIS R.Br. in W. & W. T. Aiton, Hortus Kew., 2nd ed., 109 (1812).
Erysimum barbarea L., Sp. Pl., 660 (1753).
Barbarea barbarea Miller, Gard. Dict., 8th ed. (1768).

B. taurica DC., Syst., 2: 207 (1821).

B. arcuata (Opiz ex Presl) Reichenb., Flora, 5: 296 (1822).
B. stricta Boreau, Fl. Cent. France, 3rd ed., 2: 39 (1857).
B. lyrata Ascherson, Fl. Brandenburg, 1: 35 (1864).

B. rivularis Martr., Fl. Tarn., 44 (1864).-

B. sylvestris Jord., Diagn., 1: 102 (1864).

Jackson (1916) details the synonymy of infraspecific taxa.

Common names: Wintercress, Yellow Rocket

Biennial to perennial herb up to 90(—130) cm, usually glabrous. Basal leaves pinnate, with an
elliptic to ovate terminal leaflet and (O—)2—5(—6) pairs of lateral leaflets. Upper stem leaves usually
sinuate-lobed, often with 1 or 2 pairs of small, spreading, linear lobes from base of leaflet, the

BARBAREA IN BRITAIN AND IRELAND 393

terminal lobe broad, obovate to broadly elliptic (rarely ovate) with a cuneate base. All foliage deep
green, shining, with coarsely toothed or sinuate margins. Inflorescence crowded or lax. Buds
glabrous or very rarely with a few hairs. Petals 4.9-7.0(—8.0) mm, deep to bright yellow with a
paler claw, the limb spreading, becoming reflexed with age. Fruits on thin pedicels 3-6 mm long,
spreading-erect. Siliqua (0.7—)1.5—3.2 cm, straight or curved with a slender persistent style 2.0—
3.5(—4.0) mm. Seeds (1.1—)1.2—1.6(—1.8) mm, mean approx 1.5 mm. Flowering May to July.

B. vulgaris is a very variable species within which many infraspecific taxa have been described.
In a critical study of the species in the British Isles, Jackson (1916) recognized four varieties:
var. silvestris Fries Plants usually small, with solitary stems. Lateral lobes of lower leaves very

small or absent. Fruits short, about 12 mm long, appressed.
var. campestris Fries Plant robust, fruits usually longer, up to 25 mm, obliquely erect or slightly

spreading.
var. arcuata Fries Inflorescence often laxer. Pedicels patent. Fruits arcuate- ascending, spreading

at right angles to the axis, or even deflexed.
var. transiens Druce Plant stout, robust; lower stem leaves with oblong cuneiform terminal lobe,

the lateral linear lobes well developed, up to five pairs, exceeding the terminal lobe in width.
The taxa are thus defined largely on a combination of somewhat variable characters. Since the size
of plant varies according to habitat, and the size and shape of the leaves are usually linked to the
robustness of an individual, var. transiens may just be a result of good growth conditions. The only
general correlation between characters is the density of the inflorescence and the orientation of the
pedicels and fruits — plants with laxer racemes usually have spreading fruits, and plants with dense
racemes have erect fruits. This correlation can also be seen in B. intermedia. However, in some
populations it is possible to find plants with both spreading and erect fruits, which makes the plants
look distinct although they are identical in all other respects. This may suggest there is a simple
genetic basis to this difference. The character of spreading fruits is essentially the basis on which
var. arcuata (B. arcuata (Opiz ex Presl) Reichenb.) is distinguished from var. campestris; Jackson
(1916) has shown that the other characters are unreliable, and he rightly included B. arcuata within
B. vulgaris in the British Isles. However, Bush (1939) maintained it as a separate species in Russia,
but there may be a geographical basis for this difference of opinion. The two variants are very
similar in north-western Europe, but in eastern Europe and Asia, outside the range of typical B.
vulgaris, plants with spreading fruits are much more common. There may also be clinal variation
with other characters, since B. vulgaris sensu lato is often apparently annual in the eastern part of
its range, a character not observed in any British plant.

A second character which makes some plants appear quite distinct is consistently short fruits,
typically 7-12 mm long, and such plants have been described as var. silvestris. It seems that these
are selectively collected and are far more frequent in herbaria than in the field! Other plants with a
mixture of short and long fruits can also be found, but this may be due to partial self-sterility.

Sparse hairs on the lower parts of the plant are of no taxonomic value, as pubescence is probably
environmentally determined. Material cultivated at Leicester in a constant environment room was
sparsely hairy, whilst plants grown outside from the same seed batch were glabrous. Hairy and
glabrous plants can also be observed in the field within the same population. A similar dimorphism
occurs in B. intermedia and B. verna, where sparsely hairy plants are quite frequent.

Knowles (1967) recorded that a double-flowered variant ‘Flore Pleno’ and a variant ‘Variegata’
with golden netted foliage are grown in gardens. Tilney-Bassett (1963) has shown that the
variegation in the latter is due to the action of a single recessive gene. B. vulgaris used to be
cultivated as a salad plant, but has now been replaced by B. verna.

A more detailed study of the species and its possible segregates throughout its range must be
made before it is possible to evaluate the infraspecific taxa critically. Until then the species is best
treated as a single, highly polymorphic taxon.

B. vulgaris is the commonest member of the genus in Britain and Ireland. It is a locally abundant
native species, most characteristic of places that are damp in the winter such as river levées and
shingle, ditch margins and woodland rides, but also in more disturbed habitats such as road verges,
edges of fields and waste ground. It can tolerate a wide range of soil conditions and only avoids the
more acid sites. Widespread throughout the British Isles and Ireland, it is most frequent in the
south, and is absent from much of northern and western Scotland.

394 ECs G RICH

The world distribution of B. vulgaris is Eurasia, from the Mediterranean to northern Scandina-
via, and eastwards from Portugal to central Russia and the western Himalaya. It has been
introduced to East Africa, Australasia and North America.

3. B. INTERMEDIA Boreau, Fl. Cent. France, 2nd ed., 48 (1840).
Common names: Intermediate Wintercress, Medium-flowered Wintercress

Biennial (rarely short-lived perennial) up to 60 cm, glabrous or sparsely pubescent below. Basal
leaves with a large terminal leaflet and (0O—)2—S(—6) pairs of smaller lateral leaflets. Stem leaves
pinnatifid, with (1—)2-3 pairs of lateral lobes, the terminal lobe becoming narrow, oblong-
oblanceolate. Foliage green. Inflorescence crowded. Buds glabrous. Petals 4.0-5.6 mm, pale to
bright yellow. Fruits on appressed (rarely spreading), stout pedicels 3-6 mm long, the siliqua
1.5-—3.2(-—3.6) cm, straight or slightly curved, with a stout persistent style 0.6-1.6(—1.7) mm.
Seeds 1.7—2.3(—2.4) mm, mean 2.1 mm. Flowering (March) April to July.

B. intermedia occurs on roadsides, waste ground and disturbed places throughout the lowlands
of England, Wales and Ireland, but is scarcer in Scotland. It is often associated today as a
colonist of building and road construction sites, although formerly it was most frequent as an
arable weed. It is rarely abundant, and is often found as a single individual, but is surprisingly
persistent in some localities in view of its intolerance of competition from other plants.

Britten (1864) summarized the earliest records from the British Isles. The species was first
recorded from Co. Armagh (v.c. H37) by A. J. More in 1844, although the plant was probably
present for a number of years before then but undetected due to confusion with B. vulgaris.
Lousley (1976) noted that it “has made little progress in a century”’.

It is probably native in southern and central Europe, from northern Portugal to southern
Germany and southern Yugoslavia, and also in North and East Africa, and introduced
elsewhere. Ham (1982) suggested that it is native in Holland. As with many other species that
behave as arable weeds, it is impossible to determine the original range of the species with any
certainty.

4. B. VERNA (Miller) Ascherson, Fl. Brandenburg, 1: 36 (1864).

Erysimum vernum Miller, Gard. Dict., 8th ed. (1768).

E. praecox Sm., Fl. Brit., 2: 707 (1800).

Barbarea praecox R.Br. in W. & W. T. Aiton, Hortus Kew., 2nd ed., 4: 109 (1812).
B. brevistyla Jord., Diagn., 1: 102 (1864).

Common names: Land-cress, American Wintercress, Early-flowering Yellow Rocket

Annual or biennial up to 75(—90) cm, glabrous or sparsely pubescent below. Basal rosette leaves
with a large terminal leaflet and (0O—)4—10(—11) pairs of lateral leaflets, the distal pair usually
wider than the terminal leaflet. Upper stem leaves pinnatifid, with 2—4 pairs of lateral lobes, the
terminal lobe narrow, oblong-oblanceolate. Foliage green. Buds glabrous. Petals (5.6—)6.0—8.4
mm, bright yellow. Fruits on stout pedicels (2—)3-8 mm, the siliqua (2.8—)3.5-7.1 mm,
arcuate-ascending, with a stout persistent style 0.6-2.0(—2.3) mm. Seeds (1.6—)1.7—2.3(-2.4)
mm, mean 2.0 mm. Flowering (March) April to July.

B. verna is a casual weed of waste ground, railway embankments, roadsides, bare and stony
ground, etc., usually close to habitation. It is an ancient salad crop, probably introduced as a
winter substitute for watercress in the 16th or 17th century, and commercial seed is still widely
available. White (1912) noted that some of the species of Barbarea taste different, B. verna being
a desirable salad plant, whilst B. intermedia is unpleasantly bitter. B. verna is scattered through-
out lowland Britain, although it is more common in the south, and is rare in Scotland and
Ireland. Davie (1909) suggested it might be native in Cornwall. It is more persistent in the south

BARBAREA IN BRITAIN AND IRELAND 395

and west, and it may re-appear sporadically as its seeds can remain dormant for many
years. Clement & Foster (1983) recorded B sicula C. Pres! from Britain, probably in error for
B. verna.

The species is perhaps native in the western Mediterranean region and Macaronesia, but it is
widely naturalized in western and central Europe, and its native range has been obscured by
cultivation. It has been introduced to North America, South Africa, Japan and New Zealand, and
is recorded as introduced with grain into Norway (Jgrgensen 1969).

This species is phenotypically the most plastic member of the genus in the British Isles and is
very variable in size in response to environmental conditions, as illustrated in Stace (1980). Small,
depauperate annual specimens contrast markedly with robust biennials, and it is this enormous
range of variation which has led to many problems in distinguishing B. verna from B. intermedia.

ACKNOWLEDGMENTS

I am very grateful to Dr G. Halliday who first suggested that I should study Barbarea, to Miss A.
Wightman for her help with the literature survey, and to members of the B.S.B.I. who sent
material to us at Lancaster. I should like to thank Professor C. A. Stace for comments on the
manuscript.

REFERENCES

Brewer, J. A. (1863). The flora of Surrey. London.

Brown, R. (1812). Barbarea, in W. & W. T. Aiton. Hortus Kewensis, 2nd ed. London.

BRITTEN, J. (1864). Spontaneous exotics. Naturalist, Hull, 1864: 201-207.

Busu, N. A. (1939). Barbaraea [sic] Beck., in Komarov, V. L., ed. Flora of U.S.S.R., 8: 99-102. Leningrad.

CiapHaM, A. R. (1962). Barbarea, in CLAPHAM, A. R., Tutin, T. G. & WarBurG, E. F. Flora of the British
Isles, 2nd ed., pp. 164-166. Cambridge.

CLEMENT, E. J. & Foster, M. C. (1983). Alphabetical tally-list of alien and adventive plants, 3rd ed. London.

Davie, J. H. (1909). Flora of Cornwall. Penryn. :

DvorAk, F., GRULL, F., SutorY, K. & DADAKoVA, B. (1981). In LévE, A., ed. Chromosome number reports
LXXIII. Taxon, 30: 844.

Gipzons, E. J. (1975). The flora of Lincolnshire. Lincoln.

Gopwin, H. (1975). The history of the British flora, 2nd ed. Cambridge.

Ham, R. W. J. M. VAN DER (1982). Barbarea intermedia Bor. en Barbarea verna Aschers. in Nederland.
Gorteria, 11: 36-39.

Hess, H. E., LANpDoLtT, E. & HirZEL, R. (1970). Flora der Schweiz, 2: 215-218. Basel.

Jackson, A. B. (1908). in SPRAGUE, T. A. & Hutcuinson, J. Note on Barbarea stricta Andrz. J. Bot., Lond.,
46: 106-109.

Jackson, A. B. (1916). A study of Barbarea vulgaris R.Br. J. Bot., Lond. , 54: 202-209.

JORGENSEN, P. M. (1969). Two anthropochorous species new to Norway. Blyttia, 27: 135-140.

Know es, E. (1967). Cultivars of British wild plants 7. The Cabbage family. Gdnrs Chron., 1967: 14.

LanGE, T. (1937). Sveriges Barbarea-arter. Bot. Notiser, 1937: 216-230.

LousLey, J. E. (1976). Flora of Surrey. Newton Abbot.

MacpDonaLp, M. A. & Cavers, P. B. (1974). Cauline rosettes, an asexual means of reproduction and dispersal
occurring after seed formation in Barbarea vulgaris (yellow rocket). Can. J. Bot., 52: 913-918.

Manton, I. (1932). Introduction to the general cytology of the Cruciferae. Ann. Bot., 46: 509-556.

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

Procror, M. C. F. & Yeo, P. (1973). The pollination of flowers. London.

Ricu, T. C. G. (1984). Asexual reproduction of Barbarea vulgaris R.Br. in the British Isles. Watsonia, 15:
121-122.

SPRAGUE, T. A. & Hutcuinson, J. (1908). Note on Barbarea stricta Andrz. J. Bot., Lond., 46: 106-109.

Stace, C. A. (1980). Plant taxonomy and biosystematics. London.

Titney-BasseTr, R. A. E. (1963). Selection against the variegated genotype in Barbarea vulgaris. Heredity, 18:
534-545.

TISCHLER, G. (1934). Die Bedeutungen der Polyploidie ftir die Verbreitung der Angiospermum, erlautert an
der Arten Schleswig-Holsteins, mit Ausblicken auf andere Florengebiete. Bot. Jahrb. , 67: 1-36.

396 T. C. G. RICH

Wess, D. A. (1985). What are the criteria for presuming native status? Watsonia, 15: 231-236.

Wuite, J. W. (1912). The flora of Bristol. Bristol.

WicGINGTON, M. & GRAHAM, G. G. eds. (1981). Guide to the identification of some of the more difficult vascular
plant species. N.C.C. England Field Unit, Occasional Paper no. 1, pp. 17-18. Banbury.

(Accepted June 1986)

Watsonia, 16, 397-407 (1987) 397

A morphometric study of < Orchiaceras bergonii (Nanteuil)
Camus and its parents (Aceras anthropophorum (L.) Aiton f. and
Orchis simia Lamarck) in Kent*

R. M. BATEMAN
Department of Geology, Birkbeck College, 7-15 Gresse Street, London, WIP 1PA
and
O. S. FARRINGTON

Department of Geography, Birkbeck College, 7-15 Gresse Street, London, W1P 1PA

ABSTRACT

x Orchiaceras bergonii (Nanteuil) Camus (Aceras anthropophorum (L.) Aiton f. X Orchis simia Lamarck),
recently found in Kent and new to the British Isles, is described in detail. Morphometric comparison of the
hybrid with its parent species has shown that it is intermediate in 17 of the 41 characters measured but
resembles Aceras in six and O. simia in 18. It is similar to Continental plants of x Orchiaceras bergonii; some
characters appear to be expressed consistently in hybrids between Aceras and several species of Orchis. The
status of the British x Orchiaceras bergonii as a natural hybrid, and the value of Aceras R. Brown as a genus,
are discussed. The need for accurate, quantitative diagnostic characters in Floras, and the considerable value of
multivariate methods in the study of hybrids, are emphasized.

INTRODUCTION

Bigeneric hybrids between Aceras anthropophorum (L.) Aiton f. and species of Orchis L. section
Orchis subsection Militares Parlatore are often recorded on the Continent (Nelson 1968; Peitz
1970; Davies et al. 1983), especially XOrchiaceras bergonii (Nanteuil) Camus (1892) (A.
anthropophorum x O. simia Lamarck); for synonyms see Peitz (1970). The frequent occurrence of
this hybrid, particularly in France, shows that its parents share pollen vectors and that barriers to
cross-fertilization are, at best, only partially effective. It also reflects their similar ecological
preferences and geographical distributions in mainland Europe (Baumann & Kiinkele 1982).
Aceras and O. simia both reach their northern limits in the British Isles, but curiously they have
remained almost entirely allopatric. Aceras is locally frequent along the North Downs of Kent and
Surrey, but rare elsewhere (Perring & Walters 1962); Druce (1886) reported only one Oxfordshire
site for Aceras, and only one remains (Steel & Creed 1982). O. simia has always been confined to
southern Oxfordshire, where it coexisted with O. militaris L. and was locally frequent until the
mid-nineteenth century (Godfery 1933; Summerhayes 1951), but has since declined to a single
viable population (Summerhayes 1951; Paul 1965). Consequently, in Britain, only one hybrid
combination has been recorded involving O. simia (O.xXbeyrichii A. Kerner = O. simiaxO.
militaris) (Hunt 1975a), and none involving Aceras (Hunt 1975b).

In 1920, five plants of O. simia were found at Bishopsbourne, near Canterbury, Kent. In the
same year, B. J. Brook collected two flowers of O. simia and one of Aceras, all apparently from
this locality (BM). Unfortunately, O. simia only persisted at Bishopsbourne until 1923, and only
one or two plants flowered each year (Summerhayes 1951), providing little opportunity for
hybridization. However, in 1955, H. M. Wilks discovered another Kentish locality for O. simia

* Dedicated to the late Norman R. Campbell, orchid biometrician.

398 R. M. BATEMAN AND O. S. FARRINGTON

near Faversham (Wilks 1960), and this remains the only natural population of this species south of
the Thames (Perring & Farrell 1977; Philp 1982); at present about 30-50 O. simia plants coexist
with a similar number of Aceras. When we visited the locality on the 31st May 1985, we tentatively
identified the first individual of < Orchiaceras bergonii recorded in the British Isles.

MATERIALS AND METHODS

A biometric study was performed to appraise this initial identification and to assess the

morphological variation shown by its presumed parent species. 41 characters were recorded for the

putative hybrid and for every flowering plant of O. simia (10) and Aceras (8) present:

Stem and inflorescence (7 characters).

. Stem height, above ground level.

Stem diameter, above uppermost sheathing leaf.

. Presence (1) or absence (0) of anthocyanins immediately below inflorescence.

Inflorescence length.

. Number of flowers.

. Bract length.

. Ovary length.

B. Leaves (7 characters).

Orchis and Aceras leaves are difficult to categorize. Basal leaves form a spreading rosette

immediately above ground level. Sheathing leaves arise from the rosette but surround the stem

(occasional leaves intermediate between these categories were arbitrarily classed as basal leaves).

Cauline leaves arise from the stem above its base, and are usually much smaller than the lower

leaves. Leaf shape (character 14) was assessed by determining the position of maximum width

relative to length, on a scale 1-4 (1=0-10% of length; 2=11-25%; 3=26-50%; 4=>50%).

8. Number of basal leaves.
9. Number of sheathing leaves.

10. Number of cauline leaves.

11. Length of longest leaf.

12. Width of longest leaf (often=C13).

13. Width of widest leaf.

14. Shape of longest leaf (see above).

C. Labellum (16 characters).

Fig. 1 summarizes nine measurements taken from each labellum, and explains the anthropomor-

phic terminology that we have adopted for simplicity to describe parts of the labellum. The colours

of the ‘torso’ and ‘limbs’ of each labellum, and of the reverse surfaces of the outer perianth

segments, were matched to the nearest colour block of the Royal Horticultural Society Colour

Chart (Anonymous 1966) by I. Denholm. Patches of pigmented hairs characteristic of the ‘torso’ of

O. simia are termed papillae.

15. Maximum width.

16. Width of ‘torso’.

17. Maximum length.

18. Length of ‘torso’.

19. Presence (1) or absence (0) of ‘tail’.

20. Length of ‘tail’ (if present).

21. Length of ‘arm’.

22. Width of ‘arm’, measured halfway along length.

23. Length of ‘leg’. .

24. Width of ‘leg’, measured halfway along length.

25. Colour of ‘torso’, on a scale 1-3 (1=pale straw yellow, R.H.S.11C; 2=greenish-grey,
R.H.S.157C; 3=very pale lilac, R.H.S.76D).

26. Colour of ‘limbs’, on a scale 1-4 (1=yellowish-orange, R.H.S.15C; 2=purplish-red,
R.H.S.64B; 3=reddish-purple, R.H.S.72B—C; 4=purple-violet, R.H.S.80B-C).

27. Number of papillae on ‘torso’.

28. Distribution of papillae on ‘torso’, on a scale 1-3 (1=concentrated immediately below spur
entrance, through to 3=distributed over most of ‘torso’).

STUDY OF X ORCHIACERAS BERGONII 399
MEDIAN OUTER
PERIANTH SECMENT
COLUMN

ith POLLINIA INNER
avith PERIANTH SEGMENT

LATERAL OUTER

SPUR ENTRANCE PERIANTH SEGMENT

PAPILLAE
TORSO

ARM
LABELLUM

[720

Figure 1. Nomenclature of floral parts, and labellum dimensions measured. Numbers refer to characters listed
in Materials and Methods.

29. Attitude of ‘torso’ relative to stem, on a scale 1-5 (1=parallel, through to 5=perpendicular).

30. Attitude of ‘limbs’ relative to ‘torso’, on a scale 1-4 (1=shallowly convex; 2=planar;
3=shallowly concave; 4=deeply concave).

D. Spur (3 characters).

31. Length, from entrance to apex.

32. Diameter, halfway along length when viewed laterally.

33. Shape, on a scale 1-5 (1=strongly recurved, through to 5=strongly decurved).

E. Inner perianth segments (2 characters).

34. Length.

35. Maximum width.

F. Outer perianth segments (5 characters).

36. Length.

37. Maximum width.

38. Colour of reverse surface, on a scale 1-2 (1=pale yellowish-green, R.H.S.154C—D; 2=pale/
very pale lilac, R.H.S.76C-D).

39. Presence (1) or absence (0) of peripheral and median linear markings on reverse surface.

40. Presence (1) or absence (0) of dispersed dots and/or dashes on reverse surface.

G. Phenology (1 character).

41. Percentage of flowers in inflorescence fully open.

Some of the above characters were used to calculate the following ratios, which summarize the
shapes of certain structures. The characters are numbered according to the above list and preceded
byj the letter,..€::

. Robustness of stem. C2/(C1+C2).

. Percentage of stem bearing flowers. (100 C4)/C1.

. Density of inflorescence (fls/em). C5/C4.

. Length of bract relative to length of ovary. C6/(C6+C7).
. Shape of longest leaf. C12/(C11+C12).

Roundness of labellum. C17/(C15+C17).

. Length of ‘arms’ relative to length of ‘torso’. C21/(C18+C21).
. Length of ‘legs’ relative to length of ‘torso’. C23/(C18+€C23).
Length of ‘arms’ relative to length of ‘legs’. C21/(C21+€C23).

™ oe moo Of

400 R. M. BATEMAN AND O. S. FARRINGTON

Data were analysed by multivariate methods using the Rothamsted Genstat computer program
(Alvey et al. 1977). Character 12 was discarded as it often duplicated character 13, and ratios (a) to
(i) were omitted as they duplicate their component characters. The remaining 40 characters were
used to compute a symmetrical matrix that quantified the similarities of pairs of data sets (i.e.
plants) using the Gower Similarity Coefficient (Gower 1971) on unweighted data scaled to unit
variance. This was used to construct a minimum spanning tree (Gower & Ross 1969) and
subsequently to calculate principal coordinates (Gower 1966), compound vectors that incorporate
positively or negatively correlated characters which are most variable and therefore of potential
diagnostic value. Principal coordinates have previously been used to assess morphological
relationships between orchid taxa (Bateman & Denholm 1983, 1985), but not those between
orchid hybrids and their parental taxa.

VARIATION IN SINGLE CHARACTERS

Population means (and sample standard deviations) for O. simia, x Orchiaceras and Aceras are
presented in Table 1.

CHARACTERS SEPARATING THE PARENTS

We define continuous metric characters and ratios as taxonomically useful if the parental standard
deviations do not overlap (>67% discrimination) and diagnostic if twice the parental standard
deviations do not overlap (>95% discrimination) (Fig. 2). Diagnostic scalar characters lack classes
containing individuals of both species. Eleven characters and three ratios are taxonomically
useless, and a further nine characters and two ratios are taxonomically useful but not diagnostic
(i.e. give 67-95% discrimination; Table 1). They include the majority of the floral dimensions
measured and all but one of the vegetative characters (e.g. stem height, leaf size and number),
which are potentially strongly influenced by both ontogeny and environment; the four vegetative
characters (one, stem diameter, diagnostic) classed as taxonomically useful here might be less
useful at other sites which support more vigorous populations of Aceras. However, the remaining
20 diagnostic characters and four diagnostic ratios are probably generally applicable.

Compared with Aceras, O. simia has basal leaves that are broadest closer to their tips and much
shorter bracts (about one quarter the length of the ovary rather than approximately equalling the
ovary). Its tendency to have fewer, more closely-spaced flowers results in a shorter inflorescence
forming a much smaller proportion of the total length of the stem. Its flowers open more or less
simultaneously (the lower flowers may open slightly earlier (Wilks 1960) or slightly later (Ettlinger
1976) than the upper) to form a cylindrical inflorescence, whereas those of Aceras open gradually
from the bottom upwards over a long period to form an acute conical inflorescence. The flowers of

a |
A ' ! U (
(o me © res Oi © : ©
HYBRID CATEGORY = 1 i 2 mim 3B St 4 SB --
B TAXONOMICALLY USEFUL
DIAGNOSTIC CHARACTER! : wae : o— :
4
TAXONOMICALLY USEFUL - :
NON-DIAGNOSTIC CHARACTER 1 a \ '

TAXONOMICALLY USELESS jo gee eee
CHARACTER ——____°__ ee

FicureE 2. Explanation of A) ‘hybrid categories’ (1-5) listed in Table 1, and B) character classes described in
Characters Separating the Parents. O=value for X Orchiaceras, @=mean for Orchis simia, 0=mean for Aceras;
bars indicate standard deviations of parental taxa (‘=twice the standard deviation).

STUDY OF x ORCHIACERAS BERGONII 401

TABLE 1. COMPARISON OF CHARACTER VALUES FOR xXORCHIACERAS BERGONII WITH
POPULATION MEANS (AND SAMPLE STANDARD DEVIATIONS IN PARENTHESES WHERE
APPROPRIATE) OF PARENTAL POPULATIONS
Characters are numbered as in Materials and Methods. Italicized characters are taxonomically useful, bold
characters are diagnostic (see Fig. 2). See Fig. 2 for explanation of ‘hybrid category’.

Hybrid

Character Units O. simia x Orchiaceras Aceras category
A. 1 mm 267 ~=(68) 180 223 = (46) 4
2 mm 4.28 (0.77) 3.4 2.59 (0.32) 3
3 0.9 0 0 4
4 mm 2538 1(8:9) 11 714i (25.1) i
5 17:6, (5.6) 6 32h 1 a(de5) 1
6 mm 1.8 (0.7) Ts 6.5 (0.9) 5
7. mm F395 LO) 6.8 O25 (OFS) 5
B. 8 APS (OLS) 6 4.0 (0.8) 1
9 LOei (Ol7) 1 Delos (056) a
10 0.6 (0.5) 0 1.1 (0.4) 1
Jul mm 117 (24) oS 83 (29) 3
12 mm 29 (5) 20) 18 (3) 3
13 mm 30 (6) 24 18 (3) 5)
14 4.0 4 3.0 2
(Oe 15 mm 12.2 (1.6) LEO 4.1 (0.6) 2
16 mm 1.84 (0.24) 1:7 te 234 (O.22) 2
17 , mm UPL TANG ONS) 13.0 O20 ye (222) 2
18 mm 0, (027) 7.8 Ten ale) 2
19 1.0 1 0.4 ji
20 mm 1.44 (0.57) 0.7 O27 (0-21) e}
21 mm O22 eh 1(020) 8.3 DEG Le) 2
22 mm 0.73 (0.07) 0.9 0.62 (0.08) 1
23 mm Ted icete(Ae3) 5.4 2.9 (1.1) 3
24 mm 0.92 (0.15) 0.9 0.38 (0.09) 2
25 3.0 2 1.0 3
26 4.0 3 1.3 3
27 28 (3) 6 0 3
28 3.0 2 0 3
29 2.4 2 1.0 2
30 4.0 2 1.4 “|
Dy ol mm 4.7 (0.8) 1) 0 3
32 mm 1.69 (0.35) IG 0 2
33 4.2 4 0 2
EB: 34 mm Son (LY) 6.6 ALO Ia el 2D) 3}
35 mm 1.48 (0.24) 1.4 $24 .3(0:37) 3
FE. 36 mm 11.3 (1.2) 8.0 6.4 (0.9) 3
37, mm 4.7 (0.9) 3.4 2.9 (0.4) 3
38 2.0 1 1.0 +
39 0 1 1.0 4
40 1.0 1 0 2
G. 41 % 100 =) 100 52 (24) 2
Ratio a 0.016 (0.002) 0.018 - 0.012 (0.003) 2
b % 9.5 (1.8) 6.1 32.1 (8.9) 1
c flsyem 7.0 (1.4) D5) 4.7° (1.7) 3
d 0.19 (0.05) 0.53 0.51 (0.04) 4
er 0.20 (0.02) 0.19 0.18 (0.05) 3
f 0.51 (0.04) 0.54 0.71 (0.05) 2
g 0.52 (0.03) 0.52 0.43 (0.03) 2
h 0.49 (0.04) 0.41 0.29 (0.04) 3
i 0.53 (0.03) 0.61 0.65 (0.06) =

“Values estimated for several plants lacking leaf tips. "Mean of plants possessing ‘tails’. “Value would have
decreased as flowers higher in the inflorescence opened.

