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Douglas H. Kent Memorial Issue

Volume 23 Part 1 February 2000
Editors: M. Briggs, D. L. Kelly, D. R. McKean,
D. A. Pearman, M. N. Sanford, D. A. Simpson

Botanical Society of the British Isles

Patron: Her Majesty Queen Elizabeth the Queen Mother

Applications for membership should be addressed to the Hon. General Secretary,
c/o Department of Botany, The Natural History Museum, Cromwell Road, London,
SW7 5BD, from whom copies of the Society’s Prospectus may be obtained.

Officers for 1999-2000

President, Mrs M. Briggs

President elect, Dr G. Halliday

Vice-Presidents, Dr R. J. Gornall, Mr M. Walpole, Mr D. J. McCosh
Honorary General Secretary, Mr R. G. Ellis

Honorary Treasurer, Mr M. E. Braithwaite

Editors of Watsonia

Papers and Notes, D. L. Kelly, D. R. McKean, M. N. Sanford*, D. A. Simpson
Book Reviews, D. A. Pearman
Obituaries, M. Briggs

*Receiving editor, to whom all MSS should be sent (see inside back cover).

© 2000 Botanical Society of the British Isles
The Society takes no responsibility for the views expressed by authors of Papers,
Notes, Book Reviews or Obituaries.

The cover illustration of Potentilla anglica Laich. (Trailing Tormentil) was drawn
by Rosemary Wise.

Douglas H. Kent Memorial Issue of Watsonia

This issue of Watsonia is dedicated to the memory of D. H. Kent (1920-1998). It is fitting that the
journal of the Botanical Society of the British Isles should commemorate his life and work, for
Duggie served the society with dedication for a period of 50 years.

Many of the papers in this issue are directly related to Duggie’s botanical interests, including his
life-long studies of the flora of Middlesex, his detailed accounts of the spread of Senecio squalidus
in Britain and Ireland and the important reference works he prepared. The latter include British
herbaria (1957), which was later replaced by British and Irish herbaria by D. H. Kent & D. E.
Allen (1984), the B.S.B.I. Handbook covering the Polygonaceae, Docks and knotweeds of the
British Isles, written with J. E. Lousley (1981) and the List of the vascular plants of the British
Isles (1992). We hope that this wide range of papers will be a fitting tribute to the memory of
Duggie Kent. It demonstrates that the subjects he studied are still engaging the attention of British
and Irish botanists, who find that his work provides a firm foundation on which they can build. It
is also encouraging to see how new lines of evidence such as those provided by molecular
techniques are now throwing new light on the subjects he investigated. We are grateful to
Professor C. A. Stace for originally suggesting that we produce this memorial issue, and to the
authors who have contributed to it. The fact that so many were willing and indeed anxious to
contribute papers at relatively short notice says much for the esteem in which Duggie Kent was
held by the botanical community.

C. D. Preston M. N. Sanford
Chairman, B.S.B.I. Publications Committee Receiving Editor, Watsonia

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Watsonia 23: 3—8 (2000) 3

Obituary

DOUGLAS HENRY KENT
(1920-1998)

With the death on 9 September 1998 of Douglas Kent - “Duggie”, or sometimes “Doug”, as he
was known to his friends and to British botanists more generally - the B.S.B.I. has lost the last
surviving member of the team whose devoted efforts largely sustained the Society at a high level
of efficiency throughout its great expansion of the early post-war years, and who were responsible
for putting in hand many of the activities that continue to be among its principal features today.
Though too self-effacing by nature to enjoy the prominence of the pair with whom he formed such
a memorable triumvirate, J. E. Lousley and J. G. Dony, his services to the Society were fully as
great and no less varied while also extending over the even lengthier period of 48 years without a
break.

Duggie was born on 21 March 1920 in south Hammersmith, the only child of Ernest Edmund
and Millie Louise Kent, and from the age of six lived for the rest of his life at 75 Adelaide Road in
the nearby Borough of Ealing. After attending Bordeston Secondary School in Boston Manor,
where he was a contemporary of Charles Hawtrey of ‘Carry On’ fame, he left at fifteen to join the
Gas, Light and Coke Company (subsequently, after nationalisation, to constitute the core of North
Thames Gas) and remained with that for his entire career, latterly heading a section responsible for
mains and pipelines supplies. The fact that his home and work place were both within easy reach
of the libraries and collections at Kew and South Kensington was to be an inestimable boon and
substantially influenced the pattern of his life.

A born collector, Duggie might easily have been lost to botany because of competing youthful
passions for philately and train-spotting. Luckily, however, while those persisted (he left a sizeable
stamp collection and many notebooks on the G.W.R.), they failed to suppress a developing interest
in plants. By the age of nineteen botany had taken so firm a grip that he embarked upon the
amassing of records for his home county of Middlesex, drastically altered by the relentless sprawl
of London since the days of Trimen and Dyer’s Flora of 1869. To that end he set about exploring
the promising habitats that still remained and spent many hours in libraries investigating the work
and lives of the many who had preceded him in that task. At no other time in his life were his
burrowings to take him so extensively into the manuscript domain, in particular the vast Sloane
collection in what was then the British Museum at Bloomsbury.

The outbreak of war in 1939 fortunately provided no serious interruptions to botany. Military
service was ruled out for him (due to ill-health and/or his ‘reserved’ occupation) and, apart from
serving as an air raid warden (no light responsibility during the London blitz), he was free to slip
out of the city in summer as often as he could manage. That was not without its wartime risks, as
several of his botanical contemporaries also discovered. On one occasion his fieldwork
accidentally took him a foot or two inside the perimeter of an army camp; an over-zealous N.C.O.
promptly shouted to a sentry to arrest him and he ended up with that day’s haul emptied out of his
vasculum on to the guard-room floor, to the picturesque bafflement of his captors.

Towards the end of the war his botanical activity took a marked leap forward: he began a
herbarium in 1943 and joined both the London Natural History Society and the Botanical Society
and Exchange Club (as the B.S.B.I. was then still known) in 1944. Around that time he gained his
introduction to Lousley, who was to become his friend and special mentor, and to another future
close friend and collaborator, E. B. Bangerter. He also met and was given much encouragement by
the veteran H. W. Pugsley, the most respected amateur taxonomist of British flowering plants of
his generation. Austere and rather forbidding on first acquaintance, Pugsley was ever on the look-
out for youthful promise and invited his protégé to dinners at his Wimbledon home, occasions
which Duggie still spoke of with awe half a lifetime later.

It was also in that same year, 1944, that marriage to Ivy Minnie Austen brought his one lasting
connection to somewhere other than Middlesex. They had met earlier when Ivy, a nurse, had been
attending to him during a spell in hospital in London. After marriage, Ivy moved to 75 Adelaide
Road and they lived on the upper floor over his parents. Their only child, Michael, was born two

4 OBITUARY D. H. KENT

years later. His wife came from Blean, near Canterbury, and from 1943 vice-county 15 began to
find reflection in his incipient herbarium. On his visits there from 1945 to 1947 he was also able to
give much help to F. Rose, then actively engaged on a projected Flora of Kent, and the two often
made forays together both there and in Middlesex. It must be a matter for regret that the
considerable expertise he acquired as a field botanist was never deployed more widely. Although
holidays were subsequently to take him to several parts of Britain (Devon and Cornwall,
Derbyshire, Shropshire, Yorkshire, the Lake District, mid and north Wales, and in 1971 to
Scotland), those were more or less strictly family affairs, except that plant lists from many of these
destinations found their way into his notebooks. He never once travelled overseas and very rarely
attended a field meeting or conference if it involved a night away from home.

The London N.H.S. was the body with which he principally identified to begin with. Three years
after joining, his first paper, reporting the many records of interest made in Middlesex since 1939,
appeared in its journal. The year after that he was elected Chairman of its large Botanical Section,
and subsequently he agreed as well to act as the Society’s Honorary Curator; he relinquished this
post in 1955 but continued to act as curator until 1961. His most conspicuous involvement, though,
was as co-author with Lousley of an updated list of the vascular plants recorded within the L.N.H.
S.’s official area (20 miles radius from St Paul’s Cathedral), brought out serially between 1951 and
1957 as supplements to The London Naturalist. Increasing absorption into the B.S.B.I.’s affairs,
however, compelled him to resign both his offices in 1955 and thereafter his L.N.H.S. links were
largely confined to continuing to use its journal as the outlet for his steady stream of papers on the
flora of Middlesex or Inner London.

Involvement with the B.S.B.I. followed at first a similar pattern. He was soon reporting in Plant
Notes aliens new to the British list that he and others were then turning up in the grounds of a soya
processing factory at Harefield, and following those up with papers in quick succession on the
Middlesex flora. In 1948 the first list of Local Secretaries and Recorders to appear after the war
showed him as having agreed to act in those two capacities for v.c. 21 (he was still its Recorder at
the end of his life, the longest to have served a vice-county in either capacity to date), and in
March 1950 he was rated an active enough member to be put forward for and elected to Council.

Three years later, at the instance of Lousley, who had meanwhile become Honorary General
Secretary, he was asked to step in as Honorary Assistant Secretary when W. R. Price had to resign
at short notice due to the ill health of his wife. The duties attached to this post were twofold:
dealing with applications for membership, and the much more onerous one of editing the original
Year Book, then about to be published for the fifth and last time. In his editorial to that final issue,
in September 1953, Duggie was able to announce that a twice-yearly Proceedings was taking its
place. The editorship of that now fell to him. The new journal, besides publishing papers, book
reviews and obituaries, absorbed all the matter that had appeared in the Year Book, such as reports
of field and exhibition meetings, and from Watsonia it took over Plant Records, Plant Notes and
Abstracts from Literature. It was soon apparent that his natural meticulousness made him an
effective editor and, while he interpreted that role for the most part responsively, prepared to
publish most of the material that flowed in, he refused to be dictated to by the Publications
Committee about the journal’s contents and all along insisted on his editorial prerogative to accept
what he saw fit. He likewise resisted pressure to change to a firm of printers with a less antiquated
type-face, rightly fearful of the damaging increase in cost that that would involve and personally
attuned to the relaxed way in which Buncle’s operated (which included accepting copy in his
exceptionally readable handwriting).

That twelve years’ spell as editor of, Proceedings had one outstandingly important by-product: it
confirmed his enduring commitment to abstracting. He had displayed a penchant for that as early
as 1950, in a one-off contribution to The London Naturalist. Two years later he proved willing to
take over the burden of the B.S.B.I.’s annual equivalent in Watsonia from A. H. G. Alston’s by
then sagging shoulders. As soon as Abstracts was transferred to Proceedings he took the
opportunity to make the coverage more complete and its appearances more frequent (twice yearly),
convinced that through this service alone the Society would attract considerably more overseas
members. Though a number of others, most notably E. B. Bangerter for a period, assisted him with
contributions, he realised that the bulk of the work had to be done by a single individual if the
coverage was to be other than fragmentary and over-selective.

When he eventually handed over Proceedings to E. F. Greenwood, in 1966, he was fortunately
content to carry on with abstracting. When, only three years later, the Society found it no longer

OBITUARY D. H. KENT 5

possible financially to go on supporting two journals, compelling the absorption of Proceedings by
Watsonia - a move hastened, ironically, by the ballooning of the Abstracts section consequent
upon the freeing of Duggie from his editorial reins - it was universally agreed that this much-
appreciated service that he had built up so assiduously must be preserved come what may. By
switching to a cheaper mode of production, dropping the attempt to cover plant ecology and the
flora of other western European countries, and publishing no more than bare titles in many more
cases, BSBI Abstracts was thereupon turned into a feasible proposition, makings its first
appearance in May 1971 and lasting (so far) for 28 years. This must have done much to allay the
deep disappointment he felt at the abandonment of the journal on which he had laboured so long
and so hard. When it is remembered that for 46 years Abstracts has been the work of essentially
just one person, a person moreover unable to read any language other than English, the sheer
magnitude of his effort and achievement in that direction is little short of stunning. His unrivalled
knowledge of the literature made Duggie the obvious choice to survey progress in British plant
taxonomy for two early Flora Europaea conferences.

It was Duggie’s desire for as complete a coverage as possible in his Abstracts that led him to
begin the weekly trips to the library at Kew, a habit which soon turned him into such a familiar
figure in that institution. There he made many friends and began a long association with R. K.
Brummitt, with whom he discussed matters of common nomenclatural and abstracting interest;
they were co-compilers of Index to European Taxonomic Literature for 1969 and 1970. His work
at Kew proved of such mutual benefit that Duggie was appointed to an Honorary Research
Fellowship at Kew in 1995, an honour very rarely bestowed on an amateur botanist. He also
became equally well known and formed many enduring friendships at The Natural History
Museum, in his later years visiting it once a fortnight and Kew once a week.

Despite his enormous devotion to BSBI Abstracts, this was very far from being his only activity
on behalf of the Society. Until 1965 he combined editing Proceedings with continuing to shoulder
a whole range of unglamorous ‘back-room’ chores of the kind so crucial to the efficient
functioning of any society of this kind and size. These included handling applications for
membership, keeping up the address list and dealing with orders for the Society’s publications and
enquiries about advertising in them, while also serving on Council ex officio and on three of the six
permanent working committees. And, though committees were never his forte and he was seldom
vocal on them, he was always most conscientious in attending their meetings. It should not be
overlooked either that his name featured on the Panel of Referees and Specialists for several
subjects and genera. As a further service he also compiled a card-index of books desired by
members, and he invited enquiries. The very least the Society could do, by way of recognition for
this unparalleled performance, was to elect him to Honorary Membership - and that it duly did at
the A.G.M. of 1966.

Meanwhile, quite apart from a steady flow of papers in journals, he was busily accumulating
material for several more major works. As he would have been the first to recognise, he was an
almost compulsive compiler: he liked nothing so much as hunting down and extracting
innumerable items of information and creating out of the resulting mass a useful volume of
reference. He worked fast because he was content to plough shallowly, his one failing being that
he was over-inclined to take statements on trust. It was the process of accumulation that he
particularly revelled in: arrangement and analysis came very much second and third. When he first
took over responsibility for the membership address lists, he resisted as long as he dared the
called-for regular repeats of the elaborate geographical versions of those lists produced and
published by his two immediate predecessors, in which members were listed under groups of the
vice-counties in which they lived. He did eventually produce one other such list, but never another.

British Herbaria, “an index to the location of herbaria of British vascular plants with
biographical references to their collectors”, was the first of his purely reference works to appear in
print. Embarked on in 1949, initially with A. J. Wilmott as intended co-author, it was published by
the Society in 1958. It filled a long-felt gap in the literature, and its system of assigning each
institution holding one or more collections a distinguishing set of letters, designed so as to
complement those used in Index Herbariorum (whose compilers later acknowledged much
assistance from Duggie), was promptly adopted for standard use in all the Society’s publications.
Together with its considerably larger replacement of 1984, British and Irish Herbaria, which was
compiled jointly with D. E. Allen, this was more complicated than Duggie’s other undertakings in

6 OBITUARY D. H. KENT

requiring much of the information to be collected by postal questionnaire and by the need to
distinguish between a person’s actual herbarium and the presence of merely some sheets of his or
her collecting in another institution. Of necessity, too, both volumes were the fruit of a good deal
of collaboration, but it is no secret that the bulk of the work was his in each case. At the time of the
first survey many museums were disgracefully vague about what they possessed and who had
collected their material. Between the two surveys that situation greatly improved and the mass of
fresh information that consequently mounted up rendered the second edition inevitable.

During the 1950s, while engaged on British Herbaria, Duggie simultaneously gave extensive
assistance to the American botanist S. F. Blake in connection with the latter’s second volume,
covering Western Europe, of Geographical Guide to Floras of the World, published in 1961.
Duggie’s own Index to Botanical Monographs (1967) was similar in character, if much more
narrowly focused: “a guide to monographs and taxonomic papers relating to phanerogams and
vascular cryptogams found growing wild in the British Isles”. Collected together in this are nearly
1,900 references from the year 1800 onwards.

All this time the great mountain of material he had built up on the vascular plants and
bryophytes of Middlesex had been left in limbo, with no apparent prospect of ever achieving print.
Friends at Kew now made its existence known to The Ray Society and as an avowedly historical
Flora the work was at last made publicly available in 1975. Duggie later served on the Council of
The Ray Society, from 1984 until his death. By that time a Supplement to the Historical Flora,
compiled over twenty years with characteristic meticulous attention to detail, had gone to press.
This was expected to be published by The Ray Society in early 2000.

A small flood of further books in which he had a greater or lesser hand then followed. Docks
and Knotweeds of the British Isles, which he co-authored with Lousley as the third of the B.S.B_I.
Handbooks, was the first to come, in 1981. This was originally to be a monograph by Lousley
alone, but it was rescued by Duggie after Lousley died in 1976. Lousley, being well-known as an
expert in Rumex, had largely written the (for him) more difficult accounts of all the other genera
first, which left, somewhat perversely, the account of Rumex for Duggie to compile unaided. A
second edition, being undertaken by J. R. Akeroyd, will benefit from copious notes left by Duggie
for just such a publication.

In later life, especially after retirement in 1980, he had time at last to take on editing work in
further directions. In 1976 the Richmond Publishing Company secured him as one of the six
founding editors of its periodical Natural History Book Reviews, and in each of the first nine
issues - before the publisher changed and the editorship was concentrated in one person only - up
to five and once as many as nine pieces appeared over his name. He also clearly had a hand in
enlisting just as many from botanical friends. Having been engaged by Ebury Press to vet the
nomenclature of W. Keble Martin’s The Concise British Flora in Colour, when that started out in
1965 on its astonishing climb on to the bestseller lists, Duggie was later engaged to revise the text
for a revamped new edition in 1982, after the author’s death. He was to remark ruefully that he
would have profited greatly if he had accepted the offer of a percentage royalty rather than the
lump sum for which he opted! Later, The Ray Society asked him to undertake the much more
formidable task of preparing for the press the monograph by E. S. Edees and A. Newton, Brambles
of the British Isles, a task which involved much painstaking checking of dates and references as
well as two exceptionally arduous proof readings before the book’s appearance in 1988.

As a fitting culmination, the List of Vascular Plants of the British Isles - immediately known to
everyone as the “Kent List” - was produced for the Society and published by it in 1992 after seven
years in preparation. By then it was over three decades since J. E. Dandy’s analogous volume had
appeared and numerous nomenclatural and taxonomic changes had accumulated, many new
hybrids had been detected and a marked increase in the number of established aliens had occurred.
The new list consequently extended to many more taxa than its predecessor. In one sense it was
the stepchild of BSBI Abstracts, for through his work for that Duggie had kept abreast of the
trickle of new names entering the literature and made a regular point of seeking authoritative
advice on whether or not there was a strong enough case for accepting those for B.S.B.I. purposes.
Compilation of the List proceeded in parallel to the preparation of New Flora of the British Isles
(1991) by C. A. Stace, with whom consultations on an agreed common nomenclature and
classification occupied hundreds of pages of correspondence and many days of convivial meetings
over those seven years and thereafter up to his death. The “Kent Lis?’ is merely the tip of a very

OBITUARY D. H. KENT 7

Duggie Kent (1920-1998) with a copy of W. Keble Martin’s The concise British Flora in colour (Photo. c.
1965, courtesy of North Thames Gas).

sizeable iceberg: characteristically, Duggie had compiled, as well as the places and dates of
publication of all the taxa listed, many more synonyms than the strictly limited number included in
the finished work. The product of an immense amount of meticulous library research, all this
information is safely stored on the B.S.B.I. Database at the University of Leicester, where it is kept
updated and is the object of many enquiries from researchers.

In these later years he also received further honours. In 1977 he was the recipient of the H. H.
Bloomer Medal, which The Linnean Society of London awards each year to an amateur naturalist
of particular distinction. His amateur status was in fact so well disguised by the prolific nature and
consistently high standards of his works that distinguished foreign botanists often found it hard to
believe. In 1979, after reputedly two unsuccessful attempts to persuade him to accept nomination
as President, the B.S.B.I. was able to overcome his diffidence to the extent of his agreeing to let
his name go forward for election as one of the Vice-Presidents. And finally in 1993, to the great
pleasure of his many friends, he was awarded the Order of the British Empire and personally
invested with that by Her Majesty The Queen.

This lengthy account of his botanical career must not be allowed to leave the impression that
that was the limit of his interests. Indeed, if he himself was to be believed, botany was not even his
principal one: it came a mere third, he liked to insist, to cats and Wagner, “in that order’. With the
burly frame and ruddy countenance that gave him the appearance of Farmer Giles the prime place
he accorded to cats could not but seem incongruous, but he was undoubtedly heartbroken in his
last year by the demise of his “dear friend and companion Smudger’, following not long after the
death of his beloved Ivy. Watching cricket (especially Middlesex) was another, lifelong passion.
He took a connoisseur’s delight, moreover, in forming collections of certain garden plants. He was
a dedicated member of the British Fuchsia and International Asclepiad Societies and of the
Geraniaceae Group, contributing to their journals, and he compiled a very extensive card-index of
all the Fuchsia cultivars he managed to trace.

8 OBITUARY D. H. KENT

Some of the time and energy he was able to devote to this remarkable medley of interests was
saved by not engaging in some activities that he might have been expected to pursue. He showed
little or no interest in conservation, for example, and, disliking public speaking, rarely gave a
lecture. His herbarium consisted of only about 2500 sheets, mostly from Middlesex in the 1940s
and 1950s, and was scarcely added to in his last 35 years. Living close to his place of employment
also meant he lost minimal leisure to commuting; being a non-driver, his travelling was by public
transport or by being driven by his wife.

A good idea of the way Duggie operated can be gained by a study of his botanical notes, which
were carefully listed by him in “Manuscript 24” and indexed in “Manuscript 25”. There were four
categories: notebooks, manuscripts, folders, and maps of Middlesex. His Notebooks, started in
1941 and ending in 1983, numbered 113 and contained a wide range of data. These included field
notes (principally from Middlesex but also from his family holidays); notes on Middlesex taken
from the literature, other botanists’ manuscripts and herbaria; notes that were to become the
nucleus of some of his papers (e.g. Senecio squalidus notes started in 1950, British herbaria in
1953 and Veronica filiformis in 1954); data supplementary to various papers (e.g. to Handlist of
Plants of the London Area); references to deceased botanists; and the raw data collected for BSBI
Abstracts. Interestingly, up to 1945 he mainly used English names for the plants he listed. One
notebook included the full plant list that he made on his 1971 holiday to Scotland, starting at West
Ealing and finishing in West Ross and Wick. The 24 Manuscripts included many relating to his
Historical Flora of Middlesex, including (no. 1) “Flora of Middlesex” which was intriguingly
“destroyed 6/7/1950”; a catalogue of his botanical library, including the cost of the books he
bought and their origin if second-hand; a list of British natural history journals; and a cumulative
list of the species he found within one mile of 75 Adelaide Road. Somewhat surprisingly, there
were no lists of his own herbarium or publications, but here is the abundant evidence of an
instinctive and methodical collector and collator.

One keen interest of Duggie that never came to fruition in his lifetime was his commitment to
the system of vice-county recording. One will not find mention of grid squares in his Middlesex
flora (or its Supplement), and we doubt that he ever spent much time ‘square-bashing’. He had
intended producing an updated ‘Comital Flora’ well before his paper proposing one at the 1961 B.
S.B.I. conference: notebooks 42—45 (1951) listed common species for which v.c. records were
required. As an example of lengths to which he would go to achieve his goals, and as a comment
on days before ordinary people had much spare cash and before the days of photocopiers, in the
1940s he copied out by hand the entire first and second Supplements to Watson’s Topographical
Botany (1905 & 1929-30), amounting to 217 printed pages. Duggie later became one of the most
active members of the Vice-county Census Catalogue Committee that was set up in 1988, and did
the lion’s share of the initial data-gathering. His efforts should see the light of day very early in the
new millennium.

Duggie’s final years were sadly blighted by ill health and by the death in 1988 of his only child
and in March 1997 by that of his wife. As it turned out, he outlived her by only eighteen months.
The B.S.B.I., The Natural History Museum and Kew were all officially represented at the funeral.
His herbarium has been acquired by The Natural History Museum, while his botanical books and
bound manuscripts have been purchased by Summerfield Books of Brough, Cumbria.

It is often said that no-one is irreplaceable, but also that exceptions prove the rule. Duggie was
such an exception, one of the last of the great amateurs to devote most of their lives to the study of
wild British plants and their literature, and there is sadly no-one in sight to rival his knowledge,
expertise, long memory and dedication. A man of very definite views, he was well aware of his
own limitations, and rather scornful of those who did not know theirs. Many have lost a loyal
friend and colleague, the whole B.S.B.I. an unfailing servant; at least we can be thankful that so
many of his contributions remain for all to admire and use. We shall not see his likes again.

This account would not be complete without extending our deepest sympathy to his two
granddaughters, Shelley and Nicola, to whom we are indebted for some of the key dates and
family information. Others who have made valuable contributions are Mary Briggs, Dick
Brummitt, Rodney Burton, Arthur Chater, Joan and Peter Hall, Keith Hyatt, Ron Payne and
Francis Rose.

D.E. ALLEN & C.A. STACE

Watsonia 23: 9-14 (2000) 9

Botanical publications of Douglas Henry Kent

The following list includes all the publications that we have been able to trace, with the exception
of the many book reviews in Natural History Book Reviews from 1976 onwards. Also excluded are
numerous administrative and editorial notes that have appeared in the BSBI’s publications. When
presenting an equivalent list of J. E. Lousley’s publications, Kent (1977) said that they “span more
than half a century”. In fact they spanned 51 years, exactly the same as did his own works at his
death, although in the latter case there were a few more to follow.

1947
Notes on the flora of Middlesex. London naturalist 26: 56-65.

1948

Middlesex plant records for 1947. London naturalist 27: 38-42.

Plant note - Panicum dichotomiflorum Michx. The Botanical Society and Exchange Club of the British Isles.
Report for 1946-1947 13: 278.

1949

Middlesex plant records for 1948. London naturalist 28: 31-35.

Additions and corrections to the Comital Flora for Middlesex, I. Watsonia 1: 92-101.

John Blackstone, apothecary and botanist (1712-1753). Watsonia 1: 141-148.

Excursions, 1947. September 27th. Rubbish tip, Hanwell, Middlesex. Botanical Society of the British Isles.
Year Book 1949: 45-46.

Plant note - Ipomoea lacunosa L. Watsonia 1: 120.

Plant note - Verbascum phlomoides L. var. albiflorum (Rouy) Wilmott. Watsonia 1: 120.

1950

Tothill Fields, Westminster: a lost botanical area. London naturalist 29: 3-6.
Abstracts from literature relating to botany. London naturalist 29: 20-22.

Notes on the flora of Kensington Gardens and Hyde Park. Watsonia 1: 296-300.
Herniaria glabra L. in Middlesex. London naturalist 29: 6-7.

BOS

The flora of bombed sites in Canterbury. Watsonia 2: 12-17.

Additions and corrections to the Comital Flora for Middlesex, II. Watsonia 2: 78-79.

Plant note - Lysichitum americanum Hult. & St. John. Watsonia 2: 109.

Book review - Wild flowers of chalk and limestone (J. E. Lousley). London naturalist 30: 40.

1951-1957
(with J. E. Lousley) Handlist of the plants of the London Area. London naturalist 30-36, Supplements.

1952
Book review - The study of the distribution of British plants (ed. J. E. Lousley). London naturalist 31: 94.

1953

(editor) Botanical Society of the British Isles. Year Book 1953.

(compiler) Abstracts from literature. Watsonia 2: 308-334.

Book review - Drawings of British plants, part VI (S. Ross-Craig). Watsonia 2: 425-426.

1954

Plant note - Psoralea americana L. Proceedings of the Botanical Society of the British Isles 1: 37.

Plant note - Cerinthe minor var. hispida Turrill. Proceedings of the Botanical Society of the British Isles 1: 39.

Plant note - Cypripedium calceolus L. Proceedings of the Botanical Society of the British Isles 1: 39-40.

Plant note - Cotoneaster horizontalis Decne. Proceedings of the Botanical Society of the British Isles 1: 156.

Plant note - Bromus carinatus Hook. & Arn. Proceedings of the Botanical Society of the British Isles 1: 159-
160.

Book review - The natural history of the City (R. S. R. Fitter & J. E. Lousley). Proceedings of the Botanical
Society of the British Isles 1: 104.

1954-1965
(editor) Proceedings of the Botanical Society of the British Isles 1(1) — 6(2). (22 parts)

10 BIBLIOGRAPHY

1954-1969
(compiler) Abstracts from literature. Proceedings of the Botanical Society of the British Isles 1(1) — 7(4). (28
parts)

1955

The Scottish records of Senecio squalidus. Proceedings of the Botanical Society of the British Isles 1: 312-
6 9 te

Plant note - Genistella sagittalis (L.) Gams. Proceedings of the Botanical Society of the British Isles 1: 320.

Plant note - Elodea callitrichoides (Rich.) Caspary. Proceedings of the Botanical Society of the British Isles 1:
321-322.

Plant note - Egeria densa Planch. Proceedings of the Botanical Society of the British Isles 1: 322.

Plant note - Lagarosiphon major (Ridley) C. E. Moss. Proceedings of the Botanical Society of the British Isles
1: 322-323.

1956

Senecio squalidus L. in the British Isles, 1. Early records (to 1877). Proceedings of the Botanical Society of
the British Isles 2: 115-118.

Plant note - Juncus tenuis var. anthelatus Wieg. Proceedings of the Botanical Society of the British Isles 2: 23.

Plant note - Pulsatilla vulgaris Mill. Proceedings of the Botanical Society of the British Isles 2: 134.

Plant note - Polygonum campanulatum Hook. f. Proceedings of the Botanical Society of the British Isles 2:
eye

1957

(with the assistance of E. B. Bangerter & J. E. Lousley) British Herbaria. Botanical Society of the British
Isles, London.

(with E. B. Bangerter) Veronica filiformis Sm. in the British Isles. Proceedings of the Botanical Society of the
British Isles 2: 197-217.

Senecio squalidus L. in the British Isles, 3. East Anglia. Transactions of the Norfolk and Norwich Naturalists’
Society 18(5): 30-31.

An interesting 19th century hortus siccus. Proceedings of the Botanical Society of the British Isles 2: 365-366.

Plant note - Pulsatilla vulgaris Mill. Proceedings of the Botanical Society of the British Isles 2: 236.

Plant note - Ranunculus ficaria var. aurantiacus Turrill. Proceedings of the Botanical Society of the British
Isles 2: 236.

Plant note - Silene vulgaris subsp. macrocarpa Turrill. Proceedings of the Botanical Society of the British
Isles 2: 236-237.

Plant note - Hypericum canadense L. Proceedings of the Botanical Society of the British Isles 2: 238.

Book review - Flora van Nederland (S. J. van Ooststroom). Proceedings of the Botanical Society of the British
Isles 2: 424-425.

1958

Plant note - Juncus ensifolius Wikstr. Proceedings of the Botanical Society of the British Isles 3: 49.

Plant note - Eleusine africana O’ Byrne. Proceedings of the Botanical Society of the British Isles 3: 50.

Cyperus fuscus in Middlesex. London naturalist 37: 180-181.

Book review - Handlist of the Shropshire flora (L. C. Lloyd & E. M. Rutter). Proceedings of the Botanical
Society of the British Isles 3: 128.

1959

The poisonous thorn-apple. Country life (London) 126: 489.

Plant note - Corrigiola litoralis L. Proceedings of the Botanical Society of the British Isles 3: 283-284.

Plant note - Geranium retrorsum L’ Hérit. ex. DC. Proceedings of the Botanical Society of the British Isles 3:
284.

Book review - Welsh flowering plants, 2nd ed. (H. A. Hyde & A. E. Wade). Proceedings of the Botanical
Society of the British Isles 3: 250-251.

Book review - British trees and shrubs (R. D. Meikle). Proceedings of the Botanical Society of the British Isles
3: 306.

1960

Senecio squalidus L. in the British Isles, 2. The spread from Oxford (1879-1939). Proceedings of the
Botanical Society of the British Isles 3: 375-379.

A contribution to the flora of central London. London naturalist 39: 41-62.

Short note - Bird-seed aliens at Ealing, Middlesex. Proceedings of the Botanical Society of the British Isles 3:
442. .

Plant note - Berberis manipurana Ahrendt. Proceedings of the Botanical Society of the British Isles 4: 41.

BIBLIOGRAPHY 11

Plant note - Lathyrus palustris var pilosus (Cham.) Ledeb. Proceedings of the Botanical Society of the British
Isles 4: 41.
Plant note - Prunus mahaleb. L. Proceedings of the Botanical Society of the British Isles 4: 41-42.

1961
A contribution to the flora of central London: addenda. London naturalist 40: 28.
The flora of Middlesex walls. London naturalist 40: 29-42.

1962

(with E. B. Bangerter) Further notes on Veronica filiformis. Proceedings of the Botanical Society of the British
Isles 4: 384-397.

Additions and corrections to the Flora of Central London. London naturalist 41: 16-22.

Plant note - Acaena ovalifolia Ruiz & Pav. Proceedings of the Botanical Society of the British Isles 4: 415.

1963

Progress in the study of the British flora since World War 2. Webbia 18: 129-150.

Progress in the study of the Irish flora since World War 2. Webbia 18: 303-317.

A new Comital Flora? In Wanstall, P. J., ed. Local Floras, pp. 70-80. Botanical Society of the British Isles,
London.

Senecio squalidus L. in the British Isles, 7. Wales. Nature in Wales 8: 175-178.

Plant note - Crataegus succulenta Schrad. Proceedings of the Botanical Society of the British Isies 5: 25-26.

Plant note - Holcus lanatus x mollis. Proceedings of the Botanical Society of the British Isles 5: 27.

Plant note - Pteris cretica L. Proceedings of the Botanical Society of the British Isles 5: 121.

Field meetings, 1961. October 7th. An alien hunt in north London. Proceedings of the Botanical Society of the
British Isles 5: 99.

Book review - Welsh timber trees, native and introduced, 3rd ed. (H. A. Hyde). Proceedings of the Botanical
Society of the British Isles 5: 104.

Book review - Welsh ferns, 4th ed. (H. A. Hyde & A. E. Wade). Proceedings of the Botanical Society of the
British Isles 5: 192.

Book review - Atlas of the British flora (ed. F. H. Perring & S. M. Walters). Watsonia 5: 396-397.

1964

Senecio squalidus L. in the British Isles, 4. Southern England (1940->). Proceedings of the Botanical Society
of the British Isles 5: 210-213.

Senecio squalidus L. in the British Isles, 5. The Midlands (1940->). Proceedings of the Botanical Society of
the British Isles 5: 214-216.

Senecio squalidus L. in the British Isles, 6. Northern England (1940->). Proceedings of the Botanical Society
of the British Isles 5: 217-219.

Senecio squalidus L. in the British Isles, 9. Ireiand. [rish naturalists’ journal 14: 203-204.

Notes on the flora of churchyard walls in Middlesex. London naturalist 43: 13-16.

Plant note - Senecio. Proceedings of the Botanical Society of the British Isles 5: 232.

Plant note - Elodea ernstiae St John. Proceedings of the Botanical Society of the British Isles 5: 232.

Plant note - Scleranthus perennis subsp. prostratus P. D. Sell. Proceedings of the Botanical Society of the
British Isles 5: 338.

Plant note - Salvia glutinosa L. Proceedings of the Botanical Society of the British Isles 5: 343.

Plant note - Telekia speciosa (Schreb.) Baumg. Proceedings of the Botanical Society of the British Isles 5:
343-344.

Plant note - Galanthus elwesii Hook. Proceedings of the Botanical Society of the British Isles 5: 344.

Plant note - Curtonus paniculatus (Klatt) N. E. Br. Proceedings of the Botanical Society of the British Isles 3:
344.

Book review - A Flora of Cambridgeshire (ed. F. H. Perring et al.). Proceedings of the Botanical Society of
the British Isles 5: 415-416. ;

Book review - North Atlantic biota and their history (ed. A. Love & D. Love). Watsonia 6: 82-83.

1965

(nomenclatural editor) The concise British Flora in colour, by W. K. Martin. Ebury Press & Michael Joseph,
London.

(with E. B. Bangerter) Additional notes on Veronica filiformis. Proceedings of the Botanical Society of the
British Isles 6: 113-118.

Further additions to the Flora of Central London. London naturalist 44: 18-28.

Plant note - Minuartia recurva (All.) Schinz & Thell. Proceedings of the Botanical Society of the British Isles
6: 48-49.

Plant note - Brunnera macrophylla (Adam) I. M. Johnston. Proceedings of the Botanical Society of the British
Isles 6: 50.

12 BIBLIOGRAPHY

Plant note - Senecio squalidus L. [in Northern Ireland]. Proceedings of the Botanical Society of the British
Isles 6: 51-52.

Plant note - Dactylis glomerata subsp. hispanica (Roth) Nym. Proceedings of the Botanical Society of the
British Isles 6: 52.

Plant note - Asplenium trichomanes L. subsp. quadrivalens D. E. Meyer emend. Lovis. Proceedings of the
Botanical Society of the British Isles 6: 121-122.

Book review - Joseph Dalton Hooker (W. B. Turrill). Proceedings of the Botanical Society of the British Isles
6: 197.

Book review - Introduction to botany (H. J. M. Bowen). Proceedings of the Botanical Society of the British
Isles 6: 199-200.

1966

Senecio squalidus L. in the British Isles, 8. The recent spread in Scotland. Glasgow naturalist 18: 407-408.

Book review - Glossary of the British flora (H. Gilbert-Carter). Proceedings of the Botanical Society of the
British Isles 6: 299-300.

Book review - William Turner (facsimiles by The Ray Society). Proceedings of the Botanical Society of the
British Isles 6: 300.

1967

Index to Botanical Monographs. Academic Press, London & New York.

Additions and corrections to the Flora of Central London (3). London naturalist 46: 26-35.

Notes on some Middlesex grasses. London naturalist 46: 36-39.

Buddleja, the Butterfly Bush. Wildlife and the countryside 266: 9-10.

Book review - Huntia, vol. 2. Proceedings of the Botanical Society of the British Isles 6: 416.

Book review - Companion to flowers (D. McClintock). Proceedings of the Botanical Society of the British
Isles 7: 66.

1969

(nomenclatural editor) The concise British Flora in colour, 2nd ed., by W. K. Martin. Ebury Press & Michael
Joseph, London.

Fritillaria meleagris L. in north-west Middlesex. Journal of the Ruislip and District Natural History Society
18: 15-22.

Book review - The Penguin dictionary of British natural history (R. Fitter). Proceedings of the Botanical
Society of the British Isles 7: 619.

Book review - Excursion Flora of the British Isles, 2nd ed. (A. R. Clapham, T. G. Tutin & E. F. Warburg).
Watsonia 7: 107. .

Book review - B-P-H: Botanico-Periodicum-Huntianum (ed. G. H. M. Lawrence et al.). Proceedings of the
Botanical Society of the British Isles 7: 182.

1970
Short note - Spartina glabra Muhl. - in the British isles. Watsonia 8: 163-164.

71
(with M. Kovanda & R. K. Brummitt) Index to European taxonomic literature for 1969. Regnum Vegetabile
80.

1971-1998
(compiler) BSBI Abstracts, 1-28. Botanical Society of the British Isles, London.

1972
Notes on the Scottish flora: observations in West Ross. Transactions of the Botanical Society of Edinburgh 41:
547.

Book review - The flowering plants and ferns of the Isle of Man: an annotated list (D. E. Allen). Watsonia 9:
169.

1974

Progress in the study of the British flora, 1961-1971. Memorias da Sociedade Broteriana 24: 353-375.

(nomenclatural editor) The concise British Flora in colour, 3rd ed., by W. K. Martin. Ebury Press & Michael
Joseph, London.

1975
The Historical Flora of Middlesex. An account of the wild plants found in the Watsonian vice-county 21 from
1548 to the present time. The Ray Society, London.

BIBLIOGRAPHY 13

1977

(with R. K. Brummitt) Index to European taxonomic literature for 1970. Bentham-Moxon Trustees , Kew.
J. Edward Lousley’s published works. Watsonia 11: 315-325.

Rumex X lousleyi hybr. nov. (R. cristatus DC. x R. obtusifolius L.). Watsonia 11: 313-314.

1981

(with J. E. Lousley) Docks and knotweeds of the British Isles. Botanical Society of the British Isles, London.
Notes on Hoya in cultivation, 1. Asklepios 23: 24-28.

Floras great and small. Natural history book reviews 5: 3-15.

British and Irish herbaria [appeal for information]. BSB/ news 27: 13.

Viola riviniana as a garden weed. BSBI news 28: 19.

1982

The native and introduced succulents of the British Isles [1 & 2]. Xerophyte 5: 65-72 & 97-99.
(editor) The New Concise British Flora, by W. K. Martin. Ebury Press & Michael Joseph, London.
(with M. E. Kennedy) Obituary - Thomas Gerald Collett (1911-1982). London naturalist 62: 87.
British and Irish herbaria [appeal for information]. BSB/ news 30: 23.

1983
The native and introduced succulents of the British Isles [3]. Xerophyte 6: 20-24.
Book review - A seventeenth century Flora of Cumbria (E. J. Whittaker). Watsonia 14: 290.

1984
(with D. E. Allen) British and Irish Herbaria. Botanical Society of the British Isles, London.
Two more decorative non-succulent genera. Asklepios 31: 89.

1985
Book review - A concise Flora of Dorset (R. Good, ed. H. J. M. Bowen). Watsonia 15: 413.

1987

(with R. K. Brummitt, P. S. Lusby & R. C. Palmer) The history and nomenclature of Thomas Edmondston’s
endemic Shetland Cerastium. Watsonia 16: 291-297.

Obituary - John Patrick Micklethwait Brenan (1917-1985). Watsonia 16: 211-214.

Obituary - Barbara Welch (c.1904—1986). Watsonia 16: 461-463.

Profile - Clive Anthony Stace. BSBI news 47: 6-7.