402 R. M. BATEMAN AND O. S. FARRINGTON

O. simia are generally larger (Fig. 3), especially in overall labellum width, ‘leg’ and outer perianth
segment dimensions. “Tails’ are either absent in Aceras or much shorter than those of O. simia.
The labella of Aceras are parallel to the stem and the ‘limbs’ are more-or-less parallel to the ‘torso’,
whereas the labella of O. simia are inclined upward and the ‘limbs’ arc forward. The c.5 mm long,
downward-curved spur of O. simia is represented only by a very shallow (<0.5 mm) ‘nectar pit’ in
Aceras, and Aceras lacks the 25-30 papillae that adorn the ‘torso’ of O. simia. Pigmentation
provides the most conspicuous differences between the species; O. simia has pale lilac outer
perianth segments and ‘torsos’, deepening to purple-violet on the ‘limbs’, and anthocyanins are
usually also present on the upper part of the stem, whereas Aceras has yellowish-green outer
perianth segments, a straw yellow ‘torso’ with ‘limbs’ that often open purplish-red but fade to
yellow-orange, and lacks stem anthocyanins. The outer perianth segments of O. simia bear
dispersed purple-violet dots and dashes, in contrast to the peripheral and median linear,
purplish-red markings of Aceras.

Interestingly, the ranges of labellum lengths obtained for both species in Kent are shorter than,
and do not overlap with, ranges given in Flora Europaea by Soo (1980) for O. simia, and Moore
(1980) for Aceras. So6 (1980) also described the bracts of O. simia as c.50% of the length of the
ovary (they average 23% in Kent), its labellum as longer than wide (its length approximately
equals its width in Kent), and its outer and inner perianth segments as c.10 mm long (the outer
perianth segments average 11.3 mm but the inner only 8.5 mm in Kent).

COMPARISON OF THE HYBRID WITH ITS PARENTS

Characters of the putative hybrid have been assigned to one of five ‘hybrid categories’ in Table 1
according to their values relative to the standard deviations of the parental means. Fig. 2 explains
these categories. The hybrid is morphologically more extreme than either parent (categories 1 and
5) in only six characters, and only two of these are taxonomically useful: it has a shorter
inflorescence than O. simia and slightly longer bracts than Aceras. It resembles O. simia in 13
characters (category 2), including its orbicular leaves, the inclined position and more-or-less
synchronous opening of its flowers, its wide labellum, ‘torso’ and ‘legs’, its long ‘arms’ and its small
labellum roundness index, and resembles Aceras in five characters (category 4), including having
its ‘limbs’ in the same plane as its ‘torso’ (Fig. 3), yellowish-green outer perianth segments, bracts
that equal the ovaries in length, and in lacking stem anthocyanins. The hybrid is intermediate in 16
characters (category 3). Values for 14 of these do not fall within the standard deviations
(continuous metric character) or classes (scalar characters) of either parent and therefore aid
identification of the hybrid; they include several vegetative characters, ‘leg’, ‘tail’ and spur lengths,
labellum colours, number and distribution of papillae, and outer and inner perianth segment
dimensions. Its outer perianth segments also bear both the dispersed dots and dashes of O. simia
and the peripheral and median lines of Aceras (Fig. 3).

The hybrid was a young plant flowering for the first time when measured (H. M. Wilks pers.
comm. 1985) and consequently gave small values for most vegetative characters. When this is
taken into account, the stem and leaves of the hybrid more closely resemble those of O. simia,
although it has similar bracts to Aceras. Most of its labellum dimensions are also closer to O. simia,
especially its overall width, but it has a shorter ‘tail’ and ‘legs’, and its ‘limbs’ occur in the same
plane as its ‘torso’, resembling Aceras. Its labellum is intermediate in colour, and although it has
the papillae and spur of O. simia the former are far fewer and the latter is much shorter. The outer
and inner perianth segments of the hybrid are intermediate in size and bear markings characteristic
of both parents.

This mixture of parental and intermediate characters is typical of hybrids between closely related
species (Stace 1975).

MULTIVARIATE ANALYSES

Data for individual plants were subjected to multivariate analyses for two reasons: i) to test the a
priori assignment of plants to one of the three categories (Orchis, x Orchiaceras, Aceras) in Table 1,
and ii) to investigate the potential for identifying and describing hybrids of superimposing

STUDY OF x ORCHIACERAS BERGONII 403

oe

5mm

FicurE 3. Comparison of the flowers of Orchis simia (A), XOrchiaceras bergonii (B), and Aceras
anthropophorum (C).

404 R. M. BATEMAN AND O. S. FARRINGTON

PC2

-0.6 -0.4 -0.2 0 0.2 0.4 0.6
PC1
Figure 4. Minimum spanning tree superimposed onto a principal coordinates plot of PC1:PC2. Percentages are

Gower Similarities (links within parental groups all exceed 92.5%). @=Orchis simia, O=Aceras anthro-
pophorum, ©= X Orchiaceras bergonii.

a minimum spanning tree based on Gower Similarity Coefficients onto a principal coordinates
analysis (Fig. 4).

The mean of the maximum similarities of individuals of O. simia and Aceras is only 51.6+6.4%,
so it is not surprising that the parental species form two distinct groups separated by a wide
morphological discontinuity on the very strong first coordinate (PC1). Within each parental group,
separation on PC1 is slight and minimum spanning tree links between individuals are relatively
strong, exceeding 92.5% maximum similarity. The hybrid occurs between the parental groups,
slightly closer to the nearest O. simia than to the nearest Aceras. Its predictably weak minimum
spanning tree links also suggest a greater overall similarity to O. simia.

Table 2 shows that PC1 is determined by all of the characters defined as taxonomically useful in
Table 1, with diagnostic characters taking precedence. The much weaker PC2 represents seven
vegetative characters (six taxonomically useless) that are influenced by the size of the plant; it is a
‘vigour’ coordinate. PC3 is governed by four of the six characters (five taxonomically useless) that
yield values for the hybrid more extreme than both parents; it therefore separated the hybrid from
all the other plants measured.

DISCUSSION AND CONCLUSIONS

The Kentish XOrchiaceras bergonii is very similar to plants of this parentage described by
Continental orchidologists (Camus & Camus 1932; Nelson 1968; Peitz 1970). Indeed, similar
combinations of characters are shown by hybrids of Aceras with other species of Orchis subsection
Militares (Nelson 1968, plate 23; Peitz 1970, pp. 252-3; Sundermann 1980, p. 258). Vegetative
characters are variable and often intermediate between the parents, but the hybrids usually
resemble Orchis in inflorescence density and Aceras in bract size. Floral characters are more
predictable; x Orchiaceras resembles Orchis in overall labellum shape (although ‘leg’ lengths are
often intermediate), but is intermediate in ‘tail’ and spur lengths and perianth segment dimensions.
It is also intermediate in pigmentation characters, and the papillae of Orchis, although present, are
reduced in size and number. This consistency of expression in X Orchiaceras for many characters,
irrespective of the parental species of Orchis, suggests that they are controlled by major genes
whose dominance is relatively well-established.

STUDY OF x ORCHIACERAS BERGONII 405

TABLE 2. LIST OF CHARACTERS CONTRIBUTING APPRECIABLY TO THE FIRST THREE
PRINCIPAL COORDINATES

See Materials and Methods for descriptions of numbered characters. Characters in normal typeface increase in

value with increasing value of the vector, italicized characters decrease in value with increasing value of the

vector.
Percentage of total Characters contributing to the coordinates, listed in
Coordinate variance accounted for order of decreasing contribution
PC1 76.5 ZS 8s AOI 2/62 6.51'5, 399 32-58, 39, 30,50, 255,55
24% 34;29" 6, 20,2, 16,37, 41, 13,21, 4,7
PC2 5.8 LAS OS TTL 7, 7.
PC3 Si 10, 8, 22,6

The widespread occurrence of X Orchiaceras bergonii on the Continent shows that at least some
insects, possibly bees (the pollen vectors of O. simia and Aceras have not, to our knowledge, been
studied), visit both parents. However, field observations suggested that the Kent population of O.
simia was not being naturally pollinated, so a successful programme of artificial pollination was
introduced in 1958 (Wilks 1960) and is still performed annually (H. M. Wilks pers. comm. 1985).
Thus, the Kentish x Orchiaceras could have arisen artificially, although it is more likely that
natural pollination of O. simia does occur sporadically and that an insect paid a rare visit to an O.
simia and an Aceras; two individuals, one of each taxon, have grown and flowered a few
centimetres apart for several years (H. M. Wilks pers. comm. 1985).

Some authors (Bateman 1982; Davies et al. 1983) have argued that the frequent hybridization of
Aceras with species of Orchis subsection Militares casts doubt on its usefulness as a monotypic
genus. Although much shorter, the ‘nectar pit’ of Aceras is probably homologous with the spur of
Orchis; its shallower depth and the shorter distance between the viscidia of Aceras merely indicate
adaptation to smaller pollen vectors. With these exceptions, Aceras apparently lies within the
range of variation encompassed by the eight European species of Orchis subsection Militares, and
shares with them a chromosome number of 2n=42 (Sundermann 1980; Moore 1980; Sod 1980).
Furthermore, introgression of Aceras and Orchis has been reported by several authors (Keller &
Schlechter 1928; Peitz 1970), although the feasibility of identifying such hybrids has been
questioned by Wollin (1972). Together, these facts suggest that Aceras anthropophorum would be
more appropriately placed under Orchis.

Reliable identification of putative hybrids must rest entirely on morphological discrimination
from the parents if they have identical chromosome numbers and other, less readily obtained,
biosystematic data are not available. This requires careful and detailed measurement, not only of
the hybrid but also of any potential parental taxa; our study would have been invalid if we had
relied on descriptions of the parents in the literature as these can be misleading, e.g. in Flora
Europaea (Moore 1980; So6 1980).

Principal coordinates analysis is preferred to other multivariate methods of processing such data
as it avoids the a priori identification of the parental taxa inherent in Wells hybrid distance
diagrams (Adams 1982) and of hybrids in canonical variates analysis (Neff & Smith 1979). It is also
preferable to projecting a hybrid onto a pre-ordinated canonical variates analysis (McNeill 1984),
as the minimized intra-group distances mean that the suspected hybrid would be unlikely to occupy
the same multivariate space as its putative parents even if it is merely a morphological extreme of
one of the species. Superimposing a minimum spanning tree onto a principal coordinates plot of
unweighted data scaled to unit variance has proved excellent for i) identifying the hybrid, ii)
establishing its overall similarities to the parent taxa, iii) identifying characters most suitable for
separating the parental taxa and distinguishing the hybrid, and iv) identifying characters most
strongly influenced by vigour. Moreover, good separation of taxa has been achieved by principal
coordinates analysis without maximizing the inter-taxon distances using the F-1 weighted
similarities advocated by Whiffin (1977) and Adams (1982).

In this study, PC1 is much stronger than the other coordinates and separates the parental taxa,
leaving a morphological discontinuity occupied only by the hybrid. PC2 reflects variation in vigour

406 R. M. BATEMAN AND O. S. FARRINGTON

in all the taxa measured, and PC3 identifies the few characters that separate the hybrid from both
parents. This distribution of character types among the first three coordinates may be expected in
instances of hybridization where i) the parental taxa are separated by a morphological
discontinuity, ii) the hybrid plants are less numerous than either parent, and iii) most of the
characters of the hybrid fall within or between the parental ranges (categories 2—4). As all three
criteria are fulfilled by most examples of natural hybridization, principal coordinates analysis
represents a powerful tool for investigating hybridization and deserves to be more widely used.

ACKNOWLEDGMENTS

We thank the Kent Trust for Nature Conservation for permission to visit the locality, I. Denholm
for colour-matching the flowers, Lynn Parry for help with the computing and Joan Lund for typing
the manuscript. We also thank I. Denholm, D. M. T. Ettlinger, R. H. Roberts, J. T. Temple and
H. M. Wilks for critically reading the manuscript.

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(Accepted March 1986)

?

oe)

Watsonia, 16, 409-415 (1987) 409

The current distribution and abundance of Orchis ustulata L. in
northern England

Moy FOLEY

87 Ribchester Road, Clayton-le-Dale, Blackburn, Lancs., BB1 9HT

ABSTRACT

Orchis ustulata L. was formerly thinly scattered over many of the calcareous grasslands of northern England.
One hundred and eighteen previously recorded sites have been identified, and a survey of the present status of
the species at these shows that in recent years it has become extinct at many, and that there are now only 15
colonies which definitely survive. The past and present distribution of O. ustulata is illustrated and the causes of
decline discussed. An indication is also given of current colony strengths.

INTRODUCTION

Orchis ustulata L. is distributed throughout most of central Europe, extending northwards to
southern Scandinavia and southwards to the Mediterranean area (Summerhayes 1968). In Britain
it is confined to England, increasing in abundance to the south and east, and in the past was thinly
scattered mainly in the limestone and few chalk grassland areas of northern England. Here it is
usually found on old calcareous pastures which have been subject to light grazing. It is a plant
which has an extended period between seed germination and maturity (Summerhayes 1968).
The area covered in this survey includes the whole of northern England, namely v.cc. 54 and
56-70. The species is unrecorded for v.cc. 58, 59 and 68, nor is it recorded for Scotland; the only
record for Wales was in 1953 for Caerns., v.c. 49 (de Vesian 1982) and is almost certainly an error.
The following account of the distribution and status of O. ustulata is based upon field work
carried out by the author during the period 1982-86 and from investigation into old records.

DISTRIBUTION

118 records for O. ustulata have been traced for the survey area. These have been categorised into
periods of last known occurrence, viz. pre-1930, 1930-59, 1960-81, 1982 onward, and shown in
Figure 1 on a 5 km-square basis. The progressive extinction of the species from many sites can be
seen and there are now only twelve 5-km squares (15 sites) at which the species definitely survives.
The relationship between numbers of 5-km square records and those for individual sites for the
appropriate periods is shown in Table 1, where the assumption has been made that later recorded
sites were also present earlier and are given in the cumulative totals.

The great majority of records are at sites where calcareous rocks underly. These include the
Carboniferous limestone areas of Derbyshire, Yorkshire, Cumbria, and northern Lancashire, and
also the Magnesian limestone belt running from Nottinghamshire to Durham, where the species
seems to have become extinct at many sites only recently. The chalk Wolds of northern
Lincolnshire and eastern Yorkshire also have several records.

The main strongholds for the species in northern England in present times are the upland
pastures of Derbyshire, v.c. 57, where the ground is difficult to work agriculturally, and in certain
of the riverside pastures of N.W. Yorks., v.c. 65.

410 M_ J; YY. FOREY.

Date of last record
1982+
1960-81
1930-59
pre-1930

Ficure 1. The distribution of Orchis ustulata in northern England, v.cc. 54, 56-70. Plotted in 5-km squares.

TABLE 1. NUMBER OF 5-KM SQUARE RECORDS AND NUMBER OF SITES FOR THE PERIODS
OF LAST KNOWN OCCURRENCE OF ORCHIS USTULATA; CUMULATIVE TOTALS ARE

BRACKETED
Date of Number of
last record 5-km square records individual sites
1982 onward 12°12) 157 (5)
1960-81 15:(27) 29 (44)
1930-59 11 (38) 19 (63)

pre-1930 46 (84) 55 (118)

CURRENT STATUS

The current status of O. ustulata at each site is denoted overleaf (Records) by the letters: A
typically 1-10 flowering spikes; B = 11-25; C = 26-50; D = 51-200; E = 200+; PX
possibly extinct; X = extinct. Whilst numbers of flowering plants fluctuate annually it
is felt that the above scale gives a reliable reflection of abundance, although the balance between
numbers of flowering and vegetative plants is difficult to quantify. Studies of populations where
plants in the vegetative state are relatively easily detected suggest that well over half flower in a
good season, but noticeably less in a poor season. All existing sites and many other recent and less
recent ones have been visited by the author since 1982, and estimates of status are usually based on
observations made over several seasons. For old records, the authority for the last sighting is given

ORCHIS USTULATA IN NORTHERN ENGLAND 411
TABLE 2. ABUNDANCE OF ORCHIS USTULATA IN EXTANT COLONIES IN NORTHERN

ENGLAND

Colony
10-km square Number of colonies status*
34/9.8 i B
43/2.5 5 A,A,B,D,E
44/0.8 2 A,A
44/1.8 j Cie
44/2.8 1 A
44/7.1 1 @
44/8.8 if B
44/9.8 1 B
45/4.3 1 A

* using the code described on p. 410.

wherever possible, and often, from knowledge of the site, extinction or otherwise is inferred. For
relatively recent records, it is difficult to ascribe extinction unless the site has been physically
destroyed, since prolonged over-grazing and the somewhat intermittent flowering of the plant may
give a false impression of reduced status, and flowering plants may reappear when grazing
pressures relax or other factors change. This is particularly true for small colonies which in fact
comprise the bulk of recent records for the area. Overgrown, ungrazed habitats probably also lead
to ultimate extinction by crowding out, although in at least two currently ungrazed situations,
plants have survived by adapting to compete with tall surrounding vegetation, and have resulted in
spike heights of 15—25 cm compared to the more normal heights of 9-12 cm. At one heavily grazed
colony, small plants of only approximately 5 cm survive. During this survey a very few plants of the
rare white-flowered var. albiflora Thielens were found at two sites in Derbys., v.c. 57.

At least 118 sites have been recorded for O. ustulata in northern England, but only 15 surviving
colonies, most of them small, have been refound in this survey. It is possible that there may be a
few further sites at which the species survives, but this will only be apparent if and when excessive
overgrazing is reduced. At two of the 15 extant colonies, there were respectively 200+ and 69
flowering plants in 1985 (a good flowering year); however three other colonies rarely have more
than 35 spikes, and the rest are very much smaller and, in unfavourable seasons, numbers of spikes
will be at least halved in all colonies. One of the smallest is in Durham, v.c. 66, and is the most
northerly surviving in Britain. The species is now only to be found with certainty in nine 10-km
squares of northern England, compared to the 17 post-1930 10-km squares given by Perring &
Walters (1962), and to records for 32 10-km squares identified for the same period in the course of
this survey. The colony strengths of the 15 extant sites are shown in Table 2, and details of past and
present records are listed at the end. In order to protect the sites from over-attention, precise grid
references are not given, but full details have been lodged with the Nature Conservancy Council,
York, and the Biological Records Centre, Monks Wood.

CAUSES OF DECLINE

In northern England, O. ustulata, here at the limit of its range, is steadily decreasing as its habitat is
gradually destroyed. Table 3 shows the causes of loss for the majority of extinct sites. Agricultural
upgrading of pastures is the main cause — the application of inorganic fertilisers and ploughing
appear to be fatal to the plant — and the effects of over-grazing also probably lead ultimately to
extinction. Sites have also been lost to building and industrial encroachment, and a few to
incidental causes such as quarrying and afforestation, and at least one site was deliberately
destroyed in order to counter the attention shown it by botanists. Information received from
recorders of O. ustulata sites in southern England also reflects a similar decline usually through
agricultural causes.

Of the 15 extant sites covered by this survey, one has the protection of a nature reserve whilst

412 Mii Yee FOLEY:

TABLE 3. CAUSES OF LOSS OF 103 RECORDED SITES OF ORCHIS USTULATA IN NORTHERN
ENGLAND

Cause of loss of site Number of sites lost

Definite or suspected agricultural improvement 35
Building and industrial encroachment 18
Ploughed out

Overgrazed

Scrub colonisation

Afforestation

Quarrying

Overgrazed but may possibly still survive
Unknown causes

OoONnNrnNA BN

Nr

four others are at sites having S.S.S.I. status and a further site is in the process of being so
designated. One other site is in a permanent amenity area, which may in itself provide some degree
of protection, but the remainder appear to be unprotected in any way. Other than protection as
described above, the best chance of survival appears to lie where the ground is difficult to work
agriculturally because of its configuration or the presence of outcropping rocks, but this is the case
at only a few of the extant sites.

It is quite clear that the species is now very scarce in northern England, where maybe fewer than
500 flowering plants survive. Although in the past O. ustulata has been recorded from 13 vice-
counties in the survey area, it now survives with certainty in only five of these, and is considered to
be extinct in Notts., v.c. 56, W. Lancs., v.c. 60, S.W. Yorks., v.c. 63, S. Northumb., v.c. 67, and
Cumberland, v.c. 70. It may yet be refound in S.E. Yorks., v.c. 61, Mid-W. Yorks., v.c. 64, and
Westmorland, v.c. 69.

RECORDS

N. Lines., v.c. 54: 43/9.9, Waddingham, pasture with Orchis morio. Often overgrazed, last
recorded in 1980 but still thought to be extant with status probably (A/B) (I. Weston pers.
comm.); 44/7.1, Eastoft, hay meadow with O. morio. Tall plants in ungrazed situation (C); 44/
9.1 or 54/0.1, Worlaby, formerly in three small meadows which were combined and destroyed by
ploughing in the 1970s (X); 54/1.0, Swallow, pasture with Coeloglossum viride. Known here for
many years but not seen recently (PX).

Notts., v.c. 56: 43/5.4, Bulwell (Howitt & Howitt 1963) (X); 43/5.5, near Sutton-in-Ashfield
(Howitt & Howitt 1963) (X); 43/6.4, 43/7.4, 43/7.5, Trentside, between Newark & Nottingham
(Howitt & Howitt 1963) (X).

Derbys., v.c. 57: 43/0.7, Ashwood Dale (Clapham 1969) (X); 43/2.5, Hipley Hill, south and west-
facing slopes but not seen since 1960s although site still intact. Searched for in 1985 but not
refound (PX); near Hipley Toll Bar, limestone pasture (Clapham 1969), not found recently,
including searches in 1985 and 1986 (PX); near Ballidonmoor, (B), with a possible second small
site; west of Longcliffe, two sites on heavily grazed limestone pasture near pond, status possibly
(A/B) but not seen recently; east of Rainster Rocks, pasture, two small sites (A,A); near
Brassington, grazed limestone pasturé (D); head of subsidiary valley, Griffe Grange area,
heavily grazed, not refound (PX); south-east of Harboro’ Rocks, formerly possibly (C) but
heavily grazed and not refound (PX); south-east of Ible, scrubby south-facing slopes now
appreciably overgrown and not refound (PX); Hopton/Via Gellia area, south-west facing well-
grazed slope, extensive colony possibly the strongest in northern England (E); 43/2.6, Winster,
not seen recently but formerly (A) (F. Horsman pers. comm.); 43/4.5, Alfreton (Clapham 1969)
(X); South Normanton (Clapham 1969) (X).

W. Lancs., v.c. 60: 34/4.7, Silverdale area (Wheldon & Wilson 1907) (X); 34/5.6, Over Kellett,
believed to survive here until 1940s but not seen since (L. Livermore pers. comm.) (X).

ORCHIS USTULATA IN NORTHERN ENGLAND 413

S. E. Yorks., v.c. 61: 44/6.3, Barlby (Robinson 1902) (X); 44/7.4, Allerthorpe, recorded in 1932
(Naturalist, p. 14, 1932), presumably at Allerthorpe Common but now afforested (X); also
recorded for this 10 km-square, post 1950 (Biological Records Centre records) (X); 44/7.6,
Kenneythorpe (Robinson 1902) (X); 44/9.2, Brantingham Dale (Robinson 1902) (X); Brough
(Robinson 1902) (X); 54/0.4, Arram (Robinson 1902) (X); 54/0.7, near Fordon, chalk grassland,
three plants in 1965 (F. E. Crackles pers. comm.) (PX); Cottondale, formerly on the south
facing slope of an earthwork with O. morio, Coeloglossum viride, and Spiranthes spiralis, but
site destroyed in 1965 when ploughed out because of interest shown by botanists. Formerly 30-
60 plants (T. F. Medd pers. comm.) (X); 54/1.4, Hornsea Mere (Robinson 1902) (X); 54/1.7,
Hunmanby, single plant seen in 1973 (F. E. Crackles pers. comm.) (PX).

N. E. Yorks., v.c. 62: 44/4.8, Sowerby Ings (X); Spittal Hill (X); South Kilvington (X), all
described by W. Foggitt (c. 1880); South Villa, specimen collected June 1937, (herb. G. Foggitt)
(X); Woodend, ‘“‘abundant’’, (Baines c.1840) (X); 44/5.5, Clifton Ings (Auden 1906) (X); 44/6.8,
Helmsiey, near railway station in 1945 (T. F. Medd pers. comm.) (X); 44/7.7, near Malton in
1896 (B.S.B.I. records) (X); 44/8.8, Haugh Rigg, grazed pasture with Ophrys insectifera (B);
44/9.8, Snainton, field ploughed out in the 1940s (J. H. Bolton pers. comm.) (X); Brompton,
ploughed out in the 1940s (J. H. Bolton pers. comm.) (X); near Ruston, pasture, 100+ plants in
the past but now declined, with O. morio (B); Forge Valley (old B.S.B.I. records) (PX);
Yedmandale (B.S.B.I. records) (PX); 44/9.9, Highdale north of Hackness in 1932 (B.S.B.I.
records) (PX); 45/9.0, Robin Hood’s Bay (B.S.B.I. records) (X); 54/0.8, near Scarborough race
course and Seamer in 1938 (B.S.B.I. records) (X).

S.W. Yorks., v.c. 63: 43/4.9, Rotherham, L. Langley 1828 (Lees 1888) (X); 43/5.9, Conisborough,
“near Toll-Gate”’ in 1800, herb. Salt (Lees 1888) (X); 44/5.0, Scawsby Leys (Lees 1888) (X); 44/
5.1, Hampole, Smeaton, and Brockendale, three old records all Lees (1888) (X).

Mid W. Yorks., v.c. 64: 34/8.6, Settle, waste ground (Lees 1888; Riley 1923) (X); 44/2.7, Mickley,
(Lees 1888) (X); Clotherholme, (Slater 1883) (X); 44/3.4, Woodhall, field near the river, seen in
1958 but now possibly lost (PX); Collingham, meadows by the river, recorded in 1955 by B. A.
Kirkby (Biological Records Centre record), also pre-1954 by Butcher (1954) (PX); Linton
Common, meadow now mostly built upon and overgrown, not refound in 1985 but recorded in
recent past (Nelson 1963) (PX); Wattle Syke and Wetherby, riverside meadows not seen recently
(PX); 44/3.5, Knaresborough, east of Birkham Wood, lost to ploughing in 1971 (A. Mettam
pers. comm., Jowsey 1978) (X); 44/3.6, Burton Leonard, formerly at quarries but not seen since
the 1960s (M. Sanderson and others pers. comms.) (X); (Wormald Green Quarries (D. R.
Grant pers. comm.) is almost certainly the same site as the one described above for Burton
Leonard); 44/3.7, Ripon, riverside pasture upstream from the road bridge, known for many
years and status (B) in 1974, single plant seen in 1978 is last sighting (M. Sanderson pers.
comm.) and this when land was in the process of being upgraded agriculturally, not refound in
1983 and 1985 (PX); Queen Mary’s Dubbs, seen in 1957 (Biological Records Centre records) but
not recently, heavily grazed, overgrown in places and probably improved, not refound in 1985
search (X); near Bridge Hewick (Slater 1883) (X); 44/4.3, Sherburn, old record (Lees 1888) (X);
44/4.4, Stutton, Thorp Arch (Lees 1888) (X); Tadcaster (Auden 1906) (X); 44/5.4,
Bishopthorpe area (Lees 1888; Robinson 1902) (X); 44/6.2, Barlow, near Drax (Lees 1888) (X).

N.W. Yorks., v.c. 65: 34/6.9, Sedbergh/Lower Dentdale, stated to grow with O. morio but neither
species had been seen of late (Handley 1898) and also recorded for this area by Lees (1888) (X);
34/9.8, Bishopdale, small newly found colony in pasture (B); 34/9.9, near Muker, an
unconfirmed record in the early 1980s, not refound, present status doubtful; near Crackpot,
steep sloping pasture near stream, growing with Platanthera chlorantha, occurs intermittently
and not refound in searches in 1984-1986, status probably (A/B); 44/0.8, east of Aysgarth,
pasture, often overgrazed, (A); Redmire Force, pasture to south of river. Seen in small quantity
in 1975 (W. Sledge pers. comm.), not refound in recent searches (PX); Swinithwaite, very small
colony discovered by author in 1982 on closely grazed pasture, dwarf plants (A); Wanlas,
recorded by F. M. Tayler in 1973, not refound (PX); West Witton, found on Y.N.U. Excursion
in 1957 when status (B), but not refound in recent searches although site still intact (PX); 44/1.8,
Leyburn, controlled grazed pasture protected as a nature reserve, with O. morio, (C), and
unconfirmed report nearby on disused railway line; East Witton, classic site near river, tall
plants in ungrazed situation, known here for many years but site now reduced in area with 100+

414 M. J. Y. FOLEY

plants as recently as the 1970s, but now much less (C), plants may also occur on the nearby river
banks but not refound; 44/2.7, above Tanfield, on the north bank of the Ure (Foggitt c.1880;
Slater 1883) (X); 44/2.8, near Masham, small colony in pasture (A); 45/1.0, Round Howe and
Lower Swaledale (Baker 1863) but not recorded for many years (X).

Durham, v.c. 66: 45/2.1, Baydales (Winch 1832) (X); 45/2.5, Lambton Gate (Winch 1832) (X);
45/3.1, Dinsdale, near the river (Baker & Tate 1868) (X), Middleton-One-Row (Baker & Tate
1868) (X); 45/3.3, Bishop Middleham, quarry, destroyed by working in the mid-1970s (A. F.
Porter pers. comm.) (X); 45/3.6, Fulwell and Boldon Hills (Winch 1832) (X); Cleadon Hills, on
the east side (Winch 1832) (X); 45/4.3, near Hart, coastal grassland, most northerly extant site in
Britain (A); 45/4.5, Ryhope, sea coast (Winch 1832) (X); 45/4.6, Marsden, sea coast (Winch
1832) (X).

S. Northumb., v.c. 67: 45/3.7, Cullercoats, coastal links (Baker & Tate 1868) (X), long since
extinct and formerly most northerly British record.

Westmorland, v.c. 69: 34/4.7, Arnside Knott, heathy ground (Wilson 1938) (X); New Barns,
Arnside, formerly at site of present caravan complex, known here in the 1930s. (L. Livermore
pers. comm.) (X); Black Dyke, not seen here since 1930s (M. Baecker pers. comm.) (X); 35/6.1,
Drybeck (Wilson 1938) (X); Burrells and Whirby (Wilson 1938) (X); 35/7.1, Warcop, thought to
survive here until the 1960s (X); Great Ormside, known on the banks of the Eden but ploughed
out in the 1970s and not seen since (C. Willinck pers. comm.) (X).

Cumberland, v.c. 70: 35/1.3, Woodhall (Hodgson 1898) (X); Baker (1885) gives Woodhall,
Keswick, presumably an error?; Tallentire, pasture (Hodgson 1898) (X); Blindcrake (Hodgson
1898) (X); 35/3.4, meadow below Raughton Head (Hodgson 1898) (X); 35/3.5, Stainton Banks
(Hodgson 1898), single plant seen here in 1927 (D. Blezard in records of E. Blezard) (X); 35/4.3,
Lamonby, rough pasture, site reclaimed and reseeded, not seen since early 1970s (E. H. Rhone
pers. comm.) (X); 35/4.5, Crosby-on-Eden, single plant in 1889 (Hodgson 1898) (X); 35/5.3,
Great Salkeld, known here until the 1960s but the site now a plantation (E. H. Rhone pers.
comm.) (X); Edenhall, meadows (Hodgson 1898), several plants in meadow by R. Eden in this
area in c.1930 (J. R. Parker in records of E. Blezard) (X).

A 1953 record for Caerns., v.c. 49, near Pabo (de Vesian 1982) is based upon a herbarium

specimen in NMW and is considered to be a geographical error (R. G. Ellis, W. S. Lacey, M.

Morris pers. comm.).

I would be interested to hear of any other records or of any newly discovered or refound colonies
especially within the survey area.

ACKNOWLEDGMENTS

I am grateful to the many people who have helped with information, and especially to Mrs I.
Weston and T. F. Medd.

REFERENCES

AUDEN, G. A., ed. (1906). Historical & scientific survey of York and district. York & London.

Baines, H. (c.1840). The flora of Yorkshire. London.

Baker, J. G. (1863). North Yorkshire — studies of its botany, geology, climate and physical geography. London.

BAKER, J. G. (1885). A Flora of the English Lake District. London.

Baker, J. G. & TaTE, G. (1868). A new Flora of Northumberland and Durham. Newcastle.

ButcHer, R. W. (1954). Colchicum autumnale L., in Biological Flora of the British Isles. J. Ecol., 42: 251.

CLAPHAM, A. R. ed. (1969). Flora of Derbyshire. Derby.

DE VESIAN, D. E. (1982). Plant record. Watsonia, 14: 197.

Foacaitt, W. (c.1880). Notebooks. Records held by Yorkshire Naturalists’ Union Recorder.

HANDLEY, J. (1898). Catalogue of plants growing in the Sedbergh District including the Lune Basin from
Middleton to Tebay. Leeds.

Hopecson, W. (1898). Flora of Cumberland. Carlisle.

Howrrt, R. C. L. & Howrrr, B. M. (1963). A Flora of Nottingham. Private.

Jowsey, W. H. (1978). Botanical atlas of the Harrogate District. Harrogate.

ORCHIS USTULATA IN NORTHERN ENGLAND 415

Legs, F. A. (1888). The flora of West Yorkshire. London.

NELson, G. A. (1963). A Flora of Leeds & District. Proc. Leeds Phil. Lit. Soc., 9: 153.

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

Riney, F. (1923). The Settle District & north west Yorkshire Dales. Settle.

Rosinson, J. F. (1902). The flora of East Riding of Yorkshire. Lordon & Hull.

SLATER, H. H. (1883). The flora of Ripon & neighbourhood. Trans. Yorks. Nat. Union, Part 8. London.

SUMMERHAYES, V. S. (1968). Wild orchids of Britain, 2nd ed. London.

WHELDON, J. A. & WiLson, A. (1907). The flora of West Lancashire. Private.

Witson, A. (1938). The flora of Westmorland. Arbroath.

WincHu, N. J. (1832). Flora of Northumberland & Durham. Trans. Nat. Hist. Soc. Northumberland, 2, Part 1.
Newcastle.

(Accepted October 1986)

“et

aA

Watsonia, 16, 417-426 (1987) 417

The flora of the marl-pits (ponds) in one Cheshire parish

A. D. BRIAN
Bodenham Hall East, Bodenham, Hereford, HRI 3JT

PS ERICE

Folly Farm, Holme Lacy, Hereford, HR2 6LS
B. C. REDWOOD
The Old Cider House, Kinnersley, Hereford, HR3 6QD
and
E. WHEELER
28 Forest Avenue, Goostrey, Holmes Chapel, Cheshire, CW4 8LU

ABSTRACT

All the 153 ponds present in the 13 km? area of Christleton parish (Cheshire) were assessed for various physical
characteristics and for age. Positive associations were established between degree of shading and age of pond,
alkalinity and age, and alkalinity and percentage of open water. The herbaceous, macrophyte species were
recorded for each pond. High numbers of species were associated with large, unshaded ponds with a moderate
amount of open water. Some of the common individual species showed positive associations with certain physical
characteristics of the ponds. 41% of the ponds in the parish have been lost since 1844. A simulation of the effect of
pond loss on the numbers of plant species indicated that from 1844 to the present only seven species may have
been lost, but if ponds continue to disappear this will have a much more drastic effect on the pond flora in the
future.

INTRODUCTION

This study was carried out in 1971-72 as a part of a general parish natural history survey which
involved recording the land use of the fields and all trees, hedges, etc. in the parish of Christleton, 3
km from Chester. It became apparent during the study that in this parish, as in the rest of this part of
the Cheshire plain with its intensive farming, rigorously trimmed hedges and shortage of woods, the
old marl-pits (now ponds) were much the richest areas for wild life. Indeed, the Cheshire marl-pits
have been noted for this for many years.

The origin of marl and the history of marl digging have been covered by Day etal. (1982) and many
of his comments apply to Christleton parish. Ponds in this part of Cheshire, as in Lancashire, betray
their origin by being known locally as ‘pits’ rather than ‘ponds’ and one exceptionally large one of
8400 m? in the village of Christleton itself is called ‘The Pitt’. However in this paper the word pond is
used throughout.

There have been a number of studies of the flora of flooded marl-pits (e.g. Edmondson 1967) and
in most of these the study area has been extensive and only certain ponds within the area have been
selected for study. For example, in the Fylde of Lancashire 62 ponds out of about 4000 were selected
to study flora distribution by tetrads, in the Lower Dee Valley about 400 ponds in five study areas
were selected to assess the flora and determine a method of evaluating conservation interest and in
the Wirral 52 ponds, likely to be of conservation interest, were selected (Day 1981; Day et al. 1982).

In contrast the Christleton study covered only a small area but every pond site found on any map
from the tithe map of 1844 onwards was visited and if the pond still existed the flora was recorded.
The two methods of study, sample ponds chosen from a large area and the examination of every pond
within a small area, were thus complementary and a comparison of results obtained by the two
methods may prove helpful in planning future surveys of this type.

418 A. D. BRIAN ET AL.

Alluvium

Boulder clay

Bunter sandstone

2 miles <— é ) Glacial sand
ci ea

Ches tet

scale 1 25000

Figure 1. The distribution of ponds in the parish of Christleton, Cheshire.

METHODS

The study area (12.8 km*) comprised the whole of Christleton parish, GR 33/45.66, excluding a
small part with four ponds north of the A51, GR 33/465.675. The parish is low lying and slopes to
the River Gowy on the east and towards the River Dee on the west. Most of the area is covered by
boulder clay but there is a band of New Red Sandstone across the parish on which the church and
village are built. The marl-pits were dug in the boulder clay areas (Fig. 1). Details from the five
ponds in the parish that were not marl-pits in origin have not been included in this paper.

During 1971 and 1972 every field in the parish was examined thus ensuring that every existing
pond was found, including a few which, while giving every indication of being as old as the other
ponds, were not marked on any map. Each of the 153 ponds found was given a reference number
within each tetrad thus conforming with other plant recording projects in the county. Five
characteristics were recorded in the field for each pond and a sixth, age, was estimated from maps.
A list was made of all herbaceous macrophyte species associated with aquatic habitats that were
present but their abundance was not recorded (Table 1). Trees were also recorded but have not
been included in the analysis except in so far as they may have cast shade on the ponds.

Brief notes were made on animals seen.and factors of conservation interest, extent of dumping,
if any, and use of the pond for angling or shooting. Some of the ponds were surveyed more than
once within the growing season but the differences found on successive visits were not large enough
to warrant second visits to all ponds. All the information has been deposited at the Cheshire
County Record Office, Chester, together with the other data from the Christleton Parish Survey.

The methods used to record and categorize each of the six pond characteristics are described
below and listed in Table 2.

Degree of shading. This was estimated by eye in the field to the nearest 10%, taking into account
the height of any trees, how close they were to the water and their position relative to the pond and

FLORA OF MARL-PITS IN CHESHIRE 419

TABLE 1. NUMBER AND PERCENTAGE OF PONDS IN WHICH THE DIFFERENT SPECIES

OCCURRED

Occurence in ponds
Species Number %
Alisma plantago-aquatica 109 71
Angelica sylvestris 2 1
Apium inundatum 3 2
A. nodiflorum 15 10
Berula erecta 43 28
Bidens cernua 4] AH
B. tripartita 6 4
Butomus umbellatus 1 1
Callitriche spp. 40 26
Caltha palustris 5) 3
Cardamine pratensis 32 21
Carex acutiformis 1 1
C. pseudocyperus 24 16
C. otrubae 3 2
C. riparia 1 1
Cicuta virosa 8 5
Eleocharis palustris 11 7
Elodea canadensis 17, 11
Epilobium hirsutum 64 42
E. parviflorum 7 5
Eriophorum angustifolium 1 1
Filipendula ulmaria 5 3
Galium palustre 52 34
Glyceria fluitans 2) 1
G. maxima 1 1
Hottonia palustris + 3
Hydrocharis morsus-ranae 22 14
Hydrocotyle vulgaris 8 3)
Hypericum tetrapterum 4 3
Tris pseudacorus 25 16
Juncus acutiflorus 24 16
J. bufonius 8) 3)
J. effusus 91 60
J. inflexus 70 46
Lemna gibba 2 1
L. minor 104 68
L. polyrhiza 4 3
L. trisulca 18 12
Lotus uliginosus 25 16
Lychnis flos-cuculi Tf >
Lycopus europaeus 15 10
Lythrum salicaria 5) 3
Mentha spp. 7 5
Menyanthes trifoliata 3 2.
Myosotis laxa subsp. caespitosa 8 5)
M. scorpioides 38 25
Nasturtium officinale 19 12
Nymphaea alba 2 1
Oenanthe fistulosa 41 27
Polygonum amphibium 4 3
P. hydropiper 1 1
Potamogeton berchtoldii 1 1
P. obtusifolius 1 1
P. natans 61 43
Potamogeton sp. 2 1

420

TABLE 1. continued

Species

Potentilla palustris
Ranunculus aquatilis
R. sceleratus

Rorippa amphibia

R. islandica

Rumex hydrolapathum
Scrophularia auriculata
Scutellaria galericulata
Solanum dulcamara
Sparganium emersum
S. erectum

Stachys palustris
Stellaria alsine

Typha angustifolia

T. latifolia

Veronica beccabunga
V. catenata

V. scutellata

Azolla filiculoides
Equisetum fluviatile

Aulacomnium sp.*
Calliergon cordifolium*
Fontinalis antipyretica
Polytrichum sp.*
Riccia fluitans
Ricciocarpus natans
Sphagnum spp.

A. D. BRIAN ET AL.

* Bryophytes that were only recorded when very abundant.

Occurence in ponds

Number %
14 9
V 5
58 38
il 1
1 1
2. il
14 9
il 1
84 55
5 3
88 58
2. 1
2 1
5 3
34 22
Di 14
2 il
il 1
4 3
28 18
3 p)
1 1

1 1
3 2
2 il
4 3
4 3

TABLE 2. THE PHYSICAL CHARACTERISTICS OF THE PONDS AT THE TIME OF THE SURVEY

Characteristic

Shading

% open water

pH*

Size in hectares

Age

Fence

Category

unshaded

medium shade

very shaded

91-100
10-90
0-9
below 6.0
6.0-8.9
over 9.0

below 0.04

0.04—0.08
over 0.08

dug before 1844
dug after 1844

‘marsh ponds

present
absent

IK

* The pH was not recorded for 22 (14%) ponds.
** “marsh ponds’ are those that started as ponds but at some intermediate period of their existence have been
marked on a map as marsh rather than open water.

Number of ponds

% of ponds

FLORA OF MARL-PITS IN CHESHIRE 421

especially whether any were on the south side. Ponds were finally classified as unshaded (0O-9%),
medium shade (10-90%) and very shaded (over 90%).

% of open water. This was estimated in the field by eye to the nearest 10% but for analysis the ponds
were grouped into three categories: 0-9%, 10-90% and 91-100% open water. Most of the ponds
were visited during the height of the growing season so that the results would be comparable.
pH. This was estimated in the field using Universal Indicator and for analysis the ponds were
grouped into those with acidic (below pH 6.0), neutral (pH 6.0—8.9) and alkaline (over pH 9.0)
water.

Size. The area of each pond was measured from the 1:10560 O.S. map, 1911 and 1913 editions, and
the ponds were grouped into small (below 0.04 ha), medium (0.04—0.08 ha) and large (over 0.08
ha).

Age of pond. The presence or absence of each pond recorded in the 1971/72 survey was followed
from the first map on which it appeared starting with the tithe map of 1844 and through five
editions of O.S. maps. The majority of the ponds were present on the tithe map and, since these
could have been dug very much earlier than this date, it was decided to divide ponds into two age
groups only, those made before and after 1844. However, an interesting third group emerged from
the map study. Some of the ponds were marked as open water on both an early map and on a later
one but at some intermediate period were marked as marsh. 45 ponds fell into this category and are
referred to subsequently as ‘marsh ponds’. The cause of these changes is not known but it is
unlikely to be associated with changes in the general level of the water table since this has been
falling steadily over the period in question and the water level in the ponds represents a perched
water table.

Presence or absence of a fence. Fences were found in various stages of decay. A pond was recorded
as fenced if cattle were effectively excluded from the margin of the pond and grazing thus
prevented.

RESULTS

RELATIONSHIPS BETWEEN THE VARIOUS PHYSICAL CHARACTERISTICS

Table 2 shows the various characteristics, their categories and the number and percentage of ponds
found in each. Chi-squared tests were carried out for each of the possible pairs of variables to
determine if they were associated. The results of these are shown in Table 3 and comments on the
significant associations are given below.

Degree of shading and age. A highly significant positive association shows that the older ponds dug
before 1844 were more shaded than those dug later. This result was to be expected since the longer
a pond had been present the more chance there would have been for colonization of the banks and
tree growth. It also implies that although the date chosen to divide the ponds into two age groups
was arbitrary it has real meaning and the older ponds must have been dug well before 1844 since
even the younger ones were mainly dug soon after that date.

Degree of shading and presence of a fence. This positive association was also to be expected since a
fence, by preventing cattle grazing, gives trees a chance to become established. Shaded ponds now
unfenced may well have been fenced at some earlier date.

% of open water and pH. A positive association was found between % of open water and pH; open
water tended to be alkaline whilst ponds with little open water tended to be acidic. Certainly the
most acidic ponds found were those where the centre was covered with Sphagnum. These ponds
formed a very distinct group and would seem to represent very small floating bogs (Schwingmoor).
Day (1981) found only two ponds with Sphagnum spp. whereas in the much smaller area of this
survey there were four. Sinker (1962) reported a ‘“‘miniature example” of a Schwingmoor in
Shropshire where the surface had been completely overgrown by Sphagnum in the course of 50
years. These ponds may have misled the O.S. surveyors and one at least, although quite deep at
the time of the survey, was marked on a recent map as a marsh.

Age and pH. The older ponds were significantly more alkaline than the post-1844 ponds. An
explanation of this might be that marl was first dug in areas of the parish known to be the most
lime-rich and that the practice spread to the less lime-rich areas at a later date until over half the
fields came to have their own mar! pit.

422 A. D. BRIAN ET AL.

TABLE 3. RELATIONSHIPS BETWEEN THE PHYSICAL CHARACTERISTICS OF THE PONDS
AND THE TOTAL NUMBER OF SPECIES PER POND
Chi-squared values are given in the top, right section of the Table and their probabilities, where significant, in
the bottom, left section.

% open Number
Shade water pH Size Age Fence of species
Shade — e377, 170 1.84 15.36 6.50 13.80
% open NS — 6.83 3.61 2.61 0.88 12:22
water
pH NS <0.01 _ 0.18 8.04 3.41 0.20
*
Size NS NS NS sas eae 4.44 15.70
0.74
*
Age <0.001 NS <0.01 | aR 2 0.01 0.75
Fence <0.02 NS NS NS NS — 0.54
Number <0.001 <0.001 NS <0.001 NS NS —
of species

* The top figure shows the relationship between size and ‘marsh’ or ‘non-marsh’ ponds, the lower figure the
relationship between size and pre- or post-1844 ponds.
NS Not significant.

The size of marsh ponds. The size of ponds was not associated with any of the other physical
characteristics measured, except that those ponds that at some stage in their history had been
marshes were significantly smaller than ponds that had always been open water. Probably smaller
ponds would also have been shallower and thus more likely to turn into marshes than larger ponds
if the water supply was in some way restricted.

RELATIONSHIPS BETWEEN THE NUMBER OF PLANT SPECIES IN A POND AND ITS PHYSICAL CHARACTERISTICS
There is general agreement that the more species there are present in an area the more valuable
that area is likely to be for conservation. Consequently the relationships between the total number
of herbaceous macrophyte species in a pond and each of the physical characteristics listed above
was also determined. To obtain numbers suitable for analysis the ponds were divided into only
two, roughly equal, groups, those with 0—9 species and those with 10-24 species. The results are
included in Table 3.

Shading. There was a highly significant association between the number of species and the degree
of shading, the unshaded ponds having more species. This was to be expected and agreed with the
findings of Day (1981). It was for this reason that Day et al. (1982) decided not to survey shaded
ponds.

% of open water. There was a highly significant association between the total number of species
and the % of open water but in this case most species were found in the intermediate category of
10-90% open water. The explanation of this perhaps lies in the fact that these ponds are at an
intermediate stage of the hydrosere and so have the largest range of water plants. Ponds with over
90% of open water lacked emergent water plants around the edge and ponds with under 10%
lacked submerged and floating plants.

Size. There was a highly significant association between total number of species and size, the larger
ponds having the most species. This agreed with Day’s (1981) study in Clwyd and is a relationship
found in other types of habitat.

Age of pond. No association was found here. This was in disagreement with the results of Godwin
(1923) who found a steady increase of species with age. However all his ponds had been dug more
recently than the Christleton ponds, the oldest being less than 100 years old. In contrast, all the
Christleton ponds would have had time to reach a stable number of species. In addition, any
changes in water level, for which there is some evidence, would mean that the stage in the
hydrosere reached by any one pond would not be related to the age of the pond. Periods of
changed water level have been shown to alter the plant community elsewhere in Cheshire (Willis &

FLORA OF MARL-PITS IN CHESHIRE 423

Young 1983) and in Czechoslovakia (Dykyjova & Kvét 1978). In the woodland habitat too it has
been reported that there is no evidence for an increase in plant diversity with woodland age (Reed
& Grove 1981).

To summarize, in general, the ponds with the most diverse flora were those of a good size with
only a moderate amount of open water and unshaded, with trees, if present, on the north side.

RELATIONSHIPS BETWEEN INDIVIDUAL PLANT SPECIES AND THE PHYSICAL CHARACTERISTICS OF THE PONDS
The relationship between each of the individual plant species and each of the six physical
characteristics of the ponds was determined by chi-squared tests. Only five species were found to
be positively associated with individual physical characteristics, all at the P<0.05 level.

Alisma plantago-aquatica, Lemna minor and Potamogeton natans were all positively associated
with 10-90% open water. Oenanthe fistulosa and Typha latifolia were positively associated with
large ponds. Thus many of the commoner species apparently had no preferences. There were
indications that many of the less common species favoured certain physical characteristics of the
ponds but they occurred in too few ponds for analysis. There was therefore every indication that
further work of a similar sort on a larger area with more ponds would yield worthwhile results.

THE LOSS OF PONDS AND THE LOSS OF POND FLORA

There is a good deal of information on the loss of ponds in recent years and this has been
summarized in Day et al. (1982) for different parts of the country. In Christleton parish the figures
for total number of ponds at different dates are as follows: 1844 — 263 ponds; 1908 — 237 ponds;
1972 — 156 ponds (Latham 1979). From these figures it is apparent that the rate of loss has
increased considerably. In the 64 years from 1844-1908, 26 ponds disappeared while in the same
period from 1908-1972, 81 ponds disappeared. If the trend were to continue in the future there
would be few ponds left by the year 2250 unless new ponds were made. Of the 81 lost ponds, 57
have reverted to pasture, some naturally, some after deliberate infilling, 13 are rubbish dumps and
eleven have been replaced by buildings.

From the results of the survey it is possible to estimate the effect of future loss of ponds on the
total pond flora by using the data for each pond. The effect of the loss of varying numbers of ponds
on the total pond flora of the parish was simulated by removing different numbers of ponds, taken
at random from the data set, and recording the total number of species remaining. Each simulation
was repeated four times and the average number of species ‘lost’ was recorded. The results are
shown in Fig. 2 where an empirical curve has been fitted to the points.

These results show that the number of species remaining declines more rapidly as more ponds
are lost since the gradient of the curve becomes greater as the number of ponds decreases. In 1844,
263 ponds were marked on the tithe map and it is estimated that these would have contained 97
species, while in 1908, when there were 237 ponds marked on the map, there would have been an
estimated 95 species. Since the curve flattens out very considerably, even had there been a much
larger number of ponds in the more distant past, this might not have meant very many more species
in total. Presumably there is a limit to the number of species that would grow under the prevailing
physical and climatic conditions.

Because of differences in the methods of recording it has only been possible to find where the
results of one other comparable survey of marl-pits would fall relative to the curve in Fig. 2. Day
(1981) recorded the total number of species from 405 ponds in Clwyd. When some species are
omitted from his list because, even if present, they would not have been recorded in the
Christleton survey (e.g. Salix spp.), his survey area falls quite close to the extrapolated curve.

On the other hand, Godwin’s Derbyshire ponds (1923) had a much richer flora while the ponds
surveyed by Jones (1971) in Leicestershire had a much poorer flora. Points for both these are
shown on Fig. 2. Neither of these sets of ponds owed their origin to marl digging.

If it is assumed that the extrapolated curve gives a reasonable estimate of the situation in the past
then between 1844 and the time of the survey in 1971 only about seven species may have been lost
in the parish although over 100 ponds have gone. In contrast the loss of a further 100 ponds might
mean the loss of nearly 30 species. Although these figures have been derived from a simulated
exercise they do show the urgency of preserving as many as possible of the ponds that still remain.
If action is taken now it is probably not too late to preserve the bulk of the flora of the ponds.

Total number of plant species

424 A. D. BRIAN ET AL.

O. Christleton Survey
© Figures derived from the simulation
x Actual number of ponds in 1884 and 1910, and derived number of species
[ * Day (1981)
4 Godwin (1923) Derbyshire
120
[ B® Jones (1971) Leicestershire
100 r is eS era
DC ae
| ee
| a.
60
4 yes
Al /
B
20}

= = 4 4 4 = is — —s
(0) 50 100 150 200 250 300 350 400 450 950 1000

Number of ponds

Ficure 2. Relationship, derived from a simulation, between the number of ponds and the total pond flora of
the parish.

DISCUSSION

THE SPATIAL DISTRIBUTION OF THE POND FLORA
One of the most striking features of the flora of the Christleton ponds was the great variation in
species between different ponds. Only five species occurred in over 50% of the ponds while 54
species occurred in less than 10%. One of the original aims of the survey was to try to find some of
the underlying reasons for this distribution pattern. Some of the possible contributory factors are
discussed below.

The flora available locally. This must have been an important factor in the initial colonization of
the ponds as pointed out by Day (1981). These plants would have been growing in the marshes and
ditches before the days of intensive agriculture. One such area is still present in the parish, a wet
meadow beside the River Gowy, now a nature reserve managed by the Cheshire Conservation
Trust. A comparison of the plant list for the reserve with that for the ponds showed that the two
areas had 38 wetland species in common while 37 species occurred in the ponds only and ten
species in the meadow only. Obviously the old marl-pits have indeed provided a refuge for many of
the wetland plants.

The age of the ponds. The length of time that a pond has been available for colonization might be
expected to affect the composition of the flora and Godwin (1923) has shown that for newly dug
ponds there was a steady increase of species up to 70 years. Most of the Christleton ponds are older
than this and no connection between age and flora was detected.

The rate of dispersal of water plants. A comparison was made between the number of species in
each pond and the distance of each pond from other neighbouring ponds. No association was
found, isolated ponds being often just as species-rich as ponds that had others near them. When
individual species were considered there was again little evidence that any one species occurred
mainly in neighbouring ponds and in most cases a species appeared to be scattered over the area.
The three commonest species in Christleton, Alisma plantago-aquatica, Lemna minor and Juncus
effusus, were also the three commonest in the Fylde, Wirral and Clwyd (Day et al. 1982). Similarly,
species rare in Christleton, like Lemna polyrhiza and Hydrocharis morsus-ranae, were also rare in

FLORA OF MARL-PITS IN CHESHIRE 425

the other surveys. These differences must be due in part to differences in the efficiency of their
dispersal mechanisms over land and Godwin (1923) came to the conclusion that the distribution of
the species in his seven ponds was initially a matter of chance and that the early distribution pattern
persisted because “‘land barriers are very effective in slowing down the rate of dispersal of water
plants”’.

The physical characteristics of a pond. The position of a pond, the type of soil where it was dug and
its original size must also have had some effect on the flora. Many of the earlier pond flora studies
commented that larger ponds had a richer flora and this study has shown also that individual
species were affected by the size of the pond. The original depth and steepness of the banks must
also have had their effect on the flora.