Book review - Index of collectors in the Welsh National Herbarium (S. G. Harrison). Watsonia 16: 345-346.

1988

Rubus procerus ‘Himalayan Giant’. Kew Magazine 5: 32-36.

(editor) Brambles of the British Isles, by E. S. Edees & A. Newton. The Ray Society, London.

Book review - Flora of the British Isles, 3rd ed. (A. R. Clapham, T. G. Tutin & D. M. Moore). Watsonia 17:
107-108.

Book review - Biosystematics in the Nordic flora (ed. B. & L. Jonsell). Watsonia 17: 204.

1989

Book review - The natural history of the Chew Valley (K. Allen et al.). Watsonia 17: 372.

Book review - The fauna and flora of Exmoor National Park. A provisional checklist (Exmoor Natural History
Society). Watsonia 17: 372.

Book review - Natural science collections in Scotland (botany, geology, zoology) (ed. H. E. Stace ef al.).
Watsonia 17: 373.

1990
Short note - The Shasta Daisy. Watsonia 18: 89.

199]

In defence of The Hoyan. Asklepios 53: 3-6.

(with E. M. Nic Lughadha) ““Momenclature” - a new taxonomic concept. BSBI news 57: 26.
The year of Yellow Corydalis [Pseudofumaria lutea (L.) Borkh.]. BSBI news 59: 15.

New Zealand Bittercress [Cardamine corymbosa Hook. f.]. BSBI news 59: 49.

Book review - Flora of New Zealand, vol. 4 (C.J. Webb et al.). Watsonia 18: 328-329.

1992

List of vascular plants of the British Isles. Botanical Society of the British Isles, London.
The type of Hoya imperialis Lind]. Asklepios 57: 9.

Pseudofumaria lutea. BSBI news 60: 20-21.

14 BIBLIOGRAPHY

1993
List of vascular plants of the British Isles. Errata. Botanical Society of the British Isles, London.

1994

List of vascular piants of the British Isles. Errata, List 2. Botanical Society of the British Isles, London.
Monsonia brevirostrata in England. Geraniaceae Group news 14: 9-10.

(with D. J. Goyder) Micholitzia obcordata (Asclepiadaceae - Marsdenieae) reinstated. Asklepios 62: 13-19.

1995

List of vascular plants of the British Isles. Errata, List 3. Botanical Society of the British Isles, London.

Book review - Dictionary of British and Irish botanists and horticulturists (R. Desmond with C. Elwood).
Watsonia 20: 307.

1996

List of vascular plants of the British Isles. Errata, List 4. Botanical Society of the British Isles, London.

(with P. Macpherson, J. H. Dickson, R. G. Ellis & C. A. Stace) Plant status nomenclature. BSBI news 72: 13-
16.

1997

List of vascular plants of the British Isles. Supplement I. Botanical Society of the British Isles, London.
(“1996”)

Note - The correct authority for Lesser Chickweed, Stellaria pallida (Caryophyllaceae). Watsonia 21: 364.

1998

List of vascular plants of the British Isles. Errata, List 5. Botanical Society of the British Isles, London.

1999

(with R. Wisskirchen) 1436-1437: Proposals to conserve two names in Persicaria (Polygonaceae). Taxon 48:
829-830.

2000

Supplement to the Historical Flora of Middlesex. Ray Society, London, in press.
D. E. ALLEN & C. A. STACE

Te Se

Watsonia 23: 15-21 (2000) Ls

A selective supplement to British and Irish Herbaria
D. E. ALLEN

Lesney Cottage, Middle Road, Winchester, Hampshire, SO22 5EJ

ABSTRACT

Since the publication in 1984 of British and Irish herbaria by D. H. Kent and D. E. Allen, a much-enlarged
version of Kent’s British herbaria of 1957, a considerable number of additions and amendments have come to
notice. Those rated the more important or of more especial historical interest are listed.

INTRODUCTION

After British and Irish herbaria was published (Kent & Allen 1984) both authors independently
began entering in their respective interleaved copies the additions and amendments that came to
their notice. D. H. Kent went so far as to produce a manuscript list of those he himself had
accumulated down to July 1984, adding a further page at a later date; but the disappointing sale
achieved by the volume led him to assume that the publishing of a further edition was not a
realistic proposition for the foreseeable future. As he preferred anything he compiled to be as
complete as he could make it, he resisted the suggestion of publishing a limited, selective
supplement only, content for his annotated copy to pass into other hands in due course and trusting
that its updating would be continued thereafter.

It nevertheless remains the case that with the passing of the years the information in a reference
work such as British and Irish herbaria cannot help but become less useful and reliable, as
collections are relocated, new ones discovered and known ones subjected to further research. Since
the 1960s, largely at the instigation of the Biology Curators Group, many institutions have been
publishing lists of their holdings of a valuably comprehensive and incisive kind, and hopefully that
fashion has some way to run yet; indeed, only shortly after British and Irish herbaria was
published three further, major examples of that genre (Harrison 1985; Stace et al. 1987; Webb
1988) appeared. To rectify this gradual obsolescence even if only in part would seem to be
worthwhile. The following list of just those changes judged to be the more important or the more
historically interesting has therefore been put together to that end from the two interleaved copies.
It is dedicated to the memory of D.H.K., who devoted so many years to the long-overdue task of
establishing the precise extent of the herbarium resources underpinning the study of the taxonomy
and distribution of the vascular plants of the British Isles, a task with which his name will always
be pre-eminently associated.

CRITERIA FOR INCLUSION
Priority for inclusion in this supplement has been accorded to:

1 larger collections of a non-specialised character (on the assumption that the whereabouts of
collections of critical groups will mostly be known to specialists in those and will be mostly in
the half-dozen or so principal herbaria);

2 collections which have hitherto been wholly or partly anonymous (for those are deprived of
much of their value, at least for historical purposes, as long as the person(s) responsible for
forming them remain(s) unknown);

3 older collections made by those known to have been particularly active as field botanists, if not
necessarily of national prominence;

4 collections of interest by virtue of their association with figures of historical, but not
necessarily botanical, significance - such as W. H. Fox-Talbot, the pioneer of photography.

16 DoE AEE

At the same time the opportunity has been taken to correct errors of a more than merely trivial
character. Regretfully, though, with very few exceptions, the many instances where names or life
dates have been ascertained with greater precision have had to be excluded for space reasons.

ABBREVIATIONS USED

For the initial volume, Kent (1957) devised a set of symbols and abbreviations which were
repeated for British and Irish herbaria but which are undeniably daunting to a novice reader.
These were adopted partly to minimise space, but partly also to distinguish between a person’s
actual herbarium and specimens of his or her collecting (in some cases considerably more
numerous) existing outside that context. The distinction is by no means a simple one to make. A
herbarium may have been formed by, or at least belonged to, someone else before the person with
whose name it has come to be identified: many herbaria were purchased or inherited and then
added to by the new owner, often by extensive exchanges with other collectors. Sometimes only a
fragment survives and whether that once constituted a larger whole regarded by that person as ‘my
herbarium’ can be a matter of fine judgement. In a few cases people formed more than one
herbarium, at different periods of their lives or on moving to a different area or after a first had
been destroyed. The distinction between ‘herbarium’ and ‘other specimens’ may be of little or no
moment for scientific purposes but it matters historically and biographically - and the value of so
many herbaria today is essentially as historical documents.

For the sake of consistency, Kent’s use of * to denote a herbarium identified with a named
individual has been retained in the List of Collectors below. Other symbols and abbreviations used
in the successive volumes, such as *‘D’ (donated) and ‘Pd’ (purchased), have been avoided. A new
one (+) has, however, been introduced in order to indicate names of collectors additional to those
in the analogous list in British and Irish herbaria.

INTER-INSTITUTION TRANSFERS

As schools, smaller museums and now even some universities have no further use for herbaria in
their possession or cannot adequately curate them, collections are increasingly becoming
concentrated in a limited number of larger institutions, more particularly those where research or
teaching in taxonomy is still active. The development of county museums services with centralised
facilities for the conservation of biological material, has been furthering that trend, which is surely
one to be welcomed. It is, however, the greatest cause of change, and is best noticed first and
separately:

transferred transferred

from to from to
AMBL LANC (1988) GGO GLAM (1991)
AST part to BM, part to WARMS (1988) KCK NWH
BATHG BTH KRG LSG
BRIST BRISTM except for ferns (to NMW) LIMFC DBN (1983)
CHEL LIV (1994) except for 18th century NCCE E (1993)

material (to BM)

CHRG LIV (1985) NSS LIV (1986)
CRK DBN (1989); over half had to be OHM OLDM (1995)

destroyed as irrecoverably
insect-damaged

CYN SLBI TMAL SGEL
ESNHS — GLAM (1982) WPP LSR
EXR LIV (1993) except for the Keble

Martin herbarium (to RAMM)

SUPPLEMENT TO BRITISH AND IRISH HERBARIA 17

Among relocations which have taken place within a single museums service that of THO from
Thurso to Wick (Caithness District Council, Library Annexe, Sinclair Terrace) and of BDK from
Baldock to HITCHIN (Museum Resource Centre - new code letters HITN - Bury Mead Road) are
worthy of mention. The herbaria previously held in various towns (but not cities) in Hampshire
have also been brought together at Winchester (Hampshire County Museums Service - new code
letters HCMS - Chilcomb House, Chilcomb Lane).

LIST OF COLLECTORS: ADDITIONS AND CORRECTIONS

7+ ADDISON, William, fl. 1836-52, surgeon of Malvern. *Seen by E. Lees (Naturalist, 1837, 1:
210): untraced.

+ ANDERSONIAN NATURALISTS *GLAM (1880s—1903).

+ ARBROATH HORTICULTURAL AND NATURAL HISTORY ASSOCIATION *DUE 520
(v.c. 90; 1877-90).

+ BACKHOUSE, Edward, 1808-1879. SUN under 650 (N. England) - not *.

BALFOUR, John Hutton, 1808-1884. The 8500 at PTH, though long attributed to him, is the
British Isles portion of a world-wide Edinburgh University teaching collection (1764~1863),
with numerous contributors, donated in 1880 to the Literary and Antiquarian Society of Perth.
The MSE specimens date from 1855—90 and were donated by J. C. Howden pre-1897. TCD
also has 100—200.

BARCLAY, William, 1846-1923. The collection at PTH extends to many hundreds of specimens
and is presumed to be his herbarium.

+ BATEMAN, Rev. John, 1665?—1774. *Untraced, but at least part was in *Edward JACOB. This
v.c. 15 collector is not to be confused with the London physician John BATEMAN, fl. 1663-—
1728, whose * is in PHA.

*BEILBY, M. A., later Mrs AVERY, fl. 1830s, of Moseley, Birmingham. This is the identity of
the “M.A.B.” listed on p. 193. The 400 specimens in BIRM are presumed to be the *
mentioned by E. W. Benson in Analyst, 1837, 6: 294.

BONIFACE, Ronald Archie, 1913-1985. *NMW.

BORRER, William, 1781—1862. Many specimens are or were in *H. Collins (now in *F. H.
Arnold in HSM).

BRIGGS, Thomas Richard Archer, 1836-1894. Hardly any Rubus in K, contrary to statement: c.
100 each of Rubus in BM and CGE instead.

BROMFIELD, Willim Arnold, 1801—1851. TCD 375 (v.cc. 10-12).

BROOME, Christopher Edmund, 1812-1886. * not in E but in BTH (especially 1830s).

BROWNING, F. R., fl. 1939-1970, of Pembury, Kent. * discovered in 1994 in a stable-cum-
pigsty in Swaledale, Yorks.; mostly S. E. England and 1931-1955 (P. J. Cook, B.S.B.I. News,
1994, no. 65: 30).

BUCKNELL, E. This is Edward BUCKELL (no ‘N’), a physician of Romsey, Hants.

BULL, Henry Graves, 1818?—1885. *HFD (incorporating * of his father, Henry BULL, 1816—post
1859).

BURNETT, Percy, 1903-1972. * (12 drawers) in possession of Dr R. L. Gulliver, Carraig Mhor,
Imeravale, Port Ellen, Isle of Islay, PA42 7AL.

‘C’ and ‘E.F.’ This should be ‘E.F.C.” The collection in GL relates to v.cc. 85, 87-89.

CHILDS, Kathleen Amelia, 1880-1952. Though the sheets themselves are unlocalized (and
undated), the localities are given in an accompanying catalogue. All the specimens are from
the Alton area, v.c. 12, and most date from the 1920s.

CLARK, William Andrew, 1911-1983. *BM (v.c. 110).

+ CLEAR, H. William, fl. 1880-1926, Leicester pharmacist. * untraced (“one of the largest. . . in
the Midlands” (Chemist & Druggist, 17 June 1926: 138).

COOKE, Randle Blair, 1880-1973. * in *William Andrew CLARK, q.v.

COTTON, Thomas Atkinson, 1857-1925 and Mrs Charlotte COTTON, née SPENCE, d. 1925.
*BMH.

COULTER, Thomas, 1793-1843. TCD 185 (160 Ireland, mainly Dublin area, 1820-1822; 25
Britain, 1842).

18 D. E. ALLEN

DICK, Robert, 1811—1806 (sic). For *‘500’ read ‘50’ (an album of ferns).

DICKINSON, D., fl. 1875-1902, pharmacist of Hoole, Chester. * (1 vol. out of original 3: 323,
mostly Ripon area, v.c. 64; 1878-1902) in possession of J. M. H. Shaw, 4 Albert St.,
Stapleford, Nottingham, NG9 8DB (B.S.B.I. News, 1995, no. 69: 34-35).

DIXON, Mrs (*formerly at CCR) and Mrs H. DIXON, fl. 1832-1850, are the same person. She
was Anne DIXON, née AUSTEN (a cousin of Jane Austen, the novelist), 1799-1864, wife of
the Rev. Henry Dixon, vicar of Ferring, Sussex. She was an intimate from childhood of Mrs
Anna ATKINS, née CHILDREN, 1797-1871, with whom she is known to have collected and
whose * (in BM) may thus include specimens of hers.

DORRINGTON, John, 1814?-post 1861. Delete entry.

DOWNES, Rev. John, Downes, 1810-1880. *NTN 800 (10 vols., 220 v.c. 29; 1829-1832). Listed
in Kent (1957:33) as that of an unknown collector and in Kent & Allen (1984) as that of John
DORRINGTON, this * has been identified as almost certainly that of the ‘J. DOWNES’ of the
two entries on p. 131 (Crompton 1997).

+ DRUMMOND, James Lawson, 1783-1853. GL 250 (mainly Belfast area).

+ DUNLOP, Miss, of Biggar, Lanarkshire. E 300 (mostly v.c. 77; 1890s).

EDEES, Eric Smoothy, 1902-1993. *(part: Rubus) NMW 7500. *RILSTONE (Rubus).

EHRET, Georg Dionysius, 1708—1770. *Formerly CHEL now LIV 26 (unlocalized; c. 1730).

ELDRIDGE, Ellen, fl. 1859-1868. *DZS 300 (Salisbury area).

+ EMSLEY, F. *TCD 215 (mostly Yorks.).

+ FORBES, Nanna A. B. *INV 300—400 (1 vol., v.cc. 105 & 106; pre-1914).

FORSTER, Edward, 1765-1849. Another “very large” * said to be his was auctioned by Stevens
in Dec. 1861 (Phytologist, 1861, 6 (N.S.): 16).

FOX, Brian William (d. 1999). *BON 700+ (Bolton area; adventive spp. 100+; 1948-78).

+ FRANCIS, W. QMC (Kent, Essex, Aberdeenshire; 1830s).

GIBSON, Jabez Marriage. Correct dates are 1822-1877.

GIBSON, Samuel, 1790°?-1849. Part of * (700 Pteridophyta) still in SALFM.

GORDON, Rev. George, 1801-1893. *ELN 101 (pre-1894) believed to be his at least in part.

+ GORDON, George Hamilton, Earl of Aberdeen, 1784-1860. *ABD (5 vols.; v.cc. 92 & 93;
1817).

+ GORRIE, David. *1394, donated to Alloa Society of Natural Science and Archaeology in 1875;
Society’s museum was cleared out in the 1940s and * presumed lost.

+GOUGH, Rev. William Edgar, 1864—c. 1930. * formerly at Adcote School, near Shrewsbury,
now CGE (mostly collected by Rev. Thomas STEPHENSON, 1855-1948, includes many B.E.
C. sheets).

+ GRAHAM, George John, 1803-1878, of Enfield, Middx. *formed with his friend John Stuart
Mill, later destroyed (J. Britten, Journal of Botany, 1905, 43: 318).

+ GRENFELL, Adrian Leonard, 1939-1991. *BRISTM 2000 (Bristol area; aliens).

GRIMSTON, Lady Mary Augusta Frederica, Countess of Radnor, d. 1879. The * attributed to her
in BM is most unlikely to be hers, as most specimens date from 1841 - the year in which her
first child was born (J. G. Dony, in litt., 1985).

HALL, Patrick Martin, 1894-1941. Hampshire component of * only on loan to BM pending
transfer to HCMS.

HEMSTED, Rev. John. Dates should be 1747?-1824. Delete ‘1711-1801’, which apparently
refers in part to * Joseph ANDREWS, which was once in his possession and was subsequently
donated by Hemsted descendants to BM. BM holds only a set of unlocalized Mentha
specimens collected by him (attached to the 10 vois. of *“ANDREWS). Specimens of his from
v.cc. 26 and 29 are also in * James SOWERBY in BM (Oswald 1991).

+ HENDERSON, Alistair A. R., 1937-1988. *GLAM 1300+ (Scotland; 1948—),

HILL, Sir John, 1714-1775. * acquired by the de Jussieu family, donated in recent years to P.

HOPE, John, 1725-1786. Despite the disappearance of his * its MS catalogue (1768—1780s)
survives in E.

HOWITT, Richard Crewdson Leaver, 1911-1984 and Mrs Brenda HOWITT, née CHALK, d.
1981. *CGE (v.c. 56; Salix, Potamogeton).

HUTCHINSON, Rev. Thomas, 1815-1903, nephew of Wordsworth. * awarded ‘Best Herbarium’
prize of Herefordshire Horticultural Society, 1854; not traced.

SUPPLEMENT TO BRITISH AND IRISH HERBARIA 19

I ANSON. This should be JANSON.

JAMES, Robert, LORD PETRE, 1713-1742. * (16 vols., incl. 2 of British plants) sold 1886, later
acquired by Sutro Library, San Francisco, U.S.A. Most volumes annotated by Sir Jonn Hill
(McLean 1984).

KENNETH, Archibald Graham, 1915-1989. *E (v.c. 101).

KIRBY, Rev. H., fl. 1830-1870, of Mayfield, Sussex. * (large folios; mostly c. 1840) donated in
1904 to Eastbourne Technical Institute (Journal of Botany, 1905, 43: 72). Presumably the same
person as the Rev. F. KIRBY, whose * at EBE was destroyed by enemy action in 1940.

+ LATHAM, Brian. ABDAM 800 (v.cc. 82—84, 92, 93; 1954-1957).

LEE, Sarah. Error for LEE, Susan, fl. 1850-1871. The specimens are some mere scraps from the
wall of a church, contained in a packet attached to the 1841 collection misattributed to Lady
Mary GRIMSTON, q.v. (J. G. Dony, in litt., 1985). Delete entry.

LEIGHTON, Rev. William Allport, 1805-1889. SHY 270 (v.c. 40; pre-1835).

+ LIBBEY, Richard Pearse, 1911-1987. LTR 8,000 (v.cc. 27 & 28).

MacGILLIVRAY, William, 1796-1852. The collection in ABD extends to a few thousand
specimens and thus appears to rank as *.

MADDEN, Edward, 1805-1856. Brought up in Ireland, in India 1830-1849 and a known collector
with an untraced *, possibly the former of an anonymous two-volume collection of the 1820s
acquired by SKT from a Stockport bookshop and subsequently transferred to MANCH. The
specimens are mainly from v.c. 55B (but the handwriting was unknown to the late K. G.
Messenger) and H11.

MAHER, D., fl. 1847. Believed to be a misreading of ‘Dr. MATEER’, i.e. the Dr William Mateer,
fl. 1831-1850, listed on p. 201, a contributor to the exchanges of the Botanical Society of
London in 1847-1850.

MATHEWS, William, 1828-1901. K has main part of * exclusive of v.cc. 36 & 37 specimens
(which are the 5,000 in WOS).

MOSELEY, Harriet, fl. 1836-1867. * mostly 1830s, but one specimen 1799. The 9 vols. include
material from many other collectors.

MUIRHEAD, Clara Winsome, 1915-1985. * (main) PLYP.

MURPHY, Rose J. *BM (v.cc. 1 & 2).

+ PEIRSON, Archdeacon, d. 1805/6, of York. * (1,745 British specimens) raffled in York (Sir
Thomas Frankland in litt. to J. E. Smith, 4 Feb. 1806, Smith correspondence, Linnean Society);
not traced.

+ PENNANT, Thomas, 1726-1798. * (mostly unlocalized) donated to CGE by Earl of Denbigh in
1912 (Sherborn 1940) but cannot now be traced.

mEOLEITT, A., fl. 1920. LIV 500 (v.c. 58).

+ PRATT, Anne, afterwards Mrs Pearless, 1806-1893. * (“extensive”’; 1820s—1830s; v.cc. 15 & 16
(Woodward 1896)); not traced.

PURCHAS, Rev. William Henry, 1823-1903. Many sheets (of many genera) in BMH, none of
them from exchange clubs: possibly a separate *.

RAYNER, John Frederick, 1854-1947. Only a few sheets survive of the once-extensive * in
BMH. The rest are believed to have been discarded in the 1950s.

REID, William, d. 1852. This Montrose doctor apparently collected in v.c. 90 exclusively. The 80
sheets in KRDY (v.cc. 83 & 85) were collected in 1851-1858 by a ‘Mr. Reid’ of Edinburgh,
clearly a different person.

REYNELL, Elizabeth T., fl. 1864, of Westmeath (not ‘Meath’). *TCD 166.

ROBERTS, John, of Bangor. Correct dates: 1792—1849. Apparently had 2 *, one formed from _ c.
1820, a second seen by J. E. Griffith (Jones 1998: 62). Both untraced.

ROBINSON, Gilbert Wooding, d. 1942. *formerly CHEL, now LIV; incorporates one or more
older collections, many specimens being from Botanical Society of London members and
dating back to the 1830s (see Edmondson 1994).

ROBSON, Stephen, 1741-1779. * not in SUN, as stated, but still in possession of a descendant, E.
Robson, Market Weighton, Yorks. (Davis 1981: 72; Horsman & Davis 1990: 77).

ROGERS, Rev. William Moyle, 1835-1920. * (part, except Rubus) - presumably that formerly in
THE - now in LANC.

20 D.E. ALLEN

SALMON, John Drew, 1802-1859. The * part in NWH was bequeathed, at which time it totalled
c. 2,900 (mostly Thetford area; 1830s) according to the museum’s records. Either ‘1400’ is a
miscount or much has been lost. The * part auctioned by Stevens in 1861 comprised his main
British collection minus “the Surrey folios” and totalled c. 1000 sheets (Phytologist, 1861, 6
(N.S.): 350). It must be some of these latter which NMW (via *SHOOLBRED) and WARMS
(via WAR) possess.

SANDYS, Rev. George William, 18127-1848. * (Stroud area) seen by Riddelsdell (Riddelsdell et
al. 1948: cxxi) at a private house somewhere in Gloucestershire. “H.C. WATSON presumably
contains merely duplicates from Botanical Society of London exchanges.

+ SELBY, Prideaux John, 1788-1867. A ‘dry plant book’, item 707 in the MS catalogue of his
library, may have been his * (Jackson 1992: 32); if so, not traced.

SHAW, Rev. Charles Edward, 1910-1994. *’dispersed among friends on moving house” (Hartley
1994: 28); but 63 in OLDM (“Alien flora of Oldham’, 1940-1960).

+ SIMONYYI, Erno, fl. 1853-1868, of Hungary. BP 300 (1853-1859; England and Scotland).

SMITH, Sir James Edward, 1759-1828. Also a * formerly LIVB, now LIV (c. 10,000 specimens -
fuller details in Edmondson 1985).

SMITH, William, d. pre-1836. * (Aberdeenshire?) mentioned by Murray (1836: 10, 129), his
cousin.

SPALDING GENTLEMEN’S SOCIETY. The * in SGGS listed on p. 67 is a 19th century one.
Gough & Nichols (1812: 9, f 2) record that the Society possessed a “fine hortus siccus”’ c.
1745.

STOVIN, (not ‘Mrs’) Margaret, 1756-1846. Catalogue of * in MDH published by Simmons
(1993).

SWANN, Eric Lister, 1904-1989. *NWH (v.c. 28; Carex, Gramineae).

TALBOT, William Henry Fox, 1800-1877. * (“‘large”; partly formed by his mother, Lady
Elisabeth Theresa FOX-STRANGWAYS, afterwards TALBOT, afterwards FEILING, 1773-
1846; includes sheets from a purchased collection) in Talbot Museum, Lacock, Wilts.

THOMAS, Anna. More precise dates: fl. 1849-1862. Of Bristol (*H. C. WATSON). Must thus be
the Mrs Thomas, of Bristol, listed on p. 257.

+ THOMSON, fl. 1797. “Mr Thomson continues to collect every plant near Penzance” (Davies
Giddy in litt. to William Withering, 19 Nov. 1797 (Mann 1986: 121); * untraced.

THRELKELD, Caleb, 1676-1728. *TCD 22 (only 2 localized). An anonymous collection
attributable to him on internal evidence (Doogue & Parnell 1992).

TRAILL, James William Helenus, 1851-1919. Contrary to the total of ‘304’ stated, Stace et al.
(1987) say “many thousands” of his British flowering plants are in ABD.

TRIST, Philip John Owen, 1908—1996. *CGE 3,275 (mainly British Gramineae).

VIGURS, Chambré Corker, 1867—1940. “THURSTON (in K) incorporates a second * of his.

WALLACE, Edward Charles, 1909-1986. *RNG 22,855 inclusive of foreign (v.cc.. 13-17, 37,
72, 80-88, 99, 100).

WEBB, David Allardice, 1912—1994. TCD 2,165 (Ireland, 1950-5).

WESTRUP, Alick William, 1910—1994. * formerly HIWNT, now HCMS.

WHITE, D. Presumed to be the Montrose surgeon of that name whose records are cited by the
Aberdeen physician Alexander Murray (1836). Specimens of White’s are in *E. H.
ROBERTSON, a fellow Angus collector; Robertson was not born until 1867, so White’s may
be among the specimens in that dating back to 1789 (see p. 231).

WHITEHEAD, John, 1833-1896. OLDM 100 (v.c. 59).

WILLIAMSON, William Crawford, 1816-1895. * formed in youth (E. Yorks.) (Williamson 1896:
33); not traced.

+ WILSON, Thomas Watkins, fl. 1820s — 1830s, physician of Shrewsbury. SHY 331 (200 v.c. 40,
131 Scotland).

WITHERS, Robert, 1814?-1855. Donated to TTN in 1852 “small collection of the rarer plants of
the Bath area” (Proc. Somerset Arch. & Nat. Hist. Soc. 1852, 3: 12). Survives in TTN
unidentified?

WRIGHT, Charles H., fl. 1832-1844, and his daughter. TCD 245 (v.c. 70; mostly 1839-1843).

+ YOUDALE, W.‘H., fl. 1890-1908. *LIV 314 (v.c. 70; 1891-1908) (Hofmann et al. 1990).

SUPPLEMENT TO BRITISH AND IRISH HERBARIA 21
ACKNOWLEDGMENTS

Considerably more authors and correspondents have been responsible for providing information
drawn on for this supplement than it has been feasible to acknowledge individually in the text. I
would like to take this opportunity of thanking them for their contributions and of assuring them
that everything else they have published or reported about particular collections in addition to what
appears above has been carefully noted and entered up for future reference. I would also like to
thank Dr J. R. Edmondson for reading through the paper in manuscript and commenting helpfully.

REFERENCES

CROMPTON, G. (1997). The *J.D.” herbarium at Northampton Museum. Watsonia 21: 362-364.

DAVIS, P. (1981). Stephen Robson’s hortus siccus. Naturalist 106: 67—73.

DOooGugE, D. A. & PARNELL, J. (1992). Fragments of an eighteenth century herbarium, possibly that of Caleb
Threlkeld, in Trinity College, Dublin (TCD). Glasra 1: 99-109.

EDMONDSON, J. R. (1985). The other Smith herbarium. Linnean 1(4): 17-19.

EDMONDSON, J. R. (1994). The G. W. Robinson herbarium: newly acquired by Liverpool Museum. B.S.B./.
news: 65: 29.

GOUGH, R. & NICHOLS, J. (1812). Some account of the Gentlemen’s Society at Spalding, in NICHOLS, J., ed.
Literary anecdotes of the eighteenth century 6: 1-162. Privately published, London.

HARRISON, S. G. (1985). Index of collectors in the Welsh National Herbarium. National Museum of Wales,
Cardiff.

HARTLEY, M. M. (1994). West Yorkshire herbaria, in LAVIN, J. C. & WILMORE, G. T. D., eds. The West
Yorkshire plant atlas, pp. 26—28. City of Bradford Metropolitan Council, Bradford.

HOFMANN, H., EDMONDSON, J. R. & HALLIDAY, G. (1990). The Cumbrian herbarium of W. H. Youdale.
Watsonia 18: 204-206.

HORSMAN, F. & DAVIS, P. (1990). Edward Robson’s list of the rarer plants of County Durham. Archives of
natural history 17: 67-77.

JACKSON, C. E. (1992). Prideaux John Selby, a gentleman naturalist. Spredden Press, Stocksfield.

JONES, D. (1998). The doctor and the guide: two Snowdonian plant hunters. Transactions of the
Caernarvonshire Historical Society, 1998: 55-75.

KENT, D. H. (1957). British herbaria. Botanical Society of the British Isles, London.

KENT, D. H. & ALLEN, D. E. (1984). British and Irish herbaria. Botanical Society of the British Isles,
London.

MCLEAN, E. P. (1984). A preliminary report on the 18th century herbarium of Robert James, Eighth Baron
Petre. Bartonia 50: 36-39.

MANN, R. D. (1986). William Withering and the foxglove. MTP Press, Lancaster.

MuRRAY, A. (1836). The northern flora; or a description of the wild plants belonging to the north and east of
Scotland. Part 1. Edinburgh.

OSWALD, P. H. (1991). The Revd John Hemsted (1747?-1824). Nature in Cambridgeshire 33: 26-29.

RIDDELSDELL, H. J., HEDLEY, G. W. & PRICE, W. R. (1948). Flora of Gloucestershire. Buncle, Arbroath.

SHERBORN, C. D. (1940). Where is the ? collection. Cambridge University Press, Cambridge.

SIMMONS, M. (1993). A catalogue of the herbarium of the British flora collected by Margaret Stovin (1756—
1846). Dorman Museum, Middlesborough.

STACE, H. E., PETTITT, C. W. A. & WATERSTON, C. D. (1987). Natural science collections in Scotland.
National Museums of Scotland, Edinburgh.

WEBB, D. A. (1988). The Irish and British plants in the herbarium of Trinity College, Dublin. 1. The vascular
plants. Jrish naturalists’ journal 22: 482-485.

WILLIAMSON, W. C., ed. WILLIAMSON, A. C. (1896). Reminiscences of a Yorkshire naturalist. G. Redway,
London.

WOODWARD, B. B. Anne Pratt, in LEE, S., ed. Dictionary of national biography 46: 284—285. Smith, Elder &
Co., London.

(Accepted April 1999)

fimt at

i

Watsonia 23: 23-38 (2000) 23

The herbarium of William Skrimshire (1766-1829) of Wisbech
G. CROMPTON

103 Commercial End, Swaffham Bulbeck, Cambridge, CB5 OND

and
E. C. NELSON*

c/o Wisbech & Fenland Museum, Museum Square, Wisbech PE13 1ES, Cambridgeshire

ABSTRACT

William Skrimshire (1766-1829), a surgeon of Wisbech, Cambridgeshire, assembled a substantial herbarium
before 1818 when he sold it to Viscount Milton. Subsequently Skrimshire went on collecting plants in the
vicinity of Wisbech and adjacent counties. He was in contact with other botanists, most notably the Revd
Richard Relhan, author of Flora Cantabrigiensis, and obtained specimens from them. Other members of the
Skrimshire family, especially his brother Dr Fenwick Skrimshire, also contributed to his herbarium. While a
manuscript catalogue of William Skrimshire’s extensive herbarium survives, providing valuable records of the
flora of Cambridgeshire, only about 200 specimens from his herbarium can now be traced; these are preserved
in Wisbech and Fenland Museum (WBCH) and some constitute hitherto unpublished first records of
Cambridgeshire and Northamptonshire species. The extant specimens show that William Skrimshire was an
inquisitive botanist and observant gardener. They also point to changes in the environment of Wisbech and its
hinterland during the past two centuries. Some significant specimens from the herbarium of Skrimshire’s one-
time pupil and fellow surgeon, John Rose Weatherhead (c. 1787-1849), are also noted.

KEYWORDS: Cambridgeshire, Norfolk, Northamptonshire, history, John Rose Weatherhead.

INTRODUCTION

The Wisbech Museum (now Wisbech and Fenland Museum) was founded in 1835, and today still
occupies the same building, in the town centre, that was erected in 1846 (Arthur 1985) to house the
collections. Among the earliest donations were numerous natural history specimens, including
several collections of pressed and dried plants which had been made mainly in the vicinity of the
town by local amateur naturalists; the donors were Richard Dykes Alexander and Miss Alexander,
J. R. Weatherhead, Mrs Algernon Peckover and Charles Metcalfe. In 1862, Mrs W. G. Townley
and Mrs J. R. Weatherhead donated two substantial herbaria; the Weatherhead hortus siccus
comprising seven volumes contained few specimens collected later than 1835. While the subject of
our paper, the Wisbech surgeon William Skrimshire, had died six years before the Museum was
established, specimens that he had gathered in the counties of Norfolk, Cambridgeshire,
Northamptonshire and Lincolnshire (v. cc. 27, 28, 29, 32, 53 respectively) during the 1790s and
the first three decades of the 1800s, came to Wisbech Museum because they were contained in
those separate herbaria.

The Wisbech Museum herbarium (WCBH: Kent & Allen 1984) has not been added to for many
decades, so its principal interest is as an historic collection focused on the flora of Wisbech town
and hinterland during the first half of the 19th century. For other reasons, many of the extant
specimens are by far the earliest surviving specimens known to have been collected in
Cambridgeshire (v. c. 29) and Northamptonshire (v. c. 32), and a smaller number are first records
for West Norfolk (v. c. 28; these were noted by Petch & Swann (1968: 15)). With regard to

*Address for correspondence: E-mail: tippitwitchet @ zetnet.co.uk

24 G. CROMPTON AND E. C. NELSON

Cambridgeshire (which is the special interest of G.C.), while the foundations of the herbarium in
the Department of Plant Sciences at the University of Cambridge (CGE) were made by Professor
John Henslow from 1818 onwards, Henslow’s first specimens from the Wisbech area are dated
1831. The Wisbech Museum collections therefore are of considerable interest because they predate
most of the material at Cambridge and so provide some voucher specimens for the first record of
species from Cambridgeshire.

Information about the herbarium in Wisbech Museum was incorporated in D. H. Kent & D. E.
Allen’s British and Irish herbaria (1984: 71); they employed the acronym WBCH. The Museum
Documentation Association’s acronym for the Wisbech and Fenland Museum is WISFM, and all
the herbarium specimens, which are presently being catalogued by E.C.N., now bear accession
numbers prefaced by that acronym (e.g. WISFM: 1862.17.2.86). A fully annotated list of the
specimens discussed in this paper can be obtained from The Curator, Wisbech and Fenland
Museum; this list includes transcriptions of annotations on the specimens and the original
nomenclature. In the following account all botanical names have been updated according to Stace
(1991). When the botanical names inscribed on a herbarium specimen differ from present-day
nomenclature the names in manuscript are quoted in brackets, e.g. Helianthemum nummularium
(as “Cistus Helianthemum’’).

WILLIAM SKRIMSHIRE’S SPECIMENS AND MANUSCRIPTS

We are able to record and assess the extent of William Skrimshire’s botanical interests and
expertise by examining his reliquiae, which comprise three separate items preserved in Wisbech
and Fenland Museum, namely:

| His own manuscript catalogue of his herbarium; this was first described by Perring (1956;
see also Bridson et al. 1980 (entry 435.1); Crompton 1994, 1996; Petch & Swann 1968).

2 An assortment of approximately 200 specimens, mainly representing higher plants, of
miscellaneous provenance.

3. An hortus siccus of bryophytes (see Bridson et al. 1980) (this is not discussed here).

It should be noted that Petch & Swann’s (1968) reference to “a large collection” of William
Skrimshire’s specimens in Wisbech Museum is inaccurate; Skrimshire’s material represents a very
small fraction of the Museum’s herbarium (E. C. Nelson, unpublished data).

There are also some specimens incorrectly ascribed to William Skrimshire in Norwich Museum
(NWH: see Kent & Allen 1984: 60, 244). The identity of the collector of this series of 300
specimens, all collected in 1801, is not known; despite an annotation in the Museum’s register
(23.91), William Skrimshire was not the person who assembled it, although a few specimens came
from his garden (see below).

THE SKRIMSHIRE FAMILY

William Skrimshire (1766-1829) was the oldest son of William Skrimshire (1739-1814), and he
had three brothers, Thomas, Fenwick, and George, and a sister, Elizabeth. Both Williams, father
and son, were surgeons in Wisbech, and until his father’s death William junior probably lived
with, and practised with, his father in Ship Lane. Later he moved to the outskirts of the town, to
Bushy Place — while the property’s exact location has not been discovered, Bushy Place was
almost certainly situated on the eastern bank of the River Nene, less than a mile north-north-east of
the centre of Wisbech near Horseshoe Corner. There William had a garden, including a
“peppermint plantation” where he evidently grew peppermint (Mentha xX piperita) in some
quantity presumably for distillation for medicinal use. William Skrimshire never married; he died
in Wisbech on 22 July 1829.

Fenwick Skrimshire, born at Wisbech in 1775, took his medical degree at Edinburgh, and
dedicated his thesis, De assimilatione (F. Skrimshire 1798), to his father and to his brother
William. During 1797-1798 Fenwick was President of the Natural History Society of Edinburgh

WILLIAM SKRIMSHIRE (1766-1829) 25

(this society was established in 1782, and survived until 1812 when it amalgamated with the Royal
Physical Society). During this period, Fenwick sent his brother many plant specimens from
Scotland, including Chrysosplenium alternifolium (“From Dickson’s Garden, Edinburgh ...”),
Gnaphalium sylvaticum (“Given to my Brother Fenwick by his friend Mr. Brown. 1796.”), a wild
pansy from Ben Lawers that had been named “Viola montana” by John Mackay (1772-1802), and
Campanula rapunculoides (“A very good discovery this, and as such Dr. Smith has noticed it in
his Mem. A doubt arises with me whether it be not a reyectamentum Horti, as it is cultivated in
Gardens. You will do well to be quite certain about this, as otherwise it cannot well be admitted as
a native of our Island. ... Discovered by my Brother Fenwick Skrimshire, M. D. From Blair,
Scotland. 1796.”). Dr Fenwick Skrimshire settled in Peterborough where he was physician to the
Fitzwilliam family (Fenland notes & queries 4: 211, 212 with copy from Stamford Mercury 5
November 1819), and by 1838 he was “Physician to the Peterborough Public Dispensary and
Infirmary”. He attended the poet John Clare (1793-1864), who was a native of Helpstone, a small
village between Peterborough and Stamford whence came a number of the specimens collected by
William Skrimshire (see below). In 1841 Fenwick Skrimshire filled in the application for Clare’s
admission to Northampton Lunatic Asylum, and on 29 December 1841 he certified Clare
“following years addicted to poetical prosings.” Fenwick Skrimshire wrote a number of books (see
References) including A series of essays introductory to the study of natural history, the eleventh
essay in this was entitled “The useful application of botanical knowledge” (F. Skrimshire 1805:
vol. 2, pp. 125-181). He died in 1855.

Thomas, whose date of birth has not been discovered, was admitted a pensioner at Clare
College, Cambridge, on 22 December 1791, and matriculated at Michaelmas 1793. Subsequently
he “migrated” to Magdalene College under Holmes Charity, and graduated in 1798. He was later
ordained and held a number of livings in Norfolk between 1800 and his death in 1836. While not
himself a contributor to his brother’s herbarium, Thomas’ son George, who was apprenticed for
five years (from 2 July 1817 to 31 March 1823; see Crompton 1994) to his uncle William
Skrimshire, did; a specimen of Aceras anthropophorum “From Helpstone, near Milton.
Northamptonshire. Given to me by my Nephew G. Skrimshire. 1821” is a first record for that
county (v. c. 32; G. Gent pers. comm.; see below).

The fourth brother, George, a subscriber to one of Fenwick’s books, was a farmer at Kettlestone,
Norfolk. Their only sister, Elizabeth, married Charles Metcalfe, a solicitor, and their daughter “my
niece Eliz. Metcalfe” sent specimens to William Skrimshire shortly before her early death in 1824
aged 22. Elizabeth Metcalfe’s specimens include Epilobium parviflorum and E. tetragonum from
Emneth, a village near Wisbech, an unlocalized specimen of Helianthemum nummularium (as
“Cistus Helianthemum’’), and Minuartia verna (as “Arenaria verna”’) and Parnassia palustris both
from Derbyshire (v. c. 57). As noted above, Charles Metcalfe was one of the early benefactors of
Wisbech Museum, presenting his brother-in-law’s collection of mosses.