The history of the pond post-construction. The management of the area around a pond affects the
flora by producing different conditions which favour different species. The effect of a fence has
already been described. Absence of a fence leads to trampling by cattle and a shallow edge to a
pond.

Deliberate introductions. These have probably taken place in the past, especially of attractive
plants like the white water-lily, Nymphaea alba, which was said to have been introduced by a past
land-owner into one pond near his house.

Obviously with so many different potential factors at work it would seem inevitable that there
should be large differences in the flora of the different ponds. This variation between adjacent sites
of similar habitat type is a general phenomenon and a very similar distribution of woodland species
has, for example, been found recently in a woodland survey in Herefordshire (Barfield et al. 1984).

A COMPARISON BETWEEN THE INTENSIVE CHRISTLETON SURVEY AND THE EXTENSIVE CLWYD SURVEY
The Clwyd pond survey is the only one of those described in Day et al. (1982) which gives
sufficiently detailed results for a meaningful comparison to be made with the Christleton survey.
Because of the different methods used for selecting the ponds for survey in the two areas, a
comparison of the results may prove useful in the planning of future surveys and these results are
set out in Table 4.

The Christleton ponds appear to be rather larger than those in Clwyd though this could be
caused by the different methods used to assess the areas. The number of species recorded was
fewer in Christleton, partly perhaps because fewer ponds were visited but also possibly because of
a bias in the Clwyd survey towards ponds known to be of interest. On the whole however it is
apparent that the two sets of ponds have very similar characteristics and flora and that the different
methods used to select the ponds for survey have not affected the results significantly. There is now
enough information on the physical characteristics and flora of the old marl-pits of the Cheshire
plain and surrounding areas to show that there is overall uniformity combined with a good deal of
variation in flora from pond to pond.

CONSERVATION
The importance of old flooded marl-pits as refuges for wildlife in intensively farmed areas has been
discussed by Day et al. (1982) and the Christleton survey results reinforce their conclusions.

At the time of the Christleton survey a scheme for evaluating the ponds in the parish was
prepared (Brian et al. 1975; Latham 1979). A rather broader list of criteria was used than that of
Day (1981) since, in addition to number of species per pond and rarity of species, the final pond
score was weighted by aesthetic factors, potential for educational use, accessibility and the
likelihood that for some time at least the pond would remain undisturbed. This evaluation was
more in line with that given in Spellerberg (1981) but nevertheless produced a similar proportion
(5%), of what might be called ‘first class’ ponds, to that estimated by Day (1981) in Clwyd.

In view of the variation demonstrated between different ponds there is no doubt that the general
flora, and probably the fauna as well, would be much impoverished if only the ‘first class’ ponds
survived even if transplantation was carried out. Nearly every pond has some value and
discrimination between ‘first class’ and ‘other’ ponds might hasten the loss of the ‘other ponds’.
Even if a pond has little of interest or variety from the point of view of its flora it very often
functions as a small nature reserve in an otherwise intensively managed field. Many ponds have
mature trees and patches of scrub valuable for non-aquatic birds and insects, and the banks
harbour meadow plants now eliminated from the fields around.

426 A. D. BRIAN ET AL.
TABLE 4. A COMPARISON OF THE CHRISTLETON AND CLWYD SURVEYS

Christleton Clwyd (Day 1981)

Date of survey 1971/72 1979
Number of ponds surveyed 153 406
Number of ponds per km”, mean (range) 12.0 (3-34) 11.2 (0-443)
Position of ponds in field

(i) away from hedge 57% 35%

(ii) edge of field 43% 60%
Size of ponds, mean (range), m7 490 (40-1800) 242 (23-1472)
pH mean (range) 7.6 (5-10) 7.1 (5.7-8.6)
% of ponds unshaded (0-9% shade) oy 49
% of ponds fenced Di, 14
Total number of plant species 84 112
Number of species per pond, mean (range) 9 (0-23) 15.6 (0-30)
Number of species common to both areas "3 73
Number of species not found in the other area 10 39
% of ponds with rubbish dumping 25 46
Loss of ponds 34% in last 64 years 32%

* Excluding those species that would not have been recorded in the Christleton survey even if present.

REFERENCES

BaRFIELD, T., LOVELACE, D. & BENNEWITH, O. (1984). A Herefordshire woodland survey. Herefordshire and
Radnorshire Nature Trust.

Brian, A. D., REDwoop, B. R. & WHEELER, E. (1975). How well do you know your ponds? Cheshire
Conservation Trust.

Day, P. (1981). Ponds (marl pits) in the Lower Dee Valley of Clwyd — a preliminary appraisal. Nature
Conservancy Council (North Wales Region) Report.

Day, P., DEADMAN, A. J., GREENWOOD, B. D. & E. F. (1982). A floristic appraisal of marl pits in parts of
north-western England and northern Wales. Watsonia, 14: 153-165.

DykysovA, D. & Kvet, J., eds. (1978). Pond littoral ecosystems, in Methods and results of quantitative
ecosystem research in the Czechoslovakian I.B.P. wetland project. Berlin.

Epmonpson, T. (1967). Pond flora in north Cheshire. Fld Nat., 12: 30-35.

Gopwin, H. (1923). Dispersal of pond floras. J. Ecol., 11: 160-164.

JONES, R. C. (1971). A survey of the flora, physical characteristics and distribution of field ponds in North East
Leicestershire. Trans. Leicester lit. phil. Soc., 65: 12-31.

LatuaM, F. A. (1979). Christleton. The history of a Cheshire village, pp. 116-120. Chester.

REED, T. & Grove, R. (1981). Area and isolation symposium: Monk’s Wood, October 1980. Ecos, 2 (1): 29.

SINKER, C. A. (1962). The north Shropshire meres and mosses: a background for ecologists. Fld Stud., 1: 1-38.

WILuis, M. J. & Youna, S. W. (1983). Cheshire College of Agriculture wildlife and management review 1976-
1981. Chester.

SPELLERBERG, L. F. (1981). Ecological evaluation for conservation. London.

(Accepted May 1986)

Watsonia, 16, 427-437 (1987) 427

Short Notes

ADDITIONAL RECORDS OF HALIMIONE PORTULACOIDES (L.) AELLEN ON
COASTAL ROCKS AND CLIFFS

Halimione portulacoides (L.) Aellen, Sea Purslane, is characteristically a species of salt-marshes.
Since we drew attention to its occurrence on coastal rocks and cliffs in the British Isles (Akeroyd &
Preston 1984), we have received several new records of plants in this habitat. These are
summarized below. The records from Devon and Somerset are the first reports of Halimione on
cliffs in these counties.

SOUTH DEVON, V.C.3

D. E. Bolton found Halimione, growing with Inula crithmoides and Limonium binervosum, in
crevices in Lower Devonian schist rocks at Gammon Head in September 1982. On 13th June 1985,
C.D.P. visited this site and found Halimione growing on soil and rock in four shallow gullies on the
steep, eastern side of Gammon Head, GR 20/765.356 to 20/766.357. The plants grow 10-30 m
above sea-level and are almost inaccessible. The following species were observed growing with
Halimione: Armeria maritima, Beta vulgaris subsp. maritima, Crithmum maritimum, Daucus
carota subsp. gummifer, Festuca rubra, Heracleum sphondylium, Inula crithmoides, Limonium
binervosum, Plantago coronopus, P. maritima, Sonchus oleraceus and Spergularia rupicola. With
the exception of Heracleum, all these species have been recorded as associates of Halimione on
cliffs elsewhere in the British Isles or in Brittany (Akeroyd & Preston 1984). Heracleum grew with
Halimione where the latter encroached upon a Festuca rubra sward. On the basis of the map in the
Atlas of the Devon flora (Ivimey-Cook 1984), the Gammon Head colony of Halimione is at least
12-5 km from any other Halimione plants.

NORTH DEVON, V.C.4

On 22nd June 1986, J.R.A. found Halimione growing on Upper Devonian slates at Croyde Bay,
GR 21/434.388 (RNG). A patch measuring 1 x 0-5 m occurs on a rock on the upper shore on the
south side of the bay. No other plants occurred in the immediate vicinity but nearby rocks carried
an open plant community which included Armeria maritima, Beta vulgaris subsp. maritima,
Crithmum maritimum, Desmazeria marina, Festuca rubra, Plantago coronopus and Matricaria
maritima.

SOUTH SOMERSET, V.C.5

J. H. Crothers has observed a stand of Halimione on cliffs on the eastern side of Hurlstone Point,
GR 21/899.493, over the period 1967-84. Mr Crothers (in litt. 1984) says that the colony “‘has
changed little in extent during the last 17 years, and appeared well-established when I first saw it in
1967”’.

ANGLESEY, V.C.52

On 21st July 1985, R. H. Roberts found Halimione growing on a low, Carboniferous limestone cliff
between Moelfre and Traeth Lligwy. C.D.P. visited this site on 16th June 1986. Two plants of
Halimione grow near the seaward edge of a flat, and rather bare, platform of limestone rock east of
Porth Forllwyd, GR 23/506.871. They are rooted in small depressions in the rock in which soil has
accumulated, and grow with Armeria maritima, Festuca rubra subsp. pruinosa and Puccinellia
maritima.

KIRKCUDBRIGHTSHIRE, V.C.73
Halimione was recorded on cliffs at Balcary Point in 1959 and 1977 (Akeroyd & Preston 1984). On

428 SHORT NOTES

6th July 1986, O. M. Stewart and C. D. P. found it in a gulley at this site, GR 25/828.491. A single
large plant, measuring 1 x 0-3 m, grew with Armeria maritima on a rock ledge. Aster tripolium,
Plantago maritima, Puccinellia maritima and Spergularia rupicola occurred on nearby rocks.

WIGTOWNSHIRE, V.C.74

Halimione was known on the cliffs of the Mull of Galloway in the 19th century, and was refound
there by H. A. Lang in 1980 (Akeroyd & Preston 1984). Dr Lang reported a small patch at GR 25/
154.303 that was withering and the survival of which he doubted. However, on a return visit to the
site on 19th September 1984, he found that this plant had survived, and he also discovered a further
four plants, two near the original plant and two some 50 m away (H. A. Lang in litt. 1984).

The fcur new records of Halimione on cliffs in western Britain fit the pattern of distribution that we
reported in the original paper. We had, in fact, expressed our surprise at the absence of records
from Devon. We would be interested to hear of other records from cliffs, and should particularly
like to encourage Irish botanists to look for further sites.

Halimione has recently been found on salted roadsides, one of several halophyte species which
have invaded this habitat in recent years (Kitchener 1983; Scott 1985). Kitchener (1983) found ‘“‘a
large bush spreading vegetatively on the centre reservation of the A2 near Dartford Heath
[Kent]’’. This observation also suggests that the ecological tolerance of Halimione might be
somewhat greater than is generally supposed.

ACKNOWLEDGMENTS

We are very grateful to D. E. Bolton, J. H. Crothers, H. A. Lang and R. H. Roberts for sending us
information about Halimione on cliffs.

REFERENCES

AKEROYD, J. R. & Preston, C. D. (1984). Halimione portulacoides (L.) Aellen on coastal rocks and cliffs.
Watsonia, 15: 95-103.

IvimEY-Cook, R. B. (1984). Atlas of the Devon flora, p. 28. Exeter.

KITCHENER, G. D. (1983). Maritime plants of inland roads of West Kent. Trans. Kent Fld Club, 9: 87-94.

Scott, N. E. (1985). The updated distribution of maritime species on British roadsides. Watsonia, 15: 381-386.

J. R. AKERoypD and C. D. PRESTON
Department of Botany, The University, Whiteknights, Reading RG6 2AS

BRITISH MATERIAL IN THE UNITED STATES NATIONAL HERBARIUM

The extent to which overseas herbaria contain British and Irish specimens has up to now been
known over here only at second hand, the information on these in Kent & Allen (1984) having
necessarily been taken almost wholly from the successive editions of the Index Herbariorum.

A recent stay in Washington based on the Smithsonian Institution Archives gave me the
opportunity to investigate in some depth the collections of the Department of Botany of the
National Museum of Natural History (US), one of the components of the Smithsonian complex.
These collections have been formally known since 1894 as the U.S. National Herbarium, following
the incorporation of the herbarium of the U.S. Department of Agriculture in that of the Museum
(Morton & Stern 1966).

Until the acquisition in 1972 of the herbarium of the late S. T. Jermyn, which extends to some
10,000 specimens, mainly from Essex, v.cc.18 and 19 — and of which only a small portion has as yet
been incorporated into the general collection — the flora of Britain was surprisingly scantily
represented in US, it turns out, certainly compared with the flora of many other countries in
Europe. Of Irish material I came across only two sheets in my sampling.

SHORT NOTES 429

Apart from Jermyn, only one of the British collectors listed in Kent & Allen on the strength of
the Index Herbariorum data proved identifiable in either the Herbarium itself or the Museum’s
donor files. This does not necessarily mean that none of their specimens are present, for the donor
files tend to be confined to ‘collections’, so that very small amounts of material, especially if
incorporated in some larger donation, are likely to have escaped being registered. Nevertheless, it
is worth placing on record that the many sheets bearing the labels of G. C. Joad (donated,
according to the files, by Kew in 1882) encountered in my sample runs were of Continental
specimens exclusively. There was no sign of any John Stuart Mill material, British or otherwise,
nor of anything in the names of A. B. Lambert or W. West. And the donor files identified the ‘J. F.
Rayner’ as a US. collector, not the British botanist of that name.

The exception was J. G. Baker. A contemporary internal memorandum, now in the Smithsonian
Archives, reveals that on the day of his retirement from Kew in 1899, Baker donated a part of his
personal herbarium, totalling “not less than 5000 specimens.” Most of these were lichens and
bryophytes. The vascular plants were far fewer, and the fact that these consisted in the main of
English Rubus microspecies, many of them not even named, suggests that this was unmounted
material which had long lain in his room or other working space in the Kew Herbarium and which
he had finally had to clear out. Kew at that period was following a policy of turning away (or
transferring to the British Museum) British Isles material, so presumably in desperation, not
wishing — or, perhaps, unable — to take the specimens to his home (for the donor files show the
parcel came direct from the Kew Herbarium), he included them in the offer to America. A more
inappropriate donation it would be hard to imagine, and doubtless it was only Baker’s reputation
and being told that it included an eighteenth-century set in the shape of James Dickson’s ‘Hortus
Siccus Britannicus’ that caused the Smithsonian to accept it with enthusiasm. In fact, hardly any of
the specimens are of Baker’s own gathering. Most, about 100 sheets, consist of Rubi collected
round London in 1883—91 by Eyre de Crespigny (out of respect for whose batological acumen W.
C. R. Watson was later to name R. crespignyanus in his honour). There is also a small collection of
this genus made round Bournemouth, v.cc. 9 and 11, by H. Fisher in 1885, together with the
remains respectively of one of Leighton’s widely-owned sets from Shropshire, v.c.40, and of one of
Bloxam’s from Twycross, v.c.55. Several specimens from J. T. Powell (v.c. 18) and one or two
from J. E. Bagnall (v.c. 38) and W. H. Painter (v.c. 57) complete the impression of a typical
accumulation of a late-Victorian Rubus specialist.

Apart from this collection of Baker’s, the material is very miscellaneous. The provenance of a
number of specimens, mostly collected in v.cc. 65 and 66 in 1870—72 by someone identified only by
semi-indecipherable initials (“B.M. or U.M, Jr.”’), is given as the herbarium of William H.
Seaman, donated in 1915. Several others collected by the first President of the Alpine Club, John
Ball, in mountain areas (1838-59), probably came with the much more extensive material of his
that is present from the Alps and elsewhere on the Continent, as his British herbarium is allegedly
divided between Edinburgh and Kew. There are also a number of sheets bearing Arthur Bennett's
labels and dating from the 1880s, which may have been purchased from him direct, and several of
an anonymous collector in the Bristol area, v.cc. 6 and 34, in 1878, while the pre-1894 Department
of Agriculture herbarium is the indicated source of quite a number of others. These last include
many with the distinctive blue labels, worded in Latin, of the Liverpool collector A. E. Lomax, a
few of J. T. Moggridge’s (including specimens contributed by his father, Matthew Moggridge)
which must have come with his much more numerous French Riviera material, and certainly one
and probably more acquired as duplicates from the Botanical Society of Edinburgh. Of other
collectors no more than the odd single sheet featured in my samples.

REFERENCES

Kent, D. H. & ALLEN, D. E. (1984). British and Irish herbaria. London.
Morton, C. V. & STERN, W. L. (1966). The United States National Herbarium. Plant Sci. Bull., 12: 1-4.

D. E. ALLEN
Lesney Cottage, Middle Road, Winchester, Hants., SO22 5EJ

430 SHORT NOTES
OBSERVATIONS ON STACHYS GERMANICA L. AT A NEW SITE IN OXFORDSHIRE

On 28th June 1984, I discovered several young plants of Stachys germanica L. growing close to a
stand of Carduus nutans in front of a hedge bordering an Oxfordshire lane. During a careful search
of the area over the next two weeks more plants were found — on bare ground and among grasses
and herbs — and by 18th July, 56 flowering stems were recorded, the tallest measuring c. 91 cm.
The discovery of this scheduled species, which is now confined to the oolitic limestone of
Oxfordshire, was reported to the Nature Conservancy Council and the Biological Records Centre.
The site was later declared a new one and is referred to here as Site X.

S. germanica was widely recorded by Druce (1886, 1927) but, since that time, its known sites
have dwindled to less than half-a-dozen. Acknowledged to be a capricious plant, the reason for its
unexpected appearance at Site X was at first difficult to explain. Later, from enquiries made about
the lane and its environs, it was learned that the hedge in front of which the plants were growing
had been cut back severely in the late autumn of 1982, the trimmings being burnt in situ; prior to
that date it had not been cut for 35-40 years, and possibly not since the late 1930s. In the
intervening decades it had grown very thick and wide, spreading across the lane as scrub, with
young Acer campestre, Fraxinus excelsior and Quercus robur. S. germanica being a biennial, it was
assumed that long-dormant seeds had germinated in 1983 in ground disturbed by the hedge-cutting
operations, and that over-wintering rosettes had preceded the flowering stems found in 1984. It
was seen that the plants were more or less in a row — in large or small groups, or singly — over a
distance of about 180 m, along what appeared to be the base-line of the original hedge.

In a survey of the present hedge, 17 species of shrubs and trees were recorded, including Cornus
sanguinea, Euonymus europaeus, Ilex aquifolium, Ligustrum vulgare, Rhamnus catharticus and
Salix caprea. Since the cutting-back in 1982, many Acer campestre, Cornus sanguinea, Corylus
avellana, Crataegus monogyna, Prunus spinosa and Viburnum lantana have been reduced to leafy
stools or bushes; and the presence among them of three plants of Sorbus torminalis — together with
an associated flora which includes Anemone nemorosa, Allium ursinum, Hyacinthoides non-
scripta, Lamiastrum galeobdolon, Melica uniflora, Mercurialis perennis, Poa nemoralis and Viola
riviniana — tends to support the view that the hedge is of woodland origin. The lane, in places very
wide, is believed to be an ancient trackway, and its verges in the vicinity of Site X contain many
grassland/pasture calcicoles, including Brachypodium pinnatum, Bromus erectus, Carex flacca,
Cirsium acaule, C. eriophorum, Helianthemum chamaecistus, Origanum vulgare, Pimpinella
saxifraga, Reseda lutea, Trisetum flavescens and Viola hirta; so far, 150 species have been recorded.

The site was visited regularly, and in the late summer of 1984, 20 new rosettes were found, the
majority close to the original plants. All over-wintered successfully. In 1985, each of these rosettes
produced between one and five flowering stems, of which there were 53 in total; as in 1984, most
were branched, the tallest stem measuring c. 95 cm. That autumn, more than 100 new plants, some
very small, were counted. Unfortunately, 50% were subsequently lost, possibly as a result of the
exceptionally hard winter (some of the smallest were observed to have perished from frost-lift),
followed by a cold, wet spring. During 1986, each of the remaining rosettes developed only one
flowering stem. Of the total of 56, only three were branched, and all were markedly smaller (some
only c. 10-15 cm) and less vigorous than those of the previous two summers. Flowering
commenced about the third week of July and in some plants continued into October.

Of particular interest in 1984 was an exceptionally robust plant which, late that summer,
developed a number of shoots at the base of its stem; seven of these grew during the following year
into fine flowering stems, one measuring 104 cm. Equally intriguing, in 1985, were two medium-
sized rosettes, growing close together in a more shaded situation, whose development was
completely arrested. These were marked for further observation, and it is hoped that they will
prove to be triennial by flowering in 1987.

Pollination has been mainly by large and small bumble bees, and work on the identification of
these is continuing. In August 1985, conspicuous numbers of the Hover-fly Episyrphus balteatus
visited the flowers. Seed production was good in 1984 and 1985, but poor in 1986. (In 1984, one
unbranched stem yielded c. 525 seeds; in 1986 five similar stems yielded a total of only c. 600). Ripe
seeds were found in the dried-up calyces of dead plants — most of which remained standing
throughout the winter — for up to six months. Germination appears to be continuous throughout
the growing season, which could account for the considerable difference in the size of the autumn

SHORT NOTES 431

rosettes (c. 2-0—7-5 cm in diameter). Plants which had formed the most and largest leaves by that
time had the highest survival rate.

In 1985 and 1986, several leaves and a few flower buds were damaged by caterpillars of the
leaf-rolling Tortrix Moth, Cnephasia interjectana, and the Common Frog-hopper, Philaenus
spumarius.

In a search through botanical literature for information about S. germanica it was noted that
Keble Martin (1965) described the plant as stoloniferous. No reference to this being made by
Warburg (1962), it was decided, each year, to examine the root-system of several autumn plants,
care being taken not to damage them. None was found to have stolons or rhizomes. All, in fact,
_ had surprisingly few, shallow roots for a plant which can attain a height of 76 cm or more, and
where old and new plants grew together, each was found to be independent of the other.

Management of the site has been made easier by the sympathetic interest and willing co-
operation of the owners of land adjoining the lane, and their staff, particularly at times of
hedge- and verge-cutting. Personal activities have been confined to the cutting back of vegetation
surrounding the plants (Clematis vitalba, for instance, deprived of hedgerow support, has taken
to the ground and is a menace); the maintenance of bare soil around them, the hand-weeding of
the ‘nurseries’ of young rosettes, and the protection of these from damage by rabbits and the
occasional dust-bathing pheasant. Where it has not been possible to do this, in areas where single
or small groups of flowering stems appeared in 1984 and/or 1985, there were no plants in 1986.

There would seem to be little doubt that a principal requirement for Stachys germanica is open
ground for the germination of its seeds and the formation of winter rosettes, and that its
disappearance from two recent sites has resulted from the invasion of such open areas by grasses,
mosses, coarse vegetation and scrub. If it can be assumed, from the history of Site X, that seeds
remain viable for a very long time, then the disturbance or opening-up of old sites, where known
— and where this is possible — is to be recommended.

Soon after this Note was written (November 1986), most of the 1986 seed-bearing stems of
Stachys germanica were stolen from Site X. In view of this incident and the need for stricter
protection measures, members are asked not to make requests to visit the site. It would be a
matter for profound regret if this very rare and fascinating species were ever to disappear from
the ranks of our native flora.

ACKNOWLEDGMENTS

I gratefully acknowledge the grants awarded to me by the British Ecological Society and the
Nature Conservancy Council for the protection and study of Stachys germanica at Site X. I also
wish to thank members of staff of the N.C.C. for encouragement and advice; Mr W. J. R. P.
Bishop, Mrs D. Bowyer and Mrs C. McNab for practical assistance at the site; Mr J. Campbell
(Biological Records Centre) for help with the identification of insects; and Mr J. Trinder for the
donation of several photographs.

REFERENCES

Druce, G. C. (1886). The flora of Oxfordshire, pp. 231-232. Oxford & London.

Druce, G. C. (1927). The flora of Oxfordshire, 2nd ed., pp. 342-343. Oxford.

KEBLE Martin, W. (1965). The concise British Flora in colour. London.

Waresurg, E. F. (1962). Stachys L., in CLAPHAM, A. R., Tutin, T. G. & Warpure, E. F. Flora of the British
Isles, pp. 748-750. Cambridge.

A. J. DUNN
Flat 2, Sandford Mount, Charlbury, Oxford, OX7 3TL

432 SHORT NOTES
PELORIC AND DUPLEX EXAMPLES OF ORCHIS PURPUREA HUDSON IN KENT

Peloric forms of Orchis purpurea Hudson have been found quite often in Kent (Godfery 1933;
Summerhayes 1968); the most frequent site appears to be a Kent Trust for Nature Conservation
reserve near Stelling Minnis. On 30th May 1986, I found five such plants there; four were within a
ten-metre diameter and had very similar labellum patterns, while the fifth was about 300 metres
away and had a different labellum pattern. Many normal specimens of O. purpurea were in the
vicinity of these plants.

Since I had not requested permission to take specimens for further study, I contented myself
with taking photographs. They show the following points.

1) All the specimens had additional labella in place of normal petals, i.e. they exhibited Type A
peloria sensu Bateman (1985). 2) The extra labella were smaller than the primary ones and were of
irregular shape, unlike a peloric Ophrys insectifera L. that I found in Hampshire in 1984. 3) In
three cases the primary labella were also of irregular shape. 4) The flowers were all densely packed
and were not, or only partially, resupinate. 5) The columns were malformed in all the specimens,
though rather less so in the solitary one. 6) In one specimen in the group of four, each of the three
labella had an additional small labellum in tandem (i.e. mounted immediately above its partner)
giving a total of six labella per flower. Such a ‘gardener’s double’ must be rare among European
orchids, though I have seen an Ophrys speculum Link in Mallorca that had an additional labellum
tandem-mounted on an otherwise normal flower, and an extraordinary Ophrys reinholdii Spruner
in Rhodes that had three extra labella so mounted.

Seed-set is reported to be extremely low (Summerhayes 1968), so the malformation of the
columns on the peloric flowers would seem to matter little; there does not seem to be any other
survival disadvantage since the propinquity of the group of four indicates successful vegetative
reproduction over a period of several years. Another point is that since peloric flowers are rare,
and duplex ones even more so, the chances of the two conditions occurring in the same plant
would, if they were under separate control, be incredibly small; some connection between their
causative mechanisms seems probable.

ACKNOWLEDGMENT

I am grateful to P. F. Hunt for help in the presentation of this Note.

REFERENCES

BATEMAN, R. M. (1985). Peloria and pseudopeloria in British orchids. Watsonia, 15: 357.
Goprerry, M. J. (1933). Monograph and iconograph of the native British Orchidaceae. Cambridge.
SUMMERHAYES, V. S. (1968). Wild orchids of Britain, 2nd ed. London.

D. M. T. ETTLINGER
Royden Cottage, Cliftonville, Dorking, Surrey, RH4 2JF

TERRESTRIAL AND AQUATIC VARIANTS OF LOBELIA DORTMANNA L.

Although Lobelia dortmanna L. is most commonly found as an aquatic plant, Gluck (1924)
described a terrestrial variant, forma terrestris Gluck, which occurs occasionally along the shores of
some lakes. Although the main distinguishing feature between the two variants is that of location,
Gluck also provided some measurements of leaf lengths, showing that the aquatic plant is generally
larger. The data however do show considerable overlap and my observations of populations
throughout the Scottish Highlands and Islands have suggested a complication. It seems to me that
there are at least two different variants of the aquatic plant, and that one of these is hardly
distinguishable from f. terrestris. L. dortmanna grows over a wide range of sediment types from
coarse gravel to highly organic silt and peat. Plants found on hard substrates and in wave-exposed

SHORT NOTES 433

conditions are similar to terrestrial forms. Indeed no difference can be found in lacunal size,
cortical cell size or cuticle thickness (Farmer 1985). However, plants from silty areas are much
larger and are typical of the aquatic variant described in Gluck (1924).

Studies were carried out at Lochan-na-Thuill (GR29/252.503), Sutherland, v.c. 108. Three
populations of the plant were chosen, one where net sedimentation was occurring and two in
eroded areas. In one of the latter, perspex tubes (15 cm high, 10 cm diameter) were set in place
around plants for 18 months. These acted as silt traps and so the effects of sedimentation could be
compared directly on neighbouring plants. At harvesting, plants were taken from each site. Leaf
length and length/breadth ratios were obtained. Chlorophyll content was also determined after
Hiscox & Israelstrom (1979). Samples of sediment from each of the three sites were dried and
ignited in a furnace to determine the organic content. Due to the colloidal nature of the sediment
in the silt traps, it was not possible to collect this for similar analysis.

The results are presented in Table 1 and show that plants from areas of erosion are smaller than
plants from the area of deposition. Furthermore, in eroded areas where silt traps had been erected,
after 18 months, 5 cm depth of sediment had accumulated. The plants here were seen to have
responded by increasing their leaf lengths. It should be noted that only the tips of the leaves were
protruding above the silt. This suggests that L. dortmanna may not survive sedimentation at a
much greater rate and that this may be an important factor in determining its distribution around
the shores of a lake. Sedimentation seems to have a similar effect to decreasing light levels. Aberg
(1943) grew L. dortmanna in a uniform substrate at different depths in a lake and found that it
responded to shading by extending its leaf length. It is possible that the morphological changes
noted here are the result of a shading effect of the sediment acting on the stem apex. The resulting
extension of internodes (increasing corm length) is also a common shade response. Total
chlorophyll content also decreases with sedimentation (Table 1). This explains the observation of
Gluck (1924) that terrestrial plants are greener than aquatic ones.

TABLE 1. DIMENSIONS AND CHLOROPHYLL CONTENT OF PLANTS FROM SITES IN
LOCHAN-NA-THUILL
All figures are means+S.E.; n=sample size.

%Sediment _n for n for Total leaf

organic plant Corm_ chlorophyll chlorophyll
content dimen- Leaf length Leaf length/ length dimen- content mg/g
Site n=3 sions (mm) breadth ratio (mm) sions _ fresh weight
Depositional
area 65:17+1-03 29 54:7442-15 19-5341-35 41-6+8-0 5 0-498+0-065
Eroded
area 1 1-3 +0-14 40 24:14+0-88 11-91+0-48 7:3+£0-3 5 0-631+0-119
Eroded
area 2 0-98+0-07 29 28 S027 gle 32 0:49 6:5+0-3 5 0-616+0-078
Silt traps
in eroded
area 2 — 19 599-082-311... °23-050:72 °45:847-1 5 0-320+0-027

It is, therefore, possible to distinguish two aquatic plastodemes of L. dortmanna: a small plant
from eroded areas and a larger plant from areas of deposition. The former seems to be
indistinguishable from f. terrestris, suggesting that this taxon too may be a response by the species
to terrestrial conditions, where there is also a lack of sedimentation. Further study of this is
necessary as I have evidence that at least one population of f. terrestris was formerly aquatic.

This terrestrial population of L. dortmanna was found on the shore of Loch Freuchie, Tayside,
v.c. 88. The plants were evidently not growing as emergents owing to a short-term draw-down,
because they were found amongst a range of common terrestrial species, including Agrostis tenuis
Sibth., Rumex acetosa L., Taraxacum sp., Plantago major L., Rubus fruticosus L. and Sorbus
aucuparia L. The population was also associated with two species that are commonly amphibious,
Littorella uniflora (L.) Aschers. and Juncus articulatus L. The adjacent submersed vegetation was

434 SHORT NOTES

Percentage of Plants in Each Size Class

Size class (Leaf Number)

Figure 1. The population structure of a terrestrial population of Lobelia dortmanna. For each size class is
shown the percentage of non-flowering and flowering (hatched) plants.

very sparse (due to considerable wave action), consisting of a few plants of L. dortmanna and
Littorella uniflora.

The site was visited a number of times over three years (1982-5). It was noted that during
periods of heavy rainfall the site would become flooded, though this was only temporary. The
population structure of the terrestrial population of L. dortmanna was studied. 25 cm square
quadrats were randomly distributed over the population and plant size (leaf number) and the
presence of flowers noted. Plant density was estimated for plants growing amongst other
vegetation and in open areas.

Plants were more commonly found amongst other vegetation (36-48 plants/m’) than in open
areas (11-52 plants/m7), suggesting that L. dortmanna is not necessarily excluded from shoreline
habitats by competition with terrestrial species, but rather that these species may protect the
terrestrial L. dortmanna from exposure. Periodic flooding may also reduce the productivity of
associated species and so prevent the complete exclusion of L. dortmanna from the community.
The population structure (Fig. 1) was similar to that of many aquatic populations (Farmer 1985),
i.e. there is a skewed distribution that ts extended into the larger size classes. However, this
population lacked some of the smaller size classes, indicating a reduction in recruitment. The
plants produce seed, which can survive extreme artificial drying (Farmer 1985), so the terrestrial
conditions may be acting so as to prevent seedling establishment rather than germination.

The paucity of young plants suggests that this terrestrial population of L. dortmanna may
represent a formerly aquatic population which has become marooned due to a long-term lowering
of the water-level. White (1898) noted similar plants along the shores of other Perthshire lochs in
places which he considered to be above the usual high-water mark. Such populations may,
therefore, occur in more localities throughout Scotland.

SHORT NOTES 435
ACKNOWLEDGMENTS

I thank the late Prof. David Spence for helpful discussions. This work was funded by the N.E.R.C.

REFERENCES

ABERG, B. (1943). Physiologische und okologische Studien tiber die pflanzliche Photomorphose. Symb. bot.
Upsal., 8: 1-189.

Farmer, A. M. (1985). Lobelia dortmanna: light, growth strategy and zonation. Ph.D. thesis, University of St
Andrews.

Giuck, H. (1924). Biologische und morphologische Untersuchungen tiber Wasser- und Sumpfgewachse. IV,
Untergetauchte und Schwimblattflora. Jena.

Hiscox, J. D. & IsRaELsTRom, G. F. (1979). A method for the extraction of chlorophyll from leaf tissue without
maceration. Can. J. Bot., 57: 1332-1334.

WHitE, F. B. (1898). Flora of Perthshire. Edinburgh.

A. M. FARMER
Institute of Environmental Studies, University of Wisconsin, 319 Birge Hall, 430 Lincoln Drive,
Madison, Wisconsin 53707, U.S.A.

REAPPEARANCE OF ORCHIS PURPUREA HUDSON IN OXFORDSHIRE

On 22nd May 1961, I was fortunate enough to discover one flowering plant of Orchis purpurea
Hudson amongst Mercurialis perennis L. under a canopy of beech in southern Oxfordshire whilst
accompanied by my brother. Details of the exact location are with the local Naturalists’ Trust.
Unfortunately the flowering stem was bitten off that year and we had to wait twelve months for
confirmation. On 24th May 1962, the plant had two flowers fully open, one of which was removed
by Professor Warburg and is now housed in OXF. By 8th June, the plant was in full bloom, by 7th
July eight ovaries were well swollen and by 23rd September these ovaries were ripening
satisfactorily. The occurrence of this plant was the subject of a small exhibit at a B.S.B._I.
Exhibition Meeting held in November 1962 in London (Paul & Warburg 1963).

Due to academic studies I was unable to check the site for five years, but apparently the plant did
not flower in 1963 and flowered poorly in 1964 before disappearing. From 1967 to 1985 I checked
the site with no success. The beech and nearby yew trees were felled in the late 1970s and many
open-loving plants such as Primula veris L. took over. I am pleased to record that last year, on 5th
June 1986, I rediscovered O. purpurea in the same vicinity, seven plants in an area of about six
square metres; three plants had good, flowering spikes. Associated plants were Clematis vitalba L.
(dominant), Chamaenerion angustifolium (L.) Scop., Verbena officinalis L., Senecio jacobaea L..,
Prunella vulgaris L. and Mycelis muralis (L.) Dum. Nearby was a Buddleia bush providing some
degree of shade and several plants of Linaria repens (L.) Mill., Clinopodium vulgare L., Origanum
vulgare L., Campanula glomerata L., Anacamptis pyramidalis (L.) Rich., Odontites verna (Bell.)
Dum. and planted Fagus sylvatica L. saplings.

The two tallest flowering spikes were 40 cm and 43 cm tall respectively, with the ovaries 20 mm
long at the base of the inflorescence and 12 mm long at the top. All three flowering heads had set
fruit by 31st July 1986, with respectively six, two and one swollen, ripe ovaries. These were all
nearest the top of the spike; the ovaries at the base, belonging to the larger and first-formed
flowers, did not swell.

Perhaps the significance of this reappearance after more than twenty years absence is that it
suggests this Oxfordshire site is an ancient semi-woodland habitat rather than one recently formed.
The felling of trees and soil disturbance clearly stimulated plants to reappear. The alternative view,
that the 1961 plant was the chance result of wind dispersal from Kent, now seems less likely. It is
hoped that with the co-operation of the owner the site will be suitably managed, particularly with
regard to Clematis vitalba which threatens to stifle all low-growing plants in the immediate area.

436 SHORT NOTES
REFERENCE

Pau, V. N. & WARBURG, E. F. (1963). Orchis purpurea in Oxon. Proc. bot. Soc. Br. Isl., 5: 172.

R. J. Kemp
Pipistrelles, Chapel Road, Ford, Nr. Aylesbury, Bucks., HP17 8XG

A BINOMIAL FOR THE HYBRID POTAMOGETON GRAMINEUS L.
x P. POLYGONIFOLIUS POURRET

Binomial names are available for 24 of the 25 Potamogeton hybrids recognized from the British
Isles by Dandy (1975). The exception is the rare hybrid between P. gramineus L. and P.
polygonifolius Pourret. The reason for the absence of an acceptable binomial for this plant lies in
the uncertainty about the identity of Potamogeton seemenii Ascherson & Graebner, Syn. mitteleur.
FI., 1: 335 (1897). This was based on material collected on the German island of Borkum by Otto
von Seemen. Ascherson & Graebner regarded the plant they described as the hybrid between P.
gramineus and P. polygonifolius, but this attribution was questioned by Hagstrém (1916:232).
After describing plants which he regarded as the true hybrid, which were intermediate between the
parents in morphology and in stem anatomy and possessed sterile pollen, Hagstr6m commented:
“The plants described above do not coincide with a plant from Borkum preserved in the Berlin
Museum, gathered by O. v. SEEMEN and labelled ‘P. polygonifolius x gramineus. P. Seemenii.’ ‘P.
GRAEBNER Cet.’ ‘Herb. P. ASCHERSON.’ The hybrid nature of this plant is not quite certain, as all its
pollen is fresh and fertile. At any rate it extremely approaches to P. gramineus: Leaf-margin
serrulate (ca. 10 teeth acm), apex acute, branch-leaves — like the lower stem-leaves — sessile, style
and anatomy of the stem like those of P. gramin. On the other hand the upper narrow leaf-bases
aud the narrow peduncle are really a little strange to P. gramineus; but the specimen is also a late
flowering autumnal shoot (gathered 23. VIII, with fresh pollen!), from which no certain
conclusions can be drawn. Be this as it may, the name P. Seemenii must stand for the hybrid P.
gram. X polygonif. (polygonif. X gram., ex GRAEBN.)”’.

In stating that the name P. seemenii must stand, Hagstrom was giving preference to the hybrid
formula stated by Ascherson & Graebner rather than to the actual identity of the plant they
described. By modern practice, laid down by the International Code of Botanical Nomenclature
(art. H.10), the name must be attached to the plant not to the formula. Unfortunately, there is now
no material of P. seemenii at B (M. Hakki in litt. 1986). The specimen seen by Hagstr6m was
probably destroyed, along with so many others, when the Berlin herbarium was bombed in 1943.
There are no other collections cited in the protologue, or illustrations, which might serve as a
lectotype. It is not possible to select a neotype which could be said with any degree of confidence to
represent Ascherson and Graebner’s plant. It does not appear possible, therefore, to reassess the
identity of P. seemenii. Clearly, however, in view of Hagstr6m’s comments the name cannot be
used for the hybrid P. gramineus X P. polygonifolius. It may not be irrelevant to observe that some
plants reported as this hybrid from Scotland and Ireland have proved to be P. gramineus (Dandy
1975).

No such doubt surrounds the identity of Potamogeton gramineus f. lanceolatifolius Tiselius, Pot.
Suec. Exsicc., 3: 6 (1897). This was based on Swedish specimens collected “in provincia Smaland,
Alem, Stromsrum in fluvio Alsteran’” by H. Rappe and H. Tiselius on 27th July 1896, and
distributed by G. Tiselius as no. 139 in Potamogetones Suecici Exsiccati. It was regarded by both
Hagstr6m (1916) and Dandy (1975) as the true hybrid between P. gramineus and P. polygonifolius.
I have examined specimens (in BM, CGE, E and MANCH) which were distributed by Tiselius, and
can see no reason to doubt these opinions. In their unpublished manuscript British species of
Potamogeton L., J. E. Dandy & G. Taylor recommend that, if P. seemenii proves to be referable
to P. gramineus, Tiselius’ epithet should be adopted for the hybrid. In view of the uncertainty
about the identity of P. seemenii, which it is not at present possible to resolve, it is appropriate to
make the following combination:

SHORT NOTES 437

Potamogeton xX lanceolatifolius (Tiselius) C. D. Preston, stat. nov.

Basionym: P. gramineus forma lanceolatifolius Tiselius, Pot. Suec. Exsicc. 3: 6, no. 139 (1897)
(‘lanceolatifolia’ ).

P. gramineus proles heterophyllus var. stagnalis subvar. lanceolatifolius (Tiselius) Ascherson &
Graebner in Engler, Pflanzenr. IV, 11 (Heft 31): 88 (1907).

P. seemenii forma lanceolatifolius (Tiselius) Hagstrom, K. svenska VetenskAkad. Handl., nov.
ser., 55 (5): 231 (1916).

Hybrid formula: P. gramineus L. X P. polygonifolius Pourret

ACKNOWLEDGMENTS

I am grateful to A. O. Chater, M. D. Hooper, P. D. Sell and P. F. Yeo for critically reading the
manuscript.

REFERENCES

Danpy, J. E. (1975). Potamogeton L., in Stace, C. A., ed. Hybridization and the flora of the British Isles, pp.

444—459. London.
HacstroMm, J. O. (1916). Critical researches on the Potamogetons. K. svenska VetenskAkad. Handl., n.s.,

55 (5): 1-281.

C. D. PRESTON

Institute of Terrestrial Ecology, Monks Wood Experimental Station, Abbots Ripton, Huntingdon,
Cambs., PE17 2LS

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Watsonia, 16, 439-452 (1987) 439

Plant Records

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

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

The following signs are used:

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

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

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

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

1/3. LYCOPODIUM ANNOTINUM L. 107, E. Sutherland: Ben More Assynt, GR 29/32.20.
Mountain slope. J. K. Butler & M. Murray, 1986. 2nd extant locality.

+2/2. SELAGINELLA KRAUSSIANA (Kunze) A. Braun *27, E. Norfolk: 1 km W. of Felbrigg, GR
63/19.39. Grassland. E. Beaumont & E. T. Daniels, 1985. *39, Staffs.: Patshull churchyard,
GR 33/80.00. Mown grass. C. B. Westall, 1986, herb. B.R. Fowler.

4/1. EQUISETUM HYEMALE L. *103, Mid Ebudes: By Ardow Burn near West Ardow, GR 17/
42.50. Forestry plantation. M. North, 1986, E. 1st localized record.

4/7. EQUISETUM SYLVATICUM L. 2, E. Cornwall: Goscott Wood, GR 20/22.96. Willow carr.
T. Dingle, 1986. 1st record since 1925.

9/1. CRYPTOGRAMMA CRISPA (L.) R. Br. ex Hook. *107, E. Sutherland: Ben More Assynt, GR
29/32.20. Montane scree. J. K. Butler & M. Murray, 1986.

11/1. ADIANTUM CAPILLUS-VENERIS L. +*27, E. Norfolk: Elm Hill, Norwich, GR 63/23.08.
Sheltered wall. K. E. Bull, 1986. *44, Carms.: Below Craig Ddu, GR 22/32.10. Tufa below
limestone cliff. A. Fowles, 1986. 1st native record.

15/4. ASPLENIUM MARINUM L. 107, E. Sutherland: Navidale, GR 39/05.17. Maritime cliff.
J. K. Butler, 1986. 2nd record.

21/2 bor. DRYOPTERIS AFFINIS (Lowe) Fraser-Jenkins subsp. BORRERI (Newman) Fraser-
Jenkins Locality given in Watsonia, 15: 391 (1985) is in v.c. 82, E. Lothian.

21/2 sti. DRYOPTERIS AFFINIS (Lowe) Fraser-Jenkins subsp. sTILLUPENSIS (Sabransky) Fraser-
Jenkins Locality given in Watsonia, 15: 391 (1985) is in v.c. 82, E. Lothian.

21/67. DRYOPTERIS CARTHUSIANA (Vill.) H. P. Fuchs x D. pitatata (Hoffm.) A. Gray *47,
Monts.: Gwaunynog, GR 33/0.1. Betula pubescens-Salix cinerea swamp. P. M. Benoit, 1986,
NMW.

21/9. DRYOPTERIS EXPANSA (C. Presl) Fraser-Jenkins & Jermy 73, Kirkcudbrights.: Between
Meikle Millyea and Milldown, GR 25/51.83. In shelter of boulders. J. Bevan, 1986. 2nd record.

440 PLANT RECORDS

25/int. POLYPODIUM INTERJECTUM Shivas *90, Angus: Den of Airlie, GR 37/29.51. Deciduous
woodland. H. J. Noltie, 1981, DEE, det. A. C. Jermy.

25/int.X 1. POLYPODIUM INTERJECTUM Shivas X P. VULGARE L. *77, Lanarks.: Meikle Dripps,
GR 26/57.55. P. Macpherson, E. L. S. Lindsay & E. Teasdale, 1985, herb. P.M., det. A. McG.
Stirling.

46/15. RANUNCULUS SCELERATUS L. *43, Rads.: Barland Farm, Evenjobb, GR 32/27.62.
Marshy area. J. & C. Port, 1986, NMW.

48/1. MyOSURUS MINIMUS L. 50, Denbs.: Near Worthenbury, GR 33/42.45. Farm gateway.
B. Formstone & J. A. Green, 1986. 2nd record.

+50/aqu. THALICTRUM AQUILEGIFOLIUM L. *77, Lanarks.: Braehead, GR 26/51.67. Estab-
lished in wood by abandoned railway. P. Macpherson & E. Teasdale, 1984, herb. P.M.

57/2. CERATOPHYLLUM SUBMERSUM L. 38, Warks.: Wedgnock, GR 42/26.68. Pond. J. C.
Bowra, 1986, det. P. J. Copson. 2nd record.

58/3. PAPAVER LECOQII Lamotte *81, Berwicks.: Chirnside House Hotel, GR 36/88.56.
Vegetable garden. M. E. Braithwaite, 1986, herb. M.E.B.

58/5. PAPAVER ARGEMONE L. 49, Caerns.: Tai-morfa, GR 23/28.26. Disturbed soil. A. P.
Conolly, 1986, herb. A.P.C. 1st post-1930 record.

62/1. CHELIDONIUM MAJUS L. 93, N. Aberdeen: Inverugie, GR 48/10.48. Garden wall.
D. Welch, 1986, ABD. 1st post-1930 record.

+64/exi. DICENTRA EXIMIA Torrey *90, Angus: Glen Prosen, GR 37/32.65. Woodland. S. &
E. Spreadbury, 1982, DEE, det. U. K. Duncan.

66/2. FUMARIA CAPREOLATA L. 73, Kirkcudbrights.: Maxwelltown, GR 25/97.75. Waste
ground. O. M. Stewart, 1986, E. 2nd record. *107, E. Sutherland: Bonar Bridge, GR 28/60.91.
Roadside. M. J. Marshall, 1985, herb. M.J.M., det. M. G. Daker.

+67/3. BRASSICA RAPA L. subsp. SYLVESTRIS (L.) Janchen *26, W. Suffolk: R. Brett between
Chelsworth and Semer, GR 52/9.4. River-banks. A. L. Bull, 1960.

69/1. RHYNCHOSINAPIS MONENSIS (L.) Dandy +77, Lanarks.: Cambuslang, GR 26/65.60. Old
coal bing. P. Macpherson, 1986, herb. P. M. 2nd record.

69/3. RHYNCHOSINAPIS CHEIRANTHOS (Vill.) Dandy 39, Staffs.: Bilston, GR 32/93.96.
Towpath. B. R. Fowler, 1980. 1st post-1930 record.

+70/2. SINAPIS ALBA L. 93, N. Aberdeen: Eden, GR 38/69.58. Field gateway. D. Welch,
1986, ABD. 1st post-1930 record.

+80/2. CORONOPUS DIDYMUs (L.) Sm. *50, Denbs.: Bettisfield, GR 33/45.35. Garden weed.
J. A. Green, 1986, NMW.

781/1. CARDARIA DRABA (L.) Desv. *73, Kirkcudbrights.: Castle Douglas, GR 25/77.62.
Roadside verge. O. M. Stewart, 1986, E.

84/1. THLASPI ARVENSE L. +103, Mid Ebudes: Tigh an Fhasgaidh, GR 17/40.53. Roadside.
A. A. Slack & J. W. Clark, 1986. 2nd record.

88/5. COCHLEARIA DANICA L. *75, Ayrs.: Horse Island, GR 26/21.42. Sandy soil over rocks.
B. Simpson, 1986.

+90/2. BUNIAS ORIENTALIS L. *27, E. Norfolk: N.W. of Weston Green, GR 63/09.14.
Roadside verges. A. L. Bull, 1986. 77, Lanarks.: Craigton, GR 26/53.64. Waste ground.
P. Macpherson, 1986, herb. P.M. Ist post-1930 record.

94/4. DRABA MURALIS L. +83, Midlothian: Hawthornden Castle, GR 36/28.63. Pathside.
D. R. McKean, 1986. 2nd extant locality.

PLANT RECORDS 44]

97/3. CARDAMINE IMPATIENS L. *80, Roxburghs.: Slitrig Water, Hawick, GR 36/51.06. R. W.
M. Corner, 1986, herb. R.W.M.C. Ist native locality.

+97/rap. CARDAMINE RAPHANIFOLIA Pourret 90, Angus: Dronley Farm, GR 37/34.35. Herb-
rich pasture by stream. L. Bissett, 1983. 2nd record.

+98/2. BARBAREA STRICTA Andrz. *39, Staffs.: Branston, GR 43/22.21. Stream-bank. M. E.
Smith, 1972, herb. T.C.G. Rich, det. T.C.G.R.

+98/3. BARBAREA INTERMEDIA Bor. *39, Staffs.: Brookside Farm, GR 43/11.30. Stream-bank.
B. R. Fowler, 1974. *47, Monts.: East of Aberhosan, GR 22/82.97. A. J. Morton, 1985.

102/3. RORIPPA SYLVESTRIS (L.) Bess. *103, Mid Ebudes: Cornaigbeg, Coll, GR 17/23.63.
B. & K. Cassels, 1986, E, det. A. McG. Stirling.

+102/6. RoRIPPA AUSTRIACA (Crantz) Bess. *6, N. Somerset: Hartcliffe, Bristol, GR 31/58.68.
Waste ground. R. D. Martin, 1984, herb. R.G.B. Roe.

4105/1. ERYSIMUM CHEIRANTHOIDES L. 44, Carms.: Carmarthen, GR 22/39.19. Roadside
verge. J. Bevan, R. Burton & G. Hutchinson, 1986, NMW. 2nd record. *93, N. Aberdeen:
Avochie, GR 38/53.46. Turnip field. D. Welch, 1986, ABD.

109/1. ARABIDOPSIS THALIANA (L.) Heynh. 103, Mid Ebudes: Arinagour, GR 17/22.57.
Garden. J. W. Clark, 1983, E. 1st Coll record.

113/6. VIOLA CANINA L. *81, Berwicks.: St Abbs Head N.N.R., GR 36/91.69. Grassland.
B.S.B.I. meeting, 1986, herb. M.E. Braithwaite. Ist localized record.

113/6X4. VIOLA CANINA L. X V. RIVINIANA Reichb. *80, Roxburghs.: Scaw Knowe, Ancrum,
GR 36/62.25. Grassland. R. W. M. Corner, 1986, herb. R.W.M.C.

122/1. ELATINE HEXANDRA (Lapierre) DC. 72, Dumfriess.: Mill Loch, GR 35/07.83. M. E. R.
Martin, 1986, DFSM, det. C. D. Preston. 1st post-1930 record.

123/13 X14. SILENE DioIca (L.) Clairv. X S. ALBA (Mill.) E. H. L. Krause *47, Monts.: Fan
Pool, GR 22/94.87. Old lead-mine spoil. W. A. Thompson, 1986.

133/1b. STELLARIA NEMORUM L. subsp. GLOCHIDISPERMA Murb. 46, Cards.: Cwm Clettwr,
Tre’r-dd6l, GR 22/66.92. Damp streamside in mixed woodland. I. Weston, W. M. Condry et al.,
1984. 2nd record.

133/3. STELLARIA PALLIDA (Dumort.) Piré 33, E. Gloucs.: Alderton Hill, Dumbleton, GR
42/01.34. Grassland on sandy soil. S. H. Bishop & R. Cooper, 1986. Only extant locality. *83,
Midlothian: Colinton Dell, Edinburgh, GR 36/21.69. Cut grass. O. M. Stewart, 1984, E.

135/1. MOENCHIA ERECTA (L.) Gaertn., Mey. & Scherb. 39, Staffs.: 3 km N.E. of Enville,
GR 32/83.89. Heathy grassland. W. A. Thompson, 1983. 1st post-1930 record.

136/2. SAGINA APETALA Ard. subsp. APETALA 90, Angus: Castlesea Bay, GR 37/67.43. Base-
rich turf. H. J. Noltie, 1981, DEE. 2nd extant locality.

137/4. MINUARTIA HYBRIDA (Vill.) Schischk. +*4, N. Devon: Burlescombe, GR 31/06.16.
Disused railway track. L. J. Margetts, 1986, herb. L.J.M.

141/2. ARENARIA LEPTOCLADOS (Reichb.) Guss. *77, Lanarks.: Between Glasgow and
Renfrew, GR 26/51.67. Bare ground. P. Macpherson & E. Teasdale, 1985, herb. P.M., det. J. G.
Dony.

143/3. SPERGULARIA RUPICOLA Lebel ex Le Jolis *6, N. Somerset: Anchor Head, Weston-
super-Mare, GR 31/30.62. Rocky cliff. R. S. Cropper, 1986, conf. J. A. Ratter. Ist native locality.

+149/3. MonrTIA siBIRICA (L.) Howell 103, Mid Ebudes: Acha, GR 17/18.54. J. W. Clark,
1983, E. 1st Coll record.

+149/par. MonTIA PARVIFOLIA (Mog. ex DC.) Greene *77, Lanarks.: Busby, GR 26/57.55.
Well established on river-bank. P. Macpherson, 1985, herb. P.M., det. E. J. Clement.

442 PLANT RECORDS

154/11. CHENOPODIUM MURALE L. 5, S. Somerset: Curry Rivel, GR 31/41.27. Waste ground.
S. Fisher, 1986, conf. J. M. Mullin. Only extant locality.

154/13. CHENOPODIUM HYBRIDUM L. *35, Mons.: The Nurtons, Tintern Parva, GR 32/53.01.
Cultivated ground. E. Wood, 1985, herb. T. G. Evans, det. E.W. & T.G.E.

154/14. CHENOPODIUM RUBRUM L. 39, Staffs.: Belvide Reservoir, GR 33/86.09. Waste ground
at margin of reservoir. B. R. Fowler, 1968. 1st post-1930 record. 50, Denbs.: Near Wolvesacre
Hall, GR 33/50.42. Edge of arable field. J. A. Green, 1986, NMW. 2nd record.

155/1. BETA VULGARIS L. subsp. MARITIMA (L.) Arcang. 103, Mid Ebudes: Eadar da Bheinn,
GR 17/14.54. Rocky shore. K. A. Cassels, 1985, conf. J. W. Clark. 1st Coll record.

156/5. ATRIPLEX LACINIATA L. 61, S.E. Yorks.: Spurn Point, GR 54/4.1. Shore of R.
Humber. E. Chicken, 1986, herb. E.C. 2nd record, 1st since 1798. Barmston, GR 54/17.58. Sea-
shore. Wild Flower Society Excursion, 1986. 2nd extant locality.

158/1. SUAEDA MARITIMA (L.) Dumort. *47, Monts.: R. Dovey, Dovey Junction, GR 22/6.9.
Muddy edge of tidal river. P. M. Benoit, 1986.

160/2. SALICORNIA FRAGILIS P. W. Ball & Tutin *35, Mons.: Towyn Pill, Caldicot, GR 31/
47.86. Amongst Spartina on estuarine mud. T. G. Evans & D. Upton, 1986, det. F. Rose.

+168/3. GERANIUM ENDRESSII Gay 39, Staffs.: Himley Wood, GR 32/89.90. Roadside verge.
B. R. Fowler, 1986. 1st post-1930 record.

+168/3 x4. GERANIUM ENDRESSII Gay X G. VERSICOLOR L. *73, Kirkcudbrights.: Killymingan,
GR 25/85.67. Roadside bank. O. M. Stewart, 1986.

168/14. GERANIUM PUSILLUM L. 83, Midlothian: Kirkliston, GR 36/12.73. Disused railway
line. O. M. Stewart, 1984, E. 1st post-1930 record.

+170/exi. OXALIS EXILIS A. Cunn. *39, Staffs.: 3 km S. of Tettenhall, GR 32/88.96. Between
paving slabs. C. B. Westall, 1983.