William, Thomas and Fenwick Skrimshire were keen entomologists. William and Thomas were
elected Fellows of the Entomological Society, and both published papers in the first volume of the
society's Transactions (see References). While Thomas Skrimshire “purpose[d] doing myself the
honour of transmitting the Society a letter annually upon this subject [rearing insects]”, we have
traced only one published letter from him (T. Skrimshire 1812). The large copper butterfly, a
native of the “marshes of Cambridgeshire”, was discovered by Adrian Haworth “and my very dear
friends, W. Skrimshire and F. Skrimshire, M. D.” (Haworth 1803). Tutt (1905) commented that the
Skrimshires probably saw the butterfly near Ely in 1797 or 1798 and afterwards went with
Haworth to capture it, perhaps in Bardolph Fen, a short distance east of Wisbech in West Norfolk.

WILLIAM SKRIMSHIRE AS NATURALIST

Dr Fenwick Skrimshire (1802) characterised his older brother as “‘a very accurate and attentive
observer of nature”, and, judging by William’s published articles (see References), as well as
notices of plants and animals he collected, his interests were broad-ranging from plants and insects
to the weather and phosphorescence. He reported that the roasted seeds of Iris pseudacorus could
be used as a substitute for coffee (W. Skrimshire 1809); Grigson (1955: 419) noted this, but merely
credited the discovery to a “Fenland doctor’. More than a century later, during World War II, well-

26 G. CROMPTON AND E. C. NELSON

roasted seeds from wild iris were used as a coffee substitute in the Channel Islands (see Vickery
1995: 409).

William Skrimshire assembled a substantial herbarium (hortus siccus). His manuscript catalogue
of this now-lost herbarium is extant in Wisbech and Fenland Museum. The manuscript, purchased
in 1829 by J. R. Weatherhead (see below) within two months of William Skrimshire’s death, bears
this annotation on the fly-leaf:

Catalogue of Plants contained in Mr Skrimshire’s Hortus Siccus sold some years since to Lord
Milton — and also of Plants contained in a Hortus Siccus bought by me and collected by Mr
Skrimshire after the sale of the former. The mark Y indicates those contained in this collection.
Sept. 12th 1829.

Comparison of the manuscript catalogue with specimens in Weatherhead’s herbarium, presented to
the Wisbech Museum by his widow, shows that very few of the ticked specimens have survived.

Viscount Milton (Charles William Wentworth Fitzwilliam, 1786-1857), son and heir of the
second Earl Fitzwilliam whose physician was Dr Fenwick Skrimshire, had paid seventy pounds for
William Skrimshire’s collection of minerals and his hortus siccus in February 1818 (“Feb. 2 paid
Dr [Fenwick] Skrimshire for his brothers minerals & Hortus Siccus (by draft on Snow) [£] 70.00.”
Northamptonshire Record Office Misc. Vol. 236: p. 75 (Private accounts (payments) of
Fitzwilliam. Jan. 1812—May 1822. Fitzwilliam (Milton)). Lord Milton was a patron of natural
history, “one of the oldest and most zealous friends of Natural Science in this country” (see
Desmond 1994). He was one of the few subscribers to Flora Graeca (1806-1840) to receive the
entire work (see Lack 1999: 213). No trace of Milton’s herbarium and mineral collections has been
found, and we must assume they no longer exist. However it is clear from the dates in William
Skrimshire’s manuscript catalogue and on herbarium specimens in WBCH that as soon as he had
sold his herbarium to Lord Milton, he immediately started collecting and pressing plants again.

Like any modern botanist William Skrimshire often gathered, and evidently retained, a number
of duplicate specimens, and thus there are some pre-1818 specimens in WBCH, derived from the
now-amalgamated herbaria of Skrimshire’s relatives and friends. The earliest date on an extant
specimen, among his brother-in-law Charles Metcalfe’s donation, is 1791; that year William
Skrimshire gathered Origanum vulgare at Hunstanton, West Norfolk. The series of specimens
shows that over the next 38 years Skrimshire not only collected in and around Wisbech but also
wandered through Cambridgeshire and adjacent counties in search of interesting plants.
Skrimshire’s diligence in observing and collecting plants means that he can be credited with the
first records of several species for Cambridgeshire (v. c. 29) and Northamptonshire (v. c. 32) (see
Table 1). Although his contribution to Cambridgeshire first records was published in Perring ef al.
(1964), those data were based only on Skrimshire’s manuscript catalogue and Relhan’s Flora
Cantabrigiensis (2nd edition 1802). Further research by G.C., particularly into the historical
records and the Wisbech Museum herbarium, allows us to publish a revised first records list for the
county (Table 1). The Norfolk records noted by Petch & Swann (1966: 15) all derive from
Skrimshire’s catalogue; they are Cladium mariscus (as Schoenus mariscus) and Utricularia
vulgaris, both from Bardolph Fen, and Persicaria bistorta (formerly Polygonum bistorta) from
Framingham (see Petch & Swann 1966: 165). The Northamptonshire data have not previously
been published; its flora was much less well documented in the early nineteenth century than that
of, for example, Cambridgeshire, so Skrimshire’s specimens add significantly to the tally of first
records.

One set of specimens provides a vivid record of one of Skrimshire excursions, which took place
just three months after the sale of the hortus siccus to Lord Milton. Skrimshire travelled from
Wisbech via Peterborough to Milton and Helpstone, the home-village of John Clare. On 25 May
1818, William Skrimshire foraged in Thistlemoor Wood near Milton, Northamptonshire, finding
Ajuga reptans, Anemone nemorosa, Fragaria vesca (as “Potentilla sterilis”), Ranunculus
auricomus and Viola canina. On the following day, perhaps having stayed overnight with Fenwick
in Peterborough, William found Pentaglottis sempervirens (as “Anchusa sempervirens”) in the
cathedral precincts; “there were 3 or 4 Roots, growing in the Minster Yard at Peterborough on the
E. side of the Building. May. 26. 1818”. As he often did, Skrimshire also commented about his
knowledge of the species elsewhere: “It is not found in the vicinity of Wisbech.” On 27 May, he
returned to the Milton area and visited Helpstone, gathering Helianthemum nummularium (as
“Cistus Helianthemum’’) and Hippocrepis comosa. In some old gravel pits at Helpstone, he found

WILLIAM SKRIMSHIRE (1766-1829) 2

TABLE 1. FIRST COUNTY RECORDS FOR CAMBRIDGESHIRE AND
NORTHAMPTONSHIRE

Cambridgeshire (v.c. 29).

§ Adonis annua: (as “Adonis autumnalis”’); near Bowling Green, Wisbech, c. 1820.

* Anagallis arvensis subsp. arvensis f. azurea: (as” Anagallis coerulea. Abbot ... A. arvensis var. 2”); Bushy

Place, Wisbech, 26 October 1822.

§ Atriplex littoralis: marshes below Wisbech, 1795.

§ Atriplex pedunculata: near The Black Tunnel below Bushy Place, Wisbech, 1826.

Beta vulgaris subsp. maritima: Relhan (1802).

Cardamine hirsuta: between Wisbech and Tydd Gote, 1794.

Cochlearia anglica: salt marshes below Wisbech, 1796.

Cochlearia danica: salt marshes below Wisbech, 1796.

Corydalis lutea: (as ““Fumaria lutea’) Mr Peckover’s garden wall, Wisbech 1818.

Datura stramonium: Long Drove, Wisbech, 1818.

Doronicum pardalianches: Bushy Place, Wisbech, 1818.

Limonium bellidifolium: below Wisbech, 14 September 1795 (W. Skrimshire to J. Sowerby) [NB perhaps a
Lincolnshire record; see discussion on p. 10.]

§ Mentha spicata: (as “Mentha viridis”); Bushy Place garden, Wisbech, 1813.

* Misopates orontium: Bushy Place and Crab Marsh, Wisbech, July 1819.

§ Orobanche ramosa: (Wisbech, before 20 November 1795. T. J. Woodward to J. E. Smith); hemp fields about
Wisbech, c. 1820.

Rubia tinctoria: (as “Rubia peregrina” in ms catalogue); Relhan (1802, as ‘Wild Madder’).

Ruppia cirrhosa: (as “Ruppia maritima’); salt-water ditch, east of Wisbech Tydd Gote road, 1795.

Silene uniflora: (as “Silene maritima’); see Perring (1956).

Sisymbrium irio: near Barton Lane, Wisbech 1797.

Triglochin maritima: Relhan (1802).

CHR MU KK %& UM

CR LPR % M&M

Northamptonshire (v. c. 32).

* Aceras anthropophorum: Helpstone, 1821; leg. George Skrimshire.

Ajuga reptans: Thistlemoor Wood, near Milton, 25 May 1818.

* Anemone nemorosa: Thistlemoor Wood at Milton, 25 May 1818.

Bupleurum rotundifolium: (as “Euphorbia hiberna’’); neighbourhood of Peterborough, 1821; /eg. George
Skrimshire.

Epipactis helleborine: (as “Serapias latifolia ... Epipactis latifolia’); Milton Wood near Peterborough, 1816.

Euphorbia exigua: Peterborough, between that City and Thorpe, 1821. Jeg. George Skrimshire.

Fragaria vesca: (as “‘Potentilla sterilis’”); Thistkemoor Wood at Milton, near Peterborough, 25 May 1818.

Hippocrepis comosa: Helpston, 27 May 1818.

Pentaglottis sempervirens: (as “Anchusa Sempervirens’’); Peterborough Minster-yard, 26 May 1818.

* Ranunculus auricomus: Thistlemoor Wood near Milton, 25 May 1818.

Viola canina: Thistlemoor Wood, at Milton, near Peterborough, 25 May 1818.

YC ey waieicay cua}

Unless otherwise stated, these records are attributed to William Skrimshire. * = voucher specimen
exists in WBCH; § = record in William Skrimshire manuscript catalogue.

a splendid array of orchids including Orchis ustulata, as well as Pulsatilla vulgaris (as “Anemone
Pulsatilla”). Although dated June 1818, a specimen of Ophrys apifera, which he called “Ophrys
aranifera. Spider ophrys”, was also picked from the gravel pits. Viola canina grew on “a small
Heath or Cover — on the left hand between Milton, and Helpstone”’.

Perhaps it was on one of his journeys to see his brother at Peterborough that Skrimshire stopped
at Cross Guns, a public house on the bank of the Nene at Wisbech St Mary’s parish boundary; the
site of this pub is now occupied by the Cross Guns Pumping Station (TF/347.015). The nearby
marsh yielded, for example, Schoenoplectus lacustris (as “Scirpus lacustris”) and Potamogeton
perfoliatus (there are no voucher specimens for these in WBCH). Black Boy Low was another
small pub, a short distance east of Wisbech almost on the Cambridgeshire-Norfolk border,
illustrated by Algernon Peckover in 1841 (reproduced in Crompton 1994). Skrimshire collected

28 G. CROMPTON AND E. C. NELSON

Geranium pratense “By the side of a ditch adjoining the Toll Bar between Wisbech and the Black
Boy Low at Walsoken”.

Further into West Norfolk, William Skrimshire visited Dersingham, and there collected plants
typical of heath and bog, Erica cinerea and Drosera intermedia (as “Drosera longifolia”) and D.
rotundifolia. The voucher specimens are not dated. As for the round-leaved sundew, Skrimshire
annotated it with a negative comment: “I have not found it nearer Wisbh. than Bardolph Fen.”

The specimens of Pentaglottis sempervirens and Drosera rotundifolia jast mentioned illustrate a
remarkable aspect of William Skrimshire’s herbarium. Frequently he added comments that inform
us about plants not found in the vicinity of Wisbech. In this regard, Skrimshire is most unusual
because the overwhelming majority of herbarium specimens are only labelled with the actual
locality where each specimen was obtained. Another good example, displaying his intimate
knowledge of the local flora, is a specimen of Silene latifolia labelled:

... Lychnis dioica [?] Smith L. vespertina Sibthorpe var. f. White Campion, or Cuckow flower.
In a hedge at Bushy-Place 1818. Not uncommon in hedges and by Road’s sides about
Wisbech, but I have never observed the red flowered one, within 20 miles of this place; and I
am persuaded that it is a distinct species from this Plant. W. S. 1818.

William Skrimshire was not just a gatherer of specimens. He was observant and inquisitive, and
one of the plants that intrigued him was Sagina maritima which grew abundantly in the salt
marshes on the banks of the River Nene (see Fig. 2) below Wisbech, not far from Bushy Place.
Two series of specimens, each carefully pressed and mounted, are annotated at length (see Fig. 1):

... Sagina procumbens. var. maritima. These Specimens gathered the beginning of May, on the
Salt Marsh near Bushy Place, grew upon the bare soil from which the Grass-sward had been
cut the preceding year; and as the root leaves grew in a perfectly star-like form and the
branches spread upon the bare ground as radii from the centre of the Plant the whole had a
striking appearance. I labelled it at the time as above; but from frequent and attentive
observation on the growth of this plant I have no doubt of its being the same with Dr. Smith’s
Sagina maritima. See the series of specimens on this sheet [f. 2] No. 1 Gathered the later end of
May } 1818. No. 2 Gathered the middle of June } 1818. From the Salt Marsh near Bushy
Place. f. 3 These three specimens gathered on the Salt Marsh near Bushy-Place June 30th.
1818.

These six specimens were gathered by the River side below Wisbech. Near Bushy Place. Sr. J.
E. Smith has made / this Plant / a distinct Species, under the specific name of maritima.
Dickson published this plant under the name of Sagina apetala, in his Hort. Sicc. Brit. Fasc. HI.
But I think it differs from S. apetala sufficiently to entitle it to be ranked as a distinct Species.
W.S. 1818.

James Dickson (1738-1822), one of the founders of the Royal Horticultural Society and of the
Linnean Society of London, “a most wonderful fellow” according to the Revd Richard Relhan,
issued his Hortus siccus Britannicus: being a collection of dried British plants, named on the
authority of the Linnaean herbarium ... in 19 fascicles between 1793 and 1802 (Henrey 1975: ILL,
34). Although Skrimshire evidently had an extensive library of contemporary botanical works
including William Hudson’s Flora Anglica, Richard Relhan’s Flora Cantabrigiensis, and James
Edward Smith’s Flora Britannica, he did not possess the relevant fascicle of Dickson’s Hortus
siccus. However John Nainby of King’s Lynn had a copy and generously removed the specimen
labelled Sagina apetala and gave it to Skrimshire so that he could compare it with those from his
immediate stamping grounds. And Skrimshire duly and accurately labelled it:

... Sagina apetala. S. maritima. Comp. Fl. Br. * This specimen was taken from Dickson’s Hort.
Sicc. Brit. Fasc. III. and given to me by John Nainby of Lynn. This is exactly similar to what I
find by the river side at Bushy Place, & which Dr. Smith has made a distinct species. 1818.

Plants that excited Skrimshire’s curiosity were often gathered at different seasons, so that his
herbarium specimens sometimes comprise more than one gathering. For example, he found an
odd-looking Lamium purpureum at Bushy Place and was uncertain about its exact identity:
Skrimshire labelled it “... Lamium incisum. Smith L. dissectum. With. L. purpureum [?] Huds.”.
He gathered specimens in March and April 1818, and mounted them together. Another sheet with

WILLIAM SKRIMSHIRE (1766-1829) 29

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ay led BY tibe2 7

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Jathtint e CUD LE tf AV TG lint. Lf MOGUL, Cath hb J Pe fltewl Pts (ahlen he
‘fel } y Ae ; ; / g
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Ie tot f a4 Pg Z : See
MH ‘LS 7 ‘Sr a7 ee yine mate: Lynd wl és [Ae Sex Pied Sheth <r? ton Thig She: if age

FIGURE 1. William Skrimshire’s herbarium specimens are usually meticulously documented, and neatly

presented. This sheet of Sagina maritima, a species which he observed closely, displays the care he took
(WBCH).

30 G. CROMPTON AND E. C. NELSON

NENE from neay the BLACK SLUICE tn 1525>

FIGURE 2. Alexander Peckover’s watercolour showing the River Nene in 1825 (“Nene from near the Black
Sluice in 1825”). The location was a few miles north of Wisbech town, and the tower and distinctive spire of
the parish church can be seen right of centre on the horizon. Bushy Place, William Skrimshire’s property, was
between the Black Sluice and the town, and so would lie towards the right-hand margin of this view.
(Reproduced by courtesy of Dr Peter Cave, and Wisbech and Fenland Museum; ©Wisbech and Fenland
Museum. )

four specimens of Mentha x piperita displays the stages in the growth of peppermint:

. M. piperita sylvestris. Sole. No. 1. A young Shoot, in Spring. No. 2. The Stem in Summer
before it flowers. No. 3. One of the Stolones from the main Plant in Autumn. No. 4. The Plant
in full flower. 1813.

He frequently noticed colour variants. A good example is Geranium molle: “These specimens, one
with Pink and the other with pale coloured Blossoms were gathered by the Road side at Bushy-
Place, May 15. 1819. Common about Wisbech. 1819.” The previous year on 29 June 1818 he had
obtained it “With pale coloured Blossoms”, probably from the same roadside bank.

WILLIAM SKRIMSHIRE’S GARDENS

William Skrimshire was evidently a keen gardener, and was just as interested in growing native
plants as exotics. From annotations on specimens and in his manuscript catalogue, we know that
he had a garden in Wisbech town, perhaps at Ship Lane where his father had lived, as early as
1801. It was from this garden that several of the anonymous specimens, now in the Castle
Museum, Norwich (NWH), came: Alchemilla vulgaris “May 13™ 1801 ... Archemilla [sic] -
Vulgaris ... gathered in Mr. W. Skrimshire’s Garden Wisbech ...”, Geranium lucidum and
Cymbalaria muralis (as “Antirrhinum Cymbalaria’), as well as Ajuga reptans (although the latter
specimen is not dated). John Weatherhead collected Allium ursinum “in Mr Skrimshire’s garden
Wisbech [in] May 1807” (WBCH).

One interesting annotation in his manuscript catalogue prea that he sowed seeds of Arabis

WILLIAM SKRIMSHIRE (1766-1829) 31

turrita “on my Garden Wall at Wisbech where it continued to grow for several years”; this may
have been as early as 1796 when the Revd John Hemsted (see below) gave Skrimshire a herbarium
specimen of the species.

By 1813 Skrimshire had also developed a new garden and peppermint plantation at Bushy Place.
Again using his annotations and notes, we can build up a picture of Bushy Place. It was probably
surrounded by walls: Skrimshire gathered Erodium moschatum “on an old Wall at Bushy Place,
Wisbech; May 21st. 1818. Very rare.” As well as the garden proper, there was the “peppermint
plantation’, presumably a field in which he grew a crop of Mentha x piperita. There was an
orchard, and “nut trees” (perhaps hazels) under which Doronicum pardalianches grew. The
peppermint plantation was adjacent to Long Drove. Skrimshire found Epilobium hirsutum “In
Long Drove, in the hedge of my Peppermint Plantation. Augt. Ist. 1818. It grows in a hedge at
Tip-cock-Trees about a mile below Bushy-Place. Not very common at Wisbech”. The still-house
in which he would have distilled the essential oil from the peppermint was also by Long Drove:
Vicia hirsuta (as “Ervum hirsutum’) grew in “a Cornfield opposite my Still-house in Long Drove,
Crab-Marsh. July 1. 1822”. (A house name “Mint House” is marked on the 1887 six-inch
Ordnance Survey map at TF/457.109; it may have been built on the former site of Skrimshire’s
still-house.)

He was fascinated by the weeds that grew in his garden and surrounding fields and often
gathered specimens for his herbarium. For example the “Anagallis coerulea. Abbot ... A. arvensis
var. 2. ... female pimpernel” (Anagallis arvensis subsp. arvensis) which “was wont to grow as a
weed annually in my Old Garden at Wisbech”, was also “among the Peppermint in the Orchard at
Bushy-Place; Wisbech” in October 1822. The voucher specimen represents A. arvensis L. subsp.
arvensis f. azurea Hyl. (det. G. Crompton), and is the first record of this forma from
Cambridgeshire. While he deliberately cultivated Mentha x piperita, Skrimshire found that M.
arvensis Was a “troublesome weed”, and he also gathered M. spicata in his garden at Bushy Place
as early as 1813 according to a note in his manuscript catalogue. Herniaria glabra was collected
by J. R. Weatherhead, who noted that “This came up as a weed in Mr Skrimshire’s garden”. The
manuscript catalogue contains several other weeds. By 1828 great willowherb had become a weed
at Bushy Place. Euphorbia exigua “used to come up annually as a weed in my garden ...” (the
voucher specimen came from his nephew, George Skrimshire, and was collected in Peterborough);
Tussilago farfara grew “On the Ploughed grounds by the river side, below [Wisbech]” and also
was “in my Garden as a Weed at Bushy-Place. 1822”. Other casuals were Campanula patula
“among the Peppermint in the Orchard, Bushy-Place, July 22, 1820”, and Chenopodium urbicum.
Carex spicata grew in his “Grass-field in Tinker’s Drove. Wisbh.” — presumably the pasture for
his horse (or horses) that kicked him (see below)!

Like present-day gardeners, Skrimshire battled with such weeds as Convolvulus arvensis, and
his annotated specimens vividly illustrate his frustration; at Bushy Park in June 1818 this was “one
of the most troublesome weeds that I have to contend with”. Solanum nigrum was condemned in
stronger terms: “A common weed in Gardens, about Wisbech. It is one of the pests in my
Peppermint Plantation at Bushy Place”. Less troublesome, perhaps, but no less worthy of pressing
and mounting for his herbarium, were Lamium amplexicaule “Gathered at Bushy-Place, Septr.
21st. 1818. A Common Weed both in Spring & Autumn, in Gardens & Cornfields in Crab Marsh,
Wisbech.”; Lapsana communis “From Bushy-Place, July 2nd. 1819 Common weed in Gardens and
under hedges. 1819.”; and Misopates orontium (as “Antirrhinum Orontium’’) “From my Garden at
Wisbech, where it constantly comes up as a Weed. 1. July. 1819. I have also observed it as a weed
in my Garden at Bushy-Place, and in Cornfields in Crab-Marsh, Wisbech. 1819.”

THE RIVER NENE BELOW WISBECH

The grounds of Bushy Place merged with the River Nene (Fig. 2). Although the Nene is still tidal
at Wisbech, Skrimshire’s herbarium specimens, and his manuscript catalogue, reveal the greater
extent of saline habitats, including salt marshes and flats, in the vicinity of Wisbech two centuries
ago, and thus also the extent to which maritime species penetrated inland in the early 19th century.
There were substantial areas of salt-marsh not far from Bushy Place, on both west and east banks
of the Nene, where he gathered Salicornia fragilis (as “Salicornia herbacea”), Puccinellia
maritima (as “Poa maritima, Glyceria maritima”), Centaurium pulchellum (conf. G.C. & E.C.N.

a2 G. CROMPTON AND E. C. NELSON

1999), and several scurvy-grasses (including Cochlearia danica; cont. T. C. G. Rich 1999).

Other species from saline (or maritime) habitats that Skrimshire found near Wisbech, but which
are now lost to the local flora, were Limonium bellidifolium (as “Statice reticulata”) and Atriplex
pedunculata. Limonium bellidifolium was reported as being “very common on the salt marshes
below Wisbech” by Skrimshire in a letter to James Sowerby dated 14 September 1795 (Boulger &
Britten 1918: 259-261). Sowerby later illustrated this species in English botany (1796: tab. 328)
using material collected “below Wisbech” and supplied by the Revd John Hemsted. A specimen
(WCBH; Fig. 3) gathered by Skrimshire, “Growing with the S[tatice] Limonium on the marsh
between St Peter’s Point and the Washway Aug. 14. 1821”, perhaps represents a Lincolnshire (v.
c. 53) record due to the convoluted vice-comital boundaries. The same must be said of Hemsted’s
gathering. In 1818 Skrimshire saw Silene uniflora (formerly S. maritima) “on the River-Bank on
Eastfield” and entered the locality in his manuscript catalogue; as Perring (1956) noted, this was a
new record for Cambridgeshire.

The banks of the Nene near Bushy Place were also inhabited by Arenaria serpyllifolia,
Cerastium glomerata (as “Cerast. vulgatum. Smith Cer. viscosum With. ... Cerastium viscosum”;
det G.C. & E.C.N. 1999), Mentha arvensis, Papaver rhoeas (as “Papaver Rhaeus var?”) and
Trifolium campestre (as “Trifolium procumbens”). Other “river bank” plants, noted in
Skrimshire’s manuscript catalogue, include Briza media, Valerianella locusta (as “Valeriana
locusta”) and Bupleurum tenuissimum. Carlina vulgaris grew “upon Mount Pleasant with the
Carduus nutans and upon Banks about Wisbech, otherwise not common here.” The local
hedgerows yielded a bramble, Rubus cf. corylifolius (as “Rubus [fruticosus deleted] corylifolius.
Smith. R. fruticosus var. 2. major. With[ering].”), and a “red purple variety” of Viola odorata.

WILLIAM SKRIMSHIRE’S ASSOCIATES

William Skrimshire did not botanize in isolation. He maintained contacts with many botanists both
amateur and professional. An entry in Skrimshire’s manuscript catalogue provides an important
record of one of his contacts. Blackstonia perfoliata (as “Chlora perfoliata”) was “gathered ... at
Cherry Hinton, near Cambridge; in company with my friend Relhan. 1792” (there is no
corresponding herbarium specimen in WBCH).

The Revd Richard Relhan had published his Flora Cantabrigiensis in 1785 and it contained
some records from Wisbech, but none ascribed to Skrimshire. We do not know when he became a
friend of Skrimshire, but there are five herbarium specimens in WBCH annotated by Skrimshire
as having come from Relhan. These derive from the J. R. Weatherhead hortus siccus donated to
the Museum in 1862 by Weatherhead’s widow, and are Gnaphalium luteoalbum “From Shelford,
Cambridgeshire”; Malva neglecta (as “ Malva verticillata’) “Discovered in Cambridgeshire a few
years since and sent to me by my worthy friend The Revd. R Relhan. 1818.”; Clinopodium
grandiflorum (as “Melittis grandiflora, Melissa grandiflora’) “from Mr. Relhan’s Garden, at
Cambridge. 1795.”; Pyrus communis “From Hinton near Cambridge...”; and Potamogeton
compressus (without a locality; conf. C. D. Preston).

There is one other Relhan specimen in Norwich (NWH); Trifolium subterraneum was annotated
first by Skrimshire — “Given to me by Mr. Relhan. I have not found it about Wisbh. 1795” — and
subsequently by the anonymous owner, “Mr. Skrimshire gave me this Specimen ...”. Four further
Relhan specimens in WBCH do not appear to have been in Skrimshire’s possession, rather they
were somehow acquired by J. R. Weatherhead; these are Carex cf. viridula (as “Carex Oederi?’”’),
Eleocharis acicularis (as “Scirpus pauciflorus?”; conf. C. D. Preston, G.C. and E.C.N. 1998),
Jasione montana and Liparis loeselii (as “Malaxis Loeselii, Ophrys Loeselii’).

As well as indicating a close link between these two men, the specimens reveal that Relhan may
have had a garden. As we have already noted, Skrimshire was an enthusiastic and observant
gardener, and perhaps he and Relhan exchanged living plants as well as herbarium specimens. In
the second edition of Relhan’s Flora Cantabrigiensis (1802) there are more than 30 new county
records ascribed to Skrimshire, including many for mosses, lichens and algae. Relhan (1802) paid
this tribute to his friend, acknowledging William Skrimshire’s contribution to this edition, and
stating that Skrimshire was “happily devoted to natural science and most skilled in his surgeon’s
art.”

WILLIAM SKRIMSHIRE (1766-1829) 33

j) : re | “67; -
MCplipuirtl Fr tyne,
SPE - 4
Hatice reLoutaale .

Maller, hoa Liver tL? Or S epeet
Jatt lati ted Ceivw Wiibee/
Gryiwindg swith the My pil ri 72 Whiz
; pp fips | a res

ees ; Lelwetn e/- d Lets Pyar a7

: 4 5
The Warsyird tt. « ees

FIGURE 3. This specimen of Limonium bellidifolium may have been collected in Lincolnshire (v. c. 53)
(WBCH): Skrimshire first reported the species from the Wisbech area in September 1795.

34 G. CROMPTON AND E. C. NELSON

Likewise, Skrimshire received many specimens, particularly bryophytes from the Newmarket
area, from the Rev. John Hemsted (c. 1747-1824), one-time Curate of St Mary’s, Newmarket
(Oswald 1991). Skrimshire’s acquaintance with Hemsted is confirmed by Hemsted’s
correspondence with James Sowerby (Mss in Sowerby Correspondence Vol. 26. The Natural
History Museum, London). On 13 September 1796, Hemsted informed Sowerby that:

... [have received an Invitation from my Friend Skrimshire to come and spend a few Days with
him which I propose to accept & to go down to Wisbeach next Monday, stay with him to the
latter End of the Week & then go across Country to Bedford ... after which I shall be out no
more & shall be happy to see you & Mr Skrimshire desires me to say that if you can give us
the Meeting at Wisbeach he shall be exceedingly glad of your Company... [Vol. 26: 171].

Just over a week later, on 22 September 1796 Hemsted admitted he “was disappointed of my
Excursion to Wisbeach. Our Friend Skrimshire having got a Hurt in his Leg by a Kick of a
Horse ...” (Vol. 26: 172). A later visit was more productive, as Hemsted told Sowerby: “I had
forgot to mention Conferva Capillaris found last Week in Salt-Water Ditches near
Wisbeach.” (Newmarket, 26 August 1797: Vol. 26: 174).

Apart from the numerous bryophytes, Hemsted gave Skrimshire the following specimens:
Arabis turrita “From Cambridge ... 1796.”; Groenlandia densa (as “Potamogeton densum”: conf.
C. D. Preston) “gathered near Newmarket by Mr Hempsted ...”; and Mentha x smithiana (as
“Mentha gentilis. M. rubra.a. Huds.”) “Sent to me by Mr. Hemsted, from Newmarket. I have not
met with it growing about Wisbech. 1795.”

William Skrimshire had a wide circle of botanical friends. They included a fellow surgeon John
Pitchford (c. 1737-1803), a Roman Catholic who practised in Norwich (Bull 1999: 36; Petch &
Swann 1968), one of the first British botanists to study Linnaeus’ works (Walker 1988). The two
men first met late in 1795, as Thomas Woodward (1745—1820) related in a letter to James Edward
Smith dated 20 November 1795; although Skrimshire is not mentioned by name the reference is
clear:

M' Pitchford has been at Wisbech to sell his estates, he met with a brother surgeon there who
has found very good things in that country. Orobanche ramosa in plenty, as I always supposed
likely from the soil on which I found it here, Dactylis stricta [= Spartina maritima], Riccia
glauca, as he supposed but which I [? took] at first sight (no great proof of extraordinary
sagacity) to be a much better thing — Riccia natans, also Ulva incrassata. ... [Ms in Archives,
Linnean Society, London; Smith Correspondence 18: 210]

This requires further comment. Orobanche ramosa is recorded in Skrimshire’s manuscript
catalogue from Wisbech. Spartina maritima has never been recorded for Cambridgeshire and is
now believed extinct in Norfolk (see Beckett et al. 1999): as this record is not localized, and there
is no entry in Skrimshire’s catalogue, it is impossible to assign it to a vice-county. It is not
improbable that S. maritima did grow in the lower reaches of the Nene in the 1790s. The comment
by Woodward on Riccia natans (now Ricciocarpos natans), a liverwort, is confirmed by a
specimen in Skrimshire’s moss herbarium (WBCH) labelled “found in a ditch at Outwell between
the town and the Toll-Gate 1795”. There is also a specimen labelled Riccia glauca (= Riccia
sorocarpa) that was “gathered by the side of some silt-pits, in a field opposite the Black-Boy-Low,
Walsoken” (v.c. 28) but it is dated 1796, after Pitchford’s meeting with Skrimshire.

In return, presumably, for specimens from Wisbech, Pitchford gave Skrimshire specimens of
Arabidopsis thaliana (as “Arabis thaliana 8”) from ‘ta Wall at Norwich ... 1799.”; Monotropa
hypopitys “gathered near Norwich ... 1797.°; Herminium monorchis and Orobanche elatior, both
unlocalized. Pitchford was as interested in the mints as Skrimshire (see Dawson 1934: 74-75: Bull
1999). He was a keen field-botanist, credited with the discovery of Holosteum umbellatum (Bull
1999) and the earliest Norfolk record of Liparis loeselii (Bull 1999), and he made this comment
about the contemporary problems of a field-botanist in a letter to Dawson Turner dated 15 July
1797 (Ms in Dawson Turner papers, Trinity College, Cambridge):

... but for Dianthus prolifer [= Petrorhagia prolifera; see Beckett et al. 1999: 82] — alas! What
shall I say, but that it is lost & lost I fear for ever to this Country, the unfeeling farmers, those
mortal enemies of botanists, having ploughed up the ground to the very edge so that I had

WILLIAM SKRIMSHIRE (1766-1829) 35

some difficulty forcing my way along among the barley & weeds to the very end of the hedge
row, as I was determined not to lose an inch of the precious ground where the dear creature
used to grow. [Nelson 1997a, 1997b]

Although Skrimshire did not live far from the open sea — and as we have noted the River Nene is
tidal — in 1821 he acquired a series of specimens of coastal plants from F. Coates of Brancaster,
including Salsola kali, Calystegia soldanella (as “Convolvulus Soldanella’”) and Honckenya
peploides (as “Arenaria peploides”’). Mr Lilly Wigg (1749-1829), who was variously a shoemaker,
schoolmaster and a clerk in the Yarmouth bank owned by Dawson Turner, was another of
Skrimshire’s contacts. Wigg is recorded as having instructed Dawson Turner in algae (Petch &
Swann 1968; Desmond 1994). One specimen from Wigg was Hieracium umbellatum: “This
specimen was sent to me by Mr. Wigg, from Yarmouth. — 1795. I have not found it near Wisbech.”
According to Petch & Swann (1968: 219) Wigg’s hawkweed, reported from Norfolk by Smith
(1807: tab. 1771) was “gathered on the beautiful wooded hills at the back of Thorpe, Norwich”.

The botanist and entomologist, Adrian Hardy Haworth, who was friendly with William and
Fenwick Skrimshire, also donated herbarium specimens to William’s hortus siccus. Haworth was a
succulent enthusiast and is commemorated in the southern African genus Haworthia (Aloaceae).
There are several Haworth specimens in WBCH, including Scilla verna and S. autumnalis,
Potentilla rupestris and Salix lapponum; whereas the squills could be native (the specimens are not
localized), the rock cinquefoil and downy willow specimens were from plants cultivated in
Haworth’s garden.

Individuals represented by singletons are Mr Hull — perhaps the Manchester physician, John
Hull (1761-1843). William Skrimshire was “indebted for this wild Specimen [of Polycarpon
tetraphyllum| to Mr Hull, who gave it to Fenwick. 1797”. The Rev. John Burrall presented
Skrimshire with a specimen of Dawson Turner’s discovery Tolypella glomerata (Desv.) Leonh. (as
“Chara nidifica’’), from “splashes of Salt water at Cley on the Norfolk Coast’. There is a specimen
of Silene uniflora (as “S. amoena”; conf. S. M. Walters 1999) “Given to me by Mr. Don, out of the
Physic Garden. I have not found it about Wisbeach. 1795”. Almost certainly this came from the
Botanic Garden in Cambridge, from its curator James Donn (1758-1813) (Walters 1981: Fig. 38,
p. 45).

Other evidence for contacts with fellow botanists is obtained from correspondence and
publications. Not surprisingly, William Skrimshire knew the remarkable duo of Norfolk-based
botanists, Dawson Turner and Dr (later Sir) James Edward Smith, and the naturalist and publisher
James Sowerby. There is a letter from Skrimshire to Turner, dated 10 July 1797, among Turner’s
voluminous correspondence (Trinity College, Cambridge). There is also a letter to Sowerby (Ms.
in The Natural History Museum, London) about specimens Skrimshire sent for illustrating in
Sowerby and Smith’s English botany (see e.g. Limonium bellidifolium above). Among these
illustrations was Stratiotes aloides published in 1797. Although Sowerby drew the illustration
(Sowerby & Smith 1797: tab. 379) from a living plant sent by Skrimshire, the accompanying text
does not state where he collected it. However in the manuscript catalogue of Skrimshire
herbarium, there is an undated entry for Stratiotes aloides from Bardolph Fen, West Norfolk, and
this may well have been its source.

JOHN ROSE WEATHERHEAD (C. 1787-1849)

John Rose Weatherhead was Skrimshire’s apprentice from 1803 for five years. By 1817 he was
practising as a surgeon in partnership with Skrimshire, and lived in The Crescent, Wisbech. He
became a keen botanist and corresponded with William Jackson Hooker, Professor of Botany in
the University of Glasgow (and from 1841, Director of the Royal Botanic Gardens, Kew) on both
botanical and entomological subjects. William Skrimshire credited his pupil with “probably”
discovering a new species of beetle which had been washed ashore in a flood at Wisbech in 1812:
“This pretty little Insect, which I expect is new to Britain, was first discovered this summer at
Wisbech, by my Pupil, Mr. J. R. Weatherhead, an ingenious and industrious Entomologist” (W.
Skrimshire 1812: 318).

Weatherhead’s own herbarium forms a substantial proportion of the collections in WBCH (Kent
& Allen, 1964: p. 269 as Weatherhead, I. R.), but is not the principal subject of this paper.

36 G. CROMPTON AND E. C. NELSON

However, a small number of specimens from him were evidently incorporated into William
Skrimshire’s herbarium, and sheets annotated in Skrimshire’s distinctive writing (see Figs | and 3)
include one of the hybrid Stachys x ambigua J. E. Sm. (then called “‘ambiguous woundwort’), the
progeny of S. sylvatica and S. palustris. This had been discovered by William Borrer and W. J.
Hooker in Orkney and on the mainland of northern Scotland in 1808. Weatherhead, independently,
found it near Edinburgh in August 1809 and sent a specimen to Smith who mentioned this fact in
the protologue (Sowerby & Smith 1809). The specimen that belonged to Skrimshire records other
details of Weatherhead’s discovery, the annotation on the verso by Weatherhead himself being the
more detailed:

[recto; W. Skrimshire] Found by Mr. J. R. Weatherhead in Augt. 1809 at the foot of one of the
Pentland Hills, in a moist boggy place about a Mile and a half from Hubbies How. He
presented me with this specimen. 1810.

[verso; J. R. Weatherhead] Stachys ambigua. “I found this plant in Augt. 1809 at the foot of
one of the Pentland Hills in a moist & boggy place about a mile and a half from [Edinburgh —
del.] Hubbies How, a specimen was sent to Dr Smith who considered it as new; it had however
been found during the preceding summer in the orknies & some parts of the North of Scotland.
Dr Smith has named it ambigua. It was growing inter-mixed with S. sylvatica, but no specimen
of S. palustris could be found near it.” J. R. Weatherhead.

The specimen from Skrimshire’s herbarium, and a second one, both contained in Weatherhead’ s
hortus siccus (WBCH), must be isotypes, although their precise status remains to be determined.

In September 1829, following William Skrimshire’s death, Weatherhead bought Skrimshire’s
herbarium and with it the manuscript catalogue. As already mentioned, Weatherhead then marked
with a tick all the specimens he had acquired. These specimens were presumably amalgamated
with Weatherhead’s own extensive herbarium, and many may even have been remounted and re-
labelled because most of the specimens marked by a tick are not present in the two sets of
Weatherhead specimens now in WBCH. Weatherhead himself presented four volumes of
herbarium specimens to the Wisbech Museum in 1835; there do not appear to have been any of
William Skrimshire’s specimens in this collection. Thirteen years after Weatherhead’s own death
in 1849, his widow presented seven further volumes of herbarium specimens. It was this collection
tnat contained most of the Skrimshire specimens now in WBCH, including the first confirmed and
the only record for Cambridgeshire of the rare Ruppia cirrhosa (as “Ruppia maritima’; det. C. D.
Preston 1993) collected by Skrimshire:

... in a Salt water Ditch at Walton, near the River about 3 miles below Wisbech [v. c. 28]. I
have also gathered it in a ditch which admits the tides, in a lane on the Right hand side of the
Road from Wisbech to Tidd between the 3rd and 4th. milestones [v. c. 29]. It grows plentifully
in the ditches at Long Sutton Wash-way, below Wisbech. 1795 [v. c. 53].

CONCLUSION

William Skrimshire’s manuscript catalogue and the few of his specimens that are available for
study have provided several previous authors with material for their respective Floras. The
Cambridgeshire records extracted by F. H. Perring from these reliquiae are listed on the
Cambridge Natural History Society's card index held in the herbarium, Department of Plant
Sciences, Cambridge. The Norfolk specimens are listed in the late E. L. Swann’s card index, now
with Mrs G. Beckett.

With his varied interests in natural history, and his undoubted knowledge, William Skrimshire
must have been a focus for the amateur naturalists living in the Wisbech neighbourhood in the late
18th and early 19th century. His manuscript catalogue and herbarium specimens represent an
archive of considerable value, especially as a foundation for historical studies of the flora of
Wisbech and the north-eastern sector of the old county of Cambridgeshire and the Isle of Ely, and
the adjacent parts of West Norfolk.

WILLIAM SKRIMSHIRE (1766-1829) 37
ACKNOWLEDGMENTS

We are grateful to successive Curators of Wisbech and Fenland Museum, Mr D. Devenish and Dr
R. Hanley, for allowing us to examine the herbarium collections, and we would like to express our
particular thanks to Jill Rayment and Robert Bell, not only for all their assistance but also for
finding so many sources of information and enlightenment about the Skrimshire family history.
We are also indebted to Dr Michael Lewis of Oundle for information on Dr Fenwick Skrimshire’s
achievements as a physician at Peterborough. G.C. also acknowledges the co-operation of Dr Peter
Gillam who kindly showed her relics of his ancestor Dr George Skrimshire. Dr C. D. Preston
(especially Potamogeton, Ruppia), Dr S. M. Walters (Silene) and Dr T. C. G. Rich (Cochlearia)
checked the identities of some of Skrimshire’s specimens. Mrs Gill Gent kindly confirmed
Skrimshire’s first records for Northamptonshire.