171/1. IMPATIENS NOLI-TANGERE L. +77, Lanarks.: East Kilbride, GR 26/66.55. Bank of R.
Calder. E. L. S. Lindsay & P. Macpherson, 1983. 2nd record.

+171/4. IMPATIENS GLANDULIFERA Royle 107, E. Sutherland: Blarich, GR 29/69.04. Forest
edge. M. J. Marshall, 1985, herb. M.J.M. 2nd record.

4173/2. ACER PLATANOIDES L. *77, Lanarks.: Between Glasgow and Renfrew, GR 26/51.66.
Regenerating in abandoned industrial estate. P. Macpherson & E. Teasdale, 1986, herb. P. M.

4191/1. MELILOTUS ALTISSIMA Thuill. *75, Ayrs.: Troon, GR 26/32.31. Waste ground above
shore. B. Simpson, 1985.

192/11. TRIFOLIUM SCABRUM L. 33, E. Gloucs.: Ring Hill, Haresfield, GR 32/82.08. Short dry
rough grassland. S. H. & G. M. Bishop, 1986. 2nd record.

200/3. ASTRAGALUS GLYCYPHYLLOS L. +*59, S. Lancs.: Burscough, GR 34/44.11. Waste
ground. L. Paull, 1986, LIV.

*203/1. CORONILLA VARIA L. 39, Staffs.: Wordsley, GR 32/88.86. Canal towpath. G. Bennett,
1976. 1st post-1930 record.

206/2. VICIA TETRASPERMA (L.) Schreb. 73, Kirkcudbrights.: Howell Bay, GR 25/69.43.
Grassy slope by shore. O. M. Stewart & C. D. Preston, 1986. 2nd extant locality.

206/9. VicIA oRoBUS DC. 2, E. Cornwall: Scarsick, GR 20/20.88. Disused railway
embankment. T. Dingle, 1986, conf. R. W. David. Ist post-1930 record.

206/12. VICIA LUTEA L. 73, Kirkcudbrights.: Howell Bay, GR 25/69.43. Grassy slope by
shore. O. M. Stewart & C. D. Preston, 1986. 2nd extant locality.

206/16. VICIA LATHYROIDES L. 39, Staffs.: 1 km W. of Kinver, GR 32/83.83. Neutral
grassland. B. R. Fowler, 1976, herb. B.R.F., det. I. C. Trueman. 1st post-1930 record.

PLANT RECORDS 443

207/6. LATHYRUS SYLVESTRIS L. +59, S. Lancs.: Hunts Cross, Liverpool, GR 33/43.86.
Disused railway. V. Gordon, 1986. 2nd record.

+207/gra. LATHYRUS GRANDIFLORUS Sibth. & Sm. *94, Banffs.: Netherdale, GR 38/65.49.
Grass verge. J. Edelsten, 1986, E.

4209/2. SPIRAEA DOUGLASII Hook. *59, S. Lancs.: Hunts Cross, Liverpool, GR 33/43.86.
Disused railway. V. Gordon, 1986.

7209/2 xlat. SPIRAEA DOUGLASIT Hook. xX S. LATIFOLIA (Ait.) Borck. *77, WLanarks.:
Cambuslang, GR 26/63.60. E. L. S. Lindsay & P. Macpherson, 1985, herb. P. M., conf. A. J.
Silverside.

211/11/2. Rusus scissus W. C. R. Wats. *83, Midlothian: Dalmahoy, GR 36/1.6. A. Craig-
Christie, 1900, E, det. A. Newton.

211/11/27. RUBUS TUBERCULATUS Bab. *73, Kirkcudbrights.: Clatteringshaw, GR 25/54.75.
O. M. Stewart, 1986, E, det. A. McG. Stirling.

2211/11/67. RUBUS SUBINERMOIDES Druce *103, Mid Ebudes: Ulva House, Mull, GR 17/4.3.
Old garden. J. W. Clark, 1982, E, det. A. Newton & A. McG. Stirling. Allt Mor, Coll, GR 17/
16.54. Bank of burn. J. W. Clark, 1986, E, det. A. McG. Stirling. 1st Coll record.

211/11/284. RUBUS RUFESCENS Muell. & Lefév. *75, Ayrs.: Kerelaw Burn, Stevenston, GR
26/26.42. Waste ground. B. Simpson, 1986, det. A. McG. Stirling.

211/11/297. RuBUS RADULOIDES (Rogers) Sudre *77, Lanarks.: Near Carluke, GR 26/82.50.
Roadside bank. A. McG. Stirling, 1981.

216/3. GEUM RIVALE L. 17, Surrey: Rowhill Copse, Aldershot, GR 41/85.49. Wet woodland.
C. Hall, 1986. 1st post-1930 record.

220/3/8. ALCHEMILLA XANTHOCHLORA Rothm. 34, W. Gloucs.: Caswell Wood, GR 32/54.00.
Woodland ride. S. H. Bishop, 1986, herb. S.H.B. 1st record since 1895 record from Caswell.

+225/5. ROSA RUGOSA Thunb. 39, Staffs.: Holy Austin Rock W. of Kinver, GR 32/83.83.
B. R. Fowler, 1985. 1st post-1930 record.

225/10. ROSA OBTUSIFOLIA Desv. 26, W. Suffolk: Market Weston Fen, GR 52/98.78. Scrub.
I. M. Vaughan, 1985. 2nd record.

225/cor. ROSA CORIIFOLIA Fr. *26, W. Suffolk: Market Weston Fen, GR 52/98.78. Scrub.
I. M. Vaughan, 1985, IPS.

+226/3. PRUNUS CERASIFERA Ehrh. 33, E. Gloucs.: Painswick, GR 32/87.11. Hedge. S. H. &
G. M. Bishop, 1985. 2nd record. *34, W. Gloucs.: Upleadon, GR 32/75.27. Hedge. S. H.
Bishop & S. C. Holland, 1986. Bromsberrow, GR 32/74.34. Hedge. S. C. Holland, 1986. 1st and
2nd records. 39, Staffs.: Little Gorse Wood, GR 33/90.26. I. J. Hopkins, 1983. 1st post-1930
record.

+227/bul. COTONEASTER BULLATUS Boiss. 73, Kircudbrights.: Maxwelltown, GR 25/97.74.
Wall. O. M. Stewart, 1986, E. 2nd record.

+232/4/1. SORBUS INTERMEDIA (Ehrh.) Pers. *39, Staffs.: 4 km E. of Brewood, GR 33/82.08.
Roadside wood. B. R. Fowler, 1974, herb. B.R.F., det. R. J. Pankhurst. 73, Kirkcudbrights.:
Near Cargenbridge, GR 25/94.75. Disused railway embankment. O. M. Stewart, 1985. 2nd record.

232/6/3. SORBUS DEVONIENSIS E. F. Warb. 2, E. Cornwall: Beardon, GR 20/30.93. Hedge.
R. W. David & L. J. Margetts, 1986, CGE. 1st post-1930 record.

7232/hyb. SORBUS HYBRIDA L. *93, N. Aberdeen: Eden, GR 38/70.60. Semi-natural
woodland in den. D. Welch, 1985, ABD, det. P. J. M. Nethercott.

+235/3. SEDUM SPURIUM Bieb. *103, Mid Ebudes: Lagganulva, GR 17/45.41. Rock cranny.
H. Leitch, 1986, E.

to PLANT RECORDS

237/1. CRASSULA TILLAEA L.-Garland *5, S. Somerset: Dunster Beach, GR 21/99.45. Bare
sandy ground. I. P. Green, 1986, herb. R. G. B. Roe.

+237/hel. CRASSULA HELMSII (T. Kirk) Cockayne *1, W. Cornwall: Goonhilly Downs, GR
10/ 71.21. Heathland pool. M. G. B. Hughes, 1983. *38, Warks.: Eastcote, Knowle, GR 42/
19.78. Pondside. B. Davies, 1986, det. A. Newton. Olton Reservoir, GR 42/13.81. J. C. Bowra,
1986, det. A. Newton. 1st and 2nd records. *39, Staffs.: Tettenhall, GR 33/87.00. Garden
pond. B. R. Fowler, 1968, det. C. C. Townsend.

239/6. SAXIFRAGA HIRSUTA L. +103, Mid Ebudes: Acha, Coll, GR 17/18.54. J. W. Clark,
1983, E, det. A. McG. Stirling. 1st Coll record.

4239/64. SAXIFRAGA HIRSUTA L. X S. UMBROSA L. *103, Mid Ebudes: Aros Park, GR 17/
51.53. Roadside ditch. J. W. Clark, 1986, E, det. A. McG. Stirling.

239/8. SAXIFRAGA TRIDACTYLITES L. 1, W. Cornwall: Between Wadebridge and Padstow, GR
10/95.73. Old slate quarry. L. J. Margetts, 1983. 1st post-1930 record.

254/102. EPILOBIUM PALUSTRE L. X E. PARVIFLORUM Schreb. *61, S.E. Yorksz S.W. of
Withernsea, GR 54/33.28. Marsh. F. E. Crackles, 1986, herb. F.E.C.

254/+6X9. EPILOBIUM CILIATUM Rafin. X E. oBscuURUM Schreb. *77, Lanarks.: Clyde Valley,
GR 26/84.44. Orchard. P.Macpherson, 1979, herb. P.M., det. T. D. Pennington.

262/5. CALLITRICHE HERMAPHRODITICA L. 79, Selkirks.: Clearburn Loch, GR 36/34.15. R. W.
M. Corner, 1986, herb. R.W.M.C. 2nd record.

262/bru. CALLITRICHE BRUTIA Petagna *48, Merioneth: Llyn Tegid, S.W. end, GR 23/8.3.,
and N.E. end, GR 23/9.3. Lake shore. P. M. Benoit, 1986, NMW. 1st and 2nd records.

+265/2. CORNUS SERICEA L. *77, Lanarks.: Strathclyde Park, GR 26/73.56. Waste ground by
R. Clyde. P. Macpherson, 1983, herb. P. M., det. A. C. Leslie.

4266/2. CORNUS ALBA L. cv. ELEGANTISSIMA *77, Lanarks.: Braehead, GR 26/51.67.
Woodland. P. Macpherson & E. Teasdale, 1983, herb. P. M., det. D. McClintock.

4271/1. ASTRANTIA MAJOR L. *94, Banffs.: Netherdale, GR 38/65.49. Roadside verge.
J. Edelsten, 1986, E.

301/1. AETHUSA CYNAPIUM L. *79, Selkirks.: Croft Terrace, Selkirk, GR 36/47.28. A. J.
Smith, 1986, herb. R.W.M. Corner. Ist localized record.

4311/2. HERACLEUM MANTEGAZZIANUM Somm. & Levier 39, Staffs.: Great Pool, Himley
Hall, GR 32/88.91. M. Bigwood, 1968. 1st post-1930 record.

314/1b. Daucus caROoTA L. subsp. GUMMIFER Hook. fil. *44, Carms.: Pendine, GR 22/23.07.
Limestone cliff. I. K. Morgan, 1986, NMW, det. S. B. Evans.

319/2. EUPHORBIA LATHYRIS L. +48, Merioneth: Penhelig, GR 22/6.9. Scrubby bank. S. C.
Holland, 1971, still there in 1986. 2nd record. +90, Angus: Burnside of Duntrune, GR 37/
44.34. Garden weed. W. Petrie, 1984, DEE, det. R. K. Brinklow. 1st post-1930 record.

+320/19. POLYGONUM CUSPIDATUM Sieb. & Zucc. 103, Mid Ebudes: Acha House, Coll, GR
17/ 18.54. Streamside. J. W. Clark, 1985. 1st Coll record.

+320/21. POLYGONUM POLYSTACHYUM Wall. ex Meissner *39, Staffs.: Beaudesert Wood, GR
43/05.13. B. R. Fowler, 1971. .

+320/aub. POLYGONUM AUBERTI Louis Henry *93, N. Aberdeen: Inverugie, GR 48/10.48.
Wall of ruined castle. D. Welch, 1986, ABD.

325/15. RUMEX CONGLOMERATUS Murr. 103, Mid Ebudes: Caoles, Coll, GR 17/12.52. Damp
hollow in cart-track. J. W. Clark & A. Walker, 1986, E, det. A. McG. Stirling. 1st Coll record.

325/15 X 11. RUMEX CONGLOMERATUS Murr. X R. crispus L. *11, S. Hants.: Beaulieu River
near Bailey’s Hard, Beaulieu, GR 41/39.01. Pasture. R. P. Bowman, 1985, herb. R.P.B., det. J. R.
Akeroyd.

PLANT RECORDS 445

325/11X14. RUMEXx crispuS L. X R. SANGUINEUS L. *11, S. Hants.: Otterbourne Hill
Common, GR 41/45.22. Scrub. R. P. Bowman, 1978, herb. R.P.B., det. J. R. Akeroyd.

325/12 X14. RUMEX OBTUSIFOLIUS L. X R. SANGUINEUS L. *47, Monts.: Powis Castle Park,
GR 33/2.0. By track in meadow. P. M. Benoit, 1986, NMW.

+327/1. SOLEIROLIA SOLEIROLI (Req.) Dandy *39, Staffs.: E. of Compton, GR 32/87.98.
Stonework by canal. B. R. Fowler, 1977.

343/24. SALIX ALBA L. X S. FRAGILIS L. *25, E. Suffolk: Barham Pits, GR 62/12.51.
Flooded gravel pit. P. G. Lawson, 1985.

350/1. ANDROMEDA POLIFOLIA L. *90, Angus: W. side of Mount Keen, GR 37/40.86. Blanket
peat. Ls. Birse, 1979.

372/4. ANAGALLIS MINIMA (L.) E. H. L. Krause 5, S. Somerset: Bickenhall, GR 31/28.16.
Woodland ride. C. J. Cornell, 1986. Only extant locality.

392/+2 1. SYMPHYTUM ASPERUM Lepech. X S. OFFICINALE L. 103, Mid Ebudes: Cliad Farm,
Coll, GR 17/2.5. J. W. Clark, 1986, E. 1st Coll record.

+392/3. SYMPHYTUM ORIENTALE L. *39, Staffs.: S. of Shatterford, GR 32/79.80. Roadside
verge. W. A. Thompson, 1980, still present in 1986.

4393/1. BORAGO OFFICINALIS L. *93, N. Aberdeen: Peterhead, GR 48/12.45. Waste ground
by shore. D. Welch, 1986, ABD.

+394/1. TRACHYSTEMON ORIENTALIS (L.) G. Don *39, Staffs.: Patshull Church, GR 33/80.00.
Woodland. S. R. Price, 1980. 90, Angus: Near Kinnettles Mill near Douglastown, GR 37/
42.46. Grassy bank. H. J. Noltie & R. B. Robb, 1981, DEE. 2nd record.

4406/2. CALYSTEGIA PULCHRA Brummitt & Heywood *77, Lanarks.: Easterhouse, Glasgow,
GR 26/68.65. J. H. Dickson, 1984, GL.

413/3. SOLANUM NIGRUM L. 94, Banffs.: Keith, GR 38/43.50. Garden weed. E. S. Bruce,
1986, E. 2nd record.

416/7. VERBASCUM NIGRUM L. 46, Cards.: Comins Coch, GR 22/61.82. Abandoned pasture.
A. P. Fowles, 1986, det. A. O. Chater. 2nd record. +73, Kirkcudbrights.: Near Standing Stone
of Kirroughtree, GR 25/42.65. Roadside bank. O. M. Stewart, 1984. Ist post-1930 record.
Minigaff, GR 25/41.66. Disused wood-yard. O. M. Stewart, 1985. 2nd extant locality.

420/+2 x3. LINARIA PURPUREA (L.) Mill. < L. REPENS (L.) Mill. *47, Monts.: Caersws railway
station, GR 32/0.9. Disused yard. P. M. Benoit, 1986, NMW.

420/3 x4. LINARIA REPENS (L.) Mill. x L. vULGARIS Mill. *39, Staffs.: W. of Tunstall, GR 33/
84.51. I. J. Hopkins, 1985, STO.

422/2. KICKXIA ELATINE (L.) Dumort. 39, Staffs.: N.W. of Brewood, GR 33/87.09. Bank of
field pool. B. R. Fowler, 1982. 1st post-1930 record.

7425/3. MIMULUS MOSCHATUS Dougl. ex Lindl. *27, E. Norfolk: W. of Felbrigg, GR 63/
19.39. Cracks in paving. E. Beaumont & E. T. Daniels, 1985.

426/1. LIMOSELLA AQUATICA L. *77, Lanarks.: Cambuslang, GR 26/64.61. Bank of R. Clyde.
J. Lyth, 1986, GL.

4428/1. ERINUS ALPINUS L. 103, Mid Ebudes: Craigdarroch, Coll, GR 17/22.57. Crevices in
wall. J. W. Clark, 1983 and 1986. 1st Coll record.

430/3. VERONICA CATENATA Pennell *103, Mid Ebudes: Lochan a’Chuirn, Coll, GR 17/11.52.
A. Walker, 1986, E.

7430/14. VERONICA PEREGRINA L. *35, Mons.: Raglan, GR 32/38.08. Watered gravelly paths.
C. A. Shirley, 1986, herb. T. G. Evans.

446 PLANT RECORDS

430/22. VERONICA POLITA Fr. 81, Berwicks.: Longformacus, GR 36/69.57. Vegetable garden.
M. E. Braithwaite, 1986, herb. M.E.B. 1st post-1930 record.

4430/24. VERONICA FILIFORMIS Sm. 103, Mid Ebudes: Acha, Coll, GR 17/18.54. Grass by
burn. J. W. Clark, 1983, E. 1st Coll record.

435/1/17X15. EUPHRASIA ARCTICA Lange X E. CONFUSA Pugsl. *77, Lanarks.: Bishop Loch,
GR 26/68.66. Short grassland. P. Macpherson & A. G. Carstairs, 1981, herb. P.M., det. A. J.
Silverside.

435/1/12X23. EUPHRASIA TETRAQUETRA (Bréb.) Arrond. xX E. vicursi Davey *1, W.
Cornwall: Carvannel Downs, GR 10/64.44. Maritime heath. L. J. Margetts, 1981.

440/6. OROBANCHE ELATIOR Sutton *39, Staffs.: 4 km N.E. of Oakamoor, GR 43/07.49.
Scrub. C. Hayes, 1982.

441/1. PINGUICULA LUSITANICA L. 73, Kirkcudbrights.: Below Makelston, GR 25/61.87.
Runnel in field. P. Stanley, 1985. 2nd extant locality.

- +445/7. MENTHA SUAVEOLENS Ehrh. *103, Mid Ebudes: Rubha nan Gall, GR 17/50.56.
Amongst Pteridium. A. Wright, 1984, K, det. R. M. Harley.

459/7. STACHYS SYLVATICA L. 103, Mid Ebudes: Craigdarroch, Coll, GR 17/22.57. B. Cassels,
1986, E. 1st Coll record.

461/1. LAMIASTRUM GALEOBDOLON (L.) Ehrend. & Polatschek +*90, Angus: Forfar, GR 37/
44.50. I. Ford, 1984.

465/4/2. GALEOPSIS BIFIDA Boenn. *39, Staffs.: 2 km W. of Tettenhall, GR 33/86.00. Hedge
bank. B. R. Fowler, 1980.

475/1. CAMPANULA LATIFOLIA L. 107, E. Sutherland: 2 km N. of Helmsdale, GR 39/02.17.
River bank. M. Murray, J. K. & S. I. Butler, 1984, E. 2nd record.

+475/5. CAMPANULA PERSICIFOLIA L. *107, E. Sutherland: Ferry Links, GR 28/81.97.
Roadside. M. J. Marshall, 1985, herb. M.J.M., det. D. McClintock.

485/3b. GALIUM ALBUM Miller 26, W. Suffolk: Newmarket Heath, GR 52/62.63. E. M.
Hyde, 1986, herb. E.M.H. 2nd extant locality.

4487/3. SAMBUCUS RACEMOSA L. *93, N. Aberdeen: Pitcaple, GR 38/73.26. Woodland.
D. Welch, 1985, ABD.

491/2. LONICERA JAPONICA Thunb. *39, Staffs.: S.W. of Great Haywood, GR 33/98.22. River
bank. B. R. Fowler, 1986.

+CEPHALARIA GIGANTEA (Ledeb.) Bobrov *26, W. Suffolk: Long Melford, GR 52/85.47.
Roadside bank. E. M. Coe, 1985.

502/1. BIDENS CERNUA L. 98, Main Argyll: Moine Mhor, GR 16/82.93. Edge of arable field.
B. D. Batty, 1985. 2nd record.

1502/3. BIDENS FRONDOSA L. *44, Carms.: Bynea, GR 21/55.98. Drainage ditch and edge of
marsh. B.S.B.I. field meeting, 1986, NMW, det. T. G. Evans.

+503/2. GALINSOGA QUADRIRADIATA Ruiz & Pavon 42, Brecs.: Brecon, GR 32/04.28.
Pavement. R. G. Woods, 1986. 2nd record. *81, Berwicks.: Chirnside House Hotel, GR 36/
88.56. Vegetable garden. M. E. Braithwaite, 1986, herb. M.E.B.

506/5. SENECIO CAMBRENSIS Rosser *83, Midlothian: Leith Docks, GR 36/27.76. O. M.
Stewart, 1974, det. H. J. Noltie. Carron Place, Leith, GR 36/27.76. Waste ground. H. J. Noltie,
1986, E. 1st and 2nd records.

506/2 X 1. SENECIO AQUATICUS Hill X S. JACOBAEA L. *1, W. Cornwall: Lower Bosistow Farm,
GR 10/36.24. Wet field. G. Halliday, 1985, LANC.

506/747. SENECIO SQUALIDUS L. X S. viscosus L. *73, Kirkcudbrights.: Cargenbridge, GR

PLANT RECORDS 447

25/95.74. Roadside. O. M. Stewart, 1985, E. *77, Lanarks.: Meadowside, GR 26/55.66.
Roadside. P. Macpherson, 1985, herb. P.M., det. H. J. Noltie.

506/67. SENECIO SyLvaTicus L. X S. viscosus L. *77, Lanarks.: Between Glasgow and
Renfrew, GR 26/51.66. Roadside. P. Macpherson & E. Teasdale, 1986, E, det. H. J. Noltie.

4512/1. INULA HELENIUM L. 39, Staffs.: Patshull, GR 33/80.00. Woodland. B. R. Fowler,
1981. 1st post-1930 record.

514/3. FILAGO PYRAMIDATA L. *6, N. Somerset: Fry’s Hill, Axbridge, GR 31/43.54. Stony
limestone grassland. E. J. McDonnell, 1985, herb. R.G.B. Roe, det. C. Jeffrey.

+519/sed. ASTER SEDIFOLIUS L. *17, Surrey: Ash Vale, GR 41/88.52. Established by track.
J. F. & A. C. Leslie & K. W. Page, 1986, herb. A.C.L.

7522/1. CONYZA CANADENSIS (L.) Cronq. *47, Monts.: Welshpool railway station, GR 33/
2.0. Railway yard. Mrs K. M. Benoit, 1986, NMW.

525/1. EUPATORIUM CANNABINUM L. 72, Dumfriess.: Holywood, GR 25/94.79. Bank over old
bridge parapet. A. & M. Shiells, 1986. 1st post-1930 record.

539/3. CARDUUS NUTANS L. +*75, Ayrs.: S. of Maidens, GR 26/20.07. Abandoned airfield.
A. McG. Stirling, 1985.

540/7. CIRSIUM HELENIOIDES (L.) Hill! +47, Monts.: Carno, GR 22/96.96. Roadside bank.
J. Roper, 1986, NMW, det. P. M. Benoit. 2nd record.

540/7 X3. CIRSIUM HELENIOIDES (L.) Hill X C. PALUSTRE (L.) Scop. *106, E. Ross: Contin,
GR 28/45.55. Bank of R. Blackwater. M. Barron, 1985.

540/8. CIRSIUM DISSECTUM (L.) Hill *1, W. Cornwall: Crousa Fen, GR 10/75.19. Damp field.
M. G. B. Hughes, 1983.

544/1. CENTAUREA SCABIOSA L. 77, Lanarks.: Shieldhall, GR 26/52.65. Railway bank.
E. Teasdale & P. Macpherson, 1986, herb. P.M. Ist post-1930 record.

546/1. CICHORIUM INTYBUS L. +*107, E. Sutherland: Bonar Bridge, GR 28/60.91. Roadside.
M. J. Marshall, 1985, herb. M.J.M., det. C. Jeffrey.

*552/2. TRAGOPOGON PORRIFOLIUS L. *50, Denbs.: Towyn, GR 32/97.79. Edge of footpath.
G. P. Kerfoot, 1986, NMVW.

554/2. LACTUCA VIROSA L. +90, Angus: Nicho‘ls Loch, GR 37/64.53. Disturbed ground.
D. M. Bayne & L. Farrell, 1981. 2nd record.

558/1/4. HIERACIUM ALPINUM L. *106, E. Ross: Carn Liath, GR 28/14.51. Rocky knolls.
A. G. Kenneth, 1984, det. D. J. Tennant.

558/1/6. HIERACIUM GROVESII Pugsl. *94, Banffs.: Coire Raibert, Cairngorm, GR 38/00.02.
R. Fitzgerald, 1983, det. D. J. Tennant.

558/1/13. HIERACIUM PSEUDOPETIOLATUM (Zahn) Roffey *96@, Easterness: Glen Einich, GR
27/90.98. Rocky ledge. D. J. Tennant, 1976, herb. D.J.T.

558/1/53. HIERACIUM ARGENTEUM Fr. *83, Midlothian: Arthur’s Seat, GR 36/27.72. Rock
outcrops. R. Learmouth, 1984, E, det. D. J. McCosh & P. D. Sell.

558/1/99. HiERACIUM GRANDIDENS Dahlst. *3S, Staffs.: Hall Dale, GR 43/13.53. Basic
g:assland. R. Smith, J. Bevan & W. Thompson, 1985.

5£8/1/201. HIERACIUM TRICHOCAULON (Dahlst.) Johans. *2, E. Cornwall: Forder near
Saltash, GR 20/41.57. Railway embankment. E. Griffiths, 1984, CGE, det. P. D. Sell.

558/1/204. HIERACIUM CALCARICOLA (F. J. Hanb.) Roffey *4, N. Devon: Witheridge Moor,
GR 21/85.15. Edge of moorland. L. J. Margetis, 1985, CGE, det. P. D. Sell.

558/1/222. HIERACIUM SALTICOLA (Sudre) Sell & West *33, E. Glcucs.: Dowdeswell, GR. 42/
00.20. Disused railway. S. H. Bishop, 1985, herb. S.H.B., det. J. Bevan.

448 PLANT RECORDS

+558/1/pat. HIERACIUM PATALE Norrl. *4, N. Devon: Torrington, GR 21/47.19. Old station
platform. W. H. Tucker, 1986, CGE, det. P. D. Sell.

558/2/1 eur. HIERACIUM PILOSELLA L. subsp. EURONOTUM Naegeli & Peter *77, Lanarks.:
Carmunnock, GR 26/60.56. Streamside. P. Macpherson, 1986, herb. P.M., det. D. R.
McKean. 103, Mid Ebudes: Caoles, Coll, GR 17/12.52. Rocky outcrop. J. W. Clark, 1986, E,
det. A. McG. Stirling. 1st Coll record.

558/2/1 tri. HIERACIUM PILOSELLA L. subsp. TRICHOLEPIUM Naegeli & Peter 103, Mid Ebudes:
Arileod, Coll, GR 17/15.54. Rock outcrop. J. W. Clark, 1986, E, det. A. McG. Stirling. 1st Coll
record.

558/2/1 mic. X 72/8. HIERACIUM PILOSELLA L. subsp. MICRADENIUM Naegeli & Peter x H.
AURANTIACUM L. subsp. CARPATHICOLA Naegeli & Peter *73, Kirkcudbrights.: Dun’s Wa’s near
Tongland, GR 25/70.54. Roadside bank. O. M. Stewart, 1986, CGE.

+558/2/9. HIERACIUM PRAEALTUM Vill. ex Gochnat *39, Staffs.: Fauld, GR 43/18.27. Scrub.
I. J. Hopkins, 1985, STO.

+559/2. CREPIS VESICARIA L. subsp. HAENSELERI (Boiss. ex DC.) P. D. Sell 39, Staffs.:
Draycott in the Clay, GR 43/16.30. Waste ground. I. J. Hopkins, 1985, STO. 1st post-1930 record.

567/1. HYDROCHARIS MORSUS-RANAE L. 39, Staffs.: Fishley, GR 43/00.04. Pool north of canal.
R. Hill, 1985. 1st post-1930 record.

+570/3. ELODEA NUTTALLII (Planch.) St. John *39, Staffs.: Wolverhampton, GR 32/91.99.
Canal. I. C. Trueman, 1982, det. M. Chorley. Shelton, GR 33/88.45. Shallow pool. I. J. Hopkins,
1986. 1st and 2nd records. *47, Monts.: Montgomery Canal from Aberbechan to Pentreheylin,
GR 32/1.9. J. Davidge et al., 1985. 1st and 2nd records. *72, Dumfriess.: Kirkloch Brae,
Lochmaben, GR 35/08.82. M. E. R. Martin, 1986, K, det. D. A. Simpson.

+571/1. LAGAROSIPHON MAJOR (Ridl.) Moss *29, Cambs.: Fowlmere R.S.P.B. reserve, GR
52/40.45. Pools. J. H. Bratton, 1986. *38, Warks.: Newton, GR 42/53.78. Large pool. J. C.
Bowra, 1986. Tile Hill Wood, GR 42/28.79. Pond. M. Burgoine, 1986. Both WAR, det. D. A.
Simpson. 1st and 2nd records.

577/5. POTAMOGETON LUCENS L. 34, W. Gloucs.: Thames & Severn Canal, Brimscombe, GR
32/87.02. S. H. Bishop & S. C. Holland, 1986. Only extant locality.

557/13. POTAMOGETON PUSILLUS L. 34, W. Gloucs.: Taynton, GR 32/72.22. Newly
constructed lake. S. C. Holland, 1986. Only extant locality. *98, Main Argyll: Lochan a’Bruic,
GR 17/92.10. B. H. Thompson, 1986, herb. B.H.T., det. N. T. H. Holmes.

577/17. POTAMOGETON COMPRESSUS L. *92, S. Aberdeen: Loch of Aboyne, GR 37/53.99.
B. MacK. Gerrie, 1978, ABD, conf. N. T. H. Holmes.

579/2. RUPPIA MARITIMA L. *98, Main Argyll: Estuary of R. Add below Islandadd Bridge,
GR 16/80.92. Brackish muddy ditches. A. G. Kenneth, 1986, herb. A. G. K.

580/1. ZANNICHELLIA PALUSTRIS L. *47, Monts.: R. Cam near Llanerfyl, GR 33/0.0.
Montgomery Field Society excursion, 1984. Canal between Crowther Hall and Pool Quay, GR 33/
25.11. C. Paskell, 1978. 1st and 2nd records.

+586/1. HEMEROCALLIS FULVA (L.) L. *73, Kirkcudbrights.: Below Creebridge, Minigaff, GR
25/41.65. Rocky island. O. M. Stewart, 1985, E. *90, Angus: Ninewells, GR 37/37.29. Railway
embankment. H. J. Noltie, 1981, DEE.

+591/1la. ASPARAGUS OFFICINALIS L. subsp. OFFICINALIS *49, Caerns.: Pwllheli, GR 23/3.3.
Sand-dunes. L. J. Larsen, 1986, NMW, det. R. G. Ellis. *90, Angus: Barnhill, GR 37/48.31.
H. J. Noltie, 1981, DEE.

+593/1. LILIUM MARTAGON L. *90, Angus: By Vinney Water, Friockheim, GR 37/58.49.
N. Benvie, 1981.

597/1. GAGEA LUTEA (L.) Ker.-Gawl. +26, W. Suffolk: Kedington, GR 52/70.47. Under trees
in churchyard. F. W. Simpson, 1985. 1st post-1930 record.

PLANT RECORDS 449

+605/2. JUNCUS TENUIS Willd. *50, Denbs.: Chirk, GR 33/2.3. Path by canal. B. Davies,
1986, NMW, det. R. G. Ellis.

605/4. JUNCUS COMPRESSUS Jacq. 90, Angus: Dura Street, Dundee, GR 37/41.31. Damp area
on derelict site. J. L. Thomson, 1981, DEE. 2nd extant locality.

605/amb. JUNCUS AMBIGUUS Guss. 1, W. Cornwall: Near Lowland Point, GR 10/79.19. Edge
of marsh. D. Green & A. J. Byfield, 1984, conf. C. A. Stace. 1st post-1930 record. 75259:
Somerset: Steart, GR 31/28.46. Margin of brackish pools. R. S. Cropper, 1986. *11,S. Hants.:
Hayling Island Golf Course, Sinah Common, GR 40/69.99. Damp, bare stony channel. Stanpit
Marsh, Christchurch, GR 40/16.92. Marshy drain channel. Both R. P. Bowman, 1985, herb.
R.P.B., det. C. A. Stace. 1st post-1930 and 2nd extant localities. 25, E. Suffolk: Walberswick,
GR 62/50.74. Sandy, damp areas behind dunes. P. G. Lawson, 1986. 2nd record. *47, Monts.:
Dovey Junction, GR 22/6.9. Sandy upper saltmarsh. P. M. Benoit, 1986, NMW.

605/fol. JUNcUS FOLIOSUS Desf. *4, N. Devon: Hares Down, Rackenford, GR 21/84.21.
Edge of wet heath. L. J. Margetts, 1986, herb. L.J.M. *5, S. Somerset: Sampford Arundel,
GR 31/11.17. Wet track under trees. L. J. Margetts, 1986, herb. R.G.B. Roe.

605/8. JUNCUS INFLEXUS L. *75, Ayrs.: Dean Park, Kilmarnock, GR 26/43.39. Damp
ground. B. Simpson, 1986. |

607/3. ALLIUM SCORODOPRASUM L. *44, Carms.: Laugharne Burrows, GR 22/2.0. Undistur-
bed dunes. J. Rees, 1986, NMW, det. R. G. Ellis, T. G. Evans & P. C. Hall.

611/1. LEUCOJUM VERNUM L. +*90, Angus: Letham Grange estate near Arbroath, GR 37/
62.45. Under deciduous trees. U. K. Duncan, 1981.

+615/2. SISYRINCHIUM CALIFORNICUM (Ker.-Gawl.) Ait.f. *27, E. Norfolk: Beeston Common,
Sheringham, GR 63/16.42. D. Mower & F. Farrow, 1985.

+620/1. TRITONIA X CROCOSMIFLORA (Lemoine) Nicholson *77, Lanarks.: Between Glasgow
and Renfrew, GR 26/51.67. Woodland. P. Macpherson & E. Teasdale, 1983.

625/1. EPIPACTIS PALUSTRIS (L.) Crantz 38, Warks.: Ladywalk Nature Reserve, Lea Marston,
GR 42/21.91. Damp grassland. R. J. Thomas, 1986, det. P. J. Copson. Ist post-1930
record. 46, Cards.: By Afon Mwldan N.N.W. of Penparc, GR 22/20.48. Base-rich flushes in
pasture. A. O. Chater, A. P. Fowles & D. G. Jones, 1986. 2nd record.

625/2. EPIPACTIS HELLEBORINE (L.) Crantz *90, Angus: Latch Road, Forfar, GR 37/59.60.
Shady lawn. B. Allan, 1984.

625/3. EPIPACTIS PURPURATA Sm. *39, Staffs.: Wilderness Wood, W. of Kinver, GR 32/81.83.
Woodland. B. R. Fowler, 1972.

625/6. EPIPACTIS PHYLLANTHES G. E. Sm. *27, E. Norfolk: Earlham Park, GR 63/18.07.
Woodland. I. White, 1985, conf. E. T. Daniels. Costessey, GR 63/19.10. Damp wood near R. Tud.
E. T. Daniels, 1986. 1st and 2nd records.

628/2. LISTERA CORDATA (L.) R.Br. *52, Anglesey: Newborough Forest, GR 23/4.6. Conifer
plantation. B. D. Wheeler, 1986, conf. R. H. Roberts.

629/1. NEOTTIA NIDUS-AvIS (L.) Rich. 39, Staffs.: Hinkley Wood, Ilam, GR 43/12.50.
P. Roworth, 1986. 1st post-1930 record.

635/1. COELOGLOSSUM VIRIDE (L.) Hartm. 2, E. Cornwall: Colliford Reservoir, GR 20/17.72.
Herb-rich pasture. R. Bellringer, 1986. lst record since 1926.

636/1b. GYMNADENIA CONOPSEA (L.) R.Br. subsp. DENSIFLORA (Wahlenb.) G. Camus, Bergon &
A. Camus *66, Co. Durham: Langdon Beck, GR 35/8.3. D. J. Tennant, 1958 and 1976, herb.
D.J.T., det. V. S. Summerhayes.

643/2b. DACTYLORHIZA MACULATA (L.) So6 subsp. ERICETORUM (E. F. Linton) Hunt &
Summerh. *29, Cambs.: Sedge Fen Drove, Wicken Fen, GR 52/55.70. T. J. Bennett, 1986,
CGE, conf. J. M. Lock, P. D. Sell & S. M. Walters.

450 PLANT RECORDS

643/3b X2b. DACTYLORHIZA INCARNATA (L.) So6 subsp. PULCHELLA (Druce) So6 X D. MACULATA
(L.) Sod subsp. ERICETORUM (E. F. Linton) Hunt & Summerh. *1, W. Cornwall: Between
Coverack and St Keverne, GR 10/77.19. Margin of acid bog and heath. P. D. Sell, 1974, CGE.

643/7. DACTYLORHIZA TRAUNSTEINERI (Sauter) S06 *6, N. Somerset: Winscombe, GR 31/
41.57. Damp calcareous meadow. S. Hedley, 1986, det. J. J. Wood. 65, N.W. Yorks.:
Carperby, GR 34/9.9. Marsh. D. J. Tennant, 1977. 1st post-1930 record. *101, Kintyre: A. G.
Kenneth, D. J. Tennant & M. R. Lowe, 1984, E, conf. R. H. Roberts.

643/1 x 636/1. DACTYLORHIZA FUCHSII (Druce) Sod X GYMNADENIA CONOPSEA (L.) R.Br. 529:
Cambs.: Devils Dyke, GR 52/60.62. P. D. Sell, 1961, CGE. *44, Carms.: Pant-y-llyn meadow,
Milo, GR 22/59.17. Ancient pasture. D. A. Wells, I. K. Morgan & R. D. Pryce, 1986, NMW, conf.
R. H. Roberts.

643/2 X636/1. DACTYLORHIZA MACULATA (L.) So6 subsp. ERICETORUM (E. F. Linton) Hunt &
Summerh. X GYMNADENIA CONOPSEA (L.) R.Br. *106, E. Ross: Balmaduthy Dam, GR 28/
64.57. Heath. M. Barron, 1986, conf. R. H. Roberts.

643/3 x 636/1. DACTYLORHIZA INCARNATA (L.) S06 X GYMNADENIA CONOPSEA (L.) R. Br. wat |
W. Cornwall: Crousa Downs, GR 10/76.18. Margin of wet heath and acid bog. R. Bogue, 1984,
BRIST, conf. R. M. Bateman.

643/5X636/1. DACTYLORHIZA PURPURELLA (T. & T. A. Stephenson) Sod xX GYMNADENIA
CONOPSEA (L.) R.Br. *94, Banffs.: Findochty, GR 38/46.67. Ballast of disused railway. D. C.
Shawcross, 1985, conf. D. J. Tennant & R. H. Roberts.

4646/1. ACORUS CALAMUS L. 73, Kirkcudbrights.: Cluden Water near Dumfries, GR 25/
96.77. Riverside. M. E. R. Martin, 1985. 2nd extant locality.

4648/1. LYSICHITON AMERICANUS Hultén & St John *39, Staffs.: Longdon Green, GR 43/
08.13. Stream bank. B. R. Fowler, 1971.

650/2. LEMNA TRISULCA L. *98, Main Argyll: Appin, GR 17/91.46. Edge of muddy pond.
A. Slack, 1986.

650/min. LEMNA MINUSCULA Herter *1, W. Cornwall: Penlee Gardens, Penzance, GR 10/
46.30. Pond. C. D. Preston, 1983, conf. A. C. Leslie. *35, Mons.: Mynyddislwyn, GR 31/
19.94. Small roadside pond. T. G. & U. T. Evans, 1986, conf. T. C. G. Rich.

652/2. SPARGANIUM EMERSUM Rehm. *111, Orkney: Between Muckle and Peerie Waters,
Rousay, GR 57/40.29. Boggy bank of burn. E. Charter, 1986, E. 1st confirmed record.
653/21. TYPHA ANGUSTIFOLIA L. X T. LATIFOLIA L. *21, Middlesex: Brent Reservoir, GR

51/21.87. Muddy margin. A. C. Leslie, 1983, herb. A.C.L.

654/3. ERIOPHORUM LATIFOLIUM Hoppe 46, Cards.: Llwyn-llwyd, N.N.W. of Penparc, GR
22/20.48. Base-rich flush in pasture. A. O. Chater, A. P. Fowles & D. G. Jones, 1986. 2nd record.

655/4. ScIRPUS SYLVATICUS L. 93, N. Aberdeen: Gight, GR 38/82.39. Marsh in woodland.
D. Welch, 1986, ABD. 2nd extant locality.

655/12. SCIRPUS FLUITANS L. 5, S. Somerset: Withypool, GR 21/83.35. Pond by R. Barle.
J. E. Beresford, 1986. 1st record since 1918. 39, Staffs.: Biddulphs Pool N.E. of Norton Canes,
GR 43/03.09. Pool. B. R. Fowler, 1974. 1st post-1930 record.

656/6. ELEOCHARIS UNIGLUMIS (Link) Schult. *39, Staffs.: Allimore Green, GR 33/85.19.
Marsh in ditch. K. M. Goodway, 1972, dét. S. M. Walters.

657/2. BLYSMUS RUFUS (Huds.) Link 93, N. Aberdeen: Boddam, GR 48/13.42. Rock pool on
shore. D. Welch, 1986, ABD. 1st post-1930 record.

663/2. CAREX DISTANS L. *47, Monts.: Afon Llyfnant, Dovey Junction, GR 22/6.9. Muddy
bank of tidal river. P. M. Benoit, 1953 and 1986, NMW.

663/211. CAREX DISTANS L. X C. EXTENSA Gooden. *1, W. Cornwall: Harbour Cove,
Padstow, GR 10/90.77..R. W. David, 1983, det. A. O. Chater.

PLANT RECORDS 45]

663/3. CAREX PUNCTATA Gaudin *44, Carms.: Tywyn Burrows, GR 22/36.04. Dune meadow.
R. Meade & R. D. Pryce, 1982, NMW, det. A. O. Chater & R. W. David.

663/84. CAREX DEMISSA Hornem. X C. HOSTIANA DC. *4, N. Devon: Hares Down,
Rackenford, GR 21/84.21. Wet heath. L. J. Margetts, 1986, herb. L.J.M., det. R. W. David.

663/4X7. CAREX HOSTIANA DC. x C. LEPIDOCARPA Tausch *46, Cards.: Llwyn-llwyd,
N.N.W. of Penparc, GR 22/20.48. Base-rich flush in pasture. A. O. Chater, 1986, NMW, conf.
R. W. David.

663/410. CAREX HOSTIANA DC. xX C. SEROTINA Meérat *29, Cambs.: Sedge Fen Drove,
Wicken Fen, GR 52/55.70. R. W. David, J. M. Lock & P. D. Sell, 1986, CGE.

663/7a. CAREX LEPIDOCARPA Tausch subsp. LEPIDOCARPA *46, Cards.: Mwldan valley N.N.E.
of Penparc, GR 22/20.48. Base-rich flushes in pastures. A. O. Chater, A. P. Fowles & D. G.
Jones, 1986, NMW, conf. R. W. David. Afon Mwidan, Glanilynan, GR 22/19.48. Base-rich flush.
A. O. Chater, 1986. Ist and 2nd records.

663/11. CAREX EXTENSA Gooden. *75, Ayrs.: S. of Kennedy’s Pass near Lendalfoot, GR 25/
1.9. A. Laing, 1985, det. A. C. Jermy.

663/17. CAREX VESICARIA L. 2, E. Cornwall: Near Lower Tamar Lake, Kilkhampton, GR
21/29.10. Damp ground. W. H. Tucker, 1985. 1st post-1930 record.

663/20. CAREX RIPARIA Curt. 80, Roxburghs.: Sorbietrees, GR 35/48.85. By ditch in wet
hollow. R. W. M. Corner, 1986, herb. R.W.M.C. 2nd record.

663/21. CAREX ACUTIFORMIS Ehrh. 46, Cards.: Afon Mwldan, Glanllynan, GR 22/19.48.
Flushed slopes and fen. A. O. Chater, 1986, NMW. 2nd record.

663/42. CAREX BUXBAUMII Wahlenb. *98, Main Argyll: Loch margin. P. Batty, 1986, herb.
P. B., det. D. R. McKean, conf. A. O. Chater & R. W. David.

663/49. CAREX RECTA Boott 107, E. Sutherland: Bonar Bridge, GR 29/59.30. Estuary.
J. Killick, 1984, BM, det. A. C. Jermy. 2nd record.

663/52. CAREX BIGELOWI Torr. ex Schwein. *93, N. Aberdeen: Buck of Cabrach, GR 38/
41.22. Dry moorland. D. Welch, 1986, ABD.

663/57X71. CAREX OTRUBAE Podp. X C. REMOTA L. *50, Denbs.: Sutton Green, GR 33/
40.48. Edge of pond. J. A. Green, 1986, NMW.

663/69. CAREX ELONGATA L. *11, S. Hants.: St Leonards South N.R., Hurn Forest, GR 41/
10.00. Alnus-Salix carr. Moors River N.R., Fillybrook, GR 40/12.98. Salix carr. Both R. M. Walls,
1985, herb. R.M.W., det. A. O. Chater. 1st and 2nd records.

670/11. FESTUCA TRACHYPHYLLA (Hackel) Krajina *61, S.E. Yorks.: Near Wintringham,
GR 44/9.7. Chalk bank. E. Chicken, 1986, herb. E.C., conf. P. J. O. Trist.

+670/diff. FESTUCA DIFFUSA Dumort. *77, Lanarks.: Shiels, Glasgow, GR 26/53.66. Grassy
bank in old dockland. P. Macpherson & E. L. S. Lindsay, 1985, herb. P. M., det. P. J. O. Trist.

673/cap. PUCCINELLIA CAPILLARIS (Liljebl.) Jansen *93, N. Aberdeen: Peterhead, GR 48/
13.45. Harbour wall. D. Welch, 1986, ABD, conf. H. J. M. Bowen.

676/11. POA ANGUSTIFOLIA L. 103, Mid Ebudes: Staffa, GR 17/32.35. Dense grassland. B. M.
Mull Survey, 1969, BM, det. A. Melderis. 2nd record.

4676/15. Poa cHarx Vill. *33, E. Gloucs.: Dowdeswell, GR 32/99.19. Steep bank. S. H.
Bishop, 1985.

677/1. CATABROSA AQUATICA (L.) Beauv. 90, Angus: East Brae, Turin Hill, GR 37/52.53.
Ditch. H. J. Noltie, 1981, DEE. 1st record since 1914.

683/10X13. BROMUS HORDEACEUS L. X B. LEPIDUS Holmberg *4, N. Devon: Westleigh
Quarry, GR 31/06.17. Roadside. L. J. Margetts, 1985, conf. P. J. O. Trist.

452 PLANT RECORDS

683/14. BROMUS RACEMOSUS L. *47, Monts.: A495 3 km N. of Llanfair Caereinion, GR 33/
10.09. Roadside. I. C. Trueman & W. A. Thompson, 1985.

693/2. AVENULA PUBESCENS (Hudson) Dumort. *47, Monts.: Llanymynech Hill, GR 33/
26.21. Short limestone turf. E. D. Pugh & M. Wainwright, 1983. 1st native record. 59) S:
Lancs.: Sniggery, Hightown, GR 34/29.02. Sandy field. V. Gordon, 1986. 1st post-1930 record.

695/12. HoLcus LANATUS L. X H. MOLLIS L. *44, Carms.: Near Gwyddgrug, GR 22/46.34. ,
Roadside. T. S. Crosby & A. M. Pell, 1986.

700/2. CALAMAGROSTIS CANESCENS (Weber) Roth *42, Brecs.: Llwyncus, GR 22/92.50. Carex
acutiformis swamp. R. G. Woods, 1986.

701/4. AGROSTIS GIGANTEA Roth 73, Kirkcudbrights.: Near New Abbey, GR 25/94.66.
Roadside verge. O. M. Stewart, 1986. 2nd record. *80, Roxburghs.: Yetholm Loch, GR 36/
80.27. Edge of barley field. R. W. M. Corner, 1986, herb. R.W.M.C.

707/1. PHLEUM BERTOLONI DC. *77, Lanarks.: Meikle Dripps, GR 26/57.55. Pasture. E.
Teasdale & P. Macpherson, 1985, herb. P.M., conf. P. J. O. Trist. Kittochside, GR 26/61.56. Farm
track. P. Macpherson, 1985, herb. P. M. Ist and 2nd records.

708/5X3. ALOPECURUS BULBOSUS Gouan X A. GENICULATUS L. *14, E. Sussex: Glynde
Reach, GR 51/46.09. B. Wurzell, 1977. Earliest British record.

Watsonia, 16, 453-457 (1987) 453

Book Reviews

A catalogue of natural science collections in North-east England, with biographical notes on the
collectors. Edited by P. Davis & C. Brewer. Pp. 333. North of England Museums Service, 27
Sutton St., Durham DH1 4BW. 1986. Price not stated (ISBN 0-—9510948—0-7).

Although misleadingly titled — for it is concerned exclusively with collections of natural history
(and what have local museums got against that long-established term?) — this is a valuable
compendium of all that the four counties of Northumberland, Tyne and Wear, Durham and
Cleveland have to offer to anyone in the way of specimens. Effectively the print-out of a
computerised database compiled at the Hancock Museum through the Newcastle University
facilities, it represents a notable advance on an analogous register of the collections in North-west
England that was published in 1981. In particular, a great deal of effort has gone into giving the
bare lists some human embodiment with no less than a hundred pages of biographical details.
Indexes are provided both to the geographical coverage and to the broad categories of material of
which each collection consists; 97 vascular plant collections are featured. The information is being
continuously updated, so amendments or additions can be incorporated without difficulty.

D. E. ALLEN

The secret life of an oakwood: a photographic essay. Stephen Dalton with Jill Bailey. Pp 160, with
146 colour plates. Century Hutchinson, London. 1986. Price £14.95 (ISBN 0-—7126—1278-5).

A sequence of photographs taken through a year centres this book round the illustrations, which
are grouped in the four seasons and linked by a paragraph on each picture. It has the fascination of
a study of a small area in well observed detail, highlighting the less obvious wildlife supported by
the oaks, and seven other trees, in a small oakwood forming part of a nature reserve near Ardingly
in Sussex, a relict of the forest ridge that once extended from Kent to Hampshire.

With his particular interest in insect photography, Stephen Dalton delights in pictures of these,
including a green oak-roller moth in flight and six poplar sawfly larvae munching along a leaf edge;
also dewdrops on the snare of a money spider slung on blades of grass, a jumping dormouse and a
night flashlight picture of a long-eared bat chasing an orange underwing moth. There are very few
daytime flash pictures (with black backgrounds), for example a Digitalis purpurea flower with a
foraging bumblebee, as the preference for natural light is explained in the ‘Notes on the
photography’. A stunning picture of water droplets with reflections held by capsules of moss has
natural colours, and a close-up of part of a single bluebell stem shows the flowers in all the
multishades of blue, although the haze of distant bluebells under trees does still seem to defeat
one-dimensional reproduction.

Not all the plants are given specific names; a Castanea sativa chestnut leaf is named Aesculus
hippocastanum in the text, and a lacewing on a stem with armature of assorted prickles intrigues
with speculation as to which Rubus it is. The four-page introduction summarizes the history of
English oakwoods, but this book is mainly for the eye. The wide-ranging selection of pictures
(including birds — many in flight — and the common agrion damselflies described as “‘seen whirring
in the dappled sunlight like miniature helicopters — a magical sight’’) is evocative of the richness of
the habitat, and some lighting effects used make unusual plant portraits.

M. BriGcs

The colour cauldron. The history and use of natural dyes in Scotland. Sue Grierson. Pp. 234, with
12 colour and monochrome plates, 21 line drawings and a map. Privately published and obtainable
from Sue Grierson, Mill Books, Newmiln Farm, Tibbermore, Perth PH1 1QN. Price £10.50 (ISBN
0—9510132-11).

454 BOOK REVIEWS

The author reviews the historical and practical background to the use of natural dyes in Scotland,
and also gives much useful and pragmatic advice to today’s craft dyers. Chapters cover the
history of Scottish textile dyeing (particularly tartans), the development of the dye industry,
technical information, plant conservation, native dye plants, lichens and imported dyestuffs.
Much of the information given is not otherwise easily accessible, and it is clear, well set out and
well researched. The line drawings are mostly adequate, but some are poor and one or two, e.g.
Lycopodium complanatum, are unrecognizable. The author deals at length with dyes she has
prepared in her own workshop. English names are used, but scientific names are also given, thus
avoiding the ambiguity of much current literature on this subject. She does, however, include
Sonchus asper with Cirsium palustre and C. arvense as ‘thistles’, so it is not surprising that
different colours were produced by these species. The chapter on conservation is admirable and
includes an eloquent appeal for commonsense, understanding, restraint and forethought in the
use of all natural resources, whilst the requirements of the law are clearly stated. Helpful advice
is given for achieving colours traditionally produced from the particularly vulnerable lichens by
the use of other plants, often extremely common weeds. The extensive bibliography of this
excellent book will be very helpful for further study. It is commendable for linking craft work to
scientific botany, a feature lacking in most books on dyeing from craftworkers, and it represents
very good value for money.

M. J. CANNON

Retkeilykasvio, 3rd ed. Edited by L. Hamet-Ahti et al. Suomen Luonnonsuojelun Tuki Oy,
Helsinki. 1986. Price not stated (ISBN 951—9381-14—7 hardback; 951-9381-—15-—5 softback).

This Field Flora of Finland, now in its third edition, has not previously been brought to the
attention of Watsonia readers. It is in many ways a model Flora, with keys to families, genera
and species, concise descriptions, dot-maps in which Finland is divided into 21 biogeographical
provinces, and copious illustrations (over 200 more in ed. 3 than in ed. 2). In many genera the
whole plants are illustrated, in others mostly just diagnostic details (such as leaves in Alchemilla,
calyces in Myosotis, utricles and inflorescences in Carex). There are eight subspecies in the
Ranunculus auricomus group illustrated but curiously not mentioned in the text. The symbols on
the maps differentiate between common and rare, native and archaeophyte, established and
casual alien (both before and after 1950). These symbols, and others in the text, are explained in
a brief English summary. It is interesting to see that opinion has changed on the status of a
number of species since the second edition of 1984. It is striking how many Finnish species are
largely or entirely confined to the Aland islands.

Unusually for a field Flora the critical genera are well covered (25 species of Alchemilla and
seven of Euphrasia treated in full, 89 of Taraxacum keyed and 35 with at least the leaves
illustrated, 80 of Hieracium keyed and 21 illustrated). Symbols indicating edible and poisonous
are affixed to an unusually high proportion of species, Epilobium angustifolium, Leontodon
hispidus and Elymus repens among the former, all species of Dryopteris among the latter. The
taxonomy is in general reasonably conventional. But Epilobium adenocaulon (purplish-red flow-
ers) and E. ciliatum (white) are separated, as are western and eastern races of the former, and
the two subspecies of Carex muricata are not distinguished, although subspecies and varieties are
widely adopted elsewhere in the genus. The keys prove reasonably easy to tackle with a diction-
ary, and the Flora would be essential for anyone visiting Finland. For the illustrations and maps,
though, it should be of much wider interest. The soft-covered version has a resilient, pleasantly
textured waterproof covering, but is too large for the pocket.

A. O. CHATER

Biology of plants. P. H. Raven, R. F. Evert & S. E. Eichhorn, 4th ed. Pp. xviii + 775, with
numerous illustrations in colour and black & white. Worth Publishers, Inc., New York. 1986.
Price £23-00 (ISBN 0-87901-—315-X).

BOOK REVIEWS 455

The nature and scope of this work were described in some detail in a review of the second edition
by O. Kay (Watsonia, 11: 411-412, 1977). The fourth edition is bigger and better. The increase in
size is due primarily to the provision of more, and larger, illustrations, many of them in colour, and
the further addition of very useful tables and summaries. There has also been an attempt to stress
the relevance of plants to practical human affairs, a welcome move in this day and age when
science is often taken for granted or trivialised by the newspapers and television. In this regard, an
extra chapter on the role of plants in the past, present and future development of the human race,
has been added. The coverage of topics, from molecular biology and physiology to ecology,
systematics and evolution, has been updated in accordance with recent discoveries, and I would
particularly recommend the chapter on the evolution of flowering plants to B.S.B.I. members. The
criticisms of the second edition outlined by Q. Kay have, as far as I can see, been largely remedied
and, apart from a few typographical errors (e.g. “‘hydathodes”’ misplaced in the index), this book is
a model student text and represents good value for money.