We are grateful to the Wisbech Society for allowing us to publish excerpts from G.C.’s papers
published in its annual reports (1994 and 1996).

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Linnean Society of London. Linnean Society, London.

WALTERS, S. M. (1981). The shaping of Cambridge botany. Cambridge University Press, Cambridge.

(Accepted June 1999)

Watsonia 23: 39-46 (2000) 39

John Blackstone (1713-1753); a London apothecary
and botanist of his time

E. M. and C. BOWLT

7 Croft Gardens, Ruislip, Middlesex, HA4 SEY

ABSTRACT

John Blackstone was born, educated, apprenticed and practised as an apothecary in London during the first
half of the 18th century. He made early contact with several distinguished persons, notably Sir Hans Sloane, to
further his botanical interests. His passion for plants resulted in two small Floras at a time when few were
published. One dealt with the plants around Harefield, Middlesex, in which he recorded 464 currently
acceptable flowering plant species. Comparison with the plants recorded in that area during 1965-76 indicates
that 26% of plants had become extinct in the intervening 240 years (18% if the additional surrounding 2 km is
included) in contrast to an estimated 1% for Britain as a whole in the last 300 years. He is commemorated in
having a plant and genus (Blackstonia perfoliata) named after him.

KEYWORDS: early Floras, extinctions.

INTRODUCTION

“The period from Ray’s death in 1704 until the introduction of the Linnean system into Britain
about 1760 was a lean one for the study of British plants” (Gilmour & Walters 1973). Interest in
the subject was kept alive by a few, some only minor figures in the history of botany, who in their
time made small but significant contributions to its study. Such a person was John Blackstone,
who developed a tremendous enthusiasm for plants and whose two small Floras, Fasciculus
Plantarum circa Harefield sponte nascentium (1737) and Specimen Botanicum quo Plantarum
plurium rariorum Angliae indigenarum loci natales (1746), were among the few published during
the period. Harefield, Middlesex, is remarkable in having had one of the earliest censuses of a
human population in Britain in 1699 (Cuthbertson 1992) and one of the earliest catalogues of
plants (John Blackstone 1737).

Blackstone was a predecessor in the study of plants in Middlesex in whom Douglas Kent took a
special interest. Kent’s (1949) paper stimulated our investigations into Blackstone’s life and work.
Inevitably there is much common ground between the papers of Kent and ourselves, but
corrections and new material are included here. In addition we have analysed Blackstone’s (1737)
plant records for Harefield deriving statistically meaningful numbers for extinctions during the
following years, and illustrating the continuing relevance of the work of early botanists such as
Blackstone.

EARLY LIFE

John Blackstone was born in 1713. His birth on 22 September and baptism on the 25th, are
recorded in the registers’ of the parish of St Leonard Eastcheap on the east side of the City of
London. The church of St Leonard had been destroyed in the Great Fire of 1666 and not rebuilt
and the parish joined with its neighbour St Benet Gracechurch, where he must actually have been
christened.

Blackstone was not born in humble circumstances. His family background, on both sides, whilst
not providing great wealth, gave him education and opportunity to enter a profession which seems
eventually to have provided sufficient income for him to pursue his passion for botany. It also
seems to have given him influential contacts, which he early exploited to further his interest in
plants.

40 E. M: AND C. BOWLTE

APOTHECARIES IN THE FAMILY

His father was Edward Blackstone, citizen and ironmonger’ of London, but his grandfather,
another John Blackstone, was a successful apothecary living within the precinct of Pater Noster
Row in St Faith’s parish in 1690°. Indeed the Gentleman’s Magazine’ described him as an eminent
apothecary of Newgate Street, a friend and associate of Sir Hans Sloane and a liberal benefactor of
St Bartholomew’s Hospital. He was Warden of the Apothecaries’ Company 1712-13. He had three
sons as well as Edward: William, who became Keeper of Wychwood Forest, Oxfordshire; Charles,
a silk mercer, citizen and bowyer of London; and Richard of whom nothing is known. The elder
John Blackstone apparently retired from business in 1713 “to enjoy the fruits of a competant
fortune’ and was living as a gentleman in Wandsworth, Surrey, at the time of his death in 1715.
He left £1200 to Edward to set up in business as an ironmonger. Just what Edward had been doing
until then (he was already married with a child) is unclear. His name appears neither in the list of
freemen of the Ironmongers’ Company® nor in the lists of apprentices. However, an Edward
Blackstone, son of John Blackstone, pharmacist, was apprenticed to Ambrose Crowley, a draper,
in 1703, freed in 1711 and a member of the Livery of the Drapers’ Company 1719-27’. From
these dates this could be Edward, the father of John Blackstone. If so, in preferring ironmongery to
drapery, he would have been able to set up in business easily enough, if he had capital, at a period
when the companies were not exercising so strict a control over the traders in the city as they had
in earlier years.

William, a brother of John Blackstone the elder, was also an apothecary in London and his name
appeared among those who subscribed towards the revival and improvement of the Chelsea Physic
Garden about 1700. This side of the family had property at Chipping Walden, Essex*. A William
Blackstone (apprenticed as an apothecary to a John Blackstone in 1696’) may be Edward’s elder
brother, who later became keeper of Wychwood Forest.

THE HAREFIELD CONNECTION

Edward Blackstone married Sarah, the fifth child of Francis Ashby Esq. (1671-1743) of
Breakspears, Harefield, in Middlesex, one of the lesser landed gentry with a family coat of arms.
Harefield played a large part in the life of John Blackstone. He obviously got to know it well and
had a great affection for the area. His parents were married at Harefield church on 28 February
1712'°. After John’s birth in September 1713 two brothers followed: Edward, born and baptized
(perhaps he was delicate) on the 29 December 1717. and Francis on 27 March 1723''. All the
children’s baptisms were at St Leonard’s Eastcheap. However, when the father died, in 1730, he
was buried at Harefield, and John’s mother and younger brothers moved to Breakspears to live
with her parents. John, by then apprenticed'” to Thomas Bearcroft, apothecary, was a frequent
visitor to that new home. It is possible that the family had moved to Harefield before Edward
Blackstone’s death, as the record of John’s apprenticeship’’, dating from Sth August 1729, refers
to him as the son of Edward Blackstone, “late of London, ironmonger’’.

EDUCATION

Whether John Blackstone spent most of his childhood in the city or stayed with relatives elsewhere
for long periods, perhaps with his grandparents at Harefield or with his paternal uncle, the Keeper
of Wychwood Forest in Oxfordshire, one can only guess. His early education is also unknown, but
it was sufficiently good for him to enter Merchant Taylors’ School, then situated in Suffolk Lane
not far from London Bridge, in 1727, at the age of 14'*. He must already have been reasonably
proficient in Latin, for when he left only two years later’’ he knew sufficient, not only for his
subsequent profession as an apothecary, but also to write his subsequent Floras in that language.

A contemporary at Merchant Taylors’ School was William Watson who also became an
apothecary, and ultimately a physician, with a strong interest in plants. Watson similarly entered
the school in 1727, but remained a year longer. If the two were not friends at school, they certainly
were afterwards, for Watson supplied Blackstone with herbarium material and records for his
Specimen Botanicum. He was a man of wide scientific interests, conducting electrical experiments

JOHN BLACKSTONE |

and contributing to the Philosophical Transactions of the Royal Society, becoming an F.R.S. in
1741, and eventually being knighted in 1786, the year before he died.

APPRENTICESHIP

On leaving Merchant Taylors’ John was “apprenticed and bound to Mr. Thomas Bearcroft for
eight years from this day (5 August 1729) in the sum of £100”'°. That Bearcroft was an apothecary
was probably significant in introducing him to the serious study of plants, herbs having long
played a major part in an apothecary’s pharmacopoeia. It seems that not many grew or gathered
herbs by this period (as Gerard had done in his gardens at Holborn, for instance), but they were
certainly expected to be able to identify species.

HERBARIZINGS

Simpling expeditions for apothecaries are recorded from 1620. Led by such eminent persons as
Thomas Johnson, the parties met early in the morning at St Paul’s and visited rural areas like
Hampstead Heath or Greenwich and occasionally, as in 1634, went as far afield as Bath, Bristol
and the Isle of Wight. (Hunting 1998). After the founding of the Chelsea Botanic Garden in 1675,
the Society of Apothecaries, “to excite a taste for botany among apprentices” according to
Pulteney (Pulteney 1790), encouraged private herbarizings beginning on the second Tuesday in
April, and held monthly on the same day until September, in the countryside in the immediate
neighbourhood of London, to which Apothecaries could send their apprentices. These seem to
have been in the nature of botanical field meetings with a leader from the Society until 1724 when
a Botanical Demonstrator took over the job (Hunting 1998). Isaac Rand, an apothecary and
botanist, filled that post 1724—43'’

There had been complaints about disorderly behaviour among the apprentices at the herbarizings
and henceforth they had to enter their own and their master’s names on a list kept by the leader and
bring a letter from their masters permitting their attendance. There was to be no question of the
apprentices treating the herbarizings as holidays. All had to wait to the end to attend the showing
of the plants and from thence return directly home without loss of time'®. There is no actual record
of John Blackstone having attended, but it seems highly likely that he did so. William Watson is
known to have done so because “during his apprenticeship he gained the honorary premium (a
bound copy of Ray’s Synopsis) given annually by the Apothecaries’ Company, to such young men
as exhibit a superiority in the knowledge of plants 1 in these excursions made by the Demonstrator
of the Chelsea Garden” (Pulteney 1790). There is in the British Library a copy of Ray’s Synopsis”
(bound in with other of Ray’s works) with Blackstone’s autograph and the date 1736. In 1735
Blackstone’s master died and he was ‘turned over’ to Mr Richard Stevenson for the remainder of
his apprenticeship. This concluded on 7th March 1737 “when having served his full term he was
sworn and made free”””

INFLUENTIAL CONTACTS

Before he was out of his apprenticeship the young Blackstone had already been writing to
several eminent persons, including his grandfather’s friend Sir Hans Sloane, about his botanisings
and had already written a book on the plants of Harefield. His grandfather, Francis Ashby,
apparently took an interest in plants and where they grew’', and may well have influenced the
young Blackstone’s burgeoning interest in botany and the local flora. By the end of 1736
Blackstone had been commissioned to collect plants for the Apothecaries’ garden at Chelsea as
well as the Oxford Physic garden. He mentions this in a letter written on 11 December 1736 to Dr
Richard Richardson, a well-to-do physician and well-known botanist of Bierley, Yorkshire.

“By good fortune I have discovered the Dentaria baccifera (Cardamine bulbifera) to be a native of
this island. The place where it grows is in a large wood near a village call’d Harefield, about
eighteen miles from London, where it grows in such plenty, that some acres of ground are cover’d
with it. [Old Park Wood, where it still grows]. There are several other rare plants to be found
thereabouts, of which a catalogue is now ready for the press... I have communicated this plant and
my intentions, to Sir Hans Sloane, Dr. Dillenius (from whom I had a most obliging letter) Mr
Rand, and some others... I have orders to get specimens for Chelsea and Oxford Gardens; and
wou 'd you give me directions, I shu’d be very glad to send some for your garden, if ’tis not there
already”.

42 BM. AND:C.BOWLT

A week later he had already received a reply and wrote back with some confidence, giving his
book a puff at the same time:

“Strand, Dec.18th 1736

I beg leave to return you my humble and hearty thanks for the favour of your obliging letter... I
have for these last three years been employ’d in making a collection of the native plants; and
having an opportunity of going to see my friends pretty often, I made it my business to see as
many of the adjacent places my time would permit, and to collect such plants as offer’d themselves
in the course of my walks, without ever intending to publish anything on this subject. But, being
detained last Summer by a long illness, near four months on the spot, I found so many rare plants
that I thought it worthwhile to make a catalogue of them, and show it to some skillful persons in
that science for their approbation, which succeeded to my desire; and I only beg leave to let you
know, that the plants there mentioned were gathered almost solely by myself, as also that the
catalogue is not general; being only intended as an essay for a more particular search
thereabouts. ... I purpose to print my work by subscription, tho’ the price of it cannot exceed five
shillings; and I desire no money till the book is deliver’d. I should take it as a great favour if you
cou’d procure me a specimen of Epimedium and the Christophoriana, which would be highly
acceptable. .. I expect to go to Harefield next week, and will search for some roots of Dentaria, if
you please to order it, and will (with God’s leave) send you some in perfection about the latter end
of April, when the bulbs will be ready to plant. Be pleased to accept of the compliments due to the
ensuing season””>.

The catalogue referred to was published in 1737 as Fasciculus Plantarum circa Harefield sponte
nascentium (A Little Bundle of Plants freely growing around Harefield).

His correspondence with Dr Dillenius, Professor of Botany at Oxford, continued. A letter to him
from Blackstone, dated 1744, is laid in with a specimen of Epimedium in the Sherard Herbarium
(in the Dept of Plant Sciences, Oxford): “I could not fail sending you a quotation from a letter of
that ornament to Botany, Dr Richardson,...Epimedium is a native of England. It grows in the
woods near Bingley...and not sparingly”. (Epimedium is not now, of course, considered native, but
these were the frontier days of botany with much still undiscovered.)

THE BOTANIST

With the large number of modern detailed books on plant identification available it is easy to
forget the difficulties in identifying plants in the 18th century. The relatively few books that had
been published, often long previously, seldom agreed on names and descriptions were not very
distinctive. Blackstone included several author names for each plant but in his Fasciculus
generally put the name given in Gerard’s Herbal (as revised by Johnson in 1637) first. The Herbal
is a hefty tome of more than 1600 pages and must have been difficult to procure and expensive to
buy. It is not known whether he possessed a copy or used the Apothecaries’ library at Chelsea.
Gerard’s book has woodcut illustrations to the species, which in most cases are recognisable and
hugely bolster the rather inadequate descriptions in English. Building up a herbarium, as
Blackstone did, would clearly have supplemented the rather limited Floras, and the exchange of
specimens with other botanists must have been significant in getting general agreement on the
identification of plants.

Blackstone’s Fasciculus is an early example of a local Flora. It was a remarkable achievement at
that period for a 24 year-old just out of his apprenticeship. It contains a list of 523 plants in
alphabetical order of their pre-Linnean Latin names. This is followed by their English names and
in many cases the locality (in English), with comments and time of flowering (in Latin). The book
ends with an Appendix containing some short notes relating to Harefield. To put this into
perspective, without entering into the question of species, it has been estimated that, at the time of
Ray’s death in 1704, the total number of plants recorded for the whole of Britain was about 970
(Gilmour & Walters 1954). For the whole of Middlesex, Trimen & Dyer (1869) recorded 859
species, and Kent (1975) recorded 1109 - this last including many recent aliens. A few of
Blackstone’s records are from places outside the immediate area of Harefield, such as Iver and
Chorleywood, and even as far afield as Windsor and Hampton Court (i.e. up to 15 miles distant). It
also includes a few fungi, liverwort and lichen records, but they are not always identifiable.

JOHN BLACKSTONE 43

Omitting these, and also what now appear to have been varieties, Blackstone published a list of
464 vascular plants species growing in the parish of Harefield, an area of about 1870 hectares
(4621 acres) in the north-west corner of Middlesex. There are, however, some curious apparent
omissions of plants that surely must have been common and noticeable in his day. It is difficult to
believe, for instance, that Gorse was not present in the 18th century. He does not seem to have
deliberately omitted very common plants because he lists some, such as Daisy and Dandelion. In
spite of this, including a few extra species included in his second book, the final total of 464
flowering plants recorded by Blackstone must have represented a majority of those growing
around Harefield in 1737. It is, therefore, statistically meaningful to examine the extinctions that
have occurred in this one small area since then, by comparison with the distribution maps based on
the London Natural History Society’s botanical survey 1965-1976, published in the Flora of the
London Area (Burton 1983).

The parish of Harefield is contained within a block of seven 2 x 2 km squares of the Flora maps.
This area has been assumed to be close to that used by Blackstone which was probably not too
precise. By the time of the L.N.H.S. survey, 122 flowering plant species recorded by Blackstone
were no longer present. That is, 26% of his plants had become extinct in about 240 years. If the
area is expanded to include all 2 x 2 km squares immediately surrounding the block of seven (i.e.
an additional 16 squares) the number of extinctions is reduced to 85 species, i.e. 18%. These must
rank as some of the best estimates over one of the longest periods of time of extinctions from the
flora of a relatively small area. For the British Isles as a whole it is estimated that about 15 species
(i.e. about 1% of the total) of flowering plants and ferns have been lost in the last 300 years
(Rackham 1986). The greater percentage loss in the smaller area of Harefield illustrates the
vulnerability of small populations.

Inspection of the 85 lost species indicates that most were from wetlands (25%), followed by
those from heathland (15%), corn/arable fields (13%) and woods and thickets (13%). These
findings agree with those of Kent (1975) for the extinctions in the whole of Middlesex between
1863 and 1968 and confirms many general observations.

Some of the plant species no longer growing in Harefield are only local extinctions, but others
are no longer to be found in the London area as a whole. For instance, Arnoseris minima has not
been seen within two miles of St Paul’s since 1910. Anagallis tenella was last recorded in 1951.
Parnassia palustris and Pedicularis palustris are also now extinct in the London area. Among the
orchids recorded by Blackstone at Harefield, Orchis militaris seemed to be extinct in Britain by
about 1914. Since its rediscovery in the Chilterns in 1947 it remains one of Britain’s rarest orchids.

As a Londoner botanising in Middlesex Blackstone was rather fortunate that his grandfather
lived in the parish of Harefield. In the extreme north-western corner of the county, the topography
and the outcropping geological strata gave rise to a wide range of soils and habitats within a
comparatively small area. Blackstone was aware of this, for in an Appendix to his Fasciculus he
wrote “The Soil, within this Compass, is various; that of the South and East Parts being a strong
Clay, very productive of Elm and Oak, which thrive well here. The North and West Parts,
bordering on Hertfordshire and Bucks, is generally a dry Chalk and stony Land, and The great
Variety of the Soils here, produces all Sorts of Vegetables, which our Catalogus of such Plants as
grow spontaneously in this Neighbourhood, sufficiently shews.”

At Harefield the northern edge of the London Basin rises to the surface and is cut into by the
River Colne, revealing, under the London Clay, the Reading Beds of clays and sands and the
Upper Chalk. The Colne Valley was an area of black alluvium wetlands. Unfortunately,
Harefield’s varied geology led to its rape for raw materials. Chalk pits noted by Blackstone
continue to the present day. Brickworks have used the clays, sandpits have been dug and, in this
century, the vast gravel pits have destroyed the wetlands. To add insult to injury many of the
diggings have been filled with rubbish and rubble giving the coup de grace to any plants with the
temerity to try and hang on.

These activities, along with building, are the principal reasons for the disappearance of species,
but there are others. Trimen & Dyer (1869) reproduce a quotation of Peter Collinson’s (1694—
1768) which they say helps to account for the extinction of orchids at Harefield: “There is one
Miles, a parson of Cowley, near Uxbridge, who is orchis mad, takes all up, leaves none to seed so
extirpates all wherever he comes which is cruel, and deserves chastisement.” In connection with
Orchis militaris, Kent (1975) said I “feel that unrestricted collecting by nineteenth-century
botanists contributed to the extinction of this rare and beautiful species in the vice-county.”

44 E. M. AND C. BOWLT

THE APOTHECARY

Having completed his apprenticeship in March 1737, John Blackstone seems to have been in no
hurry to set up in business, perhaps because of the ill health frequently mentioned in his letters, or
because the collecting and study of plants was proving more alluring. On 8 April 1737, he wrote to
Sir Hans Sloane: “The air proves serviceable to me in general, tho’ there are still some symptoms
remaining which a perfect state of health ought to be without, particularly the swelling of my legs
after walking, and the heat of my hands and feet towards night”

His contemporaries, like Dr Richardson, having seen the Fasciculus, certainly encouraged him
to pursue his natural history studies and he confessed to Sloane on 29 September 1737 that “the
study grows more and more agreeable to me”. Sir Hans had probably been encouraging him to
start in business, for Blackstone wrote to him on 26th October 1737, just as he was about to visit
Pembroke College, Oxford and relatives at Charlbury: “I hope, Sir, this absence will be no
prejudice to your esteem for me, but to engage in business without any foundation would not be
deem’d a prudent action, therefore as I am obliged to be depending, I must wait till those I depend
on, are pleased to make a beginning for me”*°. In December he said that if his relatives had been
willing to fix him in business he would not have left London. In fact it was almost another year
before he did set up in business at the sign of the Griffin, Salisbury Court, near St Bride’s, just off
Fleet Street, in September 1738. He was admitted to the Society of Apothecaries the same year,
and married a widow, a Mrs Webb, in 1739, but she died, probably in 17407’. A William
Blackstone, an infant, was buried at St Bride’s in April 1741 and an Elizabeth Blackstone on 11
April 1742, but they cannot be proved to be John Blackstone’s wife and child.

We know little about Blackstone’s professional life. He became a subscriber to the Laboratory
Stock of the Society of Apothecaries in January 1740, investing £107, but seems to have found
difficulty in establishing his business. He complained of trade “being very dull and money very
hard to get and seemed to be asking Sir Hans Sloane for a loan in September 1740, when he
dispatched a note with a postscript saying that his man would wait on him about noon to know his
pleasure~’. The result is unknown, but he felt, himself 1 in a position to take on an apprentice, John
Thomas Vernon of Malpas, Cheshire, in 1741°°

He married again in October 1742. His second wife was Mary Abbot, the daughter of Richard
Abbot, es stainer, and the wedding took place at St Sepulchre’s, Newgate Street on 23
October

LATER YEARS

The next few years were clouded with tragedy, for his mother and his grandfather both died in
April 1743, being buried at Harefield on the same day. The following year his wife gave birth to a
daughter, Maria, only for her to die in infancy. On a happier note a second daughter, Sophia, was
born in 1748 (and lived until 1827) and his second book, Specimen Botanicum, had been published
in 1746.

SPECIMEN BOTANICUM

The layout of this small 106-page book is the same as for the Fasciculus, but it lists only 367
plants, including 19 non-vascular cryptogams, and their sites in various places in England. A
number of records were supplied by other botanists, which he acknowledges - sometimes in a
quaint manner, such as, “found by that accurate Botanist, Dr Richardson”, and “with Dr Jarvis, an
eminent surgeon and curious Botanist”. The catalogue includes 99 vascular plants from Harefield.
All but nine of these had appeared in his earlier Fasciculus.

As in his earlier work, most of the non-vascular cryptogam identifications are dubious,
presumably a reflection of the state of their taxonomy at that period. The list is of what were
considered some of the more interesting plants growing in England. It is in no sense
comprehensive in its geographical cover, but was thought worth publishing at a time when few
botanical books were being produced. It is interesting that even in Blackstone’s time the loss of
certain species from some sites was being noted. He records that Galium cruciata, Crosswort “is

JOHN BLACKSTONE 45

lately lost in Hampstead Church-Yard, where it grew in Johnson and Parkinson’s time”. Again of
Cypripedium calceolus, Lady’s Slipper, “found by Mr Thornbeck, Surgeon, at Ingleton, who
likewise informs me that it has been lost in Helk’s Wood for some years.”

Financially, life was probably easier for Blackstone in the 1740s. His wife may have had some
money to begin with and she certainly inherited £500 from her father, Richard Abbot, who died in
1746. John inherited £30 from his mother*”, as well as a three-acre field called Maudemeade and
the four-acre Further Maudefield, running along the side of Bayhurst Wood, Harefield. He was
able to mortgage the mead in 1746 to raise £105 from Pierce Dodd of Red Lyon Square, a doctor
of physic’. Did he need the money to publish his book, the Specimen Botanicum? He sold both
fields to Anne Cane of Shiplake in 1751**. The fields are famous in botanical history, because it
was here that he noted in the Fasciculus the Snake’s Head Fritillary growing “...1n Maud Fields
observed above forty years ago by Mr Ashby of Breakspears.” This was its first record as a British
plant. In addition, overall Blackstone provided at least 103 first records for Middlesex (v.c. 21).

At this same period he was playing a role in the affairs of the Society of Apothecaries, being on
the Garden Committee of the Chelsea Physic Garden from 1746 until 1752*, when his health
began to fail. It is notable that it was to the botanical activities of the Society, rather than the
trading side, that he devoted his time and interest.

John Blackstone died on 11 March 1753, not yet 40 years old. He was buried at Harefield a
week later on the 18th March*’. He left directions in his will dated | February 1753 “to be buried
in the cheapest and most private manner possible...” To his brother, Francis he bequeathed his
cornelian seal with two heads engraved on it, the silver can with his grandfather’s coat-of-arms
and his mortuary knife. He left his largest punch bowl with a silver ladle and his seal engraved
with a man at work to Mr John Abbot, his brother-in-law, “as a mark of my gratitude for many
favours” .Everything else was left to his dear wife, Mary’ ’.

There is a collection of 44 plants (including 14 mosses and lichens) in the Sloane Herbarium, but
in 1947 three volumes of Blackstone’s plants, labelled by him, were discovered at Ripon Museum,
Yorkshire. Not all of the plants were localised and only a few were from Harefield. The collection
was transferred to the Natural History Museum, where approximately 360 specimens were laid
into the general herbarium.

BLACKSTONIA
Curiously, Blackstone did not record the plant named after him closer to Harefield than Gerrards
Cross, although we have photographed it at Harefield on some of the remaining chalkland.
Pulteney (1790) noted that when Hudson published his Flora Anglica (1762) and introduced the
Linnaean system into English botany, “he separated the Yellow Centaury from the Gentians and
gave it the name Blackstonia; which distinction Linnaeus confirmed in the Systema of 1767, but
then changed the name to Chlora, an appelation it had received from Rencaulme, in his Specimen
Historias Plantarum, published in 1611. It should seem that the discovery of the true place of this
plant in the system entitled Hudson to the dispensation of the name, or at least that Blackstone
should have been perpetuated in the trivial epithet”.

Hudson and Pulteney would be pleased to know that the plant is once more called Blackstonia
perfoliata, and it seems a fitting tribute to John Blackstone’s botanical studies, which clearly
gained the approval of his contemporaries.

REFERENCES

BLACKSTONE, J. (1737). Fasciculus plantarum circa Harefield sponte nascentium.

BLACKSTONE, J. (1746). Specimen botanicum quo plantarum rarorium Angliae indigenarum loci natales.

BurRTON, R. M. (1983). Flora of the London area. London Natural History Society, London.

CUTHBERTSON, E. (1992) Gregory King’s Harefield. Hillingdon Borough Library, London.

GERARD, J. (1633). The herbal or a general historie of plants. Enlarged and amended by Thomas Johnson.

GILMOUR, J. & WALTERS, M. (1973). Wild flowers. Collins, London.

HubDsON, W. (1762). Flora Anglica. London.

HUNTING, P. (1998). History of the Society of Apothecaries. Society of Apothecaries, London.

KENT, D. H. (1949). John Blackstone, Apothecary and Botanist (1712-53). Watsonia 1: 141-148.

KENT, D. H. (1975). A historical Flora of Middlesex. Ray Society, London.

PULTENEY, R. (1790). Historical and biographical sketches in the progress of botany in England from its
origin to the introduction of the Linnean System. London.

46

E. M. AND C. BOWLT

RACKHAM, O. (1980). Ancient woodland. Edward Arnold, London.
TRIMEN, H. & DYER, W. T. T. (1869). Flora of Middlesex. Hardwick, London.
TURNER, D., ed. (1835). Literary and scientific correspondence of Richard Richardson, MD, FRS of Bierley,

Yorkshire, pp. 351-355.

NOTES

Guildhall MS 17607: parish registers St Leonard’s East Cheap

Edward Blackstone is described as an ironmonger in the baptismal register of St Leonard’s, East Cheap -
Guildhall MS 17607 and in Chancery Proceedings - PRO C 11 1976 3, but does not appear in lists of
apprentices or freemen of the Ironmongers’ Company.

D. Kent’s Papers: Poll Tax Assessments 1678 and 1690 cxciii. 390

Gentleman’s Magazine: 1827 11 Vol 97

Ibid

Guildhall MS 16978

D. Kent’s Notes

PRO C11 1584. 7

Guildhall MS 8207: Society of Apothecaries Bindings Book 1694-1836

Harefield Parish Registers transcript: private collection.

Guildhall MS 1760718/02/93: parish registers St Leonard’s East Cheap.

Guildhall MS 8207: Society of Apothecaries Bindings Book 1694—1836.

Guildhall MS 8207: Society of Apothecaries Bindings Book 1694-1836

Guildhall Library: Merchant Taylors School Registers 1561-1934.

Guildhall Library: Ibid.

Guildhall MS 8207: Society of Apothecaries Bindings Book 1694-1836.

Guildhall MS 8228: Garden Committee Minutes 1731-70.

Ibid.

Brit Lib: 969 f 18 (2).

Guildhall MS 8206/2: Society of Apothecaries Freedom and Admissions Book.

John Blackstone: Fasciculus plantarum circa Harefield sponte nascentium 1737
Dawson Turner ed: Literary and Scientific Correspondence of Richard Richardson, MD, FRS of Bierley,
Yorkshire, pp 351-5

Ibid

Brit Lib: Sloane MS 4055 f89

Ibid 4055 ff203-4

Ibid 4055 ff223-4

Collectanea Topographica et Genealogica Vol 5, 1838.

Guildhall MS 8226: Society of Apothecaries List of subscribers to Laboratory Stock.
Brit Lib: Sloane MS 4056 £305

Guildhall MS 8207: Bindings Book 1694-1836.

Guildhall MS 7220/1: parish registers St Sepulchre’ s.

PRO: PCC Prob 11 727

London Metropolitan Archive: Acc 1085/M41

LMA: Acc 1085/M42

Guildhall MS 8228/1: Society of Apothecaries Garden Committee Minutes 1731-70.
Harefield Parish Registers transcript: private collection.

PRO Prob 11 800 71 .

(Accepted July 1999)

Watsonia 23: 47-58 (2000) AT

A history, mainly nomenclatural, of St Dabeoc’s Heath
E. C. NELSON*

Tippitiwitchet Cottage, Hall Road, Outwell, Wisbech PE14 SPE, Norfolk, UK.

ABSTRACT

St Dabeoc’s heath was first gathered in Cantabria (northern Spain) by J. P. de Tournefort before 1694, and
subsequently during 1700 by Edward Lhuyd in western Ireland. The convoluted history of the Latin names,
both pre- and post-Linnaean, for St Dabeoc’s heath is reviewed. Linnaeus was the first to employ, but not
validly publish, a binomial (Erica cantabrica) in the second edition (1759) of his Flora Anglica. William
Hudson adopted the same specific epithet, but placed St Dabeoc’s heath in Vaccinium; Hudson’s binomial
V. cantabricum was validly published and is the basionym for the current name, Daboecia cantabrica (Huds.)
K. Koch. The orthography of the generic name, first printed but not validly published by Martyn in 1807, has
been a contentious matter for over a century but as nomenclatural stability is desirable, the “corrupt” spelling
prevails. The species name is typified, the lectotype being the illustration published by Petiver in 1704.

KEYWORDsS: Daboecia cantabrica, Ericaceae, Linnaeus, typification.

INTRODUCTION

St Dabeoc’s heath is familiar to Irish, French, Spanish and Portuguese field-botanists, and to
gardeners in many other places. It belongs to a distinctive genus of the Ericaceae named Daboecia,
and that name is derived (“corruptly” to quote Smith 1791) from the vernacular Irish name for this
elegant shrub.

St Dabeoc’s heath, Daboecia cantabrica (Huds.) K. Koch, inhabits the milder, oceanic regions
of western Ireland (counties Galway and Mayo; see Webb & Scannell 1983), western and south-
western France, northern Spain (Pyrenees to Galicia) and north-western Portugal (see maps in
Woodell 1958; Dupont 1962; Small & Small 1998). There is an isolated population on the Azores,
and this is treated either as a discrete species, D. azorica Tutin & E. F. Warb. (Tutin & Warburg
1932; Sealy 1949; McClintock 1969), or as a geographically isolated subspecies of St Dabeoc’s
heath, D. cantabrica subsp. azorica (Tutin & E. F. Warb.) D. C. McClint. (McClintock 1989).

The purpose of this paper is to describe the discovery and to elucidate the nomenclatural history
of Daboecia. During the three centuries which have passed since it was first brought to the
attention of botanists, St Dabeoc’s heath has been assigned to various genera — Erica by Linnaeus
(1754), and all his predecessors back to Tournefort (1694), and by such of his successors as
Thunberg (1785); Vaccinium only by Hudson (1762); Andromeda by Linnaeus (1767, 1770), and
by Martyn (1807) who was the first author to list the generic name Daboecia; Boretta by Necker
(1791) and some subsequent authors; Menziesia by Salisbury (1781), Jussieu (1802) and de
Candolle (1805); Bryanthus only by Merino (1906); and Daboecia (or Dabeocia) by Don (1834),
Koch (1872) and most subsequent authorities.

As just indicated, there is a second matter of disagreement. The orthography of the generic name
is open to dispute (see below). Martyn (1807) and Don (1834) used Daboecia, whereas Dabeocia
was the arguably correct orthography (e.g. Babington 1888; Praeger 1925) adopted by Koch
(1872) and most authors until the early 1950s (e.g. Moore & More 1866; Colgan & Scully 1898;
Webb 1943; Gilbert-Carter 1950; Nelson 1984).

*Address for correspondence: E-mail: tippitiwitchet @ zetnet.co.uk

48 E. C. NELSON

DISCOVERY IN CANTABRIA AND IRELAND

The first mention of a heather from Cantabria (northern Spain) with large flowers and myrtle-like
leaves that were white underneath,’ referable to Erica even in pre-Linnaean times, appeared in
Joseph Pitton de Tournefort’s Elemens de botanique (1694: 475). The source of Tournefort’s
information is not exactly recorded; perhaps he gathered specimens during his exploration of the
Pyrenees in 1681, but this is unlikely given his use of the geographical epithet cantabrica.
Tournefort distributed specimens of his Cantabrian heath to fellow botanists including William
Sherard (see below) although no specimen of the heather collected by Tournefort can now be
identified in the Sherardian Herbarium (OXF; S. Marner pers. comm. 1995).

In the spring and summer of 1700, the Welsh naturalist Edward Lhuyd (1660-1709; olim Llwyd
or Lhwyd) travelled to western Ireland where he collected several hitherto unknown plants
including a heath with large purple flowers (Gunther 1945; Mitchell 1975; Nelson & Walsh 1995).
Lhuyd brought specimens to Britain and distributed duplicates to several botanical friends
including the Revd John Ray, James Petiver and William Sherard. In correspondence written
during the summer of 1700, Lhuyd also described the heath (see Mitchell 1975; Nelson 1978). He
wrote to Dr Tancred Robinson, among others, about various aspect of Ireland’s natural history;
this particular letter was published in 1712, three years after Lhuyd’s death:

In most of the Mountains of Galloway and Mayo grows an elegant sort of Heath, bearing large
Thyme-leaves, a Spike of fair purple Flowers like some Campanula, and viscous Stalks. I
know not whether it be any thing related to the Cisti Ladanifere. [Lhwyd 1712: 525].

To Dr Richard Richardson of North Brierly, near Bradford, Yorkshire, he wrote (see Owen 1922;
Dandy 1958):

In the moors of ye County of Mayo & Galloway grows a very elegant sort of Heath which so
common that ye people have given it ye name of Frych Dabedg 1. Erica [Sti.] Dabeoci &
sometimes ye women carry sprigs of it about them as a Preservative against Incontinency.
I calld it Erica maxima viscosa, rubra; Rosmarini foliis brevioribus flosculis Campanula
minoris ... .

That same year, 1700, Tournefort, Professor of Botany in the Royal Garden in Paris, published
Institutiones rei herbaria, essentially a Latin edition of Elemens de botanique, repeating the
phrase~ that he had first used in 1694 — he did not provide any new information about his
knowledge of this plant (Tournefort 1700: 603).

Meanwhile, Lhuyd’s specimens provided English authors with material for their publications.
The first to notice it was John Ray in Historia plantarum (1704: Ill, Book XXX: 98). The
particular volume was issued in the summer of 1704 — on 8 June 1704 Ray told Sir Hans Sloane
that the third volume “of my History of Plants is now finished at the press” (quoted by Raven
1950: 300). There is a further reference* to the Irish heath within a section comprising lists of
plants that Ray had received from Petiver (Ray 1704: II, Appendix, 244). The most remarkable
thing about Ray’s contribution is that he correctly equated Tournefort’s Cantabrian heather with
Lhuyd’s Irish one. Was this accurate identification just luck?

That question is answered by a letter to Lhuyd from William Sherard written on 29 May 1701
from “Badmington” (Ms Ash. 1817a, original in Bodleian Library, Oxford; Gunther 1945).
Sherard told Lhuyd: .

As to ye curious plants you have innrich’d me wth I can say little, however shall venture to
make some conjectures or queries abt them, since ye are pleas’d to comand it of me. I find (as
Mr. Ray complains to me abt my own) ‘tis hard to judg of plants by dry’d specimens,
especially where they are not perfect & well preserv’d, wch is not always practicable,
especially in travelling.

In the accompanying list, which clearly included Lhuyd’s Irish specimens, number 4 was glossed
“T had this (or some very so like it, wch upon comparing ye specimens I shall easily judg) from Dr.
Tournefort, by ye name of, Erica Cantabrica, fl. max. fol. myrti subtus incanis. Inst. r. herb.” In
other words, it was Sherard who identified Lhuyd’s gathering of St Dabeoc’s heath and equated it

HISTORY OF ST DABEOC’S HEATH 49

with Tournefort’s specimen. Lhuyd then seems to have labelled his specimens with Tournefort’s
name, including whatever material went to Ray, although from a letter written by Ray, dated 11
June 1701 (Gunther 1928: 280-281), it is evident that Lhuyd also used the name Erica S$. Dabeoci:
“The plant you intitle Erica S. Dabeoci’, responded Ray, “I am in some doubt whether it be a
genuine species of Erica, the flower falls away, & ye fruit seeming to be different.”

Shortly after Ray’s Historia plantarum was published in the summer of 1704, if we may judge
correctly from his own phrasing, Petiver published an engraving (Fig. 1, p. 53) of Erica
HIBERNICA ... S. Dabeoci in the third decade of his part-work Gazophylacium nature & artis
(Petiver 1704: plate 27), a work described as a medley of text and illustrations depicting plants,
shells, insects, birds and other animals. The unambiguous reference to Ray’s Historia plantarum in
the accompanying text’ indicates that the third decade in which the description and illustration of
St Dabeoc’s heath were issued was published late in 1704.

Jacob Dillenius, editor of the third edition of Ray’s Synopsis stirpium Britannicarum, included
St Dabeoc’s heath as the sixth member of his genus Erica, employing the long, elegantly
descriptive phrase-name Erica Cantabrica flore maximo, foliis Myrti subtus incanis of Tournefort,
and repeating the information first published by Ray (1704; Dillenius 1724). Caleb Threlkeld
(1726) listed this western heather as Erica Cantabrica flore maximo, foliis Myrti subtus incanis.
Erica Sancti Dabeoci, the name by which it was known to botanists throughout the early 18th
century. The only other Irish Flora published before 1753, Botanalogia universalis Hiberniae by
John Keogh (1735), did not include any account of St Dabeoc’s heath.

LINNAEUS AND ST DABEOC’S HEATH

The pre-Linnaean history (prior to 1753 when Linnaeus’ Species plantarum was published) of St
Dabeoc’s heath is uncomplicated and of little significance as far as, for example, the nomenclature
of the species are concerned. However, the post-1753 history is more complicated.

Linnaeus was well informed about the botanical works of earlier decades, as is clearly shown by
any page of his own publications. Thus it is not surprising that St Dabeoc’s heath should be
included in Linnaeus’ works from an early period. While inexplicably there is no entry for this
Hiberno-Cantabrian species in the first edition of Species plantarum (Linnaeus 1753), which is the
bench-mark for all modern botanical nomenclature, when Linnaeus turned his attention to the flora
of Britain and Ireland, he became aware of Erica S. Dabeoci Hibernica. Linnaeus’ chosen names
for St Dabeoc’s heath are listed in Table 1.

Erica daboeci of Flora Anglica 1754

In Flora Anglica, a work that is essentially a concordance providing Linnaean binomials for plants
described in Dillenius’ edition of Ray’s Synopsis stirpium Britannicarum (1724), and which
appeared in the guise of a thesis defended by Isaac Olaus Grufberg dated 3 April 1754, the final
entry, under “Dubia’, is for “471 Erica 6 daboeci’”. The numerals refer respectively to the page and
the paragraph number in Dillenius (1724) — in fact Linnaeus should have used either 470 because
that is the page on which the first Erica appeared, or 472 because Dillenius’ entry for St Dabeoc’s
heath is on that page. That combination of Latin words Erica and daboeci was original — neither
Dillenius nor Petiver, the two authorities cited by Linnaeus, nor any previous author had used that
particular set of words. Did Linnaeus intend this as the binomial for the heather? Why did he list it
among the doubtful (“Dubia”)? And, why did he spell it daboeci? The answer to the latter question
is perhaps linked with the many typographical errors which occurred in Flora Anglica (cf. Stearn
1973: 44; see also Sealy 1959). The first two questions are discussed below.

Stearn (1973) discussed the significance of the specific names in Linnaeus’ Flora Anglica,
stating that:

Although the nomenclature of Linnaeus’s Flora Anglica (1754) follows closely that of his
Species Piantarum (1753) it contains a number of specific names not included in that earlier
work, and these must be regarded as new names validly published by reference to the
previously published descriptive information given in the cited entries of Ray’s Synopsis, third
edition (1724).

50 EC” NELSON

TABLE 1. NAMES PUBLISHED BY LINNAEUS AND HIS CONTEMPORARY WILLIAM
HUDSON FOR ST DABEOC’S HEATH: THE RIGHT-HAND COLUMN GIVES THE
MOSTLY PRE-LINNAEAN SOURCES EXPLICITLY CITED IN EACH PUBLICATION

Linnaeus, Flora Anglica (April 1754: 29) a
Erica 6 daboeci [Dillenius: Synopsis methodica stirpium Britannicarum] 471 |recte
472|.6

Petiver: Gazophylacii ... t.27.f. 4

Linnaeus, Flora Anglica, in Amoenitates academice ... 1V: 92, 111 (November 1759)

Erica cantabrica [Dillenius: Synopsis methodica stirpium Britannicarum| 471 [recte
472].6
Ray: Dendrologia ... 98
Petiver: Gazophylacii ... t. 27. f. 4

Hudson, Flora Anglica (January—June 1762: 143)

Vaccinium cantabricum Tournefort: [/nstitutiones rei herbaria] 603
Ray [i.e. Dillenius: Synopsis methodica stirpium Britannicarum|] 472
Ray: Historia Il Dendrologia ... 98
Ray: Historia Ul Appendix ... 244
Petiver: Gazophylacii ... t. 27. f. 4

Linnaeus, Species plantarum ... editio secunda (September 1762: I, 509)

Erica daboecii Tournefort: [/nstitutiones rei herbarie| 603
Ray: Historia III Dendrologia ... 98
Petiver: Gazophylacii ... t. 27. f. 4
Vaccinium cantabricum Hudson: Flora Anglica 143

Linnaeus, Systema nature ... editio duodecima (1767: Il, 300)
Andromeda dabecia Erica daboecii [Linnaeus]: Species plantarum 509
Hudson, Flora Anglica ... edition altera (1778: I: 166-167)

Erica daboecii Linnaeus, Species plantarum [ed. 2] 509

Stearn cited examples of, for example, suppression of names by Linnaeus between the first edition
of Flora Anglica and the second edition published in Amoenitates academicae (1759), but he does
not comment on the entry for St Dabeoc’s heath.

How does the entry for “Erica 6 daboeci” stand with regard to the International code of
botanical nomenclature (Tokyo 1993) (hereinafter the JCBN (1993))? Does it fall within Art. 26.6
whereby certain “designations are not to be regarded as specific names’, including Art. 26.6 b,
such unspecified “‘other designations of species consisting of a generic name followed by one or
more words not intended as specific epithets.” The final section of Flora Anglica headed “Dubia”
is not laid out like the previous portion of the fifth section, and most entries in it contain more than
one word (a descriptive phrase) which clearly were not intended to be nomina trivialia. Moreover
the generic name and the one-word (in this case) phrase are separated by the numeral indicating
the paragraph number in Synopsis. Given the use of phrases, and given that the typography is
different, there does appear to be cause to reject this designation because it was “not intended as
[a] specific epithet.”

It is also possible to argue that because Linnaeus did not adopt binomials in the section headed
“Dubia’, he did not accept any of the apparent binomials as nomina trivialia, and thus that they are
not validly published under Art 34.1 (a): “A name is not validly published (a) when it is not
accepted by the author in the original publication ...”.°

Thus the first edition of Linnaeus’ Flora Anglica (1754) must be discounted as a source of a
binomial for St Dabeoc’s heath.

HISTORY OF ST DABEOC’S HEATH 51

Erica cantabrica of Flora Anglica 1759.

Linnaeus’ next publication of relevance is the second (1759) edition of Flora Anglica, in which he
used a different name, Erica cantabrica, and cited an additional reference, Ray's Dendrologia. In
this edition of Flora Anglica, Erica cantabrica is printed twice, once as an undoubtedly deliberate
binomial within Linnaeus’ commentary on Ireland (Linnaeus 1759: 92) and later within the section
headed “Obscura” in the form “471. Erica 6. cantabrica” (Linnaeus 1759: 111). Linnaeus’ reasons
for changing the entry from “471 Erica 6 daboeci” are not known. What is clear, however, is that
even in 1759 Linnaeus had not seen any specimens of this plant, only the crude figure in Petiver’s
Gazophylacii nature.

There can be no doubt that Linnaeus did intend Erica cantabrica to be a binomial (nomen
triviale) because he used it deliberately on p. 92, but he was not consistent, and he did not list it
with the names of four other species of Erica on p. 100. Was this binomial validly published in the
second edition of Flora Anglica? Although the case is different, because of the prior use of Erica
cantabrica on p. 92, under Art. 26.6 of the JCBN (1993) (see above) it has to be argued that this
name also is not validly published.

Erica daboecii of Species plantarum 1762.
It is noteworthy that Linnaeus reverted to Erica daboecii in the second edition of Species
plantarum (1762), citing William Hudson’s name Vaccinium cantabricum (see below) as a
synonym. Linnaeus stated that he had not seen flowers’, but his description suggests that by this
time he had been able to examine a fruiting specimen with some leaves; he correctly noted that the
capsule was erect and large and had four valves, facts not in previously published descriptions
even that by Ray (1704).°

Three years after the publication of the new edition of Species plantarum, Linnaeus received a
herbarium specimen with flowers from Peter Collinson in September 1765, and in a letter dated 27
December 1765 commented to John Ellis that “Erica Dabeoci was sent by Peter Collinson: a fine
specimen which much delighted me. It is truly an Erica, though so unlike the rest” (see O’ Neill &
Nelson 1995).

WILLIAM HUDSON AND ST DABEOC’S HEATH

The Latin binomial for St Dabeoc’s heath, Daboecia cantabrica, has as its basionym Vaccinium
cantabricum published by William Hudson F. R. S., a London apothecary and one-time Assistant
Librarian in the British Museum, in his Flora Anglica (Hudson 1762), the work which firmly
established “Linnean principles of botany in England, and their application to practical
use” (Smith 1824).

Hudson was planning to compile a British Flora as early as the Summer of 1760. On 3 July 1760
he wrote to Linnaeus (Ms in Linnaeus’ correspondence; Linnean Society of London):

I have taken the liberty to send you thro the hands of my worthy friend Mr John Ellis some few
of the Plants which are mentioned at the end of your Flora Anglica togather with some others. I
should have sent you some others but Mr. Ellis informed me that the ship was to sail in two or
three Days, and that he should put up his things the next morning, which hinderd mee from
looking out of my collection any more at present, but hope to send as many more before the
end of the summer and if there are any plants which are mentioned In R[ay’s] Syn[opsis]
which your are in want of if you will honour mee with a list off them, I will do all that laies in
my power to procure them for you. I should esteem it as a peculiar Favour if you will be so
kind as to honour me with your opinnion concerning the plants but more Especially the grasses
which have plagued me much. I should not have thought of giving you so much trouble had I
not had some intentions of Publishing a flora Britanica and not having been able to meet with
that information which I wanted here. I hope I shall soon have the honour of hearing you have
rec[eive]d the plants safe.

se

. some few of the Plants which are mentioned at the end of your Flora Anglica” would include
St Dabeoc’s heath, which was in fact last of all, but we can be fairly certain Hudson did not send a
specimen to Linnaeus as none from Hudson is in Linnaeus’ herbarium (LINN).

52 E. C. NELSON

In his Flora Anglica Hudson removed St Dabeoc’s heath from Erica and placed it in the genus
Vaccinium, but why? What was his actual role in the description and naming of St Dabeoc’s
heath? Did he examine specimens of the heather, study the species in details and then write a fresh,
original description and, finally, devise a Latin name for the plant without ever referring to
Linnaeus’ earlier Flora Anglica? Or did he merely take up a copy of Linnaeus’ Flora Anglica and
rework the text and the name to suit his own Flora? Hudson undoubtedly owned, or had direct
access to, a copy of the second edition of Linnaeus’ Flora Anglica. Within the bibliography of his
own Flora Anglica (the section of the book headed “Nomina auctorum explicata’) Hudson (1762)
cited Amoenitates academicae IV in which the second edition of Linnaeus’ Flora Anglica was
published during November 1759 (Stearn 1973).

Hudson’s description of St Dabeoc’s heath, like that published a few months later by Linnaeus
(1762), is clearly derived from Ray’s and is not in any sense original. Hudson added no new facts
to the sparse set of words employed by Ray. Thus there is no textual evidence that Hudson had a
specimen, alive or pressed, of the heath. I have never seen a herbarium specimen of St Dabeoc’s
heath bearing his name or annotations — on the other hand, most of Hudson’s herbarium was
destroyed when his house in Panton Street, London, was gutted in a fire in 1783 (Henrey 1975: II,
110; Kent & Allen 1984; Desmond 1994), and it is quite possible whatever specimens he had were
burnt then. If he had no specimen to compare with other species of Erica and Vaccinium, it is hard
to understand why he made the decision to transfer St Dabeoc’s heath into Vaccinium. I suggest he
reasoned that because the illustration published by Petiver (Fig. 1) showed a sprig with broad
leaves and large flowers, more like the large-blossomed, broad-leaved Vaccinium than the usually
small-flowered, needle-leaved heathers (Erica), this plant belonged therein, like going with like.

Thus there is no evidence that Hudson made a detailed analysis of the plant. He employed
previously published descriptions and the illustration from Petiver (1704) as the basis for his
description of Vaccinium cantabricum. He also simply took up the epithet cantabrica from
Tournefort (1694) and the second edition of Linnaeus’ Flora Anglica (1759) as the specific
epithet.

In terms of the current rules of nomenclature, Hudson’s name Vaccinium cantabricum was the
first validly published binomial for St Dabeoc’s heath and thus it has priority, but, I would argue,
that has happened by default because he omitted to be directly explicit about his sources. In his
Flora Anglica Hudson (1762) occasionally omitted to cite the original author and source of
binomials that he published. A good example, one that is discussed by Stearn (1973: 66), is
Hypericum elodes which Linnaeus (1759) had published, and there are other instances of Hudson
apparently using names from Linnaeus’ earlier Flora Anglica (1759) without acknowledgement,
including Trifolium ochroleucon which is in Linnaeus’ work. Comparable examples in the
publications of Hudson’s contemporaries, Philip Miller and John Hill, are Helianthemum
nummularium and Primula elatior respectively (see Brummitt & Meikle 1993).

In the second edition of Flora Anglica, Hudson (1778) accepted Linnaeus’ name Erica daboecii,
quoting the second edition of Species plantarum as the source (Linnaeus 1762) and so he
abandoned Vaccinium cantabricum without even mentioning it in synonymy.

A footnote may be added, by quoting an anonymous reviewer referring to Jussieu’s paper (1802)
in which yet another binomial, Menziesia polifolia, a name that held sway for many decades, was
published:

We cannot refrain from adding, that had our author [Jussieu] thought fit to have retained the
trivial name [i.e. specific epithet] before in use, he would in great measure have avoided the
confusion necessarily attendant on the change of appellation, as Dabeoci, already bandied from
one genus to another, would have been readily recognised under a new generic title.
[Anonymous 1806].

DABOECIA AND/OR DABEOCIA

The generic name Daboecia — based on the name Dabeoc but with the vowels e and 0 reversed — is
conventionally attributed to David Don (1834), but there are several previous works in which it
was printed although not validly published, the earliest being in the second part of Thomas
Martyn’s “corrected and newly arranged” edition of Miller’s The gardener’s and botanist’s
dictionary (the title page of volume | part II is dated 1807; Henrey 1975: II, 91). Martyn included

HISTORY OF ST DABEOC’S HEATH 53

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FIGURE |. St Dabeoc’s heath from Petiver’s Gazophylacii nature et artis decas tertia (1704), tab. 27, fig. 4;
this is designated (p. 55) as the lectotype of Daboecia cantabrica (Huds.) K. Koch.

54 E. C. NELSON

Daboecia as a generic name at the beginning of D, but simply added “See Andromeda” —
Andromeda Daboecia is the ninth species within Andromeda. The implication of Martyn’s edition
of The gardener’s ... dictionary is that the name Daboecia had been coined by 1807, and was
sometimes employed by gardeners and botanists as early as the first decade of the 19th century.

Another intriguing example of its use (with the vowels reversed) before Don formally published
the generic name is in a letter from Richard Duppa of Lincoln’s Inn to Sir James Edward Smith,
President of the Linnean Society, dated 16 July 1827 (Smith 1832, Britten 1916, Nelson 1978).
Duppa wrote:

Some where or other, I think, I have met with the assertion that there was but one Heath
indigenous to Ireland and that is now removed to the genus Dabeecia. My question is, whether,
of the four species of English Heath any one is found wild in Ireland. I have got into a dispute
on that point & I cannot find my difficulty solved in any book I have on the subject of Botany,
& also if our English heaths are not found in Ireland whether the bogs of Ireland are composed
of the roots of the Daboecia, the Salix herbacea, or what? ... And is ye Dabcecia which is
peculiar to Ireland, never found in any other country, that we are acquainted with?

What was Duppa remembering? Was it Martyn’s edition of Miller’s The gardener’s and botanist’s
dictionary, a conversation with a fellow botanist (perhaps David Don, as suggested by Britten
(1916: 179 fn.)), or a now-lost source? Smith’s reply is not extant, so the mystery remains. But the
more extraordinary thing about Duppa’s letter is that it was published, slightly edited, as early as
1832 by Lady Pleasance Smith in her Memoir and correspondence of the late Sir James Edward
Smith (Smith 1832: 2, p. 125). Thus Duppa’s letter was in the public domain two years before
David Don validly published the generic name Daboecia. Lady Smith (1832: 2, p. 125) added this
footnote to the name Dabecia, echoing her late husband’s opinion (Smith 1791):

Mr. Duppa probably meant Menziesia; there is no genus Dabecia. The plant was called
Andromeda Dabecia in Linn. Syst. Veg. It is named Dabeoci after St. Dabeoc, whence the
Linnean trivial name has been corruptly taken.

Given that Daboecia was derived, to quote Smith (1791) again, “corruptly” from the personal
name Dabeoc (Nelson 1984), should the spelling be corrected to Dabeocia? This question has
exercised botanists for more than a century. Article 60.7 of the JCBN (1993) states that “When
changes made in orthography by earlier authors who adopt personal, geographic, or vernacular
names in nomenclature are intentional latinizations, they are to be preserved ...” and among
examples given is Gleditsia L. (1753) which commemorates Gleditsch “... but the name is ... not to
be altered to ... “Gleditschia” ... [because] Linnaeus latinized the names of [this botanist]
deliberately as ... Gleditsius ...”. However the ICBN (1993) also states (Art. 60.1) that “The
original spelling of a name or epithet is to be retained, except for the correction of typographical or
orthographical errors ...”.

Was Daboecia an intentional latinization on the part of Martyn (1807) or even Don (1834), or a
“typographical or orthographical error’? Clearly those authors based the generic name on the
specific epithet taken up by Linnaeus (1754, 1762) from Dillenius’ edition of Synopsis stirpium
Britannicarum and thus ultimately from Ray’s Historia plantarum (1704) (see e.g. Sealy 1949).
Yet Dillenius used dabeoci, as did his predecessors (see Table 2). We cannot explain why
Linnaeus spelled the epithet daboeci (Linnaeus 1754, 1762).

Nicholson (1931) suggested that Don “Latinised [the name] to Dabcecia for convenience’, while
Sealy (1949) argued that Don “knew perfectly well that the saint’s name was Dabeoc, and ... must
have chosen to use the spelling Daboecia quite deliberately.” On the other hand Babington (1889)
stated that he did “not see why Dabeocia should be changed into Daboecia because Don’s printers
made a not unnatural blunder, and [Don] did not discover it when correcting the press.” Jackson
(1889) protested, and argued that Don “clearly meant the name to stand as he wrote it”, Daboecia.
That was not Praeger’s opinion (1925): “... when David Don in 1834 placed the plant in a separate
genus, he followed the Linnean spelling, and called the genus Dabeecia instead of Dabeocia, being
no doubt unaware of the facts of the case.” Alas, no evidence exists to show conclusively whether
Martyn or Don deliberately latinized Dabeoc’s name, or never corrected their printers’ errors.

HISTORY OF ST DABEOC’S HEATH Sp,

TABLE 2. THE TWO VARIANTS OF THE GENERIC NAME AND SPECIFIC EPITHET
DERIVED FROM DABEOC, AND SOME OF THE AUTHORS WHO HAVE USED THESE
VARIANTS; THE LISTS OF AUTHORS ARE NOT EXHAUSTIVE.

Daboecia : daboecil Dabeocia : dabeocii
Linnaeus 1754, 1762, 1767 Lhuyd [1700] 1712
Hudson 1778 Ray 1704
Smith 1791 Dillenius 1724
Jussieu 1802 Threlkeld 1726 [as Dabezoci]
Martyn 1807 More & Moore 1866
[Duppa 1827] Koch 1872
Smith 1832 Babington 1889
Don 1834 Colgan & Scully 1898
Jackson 1889 Chittenden 1931
Webb 1953: 103 [but Dabeocia in Index: 242] Webb [1943: 215]
Woodell 1958 Gilbert-Carter 1950

Webb & Scannell 1983

There is an irrefutable case for correcting the orthography of this name and reinstating Dabeocia
(with e followed by 0). Such an action does not conflict with the JCBN (1993). Indeed Dabeocia
was used by the principal Irish botanical authors (Table 2) until the early 1950s (see e.g. Webb
1943, 1953). While I argue that the name should be corrected, just as Linnaeus’s orthographic
error Stewartia has been corrected to Stuartia (in that case Linnaeus was misinformed about the
surname of the earls of Bute, it being Stuart and not Stewart (see e.g. Stearn 1996)), such a move is
unlikely to gain acceptance. Stability in nomenclature is preferable to instability.

In this context it is worth quoting Chittenden’s (1931) response to Nicholson (1931): there “can
be only one recognised way to spell a name, and we must follow the original, unless to do so
would lead to confusion. In neither of the instances given can following the original lead to
confusion, so we must write Daboecia and Diervilla |after Mons Dierville; see Stearn 1996] in
spite of the probable mistakes on the part of the originators of the names.”

PRONUNCIATION AND TYPIFICATION

Daboecia comes ultimately from an Irish name Dabeoc, and commemorates an obscure saint of
the early Irish church (see Nelson 1984). Dabeoc is pronounced as two equal syllables, da-vok.
How should Daboecia be pronounced? In one sense it is a nonsensical name, and there can be no
correct pronunciation. Webb (1943: 215) suggested DabeOd’cia, while da-b6-ee-kee-a was
suggested by Coombes (1985). Bearing in mind the root Dabeoc, da-vok-ee-a seems most
appropriate, and has the merit of being simple.

Finally, there is no record of typification of the name Daboecia cantabrica. As noted, no
herbarium specimens used by Hudson or Linnaeus in the preparation of their respective floras is
known. Both Linnaeus (1754, 1759, 1762) and Hudson (1762, 1778) cited the same figure, tab. 27,
fig. 4 from Petiver’s Gazophylacii nature et artis decas tertia (1704). This illustration (Fig. 1) can
serve as lectotype; it is the only possible choice.

Daboecia cantabrica (Huds.) K. Koch, Dendrologie vol. 2 (pt 1): 132 (1872).
basionym: Vaccinium cantabricum Huds., Flora Anglica: 143 (1762).

synonym: Erica daboecii L., Species plantarum 509. ed. 2. (1762); Hudson, Flora Anglica:
166. ed. 2. (1778), etc.

Lectotypus hic designatus: icon, tab. 27 fig. 4, in J. Petiver (1704) Gazophylacii nature et artis
decas tertia.

56 EC. NEESON

ACKNOWLEDGMENTS

My particular thanks are due to Gina Douglas, Librarian, Linnean Society, London, for making
available the many eighteenth century sources that form the basis of this paper. I am also grateful
to Dr R. K. Brummitt, Royal Botanic Gardens, Kew, for his invaluable comments and criticisms of
my opinions.

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167-169.

BABINGTON, C. C. (1888). On botanical nomenclature. Journal of botany 26: 369-371.

BRITTEN, J. (1916). An overlooked Irish botanist. Journal of botany 54: 173-180.

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MCCLINTOCK, D. C. (1989). The heathers of Europe and adjacent areas. Botanical journal of the Linnean
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HISTORY OF ST DABEOC’S HEATH 2)

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Creeting St Mary.

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124-127.

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WEBB, D. A. (1943). An Irish flora. \st ed. Dundalgan Press, Dundalk.

WEBB, D. A. (1953). An Irish flora. 2nd ed. Dundalgan Press, Dundalk:.

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NOTES
1 Erica Cantabrica flore maximo, foliis Myrti subtus incanis.
2 Erica Cantabrica, flore maximo, foliis Myrti subtus incanis.

3. 30. Erica S. Dabeoci Hibernica D. Lhwyd. Er. Cantabrica flore maximo, foliis Myrti, subtus incanis
Tournefort. Inst. rei Herb. Erice species genuina esse videtur quoad floris formam, cum Erica tenuifolia
Unedonii flore conveniens. Vasculi seminalis forma Geranium imitari videtur. Cauliculi autem ubi
florere incipiunt non minus viscosi sunt quam Muscipulz, In montibus Mayo squalido & spongioso solo
frequens est, ut & per totum Hiar-Connacht in Gallovidia.

Mulericule superstitiose surculos ejus secum circumferunt adversus incontinentiam.

58 EC. NELSON

4 which simply reads ‘Erica Hibernica foliis myrti pilosis, subtus incanis’

5 4.Erica HIBERNICA fol. Myrti pilosis subtus incanis Hort. nost. sicc. 244. E. 20. Erica S. Dabeoci D.
Lhwyd Ray Dend. V. 3. p. 98. 30. Erica Cantabrica Flore maximo, foliis Myrti subtus incanis Instit. Rei
Herb. 603 5. Elem. Botan. 475. I had a specimen of this elegant Plant from my generous Friend Mr
Edward Lhwyd, Keeper of the Museum Ashmoleanum, who gathered it in Jreland in the places Mr Ray
mentions.

6 Neither of the two editions of Linnaeus’ Flora Anglica (1754, 1759) is listed in the Appendix V (Opera
utique oppressa) of the ICBN (1993) so names published in them are not liable to be regarded as not
validly published under the same CBN (1993, Art. 32.8).

7 ‘Flores non vidi, quos describant autopte.’ In Linnaeus’ own, interleaved copy of this edition (now in the
Linnean Society, London), he has written a very detailed description of the flowers and scored out the
sentence quoted, as well as the phrase ‘Rami breviores.’, and inserted in manuscript after
‘lanceolata’ (referring to the leaves) the additional phrase ‘s[eu] elliptica’. The manuscript description in
this copy was published in Systema naturae (Linnaeus 1767) ; the printed text is almost exactly the same.

8 Quoted from the original manuscript; Smith correspondence, Linnean Society, London.

(Accepted April 1999)

Watsonia 23: 59-81 (2000) 59

Engulfed by suburbia or destroyed by the plough: the ecology of
extinction in Middlesex and Cambridgeshire

C.D, PRESTON

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

ABSTRACT

Middlesex (v.c. 21) has lost 18% of the recorded native vascular plant species through extinction; the
corresponding loss in Cambridgeshire (v.c. 29) is 13%. If species which became extinct before 1750 are
excluded, this amounts to the loss of one species every 1-7 years in Middlesex and one every 2-0 years in
Cambridgeshire. The extinct species share many ecological similarities: they tend to be small, to grow in open
habitats and to be characteristic of environments with low fertility. A disproportionate number of calcifuge
species has been lost in both areas; in Middlesex, where calcareous soils are rare, many marked calcicoles
have also been lost. In both counties the percentage extinction in the small Boreal and Oceanic groups has
been greater than that in the larger phytogeographical elements. The peak periods of extinction in
Cambridgeshire have been those characterised by major agricultural change: in Middlesex they coincide with
the spread of the London conurbation. The overall totals suggest that, judging by the number of extinctions,
agricultural intensification is almost as damaging to the native flora as urbanisation.

KEYWORDS: Life-form, Ellenberg values, phytogeography.

INTRODUCTION

One of the many ways to study the changes in the native flora of Britain during recent centuries is
to look at extinctions at a local level. Over a century ago, West (1898) suggested that no counties
had lost more native species than Middlesex (v.c. 21) and Cambridgeshire (v.c. 29), two areas
Where the landscape had been drastically modified by agricultural and urban change. A
comparison of the nature of the species which have become extinct in these counties ought
therefore to be particularly illuminating. The aim of this paper is to investigate whether the
vascular plant species which disappear when an area undergoes major environmental change have
anything in common either ecologically or phytogeographically, or whether they approximate to a
random sample of the species present in the area.

Middlesex (734 km?) is only a third the size of Cambridgeshire (2124 km7), but the counties
share important geographical similarities . Both areas are exclusively lowland: the highest point in
Middlesex scarcely exceeds 150 metres and nowhere in Cambridgeshire reaches even this altitude.
Neither county has a coastline, though a few coastal species penetrate into Cambridgeshire along
the tidal portions of the River Nene and reach Middlesex along the tidal Thames. There are,
however, important differences in the superficial geology of the two areas. Middlesex is made up
of the predominantly acidic clays, sands and gravels of the London Basin with a small area of
chalk at the edge of the county near Harefield. In Cambridgeshire calcareous soils predominate;
acidic ground is rare. Southern Cambridgeshire is dominated by chalk and calcareous boulder clay,
with only small areas of acidic, sandy soil over the Lower Greensand near Gamlingay at the
western edge of the county and on the fringe of Breckland to the east. The extensive Fenland of
northern Cambridgeshire, which falls below sea-level in places, has no parallel in Middlesex.

A comparison of extinctions in Middlesex and Cambridgeshire is particularly appropriate
because of certain similarities of botanical history. Most importantly, the counties share a long
tradition of field study, sustained in Middlesex by the presence of London with its metropolitan
population and institutions and in Cambridgeshire by the University of Cambridge. In
Cambridgeshire the first thorough investigation of the flora was Ray’s Catalogus plantarum circa

* Like Kent (1975), I have taken the area of the Watsonian vice-counties from the Agricultural Returns for 1873, the
year of publication of Watson's Topographical botany. In this paper the names Middlesex and Cambridgeshire
refer to the Watsonian vice-counties unless the administrative counties are explicitly referred to.

60 C. D. PRESTON

Cantabrigiam nascentium of 1660, followed by another detailed Flora, Relhan’s Flora
Cantabrigiensis (1785). Rather surprisingly, there was no Flora of Middlesex in this period but
very many records from sites in Middlesex were published in national works from those of Turner
(1548) onwards, and these are augmented by local publications (e.g. Blackstone 1737) and
specimens in the London herbaria. The botanical history of the two counties subsequently
converged, with the publication in the mid-19th century of Floras which included both historical
records and the results of recent fieldwork (Babington 1860; Trimen & Dyer 1869). In the 20th
century, Floras of the two counties were again published littlke more than a decade apart,
Cambridgeshire (Perring et al. 1964) again preceding Middlesex (Kent 1975). These have been
supplemented by updated checklists (Crompton & Whitehouse 1983; Kent, in press). Kent’s book
The historical flora of Middlesex (1975) was especially notable for its thorough coverage of the
voluminous historical records available from the county.

IDENTIFICATION OF NATIVE AND EXTINCT NATIVE SPECIES

In investigating the characteristics of the extinct species, I have attempted in this paper to
characterise the native flora of each county as a whole with respect to each ecological or
phytogeographical factor, then to compare this overall pattern with that shown by the species
which are regarded as extinct. Lists of native and of extinct native species for each county were
drawn up from published and unpublished sources, namely Kent (1975 and in press) for Middlesex
and Perring ef al. (1964), Crompton & Whitehouse (1983), Crompton & Wells (1996) and Preston
(1997) for Cambridgeshire. These draft lists were then submitted to the vice-county recorders,
R. M. Burton (v.c. 21) and Mrs G. Crompton & D. A. Wells (v.c. 29) for vetting. Nomenclature
follows Stace (1997).

DEFINITION OF NATIVE SPECIES

The list of British native species provided by Preston & Hill (1997) was used as the basis of the
study. Where there is doubt about whether or not a species is native in Middlesex or
Cambridgeshire, the species was accepted as native if its history, habitat and distribution in the
county is similar to that in those areas of Britain where it was assumed native by Preston & Hill
(1997). Kent (1975) classifies Middlesex plants as native, denizen, colonist, alien or introduced,
and the list of native species accepted here is very similar to the combined list of natives, denizens
and colonists. The Cambridgeshire Floras pay less attention to the question of whether or not
species are native.

IDENTIFICATION OF EXTINCT SPECIES

Species were regarded as extinct if they had not been reported from the county from 1970
onwards, or had been seen between 1970 and 1989 but were known to have disappeared
subsequently. No species seen from 1990 onwards was regarded as extinct, on the grounds that
such a judgement would be premature. Critical segregates which have been recognised recently
from herbarium material collected before 1970 (e.g. Erophila majuscula) were not regarded as
extinct if there has not yet been time to search for them in the wild. In compiling the list for
Cambridgeshire an unpublished list of extinct species prepared by Crompton & Wells (1996) was
particularly helpful, but this needed considerable modification as it includes both native and alien
taxa. The last year in which a species was recorded was noted for each species. By convention,
species which were known in Cambridgeshire to Relhan (1785, 1802, 1820) and Babington (1860)
but not to later authors were regarded as having been last recorded in 1820 and 1860 respectively
in the absence of an exact date for the last record.

In some cases species are extinct as natives but survive as aliens or introductions. These have
been treated as extinct, with the date of extinction the year when a native population was last
recorded. A particular difficulty in Middlesex was the treatment of a number of arable weeds
which are extinct as natives but persist as casuals. The species in this category were treated as
extinct (Anchusa arvensis, Anthemis cotula, Chrysanthemum segetum, Scandix pecten-veneris,
Silene noctiflora, Stachys arvensis). The last record as a persistent weed of cultivated land has
been taken as the last native record, though this is difficult to establish with certainty.

The extinct natives in Middlesex and Cambridgeshire are listed in Appendix 1, with the date of
the last native record.

EXTINCTION IN MIDDLESEX AND CAMBRIDGESHIRE 61
NUMBER AND RATE OF EXTINCTIONS

A total of 146 native species have become extinct in Middlesex since records began, representing
18% of the 816 species recorded. For Cambridgeshire the number of extinct native species is 120,
13% of the 897 species recorded. There are 45 species which have become extinct in both counties
(Table 1).

The number of species last recorded in successive 30-year intervals from 1750 is shown in Table
2. The rate of extinction, as measured by the number of species lost per decade, can be calculated
on the assumption that a species became extinct in the period when it was last recorded. The year
1750 has been chosen rather arbitrarily as the starting point for these calculations. From 1750 to
1809 the rate of extinction was low, with 2-3 species lost from Middlesex per decade and 1-5 from
Cambridgeshire. In the next 30 years the rate of extinction increased dramatically in
Cambridgeshire, with a species lost on average every |-3 years. By contrast the rate actually fell in
Middlesex during this period. This pattern was reversed between 1870 and 1929, when extinctions
in Middlesex were almost twice as numerous as those in Cambridgeshire. Extinctions continued at
approximately the same rate in Middlesex until 1989; in Cambridgeshire the rate increased from
1900 to 1959 to match that in Middlesex.

From 1750 onwards 5-8 species have been lost per decade from Middlesex and 4-9 from
Cambridgeshire; the rates for the 20th century (1900-1989) are 8-4 and 7-3 respectively. One
species has been lost from Middlesex every 1-7 years and from Cambridgeshire every 2-0 years
since 1750.

TABLE 1. ACOMPARISON OF EXTINCT AND SURVIVING SPECIES IN MIDDLESEX (V.
C. 21) AND CAMBRIDGESHIRE (V.C. 29)

Extinct in v.c.21 | Never recorded in v.c. 21 Still present in v.c. 21 Total

Extinct in v.c. 29 45 43 52. 120
Never recorded in v.c. 29 39 - 28 67
Still present in v.c. 29 62 105 610 vee
Total 146 148 670 964

TABLE 2) KATE OF EXTINCTION OF SPECIES IN MIDDLESEX (V.C. 21) AND
CAMBRIDGESHIRE (V.C. 29)

Species are assumed to have become extinct during the period when they were last recorded. The
overall rate of extinction is calculated for the period 1750-1989, and excludes species last
recorded before 1750.

Time period Number of extinct species Rate of extinction (species per decade)
V.C21 Vic. 29 Wecs 21 WeGs 20
pre-1750 8 2 - -
1750-1779 3 e] 1-0 1-0
1780-1809 11 6 3-7 2-0
1810-1839 6 24 2-0 8-0
1840-1869 14 10 4.7 3-3
1870-1899 Di 9 9-0 3-0
1900-1929 23 18 7-7 6-0
1930-1959 24 pH | 8-0 9-0
1960-1989 29 al OF7 7-0
Date unknown ] 0 = :

Total 146 120 5-8 4-9

62 C.D. PRESTON
ECOLOGICAL CHARACTERISTICS OF EXTINCT SPECIES

In this section seven ecological characteristics of the extinct species are examined. Data on plant
height and life-form are derived from the Ecological Flora Database (see Fitter & Peat 1994) with
appropriate additions and corrections. The light, moisture, pH and nitrogen requirements of the
species studied and the typical vegetation in which they grow are assessed by Ellenberg’s indicator
values (Ellenberg 1988; Lindacher 1995). Ellenberg values are available for a maximum of 778
(95%) species in Middlesex and 870 (97%) species in Cambridgeshire, and the results are
therefore based on these species. The values refer to the behaviour of species in central Europe.
Species which have a different habitat in central Europe from that in eastern England will be
misclassified from a British perspective but there are relatively few examples of such differences
(some are discussed in the section on vegetation below). A modified set of Ellenberg indicator
values, corrected where appropriate and applied to all British and Irish species, has recently been
prepared and will be available for future analyses (Hill er al. 1999).

PLANT HEIGHT

Data on the typical maximum height of a species have been used to group terrestrial species into
eight height categories. (Submerged and floating aquatics are excluded from this analysis.) These
categories are those defined for canopy height by Grime ef al. (1988). In both Middlesex and
Cambridgeshire the proportion of short species which have become extinct greatly exceeds the
proportion of taller species (Table 3, Fig. 1).

TABLE 3. EXTINCT SPECIES IN MIDDLESEX (V.C. 21) AND CAMBRIDGESHIRE (V.C.
29) RELATED TO PLANT HEIGHT AND RAUNKIAER LIFE FORM

The totals for plant height exclude aquatic species. Species which occur as two Raunkiaer life-
forms are scored as (} 5 under each.

Middlesex (v.c. 21) Cambridgeshire (v.c. 29)
no. spp. no. extinct % extinct no. spp. no. extinct % extinct

Plant height (cm)

0-10 17 5 29 16 5 3]
11-29 78 24 3] 88 ZA 24
30-59 204 46 23 234 42 18
60-99 209 30 14 224 22 10
100-300 220 28 13 243 19 8
301-600 ibe) 7 pe 0 0
601-1500 13 0 0 12 0 0
>1500 16 0 0 16 0 0
Total 772 134 iy 848 109 13
Life form

Bulbous geophytes 8 3 38 8 2 2a
Other geophytes 52 12 2 64-5 15 de)
Hemicryptophytes S275 47 14 370-5 35 9
Chamaephytes ot 8 22 46 7:5 16
Nanophanerophytes a2 7 22 32 =) 16
Larger phanerophytes 38-5 0 0 37:5 0 0
Epiphytes | 0 0 ] 0 0
Therophytes 197 43 22 ZA TS 30 14
Helophytes 52 12 23 61-5 12 20
Hydrophytes 61 14 23 64-5 13-5 ZA

Total 816 146 18 897 120 13

EXTINCTION IN MIDDLESEX AND CAMBRIDGESHIRE 63

Height Light

ho
r=)

% extinet
vy extinet
i

[=]

| a
4 2 3 4 5 6 7 8 | 2 3 4 5 6 7 8 9g
Height class Ellenberg L value
Middlesex Middlesex

Oo

% extinet
% extinet

meee CU [ 2k Sb eG

Ellenberg L value

Height class

Cambridgeshire Cambridgeshire

FIGURE |. The proportion of extinct species in Middlesex and Cambridgeshire in relation to typical maximum
plant height classes and to Ellenberg light values. Plant height increases from class | to 8 (see Table 3) and

light requirements increase from value | to 9 (see Table 4). In both cases x represents species with a wide
amplitude.

LIFE-FORM

The Raunkiaer life-form system classifies species by the height of the winter buds rather than the
summer foliage (Clapham er al. 1987). Geophytes have resting buds below ground level,
hemicryptophytes at ground level, chamaephytes up to 25 cm above the soil surface and
phanerophytes over 25 cm above the ground. For the purposes of this analysis, nanophanerophytes
(with resting buds 25 cm to 2 m above soil level) are separated from larger phanerophytes (which
have resting buds more than 2 m above the ground). In the Raunkiaer system annuals (therophytes)
form a separate category, as they pass through the unfavourable season as seeds. Marsh plants
(helophytes), water plants (hydrophytes) and epiphytes are also treated separately. The overall life-
form spectrum and the proportion of species in each life-form which have become extinct are very
similar in the two counties (Table 3). A disproportionate number of geophytes, chamaephytes and
nanophanerophytes have become extinct, and in Middlesex therophytes show a greater extinction
than average. Extinctions in the wetland groups also exceed the overall average. The proportion of
extinctions is below average for the hemicryptophytes, the preponderant life-form in both counties,
and there have been no extinctions amongst the larger phanerophytes in either county. The single
epiphyte, the hemiparasitic Viscum album, has survived in both areas.

LIGHT

Ellenberg L values for light range from 1 (plants of deep shade) to 9 (plants only found in full
light); there is also a category for species with a wide amplitude (x). The values for species in
Middlesex and Cambridgeshire (Table 4, Fig. 1) show that the light-loving species in categories 8
and 9 have suffered a disproportionate number of extinctions. In Cambridgeshire there is also a
high level of extinction in one group of shade plants (category 3), attributable to the extinction of
five woodland species (Blechnum spicant, Carex strigosa, Equisetum sylvaticum, Lathraea
squamaria, Polystichum aculeatum) in this small group. Rather surprisingly, the small group of
species of wide amplitude (x) has also lost a disproportionate number of species. This group
includes ecologically tolerant species such as Anemone nemorosa, Festuca rubra and Urtica
dioica which have survived in both counties, but also parasitic or hemiparasitic species such as
Cuscuta epithymum, Melampyrum pratense and Orobanche rapum-genistae which contribute
almost all the extinctions.

64 C2 D: PRESTON

TABLE 4. EXTINCT SPECIES IN MIDDLESEX (V.C. 21) AND CAMBRIDGESHIRE (V.C.
29) RELATED TO ELLENBERG VALUES FOR LIGHT, WATER, pH AND NITROGEN

Ellenberg value Middlesex (v.c. 21) Cambridgeshire (v.c. 29)
no. spp. no. extinct % extinct no. spp. no. extinct % extinct
Light
1 Deep shade i 0 0 1 0 0
2 8 l 13 8 1 1
3 Shade 20 ys 10 18 5 28
4 38 3 8 37 - 11
5 Half shade A7 - . 49 3 6
6 122 14 Lt 123 5 4
7 Partial shade/full light 0 be) AO 15 313 30 10
8 Light-loving 202 al 25 250 43 18
9 Full light only 51 14 ZI 69 i19/ 25
x Wide amplitude 14 3 Zz 14 o 29
Total 778 132 17 869 112. 13
Water
1 Extremely dry 0 0 - 0 0 -
2 16 . Zz 20 1 5
3 Dry 46 14 30 66 - 6
4 143 a | 19 166 18 11
5 Moist 168 15 9 169 13 8
6 68 a 6 73 12 16
7 Damp 58 15 26 65 12 18
8 78 16 21 87 12 14
9 Wet 66 20 30 80 24 30
10 Occasionally flooded 36 6 17 39 6 15
11 Emergent or floating 19 - 21 eA 2 10
12 Submerged 24 | 21 28 i, 25
x Wide amplitude 54 2 - 54 2 ~
Total 776 132 17 868 113 13
pH
| Extremely acid 8 5 63 8 a 88
2 24 5 pA 24 9 38
3 Acid 51 18 35 a2 PA AO
a 46 11 24 A8 6 13
5 Fairly acid 48 7 15 53 i 13
6 71 8 i 77 aT 9
7 Weakly acid or weakly basic 210 32 15 ZN. 24 10
8 126 24 19 160 18 11
9 Calcareous 15 - at 26 3 2
x Wide amplitude 176 16 9 183 10 5
Total TD 130 17 868 112 13
Nitrogen
| Nitrogen-poor 33 11 33 ae 13 35
2 87 29 a8 122 35 29
3 Prefers nitrogen-deficient 89 : 24 ra | 104 17 16
- 90 13 14 96 10 10
5 Average 109 ig 16 128 13 10
6 107 8 7 110 8 7.
7 Prefers nitrogen-rich 105 13 12 jig bs 7 6
8 i 5 7 70 3 4
9 Extremely nitrogen-rich 21 3 14 2A 3 14
x Wide amplitude 59 7 he 61 3 5
Total : vig 130 LZ, 863 112 13

EXTINCTION IN MIDDLESEX AND CAMBRIDGESHIRE 65

WATER

Ellenberg F values for water range from | (indicators of extreme dryness) to 12 (submerged
plants); there is also a category for species with a wide amplitude (x). The values for species in
Middlesex and Cambridgeshire are set out in Table 4. None of the species are classified as
indicators of extreme dryness. In Middlesex more extinctions than average have taken place in a
wide range of groups, especially those of dry-site (values 2, 3) and damp- or wet-site (7, 8, 9)
species. Extinct species of dry sites include some calcicoles (e.g. Helianthemum nummularium,
Scabiosa columbaria) but most are calcifuges (e.g. Potentilla argentea, Teesdalia nudicaulis). The
extinct species in groups 7-9 (e.g. Anagallis minima, Carex binervis, C. dioica, Cicuta virosa,
Cirsium dissectum, Drosera rotundifolia, Mentha pulegium, Oenanthe lachenalii, Parnassia
palustris) would have been found in a wide range of damp or wet habitats. The lowest rates of
extinction are in the species of moist sites (5, 6), large categories which include many woodland
species, and those with a wide amplitude (x). In Cambridgeshire, as in Middlesex, there is a
particularly high proportion of extinctions in the wet-site species (9) and a particularly low
proportion in the species of wide amplitude (x). However, few species in the driest sites (2, 3) have
become extinct.

pH

Ellenberg R values for pH range from | (indicators of extreme acidity) to 9 (always found on
calcareous soils); there is also a category for species with a wide amplitude (x). Both mm Middlesex
and more especially in Cambridgeshire (Table 4, Fig. 2) there has been a massive loss of species
which indicate extreme acidity (value 1). The same eight species were originally present in both
counties. Calluna vulgaris is the only one to survive in Cambridgeshire, whereas Juncus
squarrosus and Pedicularis sylvatica have also survived in Middlesex; Carex binervis, Drosera
rotundifolia, Erica tetralix, Teesdalia nudicaulis and Trichophorum cespitosum are extinct in both.
The extinctions in the other acidic categories (2, 3) are also well above the mean, especially in
Cambridgeshire. In Middlesex the species of basic habitats (9) also show a high level of extinction,
but this is not the case in Cambridgeshire. In both counties the lowest proportion of extinct species
is in the wide-amplitude category (x).