R. J. GORNALL

Plant breeding systems. A. J. Richards. Pp. xiv + 529, with numerous illustrations in black &
white, and four in colour. George Allen & Unwin, London. 1986. Price £45 (ISBN 0—04—581020-
6) hardback; £19-95 (ISBN 0-—04—581021-—4) soft cover.

Despite its title, this book deals almost exclusively with angiosperms, as the author warns in the
Preface. The subject begins with a discussion of sexual theory and a general review of sexual
reproduction, and continues with treatments of floral morphology, pollination and their effects on
gene flow. Next there are three chapters on ways in which plants enforce or promote outbreeding,
viz. by homomorphic and heteromorphic self-incompatibility and by dicliny. Lastly, the topics of
selfing, vegetative reproduction and agamospermy are covered.

The book is densely written and includes masses of detail. For this reason it has to be read
carefully in order to follow lines of argument, which may at times be complicated. One feature
whose utility I doubt is the frequent cross-referencing to other chapters. I found that if I ignored
these, I felt I might be missing something important, but if I followed them up, I could not read the
book in any ordinary way as my efforts became rather like those of a pollinator foraging hither and
thither. The topics discussed are dealt with very thoroughly, by and large; however, there is little
or no mention of the genetic control of unilateral incompatibility or interspecific incompatibility.
Nor is there mention of pollen-tube inhibition in the ovary during self-incompatibility, a
phenomenon which is probably commoner than is usually believed, and which may have
interesting evolutionary implications. However, these are relatively minor points, bearing in mind
the scope of the book and the burgeoning literature. Although some of the photographs are poor, I
would strongly recommend this book to advanced undergraduates and those seeking a modern,
condensed review of this huge subject.

R. J. GORNALL

The European garden Flora, volume 1, Pteridophyta, Gymnospermae, Angiospermae — Monocoty-
ledons (part 1). Edited by S. M. Walters and nine others. Pp. xv+ 430, with 44 figures and one map.
Cambridge University Press, Cambridge. 1986. Price £55-00 (ISBN 0—521—24859-0).

Volume one of this essential horticultural work of reference is in fact the second volume to be
published: volume two, covering the remainder of the monocotyledons (Juncaceae to Orchi-
daceae), was published in 1984 and has already been reviewed in Watsonia, 15: 284—285 (1985).
This volume is in the same format as the earlier published one with the text printed in triple
columns and has an almost identical introduction. A major difference is the inclusion of additional
major illustrations used in the Pteridophyta. These are the same xerographic reproductions as
found in The ferns of Britain and Ireland by C. N. Page (1982), very useful but not pleasant to look
at. They are not exactly silhouettes, for details of surface features are just about visible. However,

456 BOOK REVIEWS

numerous areas fading to very pale grey give the appearance that something has gone wrong with
the printing. It is a pity more money was not available for better quality illustrations, but at £55 the
book is already too expensive (volume one cost £30 in 1984, and one wonders just how much the
total six volumes will be!).

It is hoped that the book with its durable-looking cover (it has no dust jacket) is indeed hard-
wearing, so important for a frequently-consulted reference work. I am a little apprehensive about
the quality of the paper for such a large page format, also the binding: my experience is that
working copies of Flora Europaea (similar format, same publisher) have invariably to be rebound
and pages repaired.

I was disappointed to see four genera of the Liliaceae (Nectaroscordum, Bessera, Bloomeria and
Muilla) given in an addendum at the end of the book on page 392. Nectaroscordum is mentioned as
a footnote in the key to genera of the family as keying out with Allium, and the other three genera
with Nothoscordum. Differences from Allium and Nothoscordum are given in the addendum
descriptions. It is a pity these could not have been included in their correct systematic positions.
One hopes that future volumes will lack such addenda.

Only time will tell how well the keys work and how complete the treatments are. However, I am
already regularly using the work and look forward to the publication of the remaining four
volumes, such is the need for the book.

S. L. Jury

Bracken. Ecology, land use and control technology. Edited by R. T. Smith & J. A. Taylor. Pp. 464.
Parthenon Publishing, Carnforth, Lancs. 1986. Price not stated (ISBN 1—85070-128-8).

Bracken is the one fern known to almost everyone, and one of the few species of pteridophytes to
become a problem to man. It is perhaps the world’s most widespread vascular plant, and as a weed
is unusual not only in that it is a native throughout almost its entire range, but that it also possesses
all the peculiar attributes with which it is endowed by its pteridophyte life-cycle.

This book represents the proceedings of a Symposium to discuss the problems of bracken, which
was held at the University of Leeds, on 1st—Sth July 1985. It contains 49 research papers,
representing the combined endeavours of no less than 84 international contributors reporting from
disciplines as wide apart as Geography, Agriculture, Forestry, Veterinary Science, Medicine,
Pharmaceuticals, Planning, Land Use Management, Nature Conservation, Ecology and Pteridol-
ogy. Contributions also range geographically widely, from America, eastern Europe, Australia
and New Zealand, while the problem of bracken in Britain, and especially in upland Britain, is
particularly well documented. Contributions include those of both ‘pure’ and ‘applied’ approaches.

This symposium volume builds on the base established by the first two Bracken Symposia (Bot.
J. Linn. Soc., 73 (1-3), 1976 and Proc. R. Soc. Edinb., 81B (1-2), 1982). It reflects the growing
international awareness of, and concern for, the problem of bracken, whose alarming rates of
spread and menacing potential problems are becoming realized by the scientific community at
large. These aspects are particularly well reflected in this volume.

What is less clear to scientists in general, and hence equally reflected by this volume, is what best
can be done about it. Many and various ongoing research programmes to this end are reported in
the volume, reflecting (and in many cases incorporating) discussion on this aspect at, and
stimulated by, the conference. One thing that stands out clearly is that research in the bracken
problem is taking place independently in a very great number of different and unconnected
research institutes and universities, presumably funded by an equally varied array of bodies.
Another is how potentially beneficial such multi-disciplinary research can be. One danger of this
approach to the problem however, is that one scientist may not to be aware of progress by (or even
sometimes the existence of) other research groups. To a large extent, the publication of this
volume will have bridged these many gaps for the time being, and a multi-disciplinary co-
ordinating group of researchers, established as a result of the conference, should assist with better
interdisciplinary communication in future.

Bracken meanwhile, has become the world’s best known and most studied pteridophyte. It also
remains one of the world’s most extensive and enigmatic weeds, the consequences of infestation

BOOK REVIEWS 457

with which, in temperate zones, are only just becoming realized. The authors of this volume are to
be congratulated on amassing such an array of data and making it available in published form. The
book can certainly be recommended to anyone with even a peripheral interest in the diverse and
growing problem of this plant, while we can be sure that it is not the last we will hear about
bracken.

C. N. PAGE

John Hope, 1725-1786. Scottish botanist. A. G. Morton. Pp. 47, with 11 black & white
photographs. Edinburgh Botanical Garden (Sibbald) Trust, Edinburgh. 1986. Price £3-00 (ISBN
0—9504270-5-5).

The subject of this memoir was Superintendent of the Royal Garden in Edinburgh and Professor of
Botany and Materia Medica in Edinburgh University from 1761 to 1786. His activities thus co-
incided with the period of the Scottish Enlightenment, when intellectuals such as David Hume and
Adam Smith and artists such as Allan Ramsay were making Edinburgh a leading centre of culture
in Europe. Although his botanical correspondence was almost worldwide and he exchanged letters
with many famous botanists, such as Linnaeus and his son, Miller, Banks, Garden, Bernard de
Jussieu, von Haller and L’Héritier de Brutelle, he became known more as a teacher than as a
research worker; and information about him and his work is limited. Professor Morton’s vivid
picture of an important contributor to 18th century botanical thought and a much-loved and highly
influential, though not particularly articulate professor is thus all the more praiseworthy.

Hope’s interest in the Scottish flora led him to organise a botanical survey of Scotland; but it was
never completed, probably because of the publication of Lightfoot’s Flora Scotica. The collectors
whom he sent out included James Robertson, who discovered Eriocaulon aquaticum in Skye.
Hope sent specimens of this (provisionally named Nasmythia articulata) to Linnaeus in 1769 and
followed them in 1772 with material of another new species, which he named Ruppia spiralis, from
“‘a Scottish loch’’. His studies in Paris with Bernard de Jussieu led to his interest in a ‘natural
system’ of plant classification; and so, despite being one of two botanists who introduced the
(artificial) Linnaean Sexual System to Britain, he “did not admire Linnaeus so much as a
philosopher as a naturalist and systematist”’.

The account of John Hope’s life and work is preceded by an introduction by Sir George Taylor
and followed by a selection of his letters as well as notes on (i) plants especially connected with
Hope, (ii) the two catalogues of the Edinburgh botanic garden issued during his period as Regius
Keeper, and (iii) Hope’s Hortus Siccus. In all, this slim volume is a most important contribution to
our knowledge of a hitherto rather shadowy figure who, mainly through teaching and
correspondence, played a considerable part in the formation of 18th century botanical thought and
the education of some famous early 19th century botanists such as Sir J. E. Smith, W. Roxburgh,
A. Menzies and (indirectly) Robert Brown.

N. K. B. RosBson

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Watsonia, 16, 459-463 (1987) 459

Obituaries

ALEXANDERS MELDERIS
(1909—1986)

Alexanders Melderis, who died in March last year at the age of 77, was in charge of the British and
European herbaria of the British Museum (Natural History) from 1951 to 1974, and many B.S.B.I.
members will have known him well. None of us, though, knew much about his early years. He was
born and educated in Latvia and obtained his doctorate there. When the Russians invaded the
country in 1944 he escaped to Sweden and worked until 1951 in the Institute of Systematic Botany
at Uppsala. He attended the International Botanical Congress in Stockholm in 1950, and it was
there that George Taylor (now Sir George) met him. Shortly after this, Taylor, by then Keeper of
Botany at South Kensington, had to find a replacement for A. J. Wilmott and wanted for the post
someone with an interest in the flora of Europe as a whole rather than a botanist whose primary
concern was the British flora. Melderis seemed to him the most suitable candidate, and he
arranged for him to come to London and take up the post, which he filled with distinction until his
retirement, becoming also Deputy Keeper of Botany from 1966 to 1972.

Melderis’s chief interest was the grasses, and in particular the Triticeae, and he will be best
remembered for his accounts of these in various regional floras — Flora Europaea, Flora
Zambesiaca and Flora Iranica amongst others. As with most taxonomists, official retirement did
not mean stopping work, and at the time of his death Melderis was working on Asiatic grasses, for
which his knowledge of Russian, a necessary second language in Latvia even before 1939, was a
particular help. His botanical work was careful, thorough and sound, and his contributions will
stand the test of time. The botanists who knew him will, however, remember him more for his
quiet, gentle personality and his helpfulness, whether in identification, in translation from Russian
or in guidance to literature.

Apart from botany, Melderis’s other great interest was chess, and it was his enthusiasm that
popularized it in the Museum. He was so successful in this that the Museum, instead of
contributing one or two members to a combined team with the British Museum, came to have two
teams of its own which competed with considerable success in the Civil Service League. In private
life away from the Museum his chief interest and concern was the community of Latvian exiles in
London and their welfare, for which he did much in his quiet way. He will be much missed by all
who knew him.

R. Ross

JOHN SCOTT LENNOX GILMOUR
(1906—1986)

John Gilmour, former Director of the University Botanic Garden, Cambridge, and of the Royal
Horticultural Society’s Garden at Wisley, Surrey, died on 3rd June 1986, aged 79. Although he was
probably known to relatively few B.S.B.I. members, apart from the older ones or those living in
Cambridge, it was through wild flowers that he came to botany, and thence to involvement in the
Society. How he took the first step in this career is charmingly told by himself. ““When I went to a
preparatory school I knew and cared nothing about wild plants. At the end of the summer term
each boy had to produce fifty named species. On the last day but one I had not collected a single
plant. Desperation drove me to a high-speed tour of the lanes near the school, guided by a friend
who had already made his collection, and on the following day I duly presented my fifty plants.
This discreditable incident implanted in me, against every modern principle of education, a passion
for the British flora which has never been extinguished” (Gilmour & Walters, Wild flowers, 1954).

John Gilmour read Natural Sciences at Cambridge and afterwards started research there on

460 OBITUARIES

Centaurium. Soon he was appointed Curator of the Herbarium and he worked with W. T. Stearn to
produce a set of 20 exsiccatae of the British flora and a further 20 of cultivated plants. This is a bigger
undertaking than it may seem to anyone who has not done it. The accompanying ‘Schedae’ were
published with W. T. Stearn in the Journal of Botany, London, 70, suppl., 1-29 (1932)*; they are
valuable contributions on the taxonomy and nomenclature of their subjects. With A. J. Wilmott he
compiled the contemporary equivalent of B.S.B.I. Abstracts.

Gilmour’s ‘passion’ for the British flora was not an exclusive one, for he was also a philosopher,
poet, musician, bibliophile and athlete, and this involvement with the humanities surely influenced
his approach to botany. The wild-flower side of botany is, of course, one of the humanities, but his
breadth of interest was mirrored in his view of botany, and in his continual effort to stop the
professionals from isolating themselves from the amateurs and the world at large. To one with such
an aim, pragmatism was essential, and it informed his main contributions to botany. The first of these
was his advocacy of the proposition that classification is not phyiogeny and should not attempt to be.
The most useful classifications, he said, are general-purpose classifications, and to this end they must
be based on as wide a selection of characters as possible (that is, they are ‘natural classifications’ as
opposed to ‘special classifications’ based on one or a few characters). Traditional biological
classification was of this type, and Gilmour was anxious that it should not be taken over on the one
hand by phylogenists or on the other by genecologists; both groups would have made it their own
‘special classification’. To provide for the needs of genecologists he devised, in collaboration with
J. W. Gregor and J. Heslop-Harrison, the ‘deme’ terminology, a totally flexible system using
informal nomenclature. Aithough it has not been used a great deal it was probably responsible for
keeping the genecologists’ hands off the traditional system. His attempt to stave off the merger of
classification with phylogeny received a tremendous boost in the 1950s, when computers had made
possible the practice of ‘numerical taxonomy’. This, it was hoped, would bring about the era of
objective, ‘repeatable’ classifications. The numerical taxonomists knew that they must rely on facts
and exclude evolutionary speculation. Gilmour had been demanding the same thing for a different
reason (in J. Huxley, ed. The new systematics, 1940) and there ensued a period of happy mutual
support and collaboration, embracing the formation of the Classification Society.

Gilmour’s other pragmatic stand in botany was over nomenclature: rules that were too rigid were
bringing about name changes that the Code of Nomenclature was ostensibly designed to minimise, as
these might lead to a popular rejection of official nomenclature. He worked for many years towards
enactment of rules to protect specific names or epithets from what amount to nomenclatural
accidents. Some degree of protection was finally bestowed by the International Botanical Congress
in 1981, long after Gilmour had retired from the field. He had an even more central role in the
establishment and development of a workable Code of Nomenclature for Cultivated Plants. The first
edition of this was published in 1953. Its main innovation was the cultivar concept. This provided for
naming taxa important in agriculture, horticulture and forestry beyond the needs provided for in the
International Code of Botanical Nomenclature.

John Gilmour’s unfailing good nature, politeness and charm earned him a large circle of friends
and made him the natural president of the many societies that he supported and chairman of the
committees on which he worked. He did, indeed, serve as the first President of the B.S.B.I. under its
new, post-war, name (1948-51). It was a great satisfaction to him that, at the end of his career, the
Royal Horticultural Society (of which he was a Council Member) and the B.S.B.I. were able to join
forces in a conference on Plants: wild and cultivated in 1972. He, of course, contributed to the
proceedings which were published in 1973. Though Gilmour’s involvement with gardens, garden
plants and horticultural societies drew him away from the native flora and the B.S.B.I. in his later
life, many who use plant names have benefited in some way from his labours, though some may not
be aware of it. :

The circle of friends that I have alluded to was in fact the joint creation of John and Molly Gilmour,
for she was and is his equal in the qualities I have named, to which must be added hospitality. To be
received into their home was to experience a special quality of warmth, though it shone on the many
rather than the few.

P. F. YEo

* A file of Gilmour’s publications is kept in the library of the University Botanic Garden, Cambridge.

OBITUARIES 461
J. S. L. GILMOUR aAnb THE B.S.B.I.

John Gilmour’s connection with the Society began in his teens, when he had a new county record
for Galium parisiense published in the Report of the Botanical Exchange Club for 1924 by Druce
(Rep. botl Soc. Exch. Club Br. Isl., 7: 877, 1925). On Druce’s death in 1932 he was one of
several professional botanists who were recruited to the new General Committee, when the
B.E.C. was at last placed on a proper constitutional footing; and thereupon, jointly with A. J.
Wilmott, in a symbolical partnership between the British Museum and Kew, he assumed
responsibility for the abstracts from botanical literature which formed a much-valued addition to
the annual reports. At the centenary dinner in 1936, in the kind of gesture he had a noted flair
for, he had one of the huge leaves of Victoria amazonica, the B.E.C.’s emblem, brought up from
Kew for the occasion to form a striking centrepiece to the table. In those years his interest in
British field botany became largely focused on the taxonomy of Centaurium and the collecting of
local Floras. Both of these, sadly, were to be casualties of his move from Kew to Wisley and his
consequent absorption into the world of horticulture, the Floras being disposed of to N. Douglas
Simpson (and thus eventually, and appropriately, finding a resting-place in the Herbarium at
Cambridge, where he had started out on his career) and the Centaurium data passed over to a
research student working on that genus at Liverpool University. But despite that disengagement,
his suavity and savoir faire made him a very obvious choice when the B.S.B.I. (as the B.E.C. had
just become) sought a successor to its long- serving Chairman, the wise and ever-tactful H. T.
Baker, in 1947. When the office of Chairman was converted into that of President one year later,
Gilmour found himself in the difficult and distasteful role of arbitrator in the friction between the
Officers which was such an unfortunate feature of those early post-war years. In particular, it fell
to him to chair the historic, highly fraught meeting in February 1950 at which the Honorary
General Secretary, Miss Campbell, presented the Council with an ultimatum and, on finding it
unacceptable, dashed from the room in tearful fury. A few months later it was again his
misfortune to be presiding when Miss Campbell’s arch-enemy and successor, Lousley, success-
fully opposed her election as Honorary Secretary of the Development and Rules Committee in
her presence. There has never been a more trying term of office for a President, but no one could
have steered the Society through that particular patch of rough water with a surer touch and
more admirable finesse.

D. E. ALLEN

BARBARA WELCH
(c.1904—1986)

Barbara Welch, née Gullick, an honorary member of the society, who died in hospital at
Cheltenham, Gloucestershire in April 1986, was born at Salisbury, Wiltshire in c. 1904. Her
parents were the proprietors of a nursery, and it was possibly this environment that induced an
early interest in plants. On the completion of her formal education, during which time she gained
a B.Sc. degree in geology, she joined the staff of the Salisbury and South Wiltshire Museum as
an Assistant Curator, specializing in archaeology.

In 1921 she joined the Wild Flower Society, and in 1922 came second in her branch with diary
entries for 726 different species noted in the field. During the next decade she played a
prominent part in the activities of the branch, winning first prize in 1928 (the year that she joined
the B.S.B.I.) with 745 diary entries, and again in 1933 with 935 entries; in this year too she won
the southern prize for the ‘five mile radius competition’ with a record of 631 Wiltshire species.

Two plant enthusiasts residing in Salisbury who encouraged Barbara’s floristic studies were the
Hon. Mrs G. Campbell of Wilton Vicarage, with whom in 1932 she wrote her first joint botanical
paper (‘Notes on the flora of the Salisbury district’), and H. J. Goddard, a retired graminologist
formerly employed by a seed company. She also knew Major J. R. G. Gwatkin, an authority on
the flora of Potterne, and E. Marsden-Jones, who in 1923 had founded Potterne Biological
Station. Although most of her field work was undertaken in her native county, she knew Patrick
Hall, one of the leading amateur botanists of the period, and botanized with him in adjacent

462 OBITUARIES

Hampshire, as well as visiting the Bristol area, where she joined Mrs C. I. Sandwith and her son
Noel in examining the rich adventive flora of the dock areas.

No mean artist, she presented in 1936 a series of her own drawings to the Salisbury and South
Wiltshire Museum, and published in The Salisbury Times of 6th November of that year an account
of the various hoards of Roman coins located in S. Wiltshire.

In February 1939 Barbara married Dr F. B. A. Welch, a professional geologist, and in 1940 they
removed to Richmond, Surrey. Despite her move to wartime London, and activities in war work,
she still visited her beloved Wiltshire on every possible occasion, leading field meetings for the
Wiltshire Archaeological and Natural History Society, and searching out new localities for
uncommon plants. She was friendly with J. D. Grose of Swindon and contributed to his Wiltshire
plant notes published annually in the Magazine of the Wiltshire Archaeological and Natural History
Society, becoming co-author of the notes in 1942. Grose’s Flora of Wiltshire was published in 1957;
for this she not only provided many records but also wrote the account of the geology of the
county, prepared biographical summaries of several Wiltshire botanists and assisted greatly in the
preparation of the bibliography. From the early 1940 to the mid 1950s she botanized extensively in
Middlesex, often with me, but frequently alone, gathering critical material and collecting records
for my Flora of the county. Her critical eye, extensive knowledge and boundless enthusiasm were
invaluable, and her contribution to the floristics of Middlesex and my great debt to her are
reflected in the pages of The Historical Flora of Middlesex. Active, efficient and ever willing, she
soon found that her services were sought by, the B.S.B.I., and in 1953 she was appointed an
Assistant Secretary to help with conservation matters and to undertake the minuting of Council
business. In this period too she became a voluntary worker in the Department of Botany, British
Museum (Natural History), spending several days a week in the British herbarium assisting the late
Dr A. Melderis in determining and laying in material to the national collection. With the creation
of the Surrey Flora Committee in January 1957, Barbara accepted the honorary post of Secretary
and Recorder, a duty she discharged until 1963. She immediately channelled her enthusiasm and
energy into collecting and assessing records for the new Flora, and the late J. E. Lousley wrote in
the completed work, published in 1976: “It would be impossible to praise too highly the care,
patience and accuracy she devoted to the plodding spadework of the early stages’’. She contributed
records not only to the Floras of Wiltshire, Middlesex and Surrey, but also to Riddelsdell, Hedley
& Price’s Flora of Gloucestershire (1948), Kent & Lousley’s ‘Handlist of the plants of the London
Area’ (1951-57), A. E. Wade’s Flora of Monmouthshire (1970), S. T. Jermyn’s Flora of Essex
(1974) and R. G. Ellis’s Flowering Plants of Wales (1983). On the retirement of her husband in
1963 they removed to Cheltenham, and from this time, sadly, she virtually gave up her botanical
interests.

Barbara was a remarkable lady, and a brilliant field botanist in the old tradition with a great
ability for teaching. Basically reserved, she was intolerant of fools and on occasion had a very sharp
tongue, and to many appeared stern and unapproachable. Beneath this veneer she was extremely
kind and had a great affection for the B.S.B.I. and its activities, as has been evinced by her very
generous bequest to the Society.

THE BOTANICAL WRITINGS OF B. WELCH

1932

(With CAMPBELL, Mrs G.) Notes on the flora of the Salisbury district. Wilts. Arch. Nat. Hist. Mag., 46: 58-63.
1938

Cyperus longus in Wiltshire. Rep. botl Soc. Exch. Club Br. Isl., 11: 593-595.

South Wilts plant notes, 1933 to 1937. Wilts. Arch. Nat. Hist. Mag., 48: 82-86.
1942

(With Grose, J. D.) Wiltshire plant notes, 4. Wilts. Arch. Nat. Hist. Mag., 50: 71-78.

My five mile radius. Wild FI. Mag., 278: 11-12.
1949
Juniper survey. Lond. Nat. , 28: 36-37.
1952
(With BANGERTER, E. B.) The London Rocket and its allies in the London area. Lond. Nat., 31: 13-17.

OBITUARIES

(With BANGERTER, E. B.) The comfreys of the ae ee Lond. Nat. , 33: 55-58.

(With BANGERTER, E. B.) The alien Polygonums of a are area. Lond. Nat., 34: 60-65.
The herbarium of James Anson of Darlington. ue Ret Soc. Br. Isl., 3: 39-40.
Petersham Common, Surrey. Lond. Nat., 40: mg

Some plants of Ham Common, Surrey, 1941-1961. isa Nat., 41: 23-27.

(With Smitu, J. E.) Plants of Hurst Park Racecourse, Sut. Lond. Nat., 42: 13-13.

Some plants of East Sheen Common. Lond. Nat., 43: 16-18.

463

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. W. Suffolk
. E. Norfolk
. W. Norfolk
. Cambs.

. Beds.
. Hunts.
. Northants.

. E. Gloucs.
. W. Gloucs.
. Mons.

. Herefs.

. Worcs.

. Warks.

H1. S. Kerry

H2. N. Kerry
H3. W. Cork

H4. Mid Cork
HS5. E. Cork

H6. Co. Waterford
H7. S. Tipperary
H8. Co. Limerick
H9. Co. Clare
H10. N. Tipperary
H11. Co. Kilkenny
H12. Co. Wexford
H13. Co. Carlow
H14. Laois

39. Staffs.

40. Salop

41. Glam.

42. Brecs.

43. Rads.

44. Carms.

45. Pembs.

46. Cards.

47. Monts.

48. Merioneth
49. Caerns.

50. Denbs.

51. Flints.

52. Anglesey
53). 9. Lines:
54. N. Lincs.
55. Leics.

55b. Rutland
56. Notts.

57. Derbys.
58. Cheshire
59. S. Lancs.
60. W. Lancs.
61. S.E. Yorks.
62. N.E. Yorks.
63. S.W. Yorks.

64. Mid-W. Yorks.

65. N.W. Yorks.
66. Co. Durham
67. S. Northumb.
68. Cheviot

69. Westmorland
69b. Furness
70. Cumberland
71. Man .

72. Dumfriess.

73. Kirkcudbrights.

74. Wigtowns.
75. Ayrs.

IRELAND

H15. S.E. Galway

H16. W. Galway

H17. N.E. Galway

H18. Offaly
H19. Co. Kildare

H20. Co. Wicklow

H21. Co. Dublin
H22. Meath
H23. Westmeath

H24. Co. Longford
H25. Co. Roscommon

H26. E. Mayo
H27. W. Mayo
H28. Co. Sligo

NAMES OF VICE-COUNTIES IN WATSONIA
ENGLAND, WALES AND SCOTLAND

96

. Renfrews.

. Lanarks.

. Peebless.

. Selkirks.

. Roxburghs.
. Berwicks.

. E. Lothian

. Midlothian
. W. Lothian
. Fife

. Stirlings.

. W. Perth

. Mid Perth

. E. Perth

. Angus

. Kincardines.
. S. Aberdeen
. N. Aberdeen
. Banffs.

. Moray

Easterness

96b. Nairns.

. Westerness

. Main Argyll

. Dunbarton

. Clyde Is.

. Kintyre

. S. Ebudes

. Mid Ebudes

. N. Ebudes

. W. Ross

. E. Ross

. E. Sutherland
. W. Sutherland
. Caithness

. Outer Hebrides
. Orkney

. Shetland

. Co. Leitrim

. Co. Cavan

. Co. Louth

. Co. Monaghan
. Fermanagh

. E. Donegal

. W. Donegal

. Tyrone

. Co. Armagh

. Co. Down

. Co. Antrim

. Co. Londonderry

Collins
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GENERAL EDITOR

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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 are invited.

Manuscripts must be submitted in duplicate, typewritten on one side of the paper only, with wide
margins and double-spaced throughout. They should follow recent issues of Watsonia in all matters
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particularly use of capitals and italics. Only underline where italics are required.

Tables, appendices and captions to figures should be typed on separate sheets and attached at the
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Contributors are strongly advised to consult the editors before submission in any cases of doubt. 25
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‘Papers and Short Notes should be sent to Dr R. J. Gornall, Botany Department, The University,
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Ed. A. J. Wilmott, 1948. 104 pages, 18 plates. Wrappers. £4.25
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Ed. J. E. Lousley, 1951. 128 pages, illustrations and maps. £4.25
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Ed. J. E. Lousley, 1955. 189 pages, 2 plates and 23 text figs. £5.25
5. PROGRESS IN THE STUDY OF THE BRITISH FLORA
Ed. J. E. Lousley, 1957. 128 pages, 4 plates and 9 text figs. £5.25
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Ed. E. Milne-Redhead, 1963. 90 pages. £5.25
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Ed. A. J. Richards, 1978. 213 pages and 31 plates. £40.00
17. THE BIOLOGICAL ASPECTS OF RARE PLANT CONSERVATION
Ed. H. Synge, 1981. 586 pages and numerous text figs. £42.00
18. PLANT LORE STUDIES
Ed. R. Vickery, 1984. 260 pages. Proceedings of a joint conference of BSBI and Folklore Society in
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Ed. Martin Jones, 1986. Proceedings of a joint conference held in 1984.

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