NITROGEN

Ellenberg N values for nitrogen range from | (indicators of sites poor in available nitrogen) to 9
(in extremely rich situations such as areas where cattle congregate); there is also a category for
species with a wide amplitude (x). The proportion of extinct species in Cambridgeshire and
Middlesex (Table 4, Fig. 2) is highly correlated with nitrogen value, especially in Cambridgeshire:
33-35% of the indicators of sites poor in available nitrogen (value 1) have become extinct,
compared to 4—-7% of species characterised by value 8. The exception to this trend is the small
group of species characteristic of extremely rich situations (value 9), which show a higher
proportion of extinctions than those species of somewhat less rich soils. The extinct species are
Beta vulgaris, a coastal plant, and Bidens cernua, Chenopodium glaucum and C. vulvaria.

VEGETATION

Ellenberg’s (1988) classification of species by the characteristic vegetation in which they grow is
based on phytosociological categories, with each species classified by its occurrence in one of
eight major groups. The proportion of extinct species in each vegetation type is again similar in the
two counties (Table 5). The marked losses of coastal and wetland species in both counties is
evident, whereas species of wide tolerance, broadleaved woods and (in Cambridgeshire) scrub and
woodland edges show the least losses. The tiny group of conifer woodland species shows the
highest percentage loss, but of the four species which fall into this category only one (Pyrola
minor) appears from a British perspective to be correctly classified; two of the other three
(Blechnum_ spicant, Monotropa hypopitys) could equally well be classified as species of
broadleaved woodland and the fourth, Carex ericetorum, is a continental species which in Britain
is confined to calcareous grassland. As both counties have lacked native conifer woodland in
historic times they would not be expected to possess native species typical of this habitat. Pyrola
minor may have colonised the county naturally by wind-borne seed, but failed to persist.

In Table 5 the species in the three largest vegetation types are subdivided by phytosociological
class. This shows marked variation in the proportion of extinct species in different aquatic and
wetland classes. The few species of bogs (Oxycocco-Sphagnetea) are all extinct in both counties
and the larger groups typical of acidic mires (Scheuchzerio-Caricetea nigrae) and shallow, acidic

66 C. D. PRESTON

pH Nitrogen

“% extinct
% extinct

| 2 3 4 5 6 £ 8 g 1 7 3 4 5 6 7 8 9

Ellenberg R value Ellenberg N value

Middlesex Middlesex

% extinct
% extinct

{ 2: @ 4 36 °@ 7 “8 o ox 1, 2 St AL Sh, Oe ee
Ellenberg R value Ellenberg N value
Cambridgeshire Cambridgeshine
FIGURE 2. The proportion of extinct species in Middlesex and Cambridgeshire in relation to Ellenberg pH and
nitrogen values. As Ellenberg pH values increase the species become increasingly characteristic of base-rich
soils; nitrogen requirements increase from Ellenberg value | to 9 (see Table 4). In both cases x represents
species with a wide amplitude.

waters (Littorelletea) also show disproportionate extinction. In contrast, species in communities
characteristic of more basic and more eutrophic waters (Potamogetonetea, Phragmitetea) tend to
have survived. There are no marked differences between the two counties. The disturbed ground
community which has lost most species is the Isoéto-Nanojuncetea, found on seasonally flooded
terrain. Otherwise the different communities of disturbed ground show less variation in the
proportion of extinct species within each county. The main differences between counties are the
higher percentage of species in the Secalietea (arable weeds) and Agrostietea stoloniferae (pioneer
plants of damp or flooded sites) which have become extinct in Middlesex. There are also marked
differences between the two counties in the plants of heaths, meadows and pastures. In
Cambridgeshire the greatest losses have been amongst the calcifuge plants in the Nardo-
Callunetea, whereas in Middlesex the calcicolous Festuco-Brometea has lost most species. Losses
in the Molinio-Arrhenatheretea, which contains many species characteristic of moister and often
more nutrient-rich soils, have been low, especially in Cambridgeshire.

DISTRIBUTION AND PHYTOGEOGRAPHY OF EXTINCT SPECIES

DISTRIBUTION WITHIN THE COUNTIES

Have the species which have become extinct in Middlesex and Cambridgeshire always been rare,
or have more widespread species become extinct as well? Answering this question is not
straightforward, as distributional data are presented in different ways in the Floras of the two
counties. In Middlesex, Kent (1975 and in press) followed Trimen & Dyer (1869) in dividing the
county into seven divisions. In Cambridgeshire, Perring et al. (1964) and Crompton & Whitehouse
(1983) list records in 10-km squares. To make the Cambridgeshire data comparable to those
available for Middlesex, seven divisions based on blocks of 5 to 6 10-km squares were defined
(Appendix 2). Thése were broadly based on the eight divisions defined by Babington (1860), with
two of the smaller divisions amalgamated and the areas redefined in terms of 10-km squares.
Details of the number of divisions in which the native and extinct native species have been

EXTINCTION IN MIDDLESEX AND CAMBRIDGESHIRE 67

recorded are provided in Table 6. If a species is recorded as a native in some divisions and as an
alien in others, only the native occurrences have been counted, if possible. However, the
introduced or even casual occurrences of some species are not separable from the native records.

In Middlesex over 50% of extinct species were recorded from only |—2 divisions and over 80%
from 1 divisions. In Cambridgeshire the tendency of rarer species to become extinct is even
more marked, with over 50% of extinct species recorded from a single division and over 80% from
1-3 divisions. Despite this clear general trend some widespread species have become extinct,
especially in Middlesex. Seven of the nine species which were recorded from all seven divisions in
v.c. 21 but are now extinct are arable or other weeds (Anchusa arvensis, Anthemis cotula,
Chenopodium vulvaria, Chrysanthemum segetum, Lithospermum arvense, Ranunculus arvensis,
Scandix pecten-veneris), the exceptions being Ranunculus hederaceus and Spiranthes spiralis. The
only extinct Cambridgeshire species to have been recorded from all divisions is Marrubium
vulgare.

TABLE 5. EXTINCT SPECIES IN MIDDLESEX (V.C. 21) AND CAMBRIDGESHIRE (V.C.
29) RELATED TO -TYPICAL VEGETATION TYPE

Vegetation type Middlesex (v.c. 21) Cambridgeshire (v.c. 29)
no. extinct % extinct no. extinct % extinct
no. spp. spp. spp. no. spp. spp. spp.
1 Freshwater and mires 133 33 pas) 155 oy 24
1.1 Lemnetea 6 1 17 & i tps)
1.2 Utricularietea i i 100 2 Z 100
1.3 Potamogetonetea 5 f) 6 7, 33 3 9
1.4 Littorelletea | 6 Ds) 10 6 60
1.5 Phragmitetea 4§ 6 L3 D2 2 +
1.6 Montio-Cardaminetea 7 1 14 7. Z 29
1.7 Scheuchzerio-Caricetea nigrae 2 9 43 SE) IS 43
1.8 Oxycocco-Sphagnetea 3 5) 100 D 2) 100
2 Saltwater and sea coasts 5) py) 40 28 10 36
3 Frequently disturbed sites 2D 37 16 231 24 10
3.1 Isoéto-Nanojuncetea 14 4 29 1 Bi ©) 38
3.2 Bidentetea 19 2 Dy 19 3 16
3.3 Chenopodietea 52 3} 10 >)! Z d
3.4 Secalietea 34 9 26 45 ) ne
3.5 Artemisietea 59 9 tS D7: 3 e)
3.6 Agropyretea 6 1 IPE 6 if TZ
3.7 Plantaginetea 7; 0 0 ve 1 14
3.8 Agrostietea stoloniferae 24 ii 29 27. | bl
4 Stony sites and walls 8 | 13 8 | 13
5 Heaths and grasslands 189 B)5) 19 DAS 23 11
5.1 Nardo-Callunetea 25 6 24 PAE it 4]
5.2 Sedo-Scleranthetea 36 & 22 40 4 10
5.3 Festuco-Brometea 30 bk GWE 4§ 5) 6
5.4 Molinio-Arrhenatheretea 59 10 ilel 9] 5) »)
6 Scrub and wood edges 27 4 15 32 Z 6
7 Conifer woods and allied heaths 2 1 50 3 yy 67
8 Broadleaved woods 120 11 9 124 9 4
x Wide amplitude 64 6 !) 63 3 5)
Total 773 130 17 855 sila 13

68 C. D. PRESTON

TABLE 6. EXTINCT SPECIES IN MIDDLESEX (V.C. 21) AND CAMBRIDGESHIRE
(V.C. 29) RELATED TO THE NUMBER OF DIVISIONS OF THE COUNTY
FROM WHICH THEY WERE RECORDED

Number of divisions Middlesex (v.c. 21) Cambridgeshire (v.c. 29)
no. spp. no. extinct % extinct no. spp. no. extinct % extinct

Spp.- Spp. Spp. Spp.
| 71 54 76 109 61 56
2 36 20 56 56 22 39
3 37 22 59 Wp 18 25
4 66 29 38 72 11 15
5 58 7 2 88 6 I
6 83 9 ie 93 1 i
7 465 9 2 408 1 <1
Total 816 146 18 897 120 13

For details of the divisions, see Kent (1975) for Middlesex and Appendix 2 of this paper fc
Cambridgeshire.

TABLE 7. EXTINCT SPECIES IN MIDDLESEX (V.C. 21) AND CAMBRIDGESHIRE
(V.C. 29) RELATED TO NATIONAL RARITY

National rarity Middlesex (v.c. 21) Cambridgeshire (v.c. 29)
no. spp. no. extinct % extinct no. spp. no. extinct % extinct

SPP. Spp. SPP. SPP.
Extinct l l 100 3 3 100
Rare De 20 | Sal 13 42
Scarce 46 ps) 63 67 23 34
Other 747 96 13 796 81 10
Total 816 146 18 S97 120 13

There are two reasons why these results should be treated with some caution. It is doubtfi
whether recording of species in the 17th, 18th and early 19th centuries was sufficiently thorough |
ensure that species which became extinct at an early date were recorded in all the districts in whic
they grew. There may therefore be a tendency for extinct species to appear to have been le:
widespread than they were, although the habitat requirements of many of the extinct specie
suggest that they would always have been rare and restricted. In any case, the pattern in bo!
counties is so clear that one has to conclude that the rarer a species was originally, the more like.
it is to have become extinct. The species which appear to have been widespread are mainly weed
and the records in some divisions may have been only casual occurrences.

RARITY OF SPECIES IN BRITAIN :
Three terms are conventionally used to describe the rarest species in Britain: extinct, rare (prese
in | to 15 10-km squares) and scarce (present in 16 to 100 squares). The native and extinct nati
species in the two counties are classified in these categories in Table 7; details of the nation
status are taken from Stewart et al. (1994) and Wigginton (1999). One species formerly recorde
in Middlesex (Arnoseris minima) and three formerly found in Cambridgeshire (Arnoseris minim
Bromus interruptus, Tephroseris palustris) are now extinct in Britain as a whole. Of 22 ra
species which have been recorded from Middlesex only two survive (Arabis glabra, Cyper
fuscus) and over 60% of the scarce species have been lost. A smaller proportion of nationally ra
and scarce species has been lost in Cambridgeshire, although even in Cambridgeshire losses
these species have been much greater than those of species which are more frequent in Britain as
whole. |

EXTINCTION IN MIDDLESEX AND CAMBRIDGESHIRE 69

Whereas the analysis of the rarity of species within the counties is based on all records of the
species, this analysis is based on current national status. It therefore contains an element of
circularity, as species may be rare or scarce because they have declined in counties such as
Middlesex and Cambridgeshire. However, it does at least demonstrate these changes at the local
scale.

PHYTOGEOGRAPHY

Preston & Hill (1997) classify native species according to their latitudinal and longitudinal
distribution in the northern hemisphere. The extinct flora of both counties is analysed in relation to
these phytogeographical elements in Table 8.

There is a clear relationship between the phytogeographical elements and the proportion of
extinct species in both counties. The greatest proportion of extinctions has taken place in the small
Boreal and Oceanic elements. A single Boreo-arctic species (Carex dioica) occurred in both
counties, but has now been lost. In Middlesex five out of six Boreal species have been lost;
Vaccinium myrtillus survives but is reduced to a single plant on Hampstead Heath (R. M. Burton,
in litt. 1999). Similarly eight of the 13 Boreal species in Cambridgeshire have become extinct; the
surviving species are Carex lasiocarpa, Coeloglossum viride, Potamogeton alpinus, P. praelongus
and Salix myrsinifolia. Although less severe, losses of Oceanic and Suboceanic species are also
much greater than the average values.

TABLE 8. EXTINCT SPECIES IN MIDDLESEX (V.C. 21) AND CAMBRIDGESHIRE (V.C.
29) INRELATION TO PHYTOGEOGRAPHICAL AFFINITIES

The percentage of extinct species in each phytogeographical group is given. The values for groups
containing less than 11 species in total are bracketed. For details of the phytogeographical
classification, see Preston & Hill (1997).

Eastern limit category

Major | 2 3 4 5 6
biome category Oceanic Suboceanic European Eurosiberian Eurasian Circumpolar Total
Middlesex
1 Arctic-montane - - : - - - -
2 Boreo-arctic Montane - - - : - (100) (100)
3 Wide-boreal - - (O) (20) (18)

- (100) (0) - (100) (80)

4 Boreal-montane -

5 Boreo-temperate (100) (67) 14 12 19 16 17

6 Wide-temperate - - (QO) 0 (O) (O) 0
7 Temperate oi 30 16 18 14 6 18

8 Southern-temperate (20) 35 17 16 0 (10) 18

9 Mediterranean (44) 8 - - - - 20
Total 50 28 16 15 13 16 18

Cambridgeshire

1 Arctic-montane . - - - - - -

2 Boreo-arctic Montane - - . - - (100) (100)
3 Wide-boreal - - - (O) (0) 25 21

4 Boreal-montane - - (100) (O) - 64 62

5 Boreo-temperate (50) (50) 19 ») 16 24 16

6 Wide-temperate - - - 0 (0) 1 17

7 Temperate 33 By) ) 7 10 13 11

8 Southern-temperate (60) 26 8 8 0 0 11

9 Mediterranean (3) 7 - - - - 21

Total 39 Zi 10 6 10 8) 13

70 CD. PRESTON

EXTINCTIONS AND LAND-USE CHANGE

Although the ecological and phytogeographical characteristics of the extinct species of
Cambridgeshire and Middlesex are broadly similar, the above analysis revealed marked
differences in the timing of the extinctions. The differences are particularly marked in the 19th and
early 20th centuries: extinctions in Cambridgeshire exceeded those in Middlesex between 1810
and 1839, whereas the rate in Middlesex was markedly greater from 1870 to 1929. There is no
obvious explanation of these figures in terms of recording intensity: the recording history of both
counties is broadly similar, and a major Flora of each was published in the 1860s. It is therefore
worth seeking an explanation of these differences in terms of the land-use history of both counties.

CAMBRIDGESHIRE

Cambridgeshire has always been a predominantly agricultural county, and the land-use history
which is most relevant to the botanist is the history of its agriculture. The county lies in one of the
areas of Britain which are most suitable for cereal growing, and, once drained, the fenland soils
support a range of arable crops. Only the clays of the west were once more marginal as agricultural
land.

The period of just over a century from 1640 to 1750, the first for which numerous detailed
botanical records are available, was one in which agriculture was in relative decline in Britain.
Agricultural output increased much more rapidly than did the population. Agricultural prices,
which had increased six-fold between 1500 and 1640, increased by only 2% between 1640 and
1750 (Jackson 1985; Thirsk 1984). Farmers attempted to compensate for this by growing new
crops, and there was a boom in ‘alternative agriculture’ (Thirsk 1997). It was during this period
that drainage of the “Great Level’ of the Fens was accomplished in Cambridgeshire (Darby 1956;
Taylor 1973). The Bedford Level was declared drained in 1652 and Soham Mere, a large Fenland
lake, was drained, enclosed and brought into cultivation in 1664. The fact that ‘alternative’ crops
such as flax, hemp, rapeseed and vegetables could be grown in the newly drained fenland perhaps
encouraged these schemes for agricultural improvement. Continued technical problems resulted in
the deterioration of some of the drained land, especially in the first half of the 18th century when
the incentive for capital investment was low, but these problems were eventually solved by the
application of steam power to pumping engines.

From 1750 onwards the growth in agricultural output in England and Wales slowed, whereas
population grew rapidly, from 6 million in 1750 to 18 million in 1850. Improvements in transport
made it easier for farmers to get their products to the growing urban centres (Mingay 1989).
Consequently there was a prolonged agricultural boom. As agriculture prospered, farmers brought -
more and more ‘waste’ into intensive cultivation. At first the increase in the area of cultivated
grassland exceeded that of arable, but after 1800 arable expanded more rapidly and by 1840 the
acreage of arable in England and Wales actually exceeded that of pasture (Prince 1989). Interest in
alternative crops waned as the rewards of mainstream farming increased.

In Cambridgeshire the medieval open field system survived in many parishes until the end of the
18th century. The first great wave of parliamentary enclosure in the 1760s and 1770s scarcely
affected the county. However, this was not true of the second period when enclosure was
“particularly rampant’, between 1793 and 1815. During this period bad harvests and insecurity
engendered by the Revolutionary and Napoleonic Wars led to great pressure for agricultural
improvement (Turner 1980). A ‘general view’ of the agriculture of the county prepared by Charles
Vancouver (1794) for the newly established Board of Agriculture provided detailed and forceful
arguments for agricultural reform (cf. Grigg 1967). Vancouver’s report was followed by a period
in which many parishes were enclosed. Although the rate of enclosure slowed markedly after the:
end of war with France in 1815, as it did elsewhere in England, there was a period of renewed
activity in the 1820s and 1830s, a feature virtually unique to the county (Turner 1980).
Cambridgeshire is described by Turner as “the county of parliamentary enclosure”. A higher
percentage of land (53%) was enclosed than in any other county except Oxfordshire (54%), but the
Cambridgeshire figure is reduced by the fact that there was little parliamentary enclosure in the
northern, fenland areas. In the south of the county over 70% of the land was enclosed, representing
“the most concentrated agricultural organisational change that there had ever been, certainly in that
county and probably in any county” (Turner 1980).

EXTINCTION IN MIDDLESEX AND CAMBRIDGESHIRE Til

The enclosures in Cambridgeshire were followed by the widespread agricultural improvement
that Vancouver had recommended. Enclosure had a marked effect on the flora of the county
because in its immediate aftermath semi-natural habitats were taken into cultivation: fens were
drained and chalk grassland and heathland ploughed up and converted to arable land. These
reforms provide an explanation of the large number of extinctions between 1810 and 1839.
Corroborating evidence is not hard to find. The Journal of Natural History kept by the Rev.
Leonard Jenyns from 1823 to 1846 reveals “ a recurring concern about the effects of Parliamentary
Enclosures on the extent of heathland and wetland” (Crompton 1997). Babington (1860) had no
doubt why he was unable to refind some species at localities known to Relhan before 1820.
Drosera anglica at Sawston and Hinton Moors and D. intermedia at Teversham and Sawston
Moors had “not [been] found since these places were drained”, Pulsatilla vulgaris and Ulex minor
formerly grew on Barrington Hill, Linton, “a place now ploughed up”, and Vicia sylvatica had
been recorded at Hall Wood, Wood Ditton, which “does not now exist’. There is no evidence from
the study of extinctions to suggest that the intensification of agriculture on existing arable land had
a marked effect on the flora of the county at this period.

The introductory comments in Babington’s Flora (1860) are often cited. In reviewing the county
habitat by habitat, he cited major changes which had happened to each. The Chalk Country which
“until recently (within 60 years)” was “open and covered with a beautiful coating of turf...is now
converted to arable land” and “even the tumuli, entrenchments, and other interesting works of the
ancient inhabitants have seldom escaped the rapacity of the modern agriculturist”. The plants of
the Clayey District “have suffered nearly as much”, and the Fens “have undergone an equally if
not more destructive change than the Chalk district” as steam drainage had rendered the whole
level “a pattern in farming”. The statistics on extinction allow these statements to be seen in their
historical perspective. They are not the exaggerations of a congenitally conservative don, but a
reaction to the effects of a prolonged period of agricultural prosperity and expansion and in
particular a marked increase in the proportion of arable land. This had produced effects on the
native flora which no generation of botanists in Britain had hitherto experienced. Babington’s
Flora of Cambridgeshire is the first local flora in which environmental change is a major theme.

The agricultural prosperity of the 18th and 19th centuries ended relatively abruptly in the 1870s,
when an increasing amount of grain and other agricultural produce became available on world
markets (Perren 1995). The resulting agricultural depression lasted with minor interruptions until
1940. By 1880 the area of arable in England and Wales was again less than that of pasture, and it
continued to fall so that by 1914 East Anglia was the only region in which the proportion of arable
consistently exceeded that of pasture (Whetham 1978). The better land in Cambridgeshire escaped
the worst of the depression, and the agriculture of the county remained overwhelmingly arable
(Fig. 3). However, by the 1930s much of the permanent pasture in the western claylands was
“derelict or nearly so” and the view taken of farming in the area was “necessarily gloomy” (Pettit
1941). This prolonged depression ended with the Second World War and the subsequent
Agriculture Act of 1947, which guaranteed agricultural prices and markets. Further support was
offered to farmers under the Common Agricultural Policy after Britain’s entry into the European
Economic Community in 1973. Financial prosperity was combined with technological advances to
produce a prolonged period of agricultural intensification. The resulting changes in eastern
England included the expansion and intensification of arable farming, with the removal of hedges
to increase field sizes, a switch from spring to autumn sowing of crops, the use of more productive
cereal varieties and increased applications of fertilisers, herbicides and pesticides. Remaining areas
of grassland were also converted from species-rich semi-natural communities to intensively
managed, heavily fertilised, species-poor swards (Stoate 1996). A detailed account of the changes
in one Cambridgeshire parish during this period is given by Sell (1989).

The rate of extinction of species in Cambridgeshire was very low between 1840 and 1899. This
doubtless reflects the completion of agricultural enclosure followed by the end of the long period
of agricultural prosperity. It may also be influenced by the fact that there was little systematic plant
recording in southern Cambridgeshire after the completion of Babington’s Flora. The rate of
extinction increased in the period from 1900 and 1929, and it further increased from 1930
onwards. The increase from 1900 to 1929 has no clear explanation in terms of national land-use
history, but the later increase coincides with the post-war agricultural revolution. The two decades
in the 20th century in which most species were lost were the 1950s (twelve species) and the 1960s
(eleven species), although it could be argued that species which were last seen in this period are

2 C. D. PRESTON

100
50 =
&
i
= 60 7
= | Non-agricultural
op 1 Grass
= 40 @ Arable
YS
o
all
20
@) -
(oO Lo) LY) LO LO LO LO LO) LO) LO LO
ce) ~~ co Oo O = —N fo8) wT ire) ©
co co co co Oo oO) >) Oo (>) Oo [S))
Year

FIGURE 3. The percentage of arable land, permanent grassland (including rough grazing) and non-agricultural
land in the administrative county (or counties) of Cambridgeshire and the Isle of Ely, 1866-1970. Values are
plotted for 1866 and then at 5-year intervals from 1870 to 1970. They are based on official agricultural returns
extracted by Stamp (1941) for the period between 1866 and 1910 and by C.D.P. from 1915 onwards. Radical
revisions to the administrative county boundary took place in 1974.

more likely to be rediscovered than those last seen in earlier decades. In some cases the extinctions
which have occurred since 1930 may have been the long-delayed result of earlier changes:
Potamogeton polygonifolius was last seen at Gamlingay in 1948, for example, long after the
extinction of the more exacting calcifuges there. In other cases the loss of species can be attributed
directly to agricultural intensification, such as the ploughing up of the county’s only site for
Cirsium tuberosum in 1973 (Pigott 1988). However, most extinctions are probably the result of
less dramatic, gradual changes brought about by factors such as eutrophication and falling water
tables.

MIDDLESEX

The periods of maximum extinction in Cambridgeshire are closely related to periods of
agricultural improvement. Can the same relationship between the number of extinct species and
land-use history be demonstrated in Middlesex? Until it was swamped by the growth of London,
Middlesex, like Cambridgeshire, was an almost exclusively agricultural county (Tanner 1911).
The more fertile soils are suitable for arable cultivation, and in Elizabethan times this land, “fat
and fertile and full of profite’, produced “most excellent good wheate” (Norden 1593). However,
although arable cultivation persisted on the best soils, agriculture in the county became
predominantly pastoral between 1550 and 1750 in response to the insatiable demand for hay in
London. Thus Perivale, which was praised for its wheat by Norden and in Michael Drayton’s
poem Poly-Olbion (1612), was to be remembered by John Betjeman in his poem Middlesex as a
“parish of enormous hayfields” (Betjeman 1954). Hay was required for the city’s horses and for
cows kept in urban cowsheds; the name Haymarket (Piccadilly) survives from one of four new hay
markets set up in the city in the 17th century. The refuse of the city provided manure which was
returned to the countryside to sustain production. Close to London extensive areas of market
gardens developed, so that the area west of St Martin-in-the-Fields appeared to be “a continuous
garden” (Richardson 1984). Pehr Kalm, visiting London in 1748, explained that “the frightful
number of people which there crawl in the streets pays the market gardeners many fold their
labour and outlay” (Lucas 1892). A detailed account of the market gardens of Middlesex, and of
other aspects of the changing land-use of the county, is given by Bull (1957).

EXTINCTION IN MIDDLESEX AND CAMBRIDGESHIRE 13

Some of the open countryside of Middlesex was lost before Parliamentary enclosure to non-
agricultural uses such as roads, parks and buildings. Parliamentary enclosure took place earlier
than in Cambridgeshire: 28% of the total area enclosed was dealt with by 1793 and 91% by 1815,
compared to 6% and 60% in Cambridgeshire (Turner 1980). The proportion of the total area of
Middlesex subject to enclosure, 28%, was much less than the 53% figure for Cambridgeshire. A
notable feature of the enclosure of Middlesex was the conversion of commons and wastes,
including much heathland, to agricultural use. The agricultural improvers of the late 18th century
regarded the presence so close to the capital of unimproved land such as Hounslow Heath as
“disgraceful to the county, and insulting to the inhabitants of the metropolis” (Middleton 1798).
The enclosure of these commons was the culmination of a long process of attrition that reduced the
area of common land from approximately 45,000 acres in 1500 to 30,000 acres in 1760; most of
these 30,000 acres had been enclosed by 1825 (Kent 1975).

After enclosure the assured profits obtained by catering for the predictable needs of London led
to an agricultural regime which appears to have been relatively stable. In Trimen & Dyer’s (1869)
Flora the farmers of Middlesex bear no relationship to the rapacious agriculturalists of
Babington’s Cambridgeshire. Basing their account on Clutterbuck (1869), they describe the
hayfields over London clay as “small and not very convenient enclosures, sometimes overgrown
with timber in the hedgerows’. Clutterbuck (1869) applied Norden’s (1593) words to the hay
farmers of the 1860s: “they only covet to maintaine their ancient course of life and observe the
husbandrie of their fathers without adding any thing to their greater profit”.

By 1869, however, it was apparent to Trimen & Dyer that the market gardens and orchards close
to London were “giving way to the advancing wave of buildings more rapidly than they are
replaced”. The population of London was approximately 400,000 in 1650, 575,000 in 1700 (when
it overtook Paris as the largest city in Europe), 675,500 in 1700 and 900,000 in 1800 (Wrigley
1967). But in the 19th century its population more than trebled in size, reaching 3-56 million in
1901 and growing further to 5 million by 1961. It was in the second half of the 19th century and
more particularly the first half of the 20th that the large town expanded to become the “immense
continuous urban area” of Greater London (Figs 4, 5).

100 -

80

60

(1 Non-agricultural

C1 Grass
@ Arable

40

Percentage of area

20 -

0
© LY} “ly Ly} Ly Lo) Lo} is) Lt} “uy vt
i<e) hh [= @) (oy) So = MN Cy wT ite) (Oo
oO fag) i= @) i= 6) (oy) (ey) (oy) [o>] (ey) GO (ey)
Da 5 ia ba L, Tae a 5 a eee — ae 2 ae bi, ia
Year

FIGURE 4. The percentage of arable land, permanent grassland (including rough grazing) and non-agricultural
land in the administrative counties of Middlesex (1866-1891) and Middlesex plus London (1892-1964).
Values are plotted for 1866 and then at 5-year intervals from 1870 to 1960 and finally for 1964. They are
based on official agricultural returns extracted by Willatts (1937) for the period between 1866 and 1935 and
by C.D.P. from 1940 onwards. From 1949 onwards the agricultural land in the county of London was
apportioned between Kent, Middlesex and Surrey rather than included in the Middlesex total, but by then it
represented less than 1.5% of the total area of the county of London so that the change has no appreciable
effect on the diagram.

74 C. D. PRESTON

6000

50G0

4000

3000

2000

1000

People per square kilometre

0
be — se B pees —_— ie = i aad bo L ics
© N w [<e) (ee) (=) WN vr ice) ee)
(ee) (ee) [0 @) (oe) (oe) (o>) (o>) (o>) [e>) [@>)
= bs ae = ed rT raed boa geo bd b ae
Year

FIGURE 5. Population density (people per square kilometre) in Middlesex (M), extra-metropolitan Middlesex
(EM) and Cambridgeshire (C), derived from decadal census returns from 1801 onwards. There was no
enumeration in 1941 and this value was replaced by a retrospective estimate for 1939. Figures for Middlesex
refer to the ‘ancient geographical county’, in effect the vice-county, until 1901. Subsequently the Middlesex
figures are derived by adding the values for the revised administrative county of Middlesex to the figures for
those areas of the county of London which were formerly in Middlesex (the City of London and 18
metropolitan boroughs). It is not possible to derive an estimate for 1939 by this method and the value plotted
is therefore the mean of the 1931 and 1951 values. Extra-metropolitan Middlesex is the area of the county
outside the former county of London; retrospective population estimates for extra-metropolitan Middlesex
were published with the 1961 census returns. Figures for Cambridgeshire refer to the administrative county
until 1971, and subsequently to four districts (Cambridge, E. Cambs., S. Cambs., Fenland) which approximate
to the administrative county. The administrative and vice-counties are broadly similar (Crompton &
Whitehouse 1983).

The expansion of London was made possible by improved rail and then road transport, and by the
ability of engineers to solve the water supply problems which had hampered urban development
on the London Clay (Smailes 1964; Willatts 1937). The growth of London “resulted in the greater
part of the vice-county being virtually obliterated by a huge torrent of bricks and mortar and the
creation of vast new suburbs which engulfed most of the old Middlesex villages and almost
eradicated the London Clay grass plain... The rapid spread of suburbia has so reduced the
agricultural areas of the vice-county that today only a minimum remains” (Kent 1975). In addition
to the spread of suburbia, the county has been affected by other changes including gravel
extraction from the river valleys, which has been directly responsible for the extinction of some
species, and the growth of Heathrow Airport.

The land-use history of the county therefore ties in well with the statistics on extinctions. The
initially higher rate of extinction when compared to Cambridgeshire is explicable in terms of early
agricultural change, but during the period 1810—1839 the rate fell behind as the intense double
wave of enclosures in Cambridgeshire was not matched by such radical developments in
Middlesex. Thereafter the extinction rate in Middlesex picked up again, reflecting the growth of
London: the primary effect of habitat destruction and all the secondary consequences of
urbanisation.

EXTINCTION IN MIDDLESEX AND CAMBRIDGESHIRE es)

DISCUSSION

ACCURACY OF THE LISTS

The analysis in this paper rests on lists of the native and extinct native species of Middlesex and
Cambridgeshire. In some cases where the evidence of the occurrence of a species rests solely on
old records there is an element of doubt about the correct identification of the species or its
geographical location. Limonium bellidifolium, for example, is traditionally included in Floras of
Cambridgeshire on the basis of a locality discovered by W. Skrimshire, the salt marshes below
Wisbech, but three vice-counties meet in this area and there is considerable doubt as to whether the
plant actually grew in Cambridgeshire (G. Crompton, in litt. 1999); the species is therefore
excluded from this paper. The flora of both Middlesex and Cambridgeshire is very well-known,
and the rate at which new native species are discovered is now very low. In over 20 years since the
publication of Kent’s Flora in 1975, Galium constrictum and Salvia pratensis have been added to
the Middlesex flora as a result of the discovery of overlooked herbarium specimens, Erophila
glabrescens, E. majuscula and Juncus ambiguus were also found in herbaria following taxonomic
revisions, and two new native species were detected in the wild, the ferns Dryopteris affinis and
Polypodium interjectum (Kent, in press; Rich & Lewis 1999). Similarly in Cambridgeshire the
only native vascular plants to be added to those listed in the 1983 Checklist have been three
species represented by 18th or 19th century herbarium specimens (Eriophorum vaginatum, Rosa
pumpinellifolia and Ruppia cirrhosa, the last known previously from literature references which
were misinterpreted as R. maritima) and a single species discovered in the wild, Dactylorhiza
maculata. The definition of what constitutes a native species is a greater area of uncertainty than
under-recording of species in the field. In this analysis the list of native species made use of that
given by Preston & Hill (1997), which in turn largely followed Stace (1991), in order to maintain
comparability with the national study. There are certainly changes which could and perhaps should
be made to this list, but they are very unlikely to alter any of the conclusions reached in this paper.

Although the list of native species can be regarded as relatively stable, experience suggests that
the list of supposedly extinct species will change. At the vice-county level, the slogan ‘extinction
is for ever’ is manifestly false. Species may be discovered at new sites or rediscovered at sites
where they have been overlooked, they may arise from seed banks after vegetation change or they
may reinvade the county by natural means (although recolonisation 1s less likely to be a significant
factor for flowering plants than for cryptogams, which have more mobile propagules). Babington
(1860, pp. 314-315) listed 61 “lost plants” which had not been recorded in Cambridgeshire for
many years or which, although recorded more recently, were “also probably now extirpated’. Only
42 of these 61 species are natives of Cambridgeshire and, of these 42, 22 (53%) are still thought to
have become extinct before 1860, 6 (14%) were rediscovered but are now regarded as extinct and
14 (33%) are still present in the county. Similarly, Trimen & Dyer (1869, pp. 345-346) reported as
extinct 38 species which are accepted as natives of Middlesex in this paper. Of these 38, only 17
(45%) are still thought to have become extinct before 1869; the remaining 21 species were later
rediscovered and 13 (34%) of these are still present, although 8 (21%) are again thought to be
extinct. The rediscovery of supposedly extinct species has continued into modern times: 13 native
species which Kent (1975) regarded as extinct had been rediscovered by the time he prepared the
Supplement (Kent, in press). James (1997) has documented the same phenomenon in
Hertfordshire. One must therefore expect some species listed as extinct in Appendix | to be
rediscovered. There may also be species which have been seen in one or other county since 1990
and are therefore regarded as extant, but which will never be seen there again.

The extent to which the dates of the last records of extinct species reflect the actual timing of
extinctions also needs to be assessed. These dates might potentially be influenced by the history of
plant recording in each county. Even if the rate of extinction in a county was absolutely constant,
fluctuations in the intensity of recording would nevertheless lead to an apparent concentration of
last records in periods of intensive recording. It is, however, unlikely that the main fluctuations in
extinction rate suggested by the data in Table 2 can be explained in this way. In Middlesex the rate
in the 1840-1869 period was low despite the publication of Trimen & Dyer’s Flora in 1869;
subsequently the rate has been surprisingly constant. In Cambridgeshire the high rate from 1810 to
1839 cannot be explained by a subsequent falling off of recording intensity as the next period,

76 C. D. PRESTON

from 1840 to 1869, was that in which Babington prepared his Flora. There is scope for a more
detailed study of extinction in each county, which would attempt to estimate dates of extinction of
those species which have succumbed to habitat destruction by the date of the destruction of the last
site rather than by the date of the last botanical record.

Even if the date of extinction of a species is known with certainty, it will not necessarily
coincide with the date of the causal event. Plants may persist for many decades after gradual
changes which render sites unsuitable for the sexual or vegetative reproduction of a species but are
insufficient to eradicate it completely (Summerfield 1972). However, catastrophic changes such as
total habitat destruction may eliminate a species at once. Such changes were characteristic of the
agricultural revolution which accompanied parliamentary enclosure in Cambridgeshire, and
doubtless explain why this period stands out so strongly in the record of extinctions. In the absence
of radical change, the correlation between land-use history and the rate of extinction will be less
clear-cut.

ECOLOGICAL AND PHYTOGEOGRAPHICAL CHARACTERISTICS OF EXTINCT SPECIES

A high proportion of the extinct species are small, grow in open habitats, favour substrates with
low pH or are characteristic of environments with low fertility. This is true of both counties,
although the patterns are sharper in Cambridgeshire than in Middlesex (Figs 1, 2). Although these
attributes have been treated separately, there is clearly a correlation between them. Habitats which
have low pH, for example, generally have low fertility and a vegetation composed of low-growing
species. Such semi-natural habitats have often been destroyed by agricultural improvement and
urban expansion, and any remaining remnants have been indirectly modified by factors such as the
reduction in grazing in an increasingly arable or urban landscape, eutrophication, falling water
tables and increasing disturbance from a larger and more mobile population.

Twelve species combine all four attributes of the least successful species (typical maximum
height <30 cm and Ellenberg L=8—9, R=1-3, N=1-—2). Of these, four species are extinct in both
counties (Drosera intermedia, D. rotundifolia, Lycopodiella inundata, Lycopodium clavatum),
four more are extinct in the only county from which they are recorded (Antennaria dioica, Drosera
anglica, Hammarbya paludosa, Radiola linoides) and two are extinct in one county but survive in
the other (Lythrum portula extinct in Cambridgeshire and Vicia lathyroides in Middlesex).
Surprisingly, two species in this group not only survive in both counties but even persist in central
London, Aira praecox and Trifolium arvense (Kent, in press).

In an analysis of a completely different dataset, the results of the B.S.B.I. Monitoring Scheme,
Thompson (1994) demonstrated a marked correlation between low canopy height and a tendency
to decline in England. He pointed out that canopy height is correlated with competitive ability, so
the declining species tend to be the less competitive. He commented on the “increasing restriction
of slow-growing plants of infertile, relatively undisturbed habitats to fragmented islands of suitable
habitat, many of them in nature reserves, surrounded by a sea of unsuitable landscape”’.
Thompson’s conclusions about the nature of the species which have declined in the modern
landscape are strikingly similar to those of this survey.

In addition to these trends, there is a consistent indication in Cambridgeshire and Middlesex that
wetland and coastal species have suffered more than plants in other habitats. The decrease in
wetland species has been noted in other studies (e.g. Dony 1977; James 1997; Mountford 1994).
The tendency of coastal species to decrease almost certainly reflects the very small area of suitable
habitat for such species in these counties rather than a national trend. Judging by trends shown by
nationally scarce species, the distribution of coastal species is relatively stable (Stewart ef al.
1994). The persistence of the woodland flora of both counties is striking.

Some of the differences between Cambridgeshire and Middlesex reflect the differing geology of
the two counties. Only in Middlesex do the species of basic habitats show a high level of
extinction, a reflection of the small area of calcareous soil in that county. One might expect a
correspondingly greater survival of calcifuge species in Middlesex, but this is only true to a limited
extent. There has been massive extinction in calcifuge species and plants characteristic of plant
communities of acidic soils in Cambridgeshire. However, the loss of these species has also been
greater than average in Middlesex. Even though Middlesex is dominated by acidic soils, the
heathland habitats of many species have been destroyed or grossly altered.

The most striking feature of the phytogeographical analysis is the loss of species from groups at
the edge of their geographical range, the Boreal, Mediterranean-Atlantic and Oceanic plants. The

EXTINCTION IN MIDDLESEX AND CAMBRIDGESHIRE ig

Same pattern is revealed by a recent analysis of extinctions in Bedfordshire (Boon 1999). If such
changes are indeed characteristic of south-east England as a whole, they must have led to a
reduction in the phytogeographical diversity of the flora and the increasing predominance of
species in the already dominant major biome classes. However, it must be remembered that this
applies only to native species. The increase in alien species of diverse origins is an aspect of
change which has not been examined in this paper, but must be considered in an overall
assessment of the phytogeographical diversity of the area.

EFFECTS OF URBANISATION AND AGRICULTURAL IMPROVEMENT

The similarities in the extent and nature of extinctions in Cambridgeshire and Middlesex are much
more striking than the differences. This is remarkable in view of the differing land-use history of
the counties: Middlesex has become a suburban and urban county, whereas Cambridgeshire is still
rural although very intensively farmed. The fact that 18% of species have been lost from
Middlesex and 13% from Cambridgeshire suggests that agricultural intensification is almost as
damaging to the native flora as urbanisation. In making these comparisons it is difficult to account
for the effect of the differing size of the counties. If the counties were uniform, one might expect
fewer extinctions in the larger county, but the highly heterogeneous nature of Cambridgeshire with
distinct areas of chalk, clay and fenland negates such a simplistic approach. Thompson & Jones
(1999) found little relationship between the number of extinct scarce species and the area of a
vice-county as such.

An alternative explanation for the similarity of the changes in the two counties might be that the
changes in each have the same cause — that in both counties the primary cause is urbanisation.
Thompson & Jones (1999) found a strong relationship between the number of nationally scarce
species that have been lost from vice-counties and their current population density. They point out
that population density is an indicator of many aspects of land-use (including road-building,
recreation and urbanisation) which have a negative impact on plant survival. They go on to suggest
that, whereas intensive agriculture “is frequently assumed to be the main cause of decline in
Britain’s native wildlife’, these effects of population pressure may be “more pervasive and
widespread than suspected” and “a major cause of local plant extinction”. They conclude that a
more detailed search for the proximate causes of local plant extinction would be rewarding.

The study reported in this paper differs from that of Thompson & Jones (1999) in being more
limited spatially, but benefits from incorporating a temporal dimension. There has been a marked
variation in the rate of extinction in differing time periods in both Middlesex and Cambridgeshire,
and this provides a powerful insight into the possible causes of such extinctions. There is evidence
to support Thompson & Jones’ view that urbanisation has been a major cause for species loss in
Middlesex (indeed, the urbanisation of that county is so extreme that no other conclusion would be
credible). However, the analysis for Cambridgeshire has demonstrated the importance of
agricultural change as an agent of local extinction. These variations in the rate of extinction also
highlight the desirability of separating historic and recent extinctions when considering the
implications of previous changes for the conservation of the modern flora. Causal factors as well
as rates may have varied over time, and the fact that historic extinctions in Cambridgeshire have
been primarily caused by agricultural improvement does not mean that the effects of population
pressure described by Thompson & Jones (1999) are not important now, or will not become
apparent in the future.

Further detailed studies of extinction at the vice-county level can be expected to throw more
light on the changes which have taken place in the British flora. The vice-county is a convenient
unit to use in the analysis of such changes, and such analyses are completely dependent on critical
compilations of historic records such as that undertaken by Duggie Kent for The historical Flora
of Middlesex.

ACKNOWLEDGMENTS

I am very grateful to R. M. Burton, Mrs G. Crompton and D. A. Wells for help in drawing up the
lists of native and extinct native species, although I take responsibility for the many, sometimes
somewhat arbitrary, decisions which have had to be made in finalising the lists. The ecological and
phytogeographical data used in this paper have been taken from ORACLE databases held at LT.E.

78 C. D. PRESTON

Monks Wood and I thank H. R. Arnold, D. B. Roy and Dr M. G. Telfer for help with computing.
C. R. Boon, R. M. Burton, Prof. C. A. Stace and Dr K. Thompson kindly helped me obtain copies
of unpublished books or papers, and Julie Gaunt generated the diagrams. R. M. Burton, Mrs J. M.
Croft, Mrs G. Crompton, P. T. Harding, Dr M. O. Hill and P. H. Oswald kindly commented on a
draft of this paper. I am also grateful to Dr O. Rackham and Dr D. L. Kelly for their advice as
referee and editor, respectively. I appreciate the opportunity of preparing this paper for an issue
dedicated to D. H. Kent, whose detailed work on the Middlesex flora provides one basis for the
study. I was grateful for the cheerful welcome this doyen of the B.S.B.I. extended to me as a naive
newcomer, often benefited from his irreverent observations and gradually came to appreciate the
enormous amount of work he undertook for the Society with so little fuss.

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(Accepted June 1999)

80 C. D. PRESTON

APPENDIX 1. EXTINCT NATIVE SPECIES OF MIDDLESEX AND CAMBRIDGESHIRE

The extinct species are listed below, with the date of the last record.

MIDDLESEX (V.C. 21)

Agrimonia procera, 1874; Alchemilla filicaulis, 1970; Allium oleraceum, 1856; Althaea officinalis,
1760; Anagallis minima, 1800; A. tenella, 1873; Anchusa arvensis, 1966; Anthemis cotula, date
not known; Aquilegia vulgaris, 1737; Arnoseris minima, 1778; Barbarea stricta, 1965; Beta
vulgaris, 1887; Blysmus compressus, 1830; Bupleurum tenuissimum, 1860; Campanula glomerata,
1980; Carduus tenuiflorus, 1929; Carex appropinquata, 1936; C. binervis, 1983; C. dioica, 1792;
C. filiformis, 1960; C. laevigata, 1830; C. pulicaris, 1946; C. rostrata, 1792; Carlina vulgaris,
1730; Chenopodium glaucum, 1953; C. vulvaria, 1970; Chrysanthemum segetum, 1978;
Chrysosplenium alternifolium, 1838; Cicuta virosa, 1780; Cirsium dissectum, 1884; Cochlearia
anglica, 1869; Coeloglossum viride, 1913; Crepis foetida, 1873; Cuscuta epithymum, 1887; C.
europaea, 1960; Cynoglossum germanicum, 1710; C. officinale, 1955; Damasonium alisma, 1886;
Dianthus armeria, 1948; D. deltoides, 1939; Drosera intermedia, 1746; D. rotundifolia, 1935;
Eleocharis multicaulis, 1960; Erica cinerea, 1981; E. tetralix, 1970; Eriophorum angustifolium,
1980; Euphorbia platyphyllos, 1884; Euphrasia arctica, 1884; E. confusa, 1887; Filago
pyramidata, 1891; Fritillaria meleagris, 1950; Fumaria purpurea, 1892; Galeopsis angustifolia,
1945; Galium constrictum, 1780; Gentiana pneumonanthe, 1795; Gentianella amarella, 1975; G.
germanica, 1977; Gnaphalium sylvaticum, 1945; Gymnadenia conopsea, 1965; Helianthemum
nummularium, 1907; Hordelymus europaeus, 1951; Hypericum androsaemum, 1947; H. elodes,
1826; Hypochaeris glabra, 1843; Jasione montana, 1935; Juncus ambiguus, 1842; Juniperus
communis, 1746; Lactuca saligna, 1800; Legousia hybrida, 1914; Leucojum aestivum, 1845;
Lithospermum arvense, 1888; L. officinale, 1914; Littorella uniflora, 1935; Lycopodiella inundata,
1869; Lycopodium clavatum, 1865; Lythrum hyssopifolia, 1778; Maianthemum bifolium, 1924;
Marrubium vulgare, 1949; Melampyrum arvense, 1870; Mentha pulegium, 1871; Myosotis
secunda, 1847; Myosurus minimus, 1914; Myrica gale, 1901; Myriophyllum alterniflorum, 1957;
Neottia nidus-avis, 1946; Nepeta cataria, 1978; Oenanthe lachenalii, 1847; Ophrys insectifera,
1907; Orchis militaris, 1900; O. ustulata, 1737; Orobanche elatior, 1902; O. rapum-genistae,
1873; Osmunda_ regalis, 1794; Parnassia_ palustris, 1900; Pedicularis palustris, 1920;
Petroselinum segetum, 1930; Pilularia globulifera, 1800; Platanthera bifolia, 1887; P. chlorantha,
1890; Polystichum aculeatum, 1965; Potamogeton acutifolius, 1981; P. alpinus, 1886; P.
compressus, 1917; P. polygonifolius, 1884; P. trichoides, 1966; Potentilla argentea, 1916;
Pulicaria vulgaris, 1908; Pyrola minor, 1908; Radiola linoides, 1890; Ranunculus aquatilis, 1918;
R. arvensis, 1966; R. fluitans, 1977; R. hederaceus, 1974; R. parviflorus, 1885; R. sardous, 1912;
Rosa agrestis, 1818; R. micrantha, 1910; R. pimpinellifolia, 1887; Sagina nodosa, 1906; S.
subulata, 1783; Salvia pratensis, 1901; Samolus valerandi, 1866; Scabiosa columbaria, 1939;
Scandix pecten-veneris, 1888; Schoenoplectus triqueter, 1887; Scrophularia umbrosa, 1841;
Silene noctiflora, 1967; Sium latifolium, 1965; Sonchus palustris, 1835; Spiranthes spiralis, 1936;
Stachys arvensis, 1954; Stellaria palustris, 1977; Teesdalia nudicaulis, 1932; Trichophorum
cespitosum, 1780; Trifolium ornithopodioides, 1885; T. squamosum, 1721; Utricularia minor,
1746; Valeriana dioica, 1970; Valerianella dentata, 1950; V. rimosa, 1941; Vicia lathyroides,
1866; Viola canina, 1966; V. lactea, 1868; V. palustris, 1912; Vulpia unilateralis, 1969; Wolffia
arrhiza, 1898.

EXTINCTION IN MIDDLESEX AND CAMBRIDGESHIRE 81
CAMBRIDGESHIRE (V.C. 29)

Alchemilla filicaulis, 1953; Alisma gramineum, 1972; Althaea officinalis, 1959; Anagallis minima,
1820; Antennaria dioica, 1864; Aphanes inexspectata, 1977; Armeria maritima, 1930; Arnoseris
minima, 1914; Atriplex pedunculata, 1826; Beta vulgaris, 1928; Bidens cernua, 1964; Blechnum
spicant, 1660; Botrychium lunaria, 1833; Bromus interruptus, 1972; Campanula latifolia, 1852;
Carex binervis, 1961; C. curta, 1853; C. diandra, 1860; C. dioica, 1841; C. echinata, 1954; C.
pilulifera, 1949; C. rostrata, 1966; C. strigosa, 1795; Catapodium marinum, 1785; Ceterach
officinarum, 1967; Chenopodium urbicum, 1837; C. vulvaria, 1958; Cicuta virosa, 1763; Cirsium
tuberosum, 1974; Colchicum autumnale, 1926; Crassula tillaea, 1930; Cuscuta epithymum, 1975;
Drosera anglica, 1840; D. intermedia, 1820; D. rotundifolia, 1913; Eleocharis multicaulis, 1853;
Eleogiton fluitans, 1975; Epilobium palustre, 1969; Epipactis phyllanthes, 1987; E. purpurata,
1962; Egquisetum sylvaticum, 1763; Erica cinerea, 1914; E. tetralix, 1920; Eriophorum
angustifolium, 1960; E. latifolium, 1887; Festuca filiformis, 1964; Frankenia laevis, 1820;
Fritillaria meleagris, 1820; Genista anglica, 1932; Gnaphalium luteoalbum, 1802; Hammarbya
paludosa, 1855; Herminium monorchis, 1825; Hordeum marinum, 1881; Hypericum elodes, 1930;
Hypochaeris glabra, 1964; Jasione montana, 1910; Juncus squarrosus, 1833; Lactuca saligna,
1953; Lathraea squamaria, 1889; Lepidium heterophyllum, 1965; L. latifolium, 1795; L. ruderale,
1831; Limosella aquatica, 1877; Liparis loeselii, 1945; Littorella uniflora, 1820; Lycopodiella
inundata, 1853; Lycopodium clavatum, 1831; Lythrum portula, 1878; Marrubium vulgare, 1930;
Melampyrum arvense, 1862; M. pratense, 1930; Mentha pulegium, 1903; Misopates orontium,
1873; Moenchia erecta, 1928; Monotropa hypopitys, 1946; Montia fontana, 1945; Myriophyllum
alterniflorum, 1919; Narthecium ossifragum, 1833; Ophrys sphegodes, 1837; Orchis ustulata,
1955; Oreopteris limbosperma, 1820; Ornithogalum pyrenaicum, 1774; Orobanche artemisiae-
campestris, 1935; O. rapum-genistae, 1913; Osmunda regalis, 1685; Parnassia palustris, 1980;
Pedicularis sylvatica, 1912; Pinguicula vulgaris, 1984; Polygala serpyllifolia, 1954; Polystichum
aculeatum, 1820; Potamogeton compressus, 1912; P. polygonifolius, 1948; Potentilla anglica,
1945; P. palustris, 1886; Pulicaria vulgaris, 1833; Ranunculus baudotii, 1955; R. hederaceus,
1900; R. parviflorus, 1928; Rhynchospora alba, 1839; Rosa pimpinellifolia, 1827; R. sherardii,
1910; Ruppia cirrhosa, 1795; Sagina maritima, 1983; Salicornia dolichostachya, 1959; Silene
gallica, 1951; Solidago virgaurea, 1916; Sonchus palustris, 1843; Stratiotes aloides, 1960;
Teesdalia nudicaulis, 1954; Tephroseris palustris, 1830; Trichophorum cespitosum, 1820;
Triglochin maritima, 1930; Ulex minor, 1832; Utricularia australis, 1899; U. intermedia, 1898; U.
minor, 1951; Vaccinium oxycoccos, 1859; Valerianella rimosa, 1933; Vicia sylvatica, 1792;
Zostera marina, 1908.

APPENDIX 2. DIVISIONS OF CAMBRIDGESHIRE

The following seven divisions were defined to assist in comparison of data presented by Kent
(1975 and in press) for seven divisons of Middlesex. They correspond very approximately to the
divisions recognised by Babington (1860), which are named after each list of 10 km squares:

1: TL 54, 55, 64, 65, 75 (Cambridge); 2: TL 23, 24, 33, 34, 43, 44 (Royston); 3: TL 15, 25, 26, 35,
36, 45 (Cottenham and Wimpole); 4: TL 46, 56, 57, 66, 67, 76 (Burwell); 5: TL 47, 48, 58, 59, 68,
69 (Ely); 6: TL 29, 37, 38, 39, 49 (Chatteris); 7: TF 20, 30, 31, 40, 41, 51 (Wisbech).

Pie

Watsonia 23: 83—92 (2000) 83

Hybridisation between Rumex rupestris Le Gall (Polygonaceae)
and other docks

D. T. HOLYOAK

S Edward Street, Tuckingmill, Camborne, Cornwall, TR14 SPA

ABSTRACT

Hybrids of Rumex rupestris with R. conglomeratus are reported for the first time and named as R. xX
rosemurphyae D. T. Holyoak, hybr. nov. Other hybrids involving R. rupestris have been found at the same
locality in West Cornwall (v.c. la), involving R. pulcher (= R. X trimenii Camus), probably R. crispus, and
possibly R. obtusifolius. Evidence of introgressive hybridisation was found resulting from R. x rosemurphyae
backcrossing with R. conglomeratus, but there was no evidence of introgression with other coexisting Rumex
species.

KEYWORDS: conservation, Shore Dock, introgression, taxonomy.

INTRODUCTION

Shore Dock Rumex rupestris Le Gall is a rare European endemic species occurring on and near
coasts from Wales southwards to north-western Spain (Jalas & Suominen 1979; Daniels ef al.
1998). Many of its localities are on rocky sea cliffs where few other dock species grow, so that
opportunities for it to be involved in interspecific hybridisation are less prevalent than with
congeners that commonly grow together on disturbed ground inland. Indeed, the reviews by
Lousley & Williams (1975) and Lousley & Kent (1981) reported few hybrids of R. rupestris, and
those only with R. pulcher L. and R. crispus L. There do not appear to be any reports of hybrids
involving R. rupestris from outside Britain.

R. rupestris was investigated from 1994—1998 in dune-slack like habitats at Penhale Camp, West
Cornwall. During this period its population there increased from about 60 to 137 mature plants. At
this site, four other dock species (Rumex subgenus Rumex) and several of their interspecific
hybrids grow intermingled with, or close to, R. rupestris. It was expected that under these
circumstances hybrids involving R. rupestris would occur, and, over the five years of study, a total
of eight such hybrid plants was found, apparently representing four different hybrid combinations.
This paper extends and partly revises the preliminary notes (Holyoak 1995, 1996) on the hybrids at
Penhale Camp by giving descriptions of each of the hybrid taxa and naming the hybrid with R.
conglomeratus Murray which has not been reported from elsewhere. In addition, previous records
of hybrids involving R. rupestris are reviewed.

Results are also described of biometric investigation of apparent introgression of R. rupestris
and R. conglomeratus at Penhale Camp. Potential threats to the survival of R. rupestris from
introgressive hybridisation are discussed on the basis of these data.

METHODS

Penhale Camp (West Cornwall; c. SW/770.570) occupies an extensive area of coastal sand-dunes.
In most winters water stands in several large and small pools and flows along an ephemeral
stream, but all of these areas are usually dry in summer. These dune-slack like areas support
vegetation characteristic of dune-slacks, fens and pool-margins, with locally dominant plants of
different areas including Eleocharis palustris (L.) Roemer & Schultes, Epilobium hirsutum L.,
Equisetum palustre L., Mentha aquatica L., Pulicaria dysenterica (L.) Bernh. and Rorippa
nasturtium-aquaticum (L.) Hayek. Rumex rupestris grows intermixed with these wetland plants in
four separate small colonies.

84 D. T. HOLYOAK

TABLE 1. HYBRID FREQUENCY INDEX MEASUREMENTS OF RUMEX RUPESTRIS,
R. CONGLOMERATUS AND INTERMEDIATE PLANTS FROM LOCALITIES IN
WEST CORNWALL, AUGUST 1996

Hybrid-index score

Site 0 1 2 22 4 3 (6 7 8 9 0D Pity ibrsAais

Penhale Camp Ze Be 2 - - - 1 - = 2A oO 22) 28
SW/769.569

Mount Field - - - - - - - - - - . - (ea ela) 20
SW/781.571

near Ventongimps - - - - - - - - - - - - Sesto. 20
Moor SW/779.511

Bonython Estate - - - - - - - - - - - - SU Ome Mees
SW/696.207

See Appendix | for details of the five characters used and the scoring system, with which a typical
plant of R. rupestris scores 0 and a typical plant of R. conglomeratus scores 15. R. conglomeratus
occurred at all four of these sites; R. rupestris was present with it only at Penhale Camp.

Other Rumex species present in the same wet areas are R. conglomeratus (hundreds of plants), R.
crispus subsp. littoreus (J. Hardy) Akeroyd (many hundreds of plants, but most of them growing in
drier edges of wetland vegetation) and R. obtusifolius L. var. obtusifolius (2 plants seen). In
addition, R. pulcher occurs in very small quantity on dry slopes nearby. Other dock hybrids
recorded in the same area were R. conglomeratus X R. crispus (5), R. conglomeratus X R. pulcher
(1) and R. crispus X R. obtusifolius (c. 47) (Holyoak 1996 and subsequent pers. obs.).

The area was visited several times in August of each of the years 1994 to 1998, so that virtually
all of the docks present could be identified and counted as their fruits ripened. Some plants were
individually marked from 1995 onwards and by 1998 all plants of R. rupestris and its hybrids had
been individually marked. In 1995, 1996 and 1998 specimens were collected from each of the
marked hybrids involving R. rupestris at times when they had at least some mature fruits.

During August 1996 it was noticed that some fruiting plants of R. conglomeratus showed
characters approaching those of the R. rupestris growing near them. Because these intermediate
characters seemed likely to have resulted from hybridisation, their morphology and those of the
closest plants of R. conglomeratus and R. rupestris were investigated using the “hybrid frequency
index” technique of Anderson (1936). The scoring system used is explained in Appendix | and
other details are given with the results in Table 1. Comparative data were obtained from three
populations of R. conglomeratus growing at localities in West Cornwall that lacked R. rupestris.

Several counts of chromosomes at mitotic metaphase were obtained from root tips of seedlings
germinated on moist filter paper in petri dishes. Excised root tips were fixed overnight in Farmer’ s
fluid before squash preparations were prepared using acetic orcein stain.

RESULTS

Rumex X rosemurphyae D. T. Holyoak, hybr. nov.
(Rumex conglomeratus Murray X R. rupestris Le Gall) (Fig. 1)
Hybrida a Rumice conglomerato Murray et R. rupestri Le Gall genita et characteribus plerlsque

intermedia; ab ambobus fructibus pro parte maxima abortivis et statura nonnunquam multo majore
differt.

A hybrid between Rumex conglomeratus and R. rupestris, found within 2 m of plants of those two
species. It is intermediate between them in most characters but differs from both in being mostly
but not completely infertile and sometimes in its much greater size.

A robust perennial growing in a compact clump from a stout rootstock. The holotype was much
larger than accompanying plants of R. conglomeratus and R. rupestris, on 8 August 1995 it had 35

RUMEX RUPESTRIS HYBRIDS 85

(G; iy
7 Q ORD
(Cy (ONY
By AY ce
ow N NY

NY \"
ee
Gf
6 Hye
|
OD
Ay. ys

AD
BY) i)

FIGURE 1. Rumex x rosemurphyae. A. Fruit (i.e. perianth enclosing nutlet) B. Diagrammatic section through
fruit. C. Single whorl of inflorescence. D. Inflorescence. E. Leaf from lower part of stem. Scale bars represent

1 mm (A-C) or 10 mm (D, E).

86 Di T.HOLYOAK

stems up to 1-4 m long, the same plant on 18 July 1998 had 77 flowering stems up to 1-5 m long,
in both years most stems had become decumbent before fruits ripened. Lower stem leaves with
lamina at flowering up to 20 x 8 cm or more, but most lower leaves wither before fruits ripen.
Upper stem leaves smaller, oblong-lanceolate to lanceolate. At least some leaves thicker than in
accompanying plants of R. conglomeratus, but less thick than in some R. rupestris.

Panicles with many branches that mostly arise at about 45° from main stem, the branches more
numerous than is usual in R. rupestris, but with branching at more acute angles than typical of R.
conglomeratus. Each of the lower whorls of flowers on each branch subtended by an ovate-
lanceolate to narrowly lanceolate leafy bract, the bracts becoming abruptly smaller towards the
middle of each branch and absent near the branch apex, as in R. conglomeratus. On those parts of
the inflorescence where fertile fruits occur, whorls of flowers appear less close and congested than
in R. rupestris, more like those of R. conglomeratus. Where nutlets develop, inner perianth-
segments oblong to oblong-lanceolate with bluntly rounded apex and sides subparallel in upper
part, always entire. Where nutlets develop, inner perianth-segments very variable in size, some as
small as in typical R. conglomeratus (length 1-7—2-5 mm) others as long as in typical R. rupestris
(2-8-3-7 mm), but with many of intermediate lengths. Where nutlets develop, each of the inner
perianth-segments has a swollen, rounded tubercle that varies from 70-120% of the maximum
width of perianth segment and 40-65% of its length. The few well developed nutlets seen were
trigonous, 1-4—1-6 mm long, brown, glossy, with acute angles.

Counts of mitotic chromosomes from three seedlings grown from seed collected from the
Holotype were all n = 20. Identical counts were obtained from seedlings of R. conglomeratus (4)
and R. rupestris (5) grown from seed collected from plants growing within 10 m of the holotype.
Counts of n = 20 have been reported previously for both of these species (e.g. Degraeve 1975;
Rechinger 1993; Kay 1996).

Named for Miss Rosaline J. Murphy in recognition of her work on the Cornish flora and as
thanks for introducing the author to Penhale Camp.

HOLOTYPUS: W. Cornwall, v.c. la, Penhale Camp (SW/768.569), among Epilobium hirsutum in
fen along course of ephemeral stream, 7-8 August 1995 and 18 July 1998, D. T. Holyoak, field
labels C and 28 (RNG).

A smaller plant growing 6 m away from the holotype (field label D) was also identified as R. x
rosemurphyae. On 8 August 1995 this plant had only two flowering stems, the longest 70 cm tall.
It resembled the Holotype closely in other respects, including its low fertility.

In addition to the two plants described above as R. X rosemurphyae and interpreted as F; hybrids
between R. conglomeratus and R. rupestris, at least six (and perhaps as many as 13) of the 40
plants of R. conglomeratus growing within 20 m of them in August 1996 showed characters that
somewhat approached those of R. rupestris (Table 1, Appendix 1). As discussed below, these are
believed to represent back-crosses between R. x rosemurphyae and R. conglomeratus.

On 13 September 1998 two more plants attributed to R. x rosemurphyae were seen in a dune-
slack at Gear Sands (SW/7.5), about 700 m from the Penhale plants (marked by C. J. Neil as
numbers 117 and 122; vouchers were given field labels DTH 4 and 6 respectively). They were
again close to plants of both R. conglomeratus and R. rupestris. Both of these hybrids also had low
fertility (<20% of nutlets developed) and both showed evidence of “hybrid vigour’, one plant
having about 20 flowering stems up to 1-3 m tall, the other 39 flowering stems up to 1-4 m tall.
Details of the inflorescence and inner perianth-segments were similar to those described for the
Holotype from Penhale Camp. Their (fertile) inner perianth-segments measured (2-0)3-0—-3-3(3-7)
mm on one plant and (2-2)3-0—3-2(4-0) mm on the other.

Probable Rumex crispus L. x R. rupestris Le Gall

A single plant tentatively attributed to this hybrid was found at Penhale Camp on 24 July 1996,
close to both of the supposed parent species. It was about 60 cm tall with a single main stem. The
lower stem leaves were lanceolate, up to 23 x 5 cm, rather thick and with somewhat crisped
margins. The panicle had nine rather upright branches diverging at 10—20° from the main stem,
several of the longer branches having several whorls in the lower half of the branch subtended by a
narrowly lanceolate, petiolate bract. The whorls of the inflorescence appeared less crowded than in
R. crispus, but this impression apparently resulted from low fertility, with many of the inner

RUMEX RUPESTRIS HYBRIDS 87

perianth-segments failing to enlarge after flowering. The minority of perianth segments that had
enlarged were smaller and narrower than in R. crispus, but broader than in R. rupestris (reaching
3-7 mm long X 2-8 mm wide), with wider apices, three swollen tubercles and entire margins
(occasionally with a few short or indistinct teeth).

While the balance of probabilities would suggest that this is a hybrid between R. crispus and R.
rupestris, it may be impossible to discount other hybrid combinations using morphological
characters. In particular, the parent with wide inner perianth segments might have been R. crispus
x R. obtusifolius (= R. x pratensis Mert. & Koch) rather than R. crispus, a possibility strengthened
by the presence of short teeth on some inner perianth segments of the hybrid. Varied forms of R. x
pratensis were growing nearby, some of them with up to 40% of nutlets well developed. It is even
possible that this hybrid plant represents an extreme form of R. X pratensis or a backcross between
it and R. crispus, but some of its enlarged inner perianth segments appear too narrow for that to be
likely. The second parent with narrow inner perianth segments might have been R. conglomeratus
rather than R. rupestris, but that seems less likely in view of the upright branches of the panicle in
the hybrid and the large size of some inner perianth segments and their tubercles. However,
involvement of R. conglomeratus would explain the rather prominent bracts on some branches of
the panicle. Because of these doubts about identification a new name for the hybrid combination of
R. crispus x R. rupestris is not introduced here. The possible occurrence of “triple” hybrids
involving R. X pratensis is discussed further below.

Rumex crispus X R. rupestris has been reported from the Isles of Scilly (v.c. 1b) and Kenfig,
Glamorgan (v.c. 41) (Lousley & Williams 1975; Lousley & Kent 1981; Stace 1991); there are
specimens from both vice-counties at RNG. Dr J. R. Akeroyd (pers. comm.) has located an
additional specimen collected above rocks just above HWM at Pendower Beach, E. Cornwall (v.c.
2) by Olga Stewart 277/82 on 13 September 1982 (FE).

Kay (1996) mentions instances of R. crispus subsp. littoreus being mistaken for this hybrid.
However, the Penhale plant and those discussed by Lousley & Williams (/oc. cit.) differed from R.
crispus subsp. littoreus not only in being largely infertile but also in having at least some inner
perianth-segments narrower overall, or narrower apically, than in R. crispus.

Possible Rumex obtusifolius L. x R. rupestris Le Gall

Three dock plants growing close together in the edge of a fen area at Penhale Camp were studied
on 8 August 1995 and on 24 July 1996. Two of them that survived were studied again on several
visits during July-September 1998, allowing herbarium material to be collected at various stages
of development. Their puzzling combination of morphological characters and consistently low
fertility (with less than 20% of nutlets developing) implied they were hybrids, but although the
three plants are rather similar to each other, they show an odd mixture of features that has
prevented confident inference of the parent species.

All three plants grew as compact patches from stout rootstocks, with strong, erect flowering
stems. In August 1995 one plant (field label E) had about 25 flowering stems up to 1-1 m tall, the
other plant (G) had 11 stems up to 1-2 m tall. The basal and lower stem leaves were thick and
fleshy, with strongly undulate margins; an immature basal leaf had petiole 7 cm, lamina 11 cm; the
longest stem leaves were oblong-lanceolate and had petiole 6:5 cm, lamina 20-5 cm. The underside
of the leaf midrib and some of its strongest veins had low conical papillae, recalling those in R.
obtusifolius, but much smaller and less developed. The panicles were similar in habit to those of R.
rupestris, with branches mainly rather erect (at 20—30° from main stem) and none widely
divaricate. The inner perianth segments of the minority of fruits that ripen were wider than in R.
rupestris, but narrower than in R. crispus (reaching 4-6 mm long and 3-0 mm wide), with a longer
and more attenuate apex than in R. crispus, mostly with one or two short teeth on the basal
margins. All three inner perianth segments on each well-developed fruit had a large swollen
tubercle, that on one perianth segment being larger than those on the other two perianth segments.

Dock species growing within 20 m of these plants were R. crispus, R. rupestris and R.
conglomeratus, along with numerous R. X pratensis and two R. X rosemurphyae; the only other
dock species within many hundreds of metres being two plants of R. obtusifolius and a few of R.
pulcher. Nevertheless, the distinct teeth on the inner perianth segments imply that among the dock
species occurring nearby, either R. obtusifolius or R. pulcher was one of the likely parents of the
hybrids. However, involvement of R. pulcher seems unlikely, as the hybrids gave no evidence of

88 DoT HOLYOSK

the divaricate branching, warty tubercles or other characters of that species. On the other hand,
involvement of R. obtusifolius might be deduced from the presence of papillae on the back of the
leaf midribs, albeit that these and other features of R. obtusifolius appear poorly developed.

Other features of these plants imply that R. rupestris was one parent, including the thick leaves,
large to very large tubercles and rather narrow inner perianth segments with more or less attenuate
apex. Nevertheless, a supposed parentage of R. obtusifolius x R. rupestris does not account for the
strongly undulate leaf margins, for which it is tempting to infer some involvement of R. crispus.
Since numerous plants of R. crispus x R. obtusifolius (R. X pratensis) were present nearby.
whereas the only two plants of R. obtusifolius found were several hundreds of metres distant, the
characters of the three hybrids might therefore be best explained by inferring their parentage as (R.
crispus X R. obtusifolius) X R. rupestris.

Hybrids between R. obtusifolius or R. X pratensis and R. rupestris have not been reported before.
Indeed, no “triple” hybrids have been reported for wild docks in Britain, although crosses
involving three species have been produced experimentally and they are known in Europe
(Lousley & Williams 1975). The absence of British reports of “triple” hybrids in Rumex might
therefore result not from their absence but from the almost insuperable difficulties in identifying
them from morphological characters.

Williams (1971) suspected from field observations that R. x pratensis back-crosses with both
parental species and this suspicion was strengthened because R. crispus X R. X pratensis has been
produced in cultivation. Holyoak (1996) noted that the numerous R. x pratensis at Penhale Camp
vary widely in fertility and in characters of the inner perianth segments, concluding that it is
uncertain to what extent their marked variability is due to back-crossing or merely the expression
in F,; hybrids of an independent assortment of varied characters from the parental genotypes.

Overall, it seems likely that the three puzzling plants described above originated either from R.
obtusifolius or R. xX pratensis hybridising with R. rupestris. Because analysis of their
morphological characters alone may provide an insufficient basis to choose between these
alternatives the hybrid combination is not named here.

Rumex x trimenii Camus

(Rumex pulcher L. xX R. rupestris Le Gall)

A single plant of this hybrid grew close to numerous plants of R. rupestris at Penhale Camp from
1994-1996 (RNG); it was described and illustrated by Holyoak (1995). A similar, but smaller,
plant was found in 1998 close to a different colony of R. rupestris. The nearest plants of R. pulcher
to both of these hybrids were 200 m away and few in number.

Both of the Penhale hybrids had low fertility although at least some apparently fertile fruits were
surrounded by perianth-segments that enlarged after flowering. Their widely divaricate branches
resembled those of R. pulcher and the influence of that species was also apparent from the
reticulate venation of the perianth segments, the presence on them of marginal teeth and the warty
surface of their tubercles. Influence of R. rupestris was apparent in the hybrids from the strong
stems, the rather thick leaves of broadly lanceolate shape and the narrowly lingulate inner
perianth-segments. Lousley & Kent (1981) describe a similar combination of characters in R. Xx
trimenii. There appear to be three previous records of this hybrid in the wild, each of single plants:
in v.c. 1b from east coast of Samson, Isles of Scilly (RNG) and New Grimsby, Tresco, Isles of
Scilly (RNG) , and in v.c. 2 at Whitesand Bay, E. Cornwall (BM); it also arose spontaneously in
the garden of the South London Botanical Institute (Lousley 1971, 1983; Lousley & Williams
1975; Lousley & Kent 1981; Margetts & David 1981; Holyoak 1995, 1996).

DISCUSSION

Rumex rupestris at Penhale Camp has apparently produced hybrids involving all four of the other
dock species that grow in the same area. The total population of R. rupestris there was about 60
mature (fruiting) plants in 1994, but it had increased to 132 by 1998. Although the number of
hybrids involving this species at Penhale was small (eight plants), they apparently exceeded 10 %
of the total R. rupestris population during 1994-1996. Moreover, six of the eight hybrids were
found in one small fen area (SW/768.569) that supported a maximum of 18 plants of “pure” R.

RUMEX RUPESTRIS HYBRIDS 89

rupestris. Two more hybrids were found at Gear Sands alongside a colony of about 27 plants of R.
rupestris.

Most, if not all, of these hybrids produce at least some pollen and at least small amounts of
viable seed. Hence, given the rather high frequencies of F; hybrids that have been found, there
may be opportunities for introgression to occur between the dock species involved. The following
discussion explores various possibilities of introgression between Rumex species at Penhale.

POSSIBLE INTROGRESSION OF R. RUPESTRIS GENES INTO R. CRISPUS

Lousley and Kent (1981) reported possible introgression of R. rupestris into R. crispus in the Isles
of Scilly and at Kenfig, Glamorgan, noting that “in both these localities plants referred to R.
crispus occur with exceptionally large, elongate tubercles recalling those of R. rupestris and
indicating possible introgression”. However, these may have been merely the coastal taxon now
treated as R. crispus subsp. littoreus (e.g. in Stace 1991). Nevertheless, plants of R. crispus subsp.
littoreus growing at Penhale Camp in the same areas as R. rupestris also include some with
unusually elongate tubercles and it was tempting at first sight to suspect introgression had occurred
between these species. However, the Penhale plants also grew alongside numerous R. X pratensis
(26 were counted in 1995) and possible back-crosses of these to R. crispus, the plants showing
considerable variability in morphology and in fertility (cf. Holyoak 1996). Hence there is no need
to invoke introgression from R. rupestris to explain variability in the R. crispus growing close to it
at Penhale Camp.

INTROGRESSION OF R. RUPESTRIS GENES INTO R. CONGLOMERATUS
In August 1996 the small fen area at Penhale Camp with the two plants of R. x rosemurphyae also
had about 40 plants of R. conglomeratus and at least six and perhaps as many as 13 of these
showed characters approaching those of R. rupestris (Table 1). Because no evidence of R.
conglomeratus showing similar characters was found in 39 plants studied from other parts of
Penhale Camp or in a total of 125 plants from three localities elsewhere in West Cornwall, the
presence of R. rupestris seems to be associated with occurrence of some of its characters in the
coexisting population of R. conglomeratus (Table 1). Since two F, hybrid plants (R. x
rosemurphyae) were present at the site it seems likely that some of the plants of R. conglomeratus
had acquired genes from R. rupestris, presumably as a result of back-crossing from F, hybrids.
Although fully adequate data on pollen fertility of R. x rosemurphyae are not available, freshly
collected pollen from its Holotype mainly appeared well formed when examined microscopically
in July 1998. The F, hybrid (R. x rosemurphyae) produces few fertile fruits so that few F, plants
would be expected to occur, although some of its fruit has been successfully germinated in
cultivation. In contrast, the plants of R. conglomeratus putatively introgressed with R. rupestris
appear to produce mainly fertile fruits so these back-crosses would be expected to persist once
established and this may explain the occurrence of at least six and perhaps as many as 13 such
plants in one small fen area at Penhale Camp.

RISK OF INTROGRESSION OF R. CONGLOMERATUS GENES INTO R. RUPESTRIS

R. rupestris is regarded as a globally threatened species, which is included in the British Red Data
Book (Wigginton 1999), placed on the “Biodiversity Short List” (B.S.G.R. 1995) and included in
Schedule 8 of the Wildlife and Countryside Act, 1981. It is included in English Nature’s “Species
Recovery Programme” which is being undertaken collaboratively with Plantlife’s “Back from the
Brink” Project. Its population at Penhale Camp (137 fruiting plants in 1998) apparently includes at
least 20 % of all R. rupestris plants currently known in the British Isles (cf. Daniels et al. 1998).

In these circumstances any “leakage” of genes from other docks into R. rupestris populations
would complicate attempts to maintain a favourable conservation status for “genotypically pure”
R. rupestris. The potential danger is evident from several well documented instances of the loss of
flowering plant taxa through introgressive hybridisation, either locally (DePamphilis & Wyatt
1990; Klier et al. 1991), or over the whole range of geographically restricted taxa (Reiseberg et al.
1989). However, although the data presented in this paper appear to show “leakage” of genes out
of R. rupestris into R. conglomeratus there is no direct evidence for genes of R. conglomeratus
entering the R. rupestris genotype. This asymmetry might be genuine and have arisen because
hybridisation was asymmetrical with respect to male and female parentage, or because of
differential mortality in hybrid products.

90 D. T. HOLYOAK:

However, this apparently fortunate result might be seriously misleading, both regarding the
situation at Penhale Camp and elsewhere. Some doubts arise because few R. rupestris plants could
be studied (although at least 130 have now been studied in detail, of which 35 were scored for
hybrid index frequency in 1996), despite the Penhale Camp population being one of the largest
known. Thus, relatively rare, introgressed, fruiting plants of R. rupestris might be absent merely as
a result of stochastic processes operating during germination or causing mortality during growth,
so that viable seed with an introgressed genotype might nonetheless be present at low frequency in
the local seed-bank.

It is also possible that the mode of inheritance or mode of phenotypic expression of the
characters used in deriving the hybrid frequency index means it is easier to recognise introgressed
R. conglomeratus than introgressed R. rupestris. Although polygenic traits can be expected to
show intermediate expression in hybrids, traits governed by one or two genes are more likely to
show parental expression, which could include strong maternal effects. This may explain why, in a
survey of morphological patterns in natural and experimental hybrids of flowering plants,
Reiseberg & Ellstrand (1993) found that hybrids are no more likely to display intermediate
morphological features than parental ones. The classic “hybrid frequency index” technique of
Anderson (1936) can thus be viewed as applicable in only the minority of situations where hybrids
are intermediate.

Considered against this background, the present lack of morphological evidence for
introgression of R. conglomeratus genes into R. rupestris might well be misleading. It seems likely
that if a theoretical introgressed R. rupestris was once established in a population, further back-
crossing and introgression into that species could easily occur, much as appears to have happened
with introgressed R. conglomeratus. In view of such dangers, further work on hybridisation of R.
rupestris with other docks is desirable, for which morphological studies might usefully be
supplemented by genetic studies that should provide additional characters.

Daniels et al. (1998) reported “preliminary” results of isozyme electrophoresis on samples of R.
rupestris from south-western England, disclosing a high level of genetic diversity within some
populations and significant differences between certain groups of populations. Nevertheless, it
remains unclear how much of that genetic variation is intrinsic to R. rupestris and how much of it
might be derived from introgressive hybridisation with congeners. This doubt becomes important
if the species is to be introduced or reintroduced into the wild since it is then desirable to establish
that genotypically “pure” R. rupestris plants are used. Otherwise, attempts to maximise genetic
variation in the introduced populations might result in introgressed plants being chosen for
introduction attempts.

ACKNOWLEDGMENTS

Thanks are due to the Ministry of Defence for access to Penhale Training Camp and to Major B.
Andrews and Lt Col R. C. Taylor for assistance and information. The following botanists helped in
field surveys of docks at Penhale or commented on specimens: Dr J. R. Akeroyd, M. Atkinson, T.
Atkinson, I. Bennallick, Dr C. N. French, Dr P. A. Gainey, E. C. M. Haes, G. A. Holyoak, G.
Kitchener, E. J. McDonnell, H. M. Meredith, R. J. Murphy, C. J. Neil and J. Stewart. Dr Akeroyd
and Dr S. L. Jury (RNG) provided information on specimens in herbaria and A. Jones and Dr Q.
O. N. Kay provided other unpublished information. T. L. Blockeel kindly prepared the Latin
diagnosis. Thanks are also due to Zowie Keating for drawing the figures of Rumex xX
rosemurphyae and to the B.S.B.I. for assisting with the cost of the drawings.

REFERENCES

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London.

DANIELS, R. E., MCDONNELL, E. J. & RAYBOULD, A. F. (1998). The current status of Rumex rupestris Le Gall
(Polygonaceae) in England and Wales, and threats to its survival and genetic diversity. Watsonia 22: 33—

RUMEX RUPESTRIS HYBRIDS 91

39. ;

DEGRAEVE, N. (1975). Contribution a l'étude cytotaxonomique des Rumex. 1. Le genre Rumex L. sensu
stricto. Caryologia 28: 187-201. s

DEPAMPHILIS, C. W. & WYATT, R. (1990). Electrophoretic confirmation of interspecific hybridization in
Aesculus (Hippocastanaceae) and the genetic structure of a broad hybrid zone. Evolution 44: 1295-1317.

HOLYOAK, D. T. (1995). A rare hybrid dock Rumex xX trimenii Camus at Penhale Camp, West Cornwall.
Sanctuary 24: 35. a

HOLYOAK, D. T. (1996). Hybridization in docks (Rumex subgenus Rumex) in Cornwall. Botanical Cornwall 7:
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Polygonaceae. The Committee for Mapping the Flora of Europe & Societas Biologica Fennica Vanamo,
Helsinki. .

Kay, Q. O. N. (1996). The conservation of Rumex rupestris (Shore Dock) in Wales. Past, present and possible
future sites and habitats for Rumex rupestris in South and West Wales. Unpublished Report of
Countryside Council for Wales Contract Survey.

KLIER, K., LEOSCHKE, M. J. & WENDEL, J. F. (1991). Hybridization and introgression in white and yellow
ladyslipper orchids (Cypripedium candidum and C. pubescens). Journal of heredity 82: 305-319.

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Harvey (1983). Isles of Scilly Museum Publication no. 4. St Mary’s.

LOUSLEY, J. E. & KENT, D. H. (1981). Docks and knotweeds of the British Isles. Botanical Society of the
British Isles, London.

LOUSLEY, J. E. & WILLIAMS, J. T. (1975). Rumex L., in STACE, C. A., ed. Hybridization and the flora of the
British Isles. Academic Press, London.

MARGETTS, L. & DAVID, R. W. (1981). A review of the Cornish Flora, 1980. Institute of Cornish Studies,
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RECHINGER, K. H. (1993). Rumex L. (revised by J. R. AKEROYD), in TUTIN, T. G. et al., eds. Flora Europaea
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REISEBERG, L. H. & ELLSTRAND, N. C. (1993). What can molecular and morphological markers tell us about
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REISEBERG, L. H., ZONA, S., ABERBOM, L. & MARTIN, T. D. (1989). Hybridization in the island endemic,
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(Accepted October 1999)

o2 D. T. HOLYOAK

APPENDIX 1

Methods used for hybrid frequency index measurements of Rumex rupestris, R. conglomeratus and
intermediate plants from localities in West Cornwall, August 1996.

The hybrid index was derived by scoring each of five characters on each plant. A score of 0 was
assigned for character-states typical of R. rupestris, 3 for those typical of R. conglomeratus and 1—
2 for intermediate states. Hence, summing data for five characters, a typical plant of R. rupestris
would score 0, a typical plant of R. conglomeratus would score 15. Data were scored only from
undamaged plants with mature (drying) perianths. The characters and scoring systems were as
follows:

Length of longest inner-perianth segment: mean of 10 perianths examined from middle part of
inflorescence; measurements made with eyepiece graticule to accuracy of + 0-05 mm; 0 = > 3-0
mm, | = 2-5—3-0 mm, 2 = 2-0—2-:5 mm, 3 = < 2:0 mm;

Length of tubercle as per cent of length of longest inner-perianth segment: mean of 10
perianths examined from middle part of inflorescence (same perianths as for preceding character);
measurements made with eyepiece graticule to accuracy of + 0-05 mm; 0 = > 60%, 1 = 55-60%, 2
= 50-55%, 3 =< 50%:

Angle of main branches of inflorescence: modal value; angle measured from main stem (not
from vertical); measured only for branches > 5 cm long; 0 = < 50°, 1 = 50-60°, 2 = 60-70°, 3 = >
70°;

Number of bracts on longest three branches of inflorescence: using only branches > 10 cm
long; 0 = 1-5 bracts, 1 = 6 or 7 bracts, 2 = 8 or 9 bracts, 3 = > 9 bracts;

Crowding of whorls on main branches of inflorescence: estimated as ratio of inflorescence
whorl width (flowers + their pedicels)/ interwhorl width along the inflorescence axis (length of
stem clear of all flowers + pedicels); modal value for all of main inflorescences: 0 = ratio <1-0/1, |
= ratio 1 -0—1-25/1, 2 = ratio 1-25—1-5/1, 3 = ratio > 1-5/1.

Watsonia 23: 93—110 (2000) 93

Prize-winners to pariahs - A history of Japanese Knotweed s./.
(Polygonaceae) in the British Isles

J.P. BAILEY and A. P. CONOLLY

Biology Department, University of Leicester, Leicester, LE] 7RH

ABSTRACT

The early history of the discovery and introduction to the British Isles of the alien invasive perennials
belonging to Fallopia section Reynoutria (hereafter referred to as Japanese Knotweed s./.) are described from
a study of documentary evidence and herbarium material. The role of the various agencies involved in the
dispersal of these plants is discussed, as is the influence that contemporaneous gardening trends had on the
ornamental use of Japanese Knotweed s. /.

KEYWORDS: Fallopia japonica, Fallopia x bohemica, Fallopia sachalinensis, invasive plants, garden escapes,
plant collectors.

INTRODUCTION

The group of plants referred to here as Japanese Knotweed sensu lato needs little introduction.
Details of reproductive biology, cytology and distribution are given in Conolly (1977), Bailey &
Conolly (1985, 1991), Bailey (1989, 1994), Bailey & Stace (1992), Beerling et al. (1994) and
Bailey et al. (1996). The term Japanese Knotweed s./., may need further explanation. Since the
amalgamation of Reynoutria and Fallopia there is no convenient way of referring to the complex
of taxa formerly circumscribed by the name Reynoutria, which comprises F. japonica var.
Japonica, F. japonica var. compacta, F. sachalinensis, the hybrids between F. japonica and F.
sachalinensis (F. x bohemica) and between var. japonica and var. compacta as well as any
backcrosses between the hybrids and their parents.

It is generally known that these plants were originally garden plants, but it is not so well
appreciated just how valuable an addition to the garden they were once considered to be. This
paper seeks to build on the initial contribution by Conolly (1977), by giving a more detailed
examination of just those taxa transferred from Reynoutria to Fallopia and using the year 1914 as
a general cut-off point.

It is our intention to deal with each taxon in turn, to detail its original discovery and the most
probable source of the accessions that were introduced to Europe, and to try and unravel the
somewhat bewildering array of synonyms that litter the scientific and horticultural literature. Using
a combination of type and other early herbarium specimens, archive material, scientific and
horticultural literature, nursery garden catalogues and accounts of plant collecting expeditions, the
discovery and spread of each taxon is given in as much detail as possible. There follows a
consideration of the gardening movements that contributed to the dissemination of Japanese
Knotweeds to every corner of the British Isles. Since we are dealing with long-lived perennials
spreading primarily by vegetative means, and easily outliving gardener and garden alike, it is
important to have records of where these plants were once grown or available for sale. Large
private gardens or estates, nursery gardens and botanic gardens are likely to constitute the initial
loci from which further dispersal by exchange, gift, sale or disposal took place.

With regard to the hybrid F. x bohemica, it is useful to know where both parental taxa were
grown together, especially where the F. sachalinensis involved is male-fertile. We will now
consider in detail, by taxon and in chronological order, the early history of these plants as aliens in
the British Isles, as we track the riotous career of F. japonica from gold medal holder in 1847 to
proscription in the 1981 Wildlife & Countryside Act.

94 J.P. BAILEY AND-AjP2CONOELY

FALLOPIA JAPONICA VAR. JAPONICA (HOUTT.) RONSE DECRAENE

A MOST PROMISING START

Fallopia japonica was awarded a gold medal in 1847 by the Society of Agriculture & Horticulture
at Utrecht for the most interesting new ornamental plant of the year, according to the 1848
catalogue of Von Siebold & Company of Leiden, the specialist importers of plants from the East
(Siebold 1848). An accompanying seven line foot-note in this price-list extols its great vigour, its
combination of ornamental and medicinal use, its value in protecting young plantations from wind
and sun, its edible young stems and leaves and a rhizome highly valued in Japanese and Chinese
medicine. Such a plant, newly imported from Japan, would obviously command a high price. 500
francs would purchase a “mother” plant and 25 strong plants. In order to put this sum into
perspective, the same catalogue offered 250 plants of Hydrangea japonica or 500 plants of
Wisteria sinensis for the same amount! If, by the time of Siebold’s 1856 catalogue, the price had
plummeted (100 plants for 25 francs), the rhetoric certainly had not. In addition to the benefits
listed above were added stabilisation of sand dunes, gracious flowers suitable for bouquets, a
forage which cattle loved to eat, melliferous flowers which gave bees their winter sustenance and,
last but not least, dead stems that could be used to make matches! There was, however, no element
of exaggeration in the claim that the plant was “inextirpable”.

SIEBOLD, HIS EXPEDITION AND HIS COMPANY

Phillipe von Siebold had for his time absolutely unparalleled access to the then closed country of
Japan. Appointed as doctor in residence to the Dutch trading settlement in Japan, he stayed there
from 1823-1829. His medical knowledge gained him many friends and high level contacts in
Japan. Siebold used every opportunity to collect plant and animal specimens, and had friends and
students collecting material from all over Japan. After a series of fascinating peregrinations
(shipwrecks, imprisonment and civil war), he was not finally reunited with all his live Japanese
plant specimens back in Leiden until 1841 (Kimura & Kitamura 1977). Here he founded, in 1842,
a Royal Society for Encouragement of Horticulture (Anderson 1951) - apparently just a
respectable cover for the commercial activities of his Von Siebold & Co. The horticultural
company owned not only the specimens brought back from his sojourn in Japan but also a
governmental monopoly on any subsequent introductions from Japan and Java. By 1847 this Royal
Society had severed its connections with Von Siebold & Co. Fortuitously, this connection allowed
the survival of a more-or-less complete collection of sales lists in the pages of their journal:
Jaarboek van de Koninklijke Nederlandsche Maatschappij tot Aanmoediging van den Tuinbouw.
From a study of these lists it is possible to reconstruct the very early days of this plant in Europe.
The name Polygonum sieboldii Reinw. first appears in 1848, and this name is given as a synonym
of Polygonum cuspidatum Sieb. & Zucc. by De Vriese (1849) (see Table 1). Von Siebold & Co.
presumably considered the eponymous P. sieboldii to be a better advertisement than the name
given to it by Siebold and Zuccarini in 1846. In the 19th century gardening literature the plant is
generally known as P. sieboldii.

UNTANGLING THE KNOTWEED NOMENCLATURE

The history of Fallopia japonica really begins in the 18th century, when it was described as
Reynoutria japonica by Houttuyn (1777), presumably from dried material brought back from
Japan by Thunberg. Considering that both Houttuyn and Thunberg wrote substantial botanical
books, it is surprising that the world in general was unaware of the plant until it was described by
Siebold & Zuccarini in 1846. Even as late as 1895 the first edition of Index Kewensis lists
Reynoutria as incertae sedis. It was not until 1901 that Makino realised that Reynoutria japonica
of Houttuyn and Polygonum cuspidatum of Siebold and Zuccarini were one and the same; on these
grounds he made the new combination Polygonum reynoutria, but without sufficient grounds for
replacing P. cuspidatum.

There are various reasons why this important taxon was lost to science for seven decades: one
was that Houttuyn’s Natuurlijke Historie (1777) was written in Dutch, not the most accessible of
languages. He also erroneously described the plant as having ten stamens, thus putting Reynoutria
under “Decandria Trigyna”’, isolated from the other Polygonums. However, Houttuyn does have a
good clear illustration of the plant in his work. Thunberg (1784) does not recognise Reynoutria in

95

HISTORY OF JAPANESE KNOTWEED

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FIGURE |. Engraving from De Vriese (1849) of Siebold’s plant.

96 J.P. BAILEY AND A. P. CONOLLY

TABLE |. EARLY SYNONOMY OF FALLOPIA JAPONICA

Name Authority/user Current name

Reynoutria japonica Houttuyn 1777 Fallopia japonica
Polygonum multiflorum © Thunberg 1784 Fallopia japonica

P. pictum Siebold 1841 = F. japonica var. compacta ?
P. cuspidatum Siebold & Zuccarini 1846 F. japonica var. japonica

P. sieboldii used by de Vriese 1849 F. japonica var. japonica

P. reynoutria Makino 1901 F. japonica

his Flora Japonica, but has his own taxon, Polygonum multiflorum (Thunb.), which is obviously
not in Natuurlijke Historie. This is a perfectly good species, now known as Fallopia multiflora,
and one of the link species between the climbing and non-climbing members of the genus
Fallopia. Photocopies of some of the Thunberg specimens from Uppsala (UPS) have been seen.
Herb. Thunberg 9706 is labelled Polygonum multiflorum (probably in Thunberg’s hand) and is
indeed that taxon. However, specimen 9705 bears the label “Polygonum multiflorum a’, but is
clearly a specimen of Fallopia japonica. Ironically, the Houttuyn specimen of F. multiflora (Herb.
Delessert collection Burman s.n., G) is labelled “Polygonum (Helxine) Japonicum Mihi. Vid Nat
His. VII Pl. 49 Fig 3” in Houttuyn’s hand (?). However, the name does not tally with the text
reference, which refers to the “Japansch” version of Polygonum chinense L., whereas the figure
depicts F. multiflora. In summary, although Houttuyn and Thunberg both handled specimens of F.
Japonica and F. multiflora, the former regarded F. multiflora as the Japanese form of P. chinense,
whilst the latter regarded F. japonica as a variant of F. multiflora. Further, it was only possible to
appreciate Thunberg’s circumscription of P. multiflorum by reference to his herbarium specimens!

Since there is no evidence of live F. japonica being in Europe in the 18th century, in order to
find the source of the live introductions to Europe, we need to look at the Siebold’s collections.
The Rijksherbarium at Leiden sent us four specimens; the one labelled “Herb. Lugd. Batav. No.
908.174 826 Polygonum cuspidatum S. & Z. Legit v.S.” is probably the holotype of P.
cuspidatum. The specimen is male-sterile and very similar to our British plants. Further evidence
that this is the source of our plants is provided by De Vriese (1849) and the excellent detailed
engraving (see Fig. 1). De Vriese heaps praises upon the plant and states that it is the sole property
of Siebold who has had it growing for many years [at Leiden]. The red spotted stem and the zig-
zag appearance of the stem are also mentioned. The figure (Fig. 1), drawn from life from Siebold’s
plant, shows leaves with truncate bases closely matching the appearance of our plants. It is
interesting to note that it is also male-sterile and bears good seed with embryos; this raises the
question of what could have pollinated it in the absence of F. sachalinensis? The most likely
source is Polygonum pictum (= F. japonica var. compacta?) which had been in cultivation at
Leiden for some time.

THE ARRIVAL OF FALLOPIA JAPONICA IN BRITAIN

Although the first date that the plant was commercially available in Europe is undeniably 1848, the
date that it actually crossed the Channel to Britain was not known until now. The literature
contains two conflicting dates: 1825 (Conolly 1977 ex Synge 1956) and the late 1840s (Bailey
1994). It appears that in a sense, both dates are correct! Morren (1849) gives a résumé of the
DeVriese paper of 1849 which describes Polygonum cuspidatum Sieb. & Zucc., and says that it
was introduced by Phillipe von Siebold and that it was his sole property. Lindley & Paxton (1850),
however, in reply to the Morren article, declare that the plant was actually introduced by the
Horticultural Society of London around 1825. The plant had been thought to be Houttuynia
cordata, and had been planted in an artificial swamp at their Chiswick garden; not surprisingly, it
had failed to thrive, though an illustration is provided of the one time that it did manage to flower.
Lindley & Paxton ended with the comment that they considered it unlikely that such a plant would
bind blowing sand together in the manner described by Dr Siebold. Significantly, they record that
the Chiswick plant came from China and indeed their illustration shows quite clearly the
acuminate leaf base so characteristic of Chinese accessions of F. japonica. This was not the end of

HISTORY OF JAPANESE KNOTWEED 97

the cross-channel dialogue. Groenland (1858) writing in La Revue Horticole (Paris), expresses
surprise that “Fallopia japonica” could remain unnoticed for 20 years in the garden at Chiswick,
without even flowering, adding that in Japan the plant is used to bind sand-dunes.

With the knowledge that F. japonica had no name in 1825, since this predates the Sieb. & Zucc.
name, and the earlier Reynoutria combination had been lost until 1901, one of the authors (J.P.B.)
had severe reservations regarding an introduction date of 1825. In the event, the mis-identification
of the Chiswick plant as Houttuynia neatly side-stepped this objection; (that it should have been
confounded with a Houttuyn eponym is rather a nice irony!).

So, although the earliest recorded live plant of F. japonica in Europe was in the Chiswick
garden of the Horticultural Society, this was a Chinese accession. Chinese plants are
morphologically distinguishable from the Siebold clone of F. japonica, and, even if the Chiswick
plant was ever distributed to any extent, it is doubtful that any has survived; it has had no
discernible role in the Japanese Knotweed invasion of Europe. Siebold’s nursery at Leiden, where
the plant was actively sold and heavily promoted, even to the extent of offering a 25% discount to
botanical gardens, is considered the initial locus of the F. japonica clone that has so effectively
invaded much of Europe.

The manuscript “Inwards Book” 1848-1858 at the Royal Botanic Gardens Kew records that
Polygonum sieboldii was received on 9 August 1850 as part of a large batch of plants from M.
Siebold of Leyden. There is appended the following note: “They are intended to be in exchange for
new China and Japan plants. But on account of the bad selection he is written to, telling him that
only 6 of them are probably new to us”. The plants were probably sent unsolicited in the hope of
receiving further novelties in return. The Kew herbarium has a male-sterile F. japonica var.
Japonica specimen labelled “Hort, Kew 1857”, which is presumably the Siebold introduction. The
Kew archive also contains sets of “outward Books” which record a commerce absolutely
unimaginable to modern-day gardeners attempting to obtain plants from the Kew collection.
Collections and cabinets of plants were being dispatched alike to Honourables, military men and
commercial nursery gardens apparently without distinction. However, a search of the Outward
Book 1853-1856 has failed to reveal a single reference to Japanese Knotweed. This does not mean
that Kew took no further role in the distribution of F. japonica, but that only the choicer plants
were listed individually, there being plenty of examples of collections of un-named herbaceous
plants being sent out. The archive at the Royal Botanic Garden Edinburgh also has a number of
manuscript accession ledgers known as “Plant Books”. The 1849-55 Plant Book revealed that, on
26 April 1854, Polygonum sieboldii was received from Messrs Jackson & Son Nurserymen of
Kingston. This, incidentally, is the earliest record of a British nurseryman offering Japanese
Knotweed for sale by a very long way (see Table 2), and it is tempting to think that, with the
proximity of Kew and Kingston upon Thames, the plant was supplied to Jacksons by Kew. Thus in
the year 1850 began the inexorable spread of Japanese Knotweed throughout the British Isles.

Before moving on, it is worth considering the fate of Siebold’s Garden of Acclimatisation at
Leiden. In the year of his death, 1866, it boasted nearly a thousand different species and varieties.
By the time the English horticulturists F. W. Burbidge and P. Barr visited in 1883 they found a
neglected jungle, overrun by none other than - Japanese Knotweed (Sharman 1990).

F, JAPONICA VAR. COMPACTA (HOOK. F.) J. P. BAILEY

The 1844 catalogue of Von Siebold & Company records that a taxon called Polygonum pictum
Sieb. was first introduced to Europe in 1841. It is thought that Siebold’s Polygonum pictum is
actually the dwarf variant of F. japonica. This is based on two supporting pieces of evidence.
Firstly its height is given by Siebold as 60 cm, and secondly the name is given as a synonym of P.
cuspidatum by De Vriese in 1849. That aside, little is known about the early history of var.
compacta - it is certain that the name P. pictum has not yet been found in the British botanical
literature. Hooker (1880) gives the taxon specific rank as P. compactum, but this treatment has not
been widely followed. It is a decorative garden plant with a dwarf habit, crimped leathery “square
shaped” leaves, and usually growing up to about one metre, but with occasional records of much
shorter plants in cultivation (c. 30 cm high). Male-sterile plants often have a conspicuous red
pigmentation, making them more desirable in the garden. Unlike var. japonica, both male-sterile
and hermaphrodite individuals are found. It was recommended by Gertrude Jekyll, who included

98 J. P=BAILEY AND A. -PZCONOLEY.

“this dwarf Japanese Knotweed” in some of her planting schemes, such as the garden at Long
Aston, Birmingham (Bisgrove 1992).

The earliest record for cultivation in the British Isles is from John Wood’s garden at Kirkstall
Abbey, Leeds in 1881. In 1883 it was grown in Mr Scott’s garden, Denzell (Dunham Massey, near
Altrincham, Cheshire). Two 1911 herbarium specimens of plants grown at Charles Bailey’s garden
at Cleeve Hill near Cheltenham (v.c. 33) are labelled as having been supplied by Smith’s nursery.
Richard Smith of Worcester was a well known nurseryman at this time (see Table 2). These latter
are labelled as P. alpinum, but it is not clear whether this error was Bailey’s or Smith’s. (All these
herbarium specimens are now at MANCH).

It is now thought that the plants we refer to as var. compacta are actually particular alpine
variants of F. japonica. Shiosaka & Shibata (1993) in their transplant experiments clearly show
that there is a genetic component governing both height and time of flower initiation in this taxon.
As the altitude from which the plant was collected increased, so the final stature of the plant
diminished and the initiation of flowering became earlier. Plants from 2,400 m on Mt Fuji only
grew 10 cm tall and initiated flower buds in May. We have noted very small var. compacta plants
in Britain at Arno’s Castle, Bristol (LTR) and Wisley. L. H. Bailey (1949) lists “Polygonum
Reynoutria Makino” as a ground cover plant 4-6 inches high. Synge (1956) distinguishes
“Polygonum Reynoutria Makino”, a synonym of P. cuspidatum, from “Polygonum ‘Reynoutria’ of
gardens, non Makino”, a male-sterile var. compactum with red fruiting perianths. Amos Perry in
1936 received an R.H.S. Award of Merit for his “P. Reynoutria”, 18 inches high (according to his
1937 catalogue). Its crimson fruiting sprays were much admired in the Wisley September notes in
the 1940s (Anon. 1943). The “P. Reynoutria” at E. A. Bowles’ rockery at Myddleton House,
Enfield is reported as having come from this source. It is quite possible that there were multiple
introductions, as these dwarf variants would be encountered by any botanical expedition to the
mountains.

The earliest recorded escape was 1915 Melrose Abbey (E). The plant is still rather rare, and
there are few instances of naturalisation; to date we have only mapped 30 records on a 10 km
square basis, including garden plants. When found it is usually only in small stands and is not
considered to be very invasive.

F. SACHALINENSIS (F. SCHMIDT EX MAXIM.) RONSE DECR.

F. sachalinensis is a native of Japan, Sakhalin, and possibly also the isolated Ullung-do Island
between Korea and Japan. Early collections were made in the 19th century from both areas, but the
exact dates and mode of entry to Europe are not always clear. Apart from the collectors own
accounts, Bretschneider (1898) gives the most reliable account of expeditions to these areas. The
Russian expeditions were sent from St Petersburg, and the British were sponsored by Kew,
privately, or by nursery gardeners such as James Veitch & Sons (Veitch 1906).

F. sachalinensis was discovered on Sakhalin by Dr H. Weyrich. He was the surgeon on the
Vostok under the command of Capt. Lieut. Rimsky-Korsakov, on the Russian Naval Expedition to
eastern Asia (1852-55). [Incidentally the composer N. A. Rimsky-Korsakov was the younger
brother of the commander of the Vostok; initially his “..heart was set not on music but on a career
in the navy in emulation of his brother 22 years older than himself’, (Grove, ed. Sadie 1980)].
During the survey of the west coast of Sakhalin, in early September 1853, Weyrich landed at
Noto-sama, where he collected F. sachalinensis from damp river banks. The Russian Naval
Expedition returned to St Petersburg in 1855 and Weyrich’s plants went to the St. Petersburg
Botanic Gardens (Bretschneider 1898); his F. sachalinensis was described and named by F.
Schmidt in Maximowicz’s Primitiae florae amurensis (1859). The type description records that the
fruits were immature, though our examination of the type specimen (LE) contradicts this.

Further collections of F. sachalinensis (BM; K) were made from Sakhalin by P. von Glehn in
1861, who, with F. Schmidt, made the first extensive survey of the flora of Sakhalin, during 1860-
1862. They arrived back at St Petersburg in early 1863 (Schmidt 1868).

Material of Japanese origin was collected by C. J. Maximovicz on his second expedition to
eastern Asia 1859-64 (1861 Hakodate, Japan BM; K). He arrived back at St Petersburg in July
1864 with an enormous collection of living plants, seeds and herbarium specimens.

HISTORY OF JAPANESE KNOTWEED 99

Regel (1864) reported that F. sachalinensis was growing in the Botanic Garden at St Petersburg
by 1864 and that it was able to overwinter there. At this date he can only have been referring to
plants of Sakhalin origin. Regel (1874) further suggested that the St Petersburg F. sachalinensis
originated from seed collected by Schmidt, and was then distributed widely to European botanic
gardens. Certainly the herbarium specimen from the Glehn and Schmidt expedition (K) has good
seed. So, whilst it is certain that the first F. sachalinensis at St Petersburg was indeed from
Sakhalin, it is not entirely clear whether it was from the collection of Weyrich or Schmidt or both.
Certainly by the time of Mr Bull’s importing it to Britain (see below), both Japanese and Sakhalin
plants could well have been available in St Petersburg.

Edouard André (1893) claims to have been the first person to have brought F. sachalinensis to
the notice of the horticultural world. André relates (André 1870) that when in the Garden of
Acclimatisation in Moscow in 1869 he was struck by various plants with big foliage recently
introduced from the most distant parts of the Russian empire. One such plant that particularly took
his interest was P. sachalinense, which the head gardener, Monsieur Desmurs, informed him had
come from Sakhalin Island (André 1893).

Concerning the introduction to Britain, Hooker (1881) recalls that it had been in cultivation at
Kew for about 20 years and that it had probably come from the collections of Wilford or Oldham.
This is indeed possible, as there is an 1859 Wilford specimen of F. sachalinensis (labelled as P.
cuspidatum) from Hakodadi (Japan) at K. The Kew “Plants Inwards” book for 1859-67 records
two sendings from Wilford in early 1860, but neither mentions P. cuspidatum or P. sachalinense
by name. The same ledger records the sendings of Oldham, also from Japan, in 1862-3. These are
not generally itemised, but seem to be primarily seed and spores; the April 1862 package being
dismissed as *..mostly common and well known species scarcely any germinated”. We have not
located any Oldham herbarium specimens of F. sachalinensis, but he cannot be ruled out as a
potential source of seed of this taxon.

Whatever the complexities and confusions of the early history of the introduction of live plants
or seed from Sakhalin and Japan to the West, whether via Russia or Kew, we do have good
documentary evidence of when this species was first available to the gardening public in Britain.
The 1869-70 catalogue (No. 48) of William Bull of Chelsea records P. sachalinense under “New
plants offered for the first time”. It was offered at the then considerable price of 7/6d per plant,
along with the assurance that “....coming from the Amoor country will be perfectly hardy”. [At this
period the “Amoor country” (Amur) included Sakhalin Island; F. sachalinensis is not known to
occur on mainland Amur]. The 1871 (No 59) catalogue notes that it “..is perfectly hardy”, which
demonstrates that they had overwintered it in Britain. That it was available in commercial
quantities to unleash on an unwary public by 1869 suggests that it had been in cultivation for a
year or more by then. Charles Maries, when in Japan in 1878-9, collected further material from the
central mountains of Japan, whilst employed by James Veitch & Son (Veitch 1906). We have seen
a herbarium specimen dated 1880 at Kew from this collection. If live material was brought back, it
would have added further to the genetic diversity of this taxon in Britain.

F. sachalinensis was also recommended, particularly on the continent, as a forage plant for cattle
(André 1893). In Britain it was planted for this purpose on “oozy hillsides in Yorkshire” (Davies
1896), though there is no record of any such plantings surviving there. It was even introduced to
Bengal for this purpose, and as a riverbank stabiliser, in the late 19th century, where it was known
as Knot-grass or Machute (Christy 1893). We have recently heard that it still survives along
rivercourses in Bengal (W. Griffin, pers. comm.). The Yokohama Nursery Company was in 1907
offering 100 lb of cleaned F. sachalinensis seed for $100, though we have not heard of any
plantings on this scale. F. sachalinensis also has certain pharmaceutical properties, and is used
commercially in Germany for the manufacture of a fungicide (H. H. Heine, pers. comm.).

Early plantings in the British Isles are known from Mr Garnett’s Penketh (near Warrington)
1892, Eastnor Castle (Herefordshire) 1886, Cosmo Melvill’s garden at Brook House, Prestwich
(Lancs.) 1901 and from a horticultural specimen of 1874; all Grindon (MANCH). In the 1890s
Surgeon-General Cook grew both sexes of F. sachalinensis in his garden near Lydney (Gloucs.)
(K).

F. sachalinensis plants in Europe possess much more genetic diversity than F. japonica var.
japonica (Hollingsworth 1998). This could be the result of multiple introductions from widely
separated areas or the introduction of seed rather than rhizomes, or a combination of the two.

100 J.P BAILEY: AND Az P_..CONG@ELY

FALLOPIA x BOHEMICA (CHRTEK & CHRTKOVA) J. P. BAILEY

F. x bohemica is the name given to hybrids between F. sachalinensis and F. japonica var.
Japonica or F. japonica var. compacta. It was first described in 1983 in a Czechoslovakian wildlife
magazine (Chrtek & Chrtkova 1983). It was not until 1986 that, at Leicester University, we
became aware of this; we had been independently describing the hybrids on morphological and
cytological grounds since the early 1980s (Bailey & Conolly 1985). A.P.C. had, however, earlier
been suspicious of certain specimens from Par (Cornwall) and Marston (near Oxford), the
determinations of which had over the years alternated between F. sachalinensis and F. japonica.
These suspicions were heightened by plants grown from seed sent by European botanic gardens,
which morphologically did not match the taxon they were sent as, and which were subsequently
found to be hybrids. Concentration on plants previously thought to be male-fertile F. japonica var.
Japonica, and which were also subsequently found to be hybrids, highlighted the morphological
features of the hybrid to us. Bailey er al. (1996) summarise the current knowledge of the hybrid’s
history and distribution in the British Isles. None of the locations listed there predates the South
Wylam, Co. Durham (v.c. 66) plant which has been established since at least 1954 (Sowerby, B.
BM). For the earliest British record, 100 years prior to its description, we are again indebted to the
Grindon Collection at MANCH. Here there are two collections (dated October 1872 and
September 1876) from Manchester Botanic Garden, on a single sheet labelled P. sieboldii, both of
which we have determined as F. x bohemica. A 1911 male-fertile specimen of F. x bohemica from
Charles Bailey’s garden “Haymesgarth” at Cleeve Hill near Cheltenham (v.c. 33) is especially
interesting as 1t came as a root from Smith’s Nursery, Worcester (see var. compacta section). This
is direct evidence that a nursery garden was supplying this taxon at an early date. The initial
hybridisation could also have occurred there, and such nursery establishments are likely origins of
the hybrid in general. The plants certainly had plenty of opportunity for hybridisation, as the
parental taxa had been growing together there since at least 1880 (Table 2). If Smith’s were
sending out F. x bohemica widely, one would expect that the same male-fertile clone could be
well-established in widely separated geographical locations; this is being tested by molecular work
currently underway at Leicester. Hybridisation must have occurred on a number of occasions,
whether in nursery, botanic or private gardens or even spontaneously from naturalised plants. This
is borne out by the fact that there are at least five genotypes of F. x bohemica known from Britain,
4x, 6x and 8x male-fertile plants and 4x and 6x male-sterile plants (Bailey et al. 1996).

THE “COOK MAMMOTH” ENIGMA
Catalogues of V. N. Gauntlett & Co. at Redruth from the early 20th century proudly announce:

“Polygonum ‘cooki’ - new mammoth species, introduced by Surgeon-General Henry Cook,
from North America some years ago, who writes us that this variety has formed an enormous
clump in his garden in Gloucestershire, of quite tropical appearance, with culms 16 feet high
and 5 inches in circumference. It is perfectly hardy and, like the species, spreads rapidly,
making a splendid plant for the wild garden or woodland drives, etc. We have secured the
whole stock of this plant. 7/6d.” (Gauntlett’s Hardy Plants Worth Growing, Cat. 88 p. 61).

This is the earliest record that we have located. Catalogue 88 is the earliest from the Japanese
Nurseries, Redruth which lists this taxon, and it predates the move of the company to
Chiddingfold, Surrey in 1906/7 (Willson 1988). Catalogue 88 cannot be earlier than 1901 since it
refers to His Majesty’s Government. At Chiddingfold they advertised their “P. Cookii” up to the
late 1920s/early 30s. From 1909 onwards, a photograph of “P. Cookii” is added. This photograph
is taken from some distance and is not very clear, but depicts either F. sachalinensis or F. x
bohemica. Were this plant actually to be F. x bohemica we have the fascinating possibility that the
catalogue entry constitutes valid publication of an earlier name for F. x bohemica! Perry’s of
Enfield were also selling P. “Cookii” between 1935 and 1937.

Henry Cook, in A Gloucestershire Wild Garden (1903), describes his garden “Priors Mesne”’,
near Lydney. Here he grew four large “Polygonums” in several places, including both male and
female Polygonum sachalinense on either side of a pond, with P. cuspidatum/sieboldii close by.
Thus there was potential here for on-site hybridisation and production of F. x bohemica. Cook also
notes that Kew had told him that they did not have the male P. sachalinense at the time of writing

HISTORY OF JAPANESE KNOTWEED 101

(1899): he swiftly rectified this by sending them a specimen the same year. This was followed in
1901 by a specimen (K) that we have now determined as F. x bohemica.

Whilst we have established that Cook had both ?. sachalinensis and F. x bohemica in his garden
prior to 1904, when Gauntlett’s would have collected their mammoth Polygonum, there is
unfortunately nothing to link either taxon to the Polygonum being distributed by Gauntlett’s as
“Cookii”. Thus it is not known whether the plant taken by Gauntlett’s from Cook’s garden was F.
x bohemica or just a large clone of F. sachalinensis. In any case the herbarium specimen (K) is
good evidence of F. x bohemica being a valued garden plant. Whether it originated in his garden,
spontaneously or from deliberate planting of seed, or even in North America may never be known.

PROMOTERS AND PERPETRATORS

Having once established the probable date and place of arrival of these taxa in the British Isles, we
are then set the task of explaining the current widespread distributions of F. japonica var.
japonica, F. sachalinensis and F. x bohemica. For this we must turn our attention to the
enthusiastic gardeners of the time, the commercial nurseries that pandered to their insatiable
demand for novelty and the botanic gardens which were involved in widespread exchange with the
former.

THE ROLE OF COMMERCIAL AND BOTANIC GARDENS

When looking at the dispersal of a garden plant an obvious place to start is the gardening literature.
When available, herbarium specimens are of course direct evidence. The Nursery garden
catalogues themselves are good primary sources of data. Although some catalogues were quite
substantial publications (e.g. Gauntlett’s, Veitch’s), they are quintessentially ephemeral items, and
interest would generally be restricted to the current issue. Hence it is not unusual to come across
undated and unnumbered catalogues. Another difficulty is, of course, that few people had the
foresight to save them. The Royal Botanic Gardens at Kew and Edinburgh and the Royal
Horticultural Society's Lindley Library have substantial collections, but few other than the most
famous nurseries are represented in any quantity, and complete runs appear virtually unknown.
This is hardly surprising when considering the vast numbers of nurseries in business in the
Victorian period. Even scarcer are catalogues from the 1850s and 60s - the period of the initial
dispersal. Apart from those of Von Siebold & Co., which were fortuitously republished in a
contemporary journal, we have seen no catalogues dated earlier than 1869. However, in spite of
the fragmentary nature of the resources available, we have gained evidence of a substantial
nationwide trade in these plants (Table 2). Nurseries offering them are well distributed throughout
the British Isles - not that this was a prerequisite for effective dispersal, since a highly efficient
postal service existed during the period under consideration. This list also provides a historic
record of where and when F. japonica and F. sachalinensis were being grown together, giving the
potential for hybridization.

The prices generally follow a similar pattern, a high price for novelty followed by a sharp drop,
as illustrated by F. sachalinensis in the Bull catalogues and Cook’s mysterious Polygonum offered
for over 30 years by Gauntlett’s. Another point of interest is the scale of the trade in these plants:
Sieboid, for instance, giving a price per hundred F. japonica plants and the Yokohama nursery
offering F. sachalinensis seed in 100 Ib lots. Gauntlett’s and others were offering F. sachalinensis
by the dozen - clearly not aimed at the ordinary suburban garden. Another point is the sheer length
of time that these plants were retained on sales lists in spite of their true nature being known.
Though we have not been searching actively for latest dates, Table 2 shows that F. japonica was
on sale for at least 87 years! Indeed, the mottled variant of F. japonica, “P. spectabile”, was still in
Beth Chatto’s list in the 1980s.

GARDEN NOMENCLATURE

Due to the mixed usage of horticultural and scientific names and the idiosyncratic application of
some epithets, a certain amount of botanical licence has been employed in the interpretation of
these taxa in the Nursery catalogues. Although Siebold was one of the authorities for P.
cuspidatum, he used the eponymous P. sieboldii Reinw. in his catalogues, and it is under this
combination that all the earliest references occur. As the compacta epithet was not published until
1880, another name was needed to differentiate the two varieties of F. japonica in the earlier
catalogues. One way of doing this was by keeping sieboldii for the normal plant and using

J.P. BAILEY AND A. P. CONOLLY

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HISTORY OF JAPANESE KNOTWEED 103

cuspidatum for the dwarf, as was done in Smith’s catalogue 1880/81. The name P. reynoutria was
also applied to a dwarf variant earlier this century. The Giant Knotweed did not provide any such
problems, as it was always listed under its correct name (even though spelt in a number of creative
ways).

There is considerable evidence from the literature and herbarium specimens that plants were sent
out under incorrect names. C. Wolley-Dod noted that Standish’s nursery (Ascot) were sending out
F. japonica var. japonica under the name P. japonicum. The Grindon collection at Manchester
Museum (MANCH) has an impressive collection of early Japanese Knotweed specimens, with
interesting comments and press-cuttings attached. One such specimen implies that Smith’s of
Worcester may have been sending out var. compacta labelled as P. alpinum. Another specimen
cultivated at the gardens of the Royal Botanic Society London in 1867 is labelled P. sibericum.
These names are not synonyms. A sheet with specimens of F. japonica var. japonica and F.
sachalinensis (1878) (MANCH) has an 1894 letter attached from a seed company to Leo Grindon
requesting “the specific name of the enclosed Polygonum”.

Such errors are perhaps inevitable during a period of rapid expansion of the exotic taxa available
to gardeners. Difficulties of identification of naturalised alien taxa are to be expected, as there is
inevitably a time lag between naturalisation and the incorporation of adequate accounts in the local
Floras. Even up to the first edition of Clapham et al. (1952), P. polystachyum and F. sachalinensis
were not readily separable.

THE WILD GARDEN MOVEMENT

It was William Robinson (1838-1935), one of the most influential of the late Victorian gardeners,
who advocated replacing the regimented “carpet-bedding” of High Victorian practice (cf. Carter
1984; Thacker 1994) by a more informal “lay-out” and a more natural setting enhanced by the
“new” exotic species then becoming available . He promoted his ideas through articles in The
Gardeners’ Chronicle and other journals, culminating with the publication in 1870 of the highly
influential book The Wild Garden. This book was subtitled “Our Groves and Shrubberies made
beautiful by the Naturalisation of hardy exotic plants”. He described Polygonum sieboldii
enthusiastically, and recommended P. cuspidatum under “Plants with large or graceful foliage
suitable for naturalization”. The enormous influence of Robinson’s writings led, for example,
Surgeon-General Henry Cook, whilst in Bombay with the Indian Medical Service, to dream of
making just such a garden of his own when he retired. The resultant Gloucestershire garden, where
he grew many of the alien species of Polygonum, is described by Cook (1903) and is an important
record of early plantings.

In The English Flower Garden (1898), Robinson states that “the great Japan Knotworts
(Polygonum) are handsome in rough places in the wild garden’, and that there is “no better plant
for semi-wild places, or for association with vigorous things on the turf in the pleasure ground”
than Polygonum sachalinense.

The influence of Gertrude Jekyll (1843-1932) was just as far-ranging, abroad as well as in
England, though less concerned with naturalisation and woodland. However, in A Gardener’s
Testament (Jekyll, F. & Taylor 1982) she is quoted as envisaging a woodland-walk flanked by
groups of “plants of rather large stature ... bamboos and the great Knotweeds of Japan”. In Home
and Garden (1900) “we ought not forget the quick growing ways of the great Japan Knotweeds
(Polygonum) growing fast and tall”. At least two of her garden plans include Fallopia japonica
var. compacta: the wild garden at Little Aston, Birmingham, and Drayton Wood, Norfolk
(Bisgrove 1992). Mrs C. W. Earle, a Surrey enthusiast of the “wild garden” (1897), is ecstatic in
her praise of her “favourite Polygonum cuspidatum” - the “‘..handsomest, easiest-grown, hardiest,
most useful plant for London gardens”. F' sachalinensis is also recommended, but said to lack the
“..beautiful up-standing and yet graceful growth.” of F. japonica.

THE ROLE OF PRIVATE GARDENS AND THE KNOTWEED CIRCLES

Canon Ellacombe of Bitton (Gloucs.), Miss Gertrude Jekyll in west Surrey, E. A. Bowles and Miss
Ellen Willmott in Essex, and the Rev. C. Wolley-Dod in Cheshire formed a “circle” of specialist
gardeners who corresponded, visited each other and exchanged plants, ideas and expertise. They
all had exceptionally rich gardens: Ellacombe’s one and a half acres contained a “unique gathering
of out of the way plants” (see Hill 1919). Apart from British nursery gardens they could have also

104 J.P. BAILEY AND A. P. CONOLLY

ordered from continental nursery gardens such as Von Siebold & Co. or Haage & Schmidt, both
early suppliers of F. japonica (Table 2). At least some were also in regular receipt of plants from
the major botanic gardens including those on the continent. There was regular two-way traffic in
new plants between Kew Gardens and these wealthy amateurs - Ellacombe was recorded as
supplying Kew with some 20 species for description in Curtis’s Botanical Magazine. He was also
in correspondence with Kew directors from J. D. Hooker onwards: the 1881-95 Kew ‘Plants
Outward” book recording 49 dispatches of plants to him.

An 1882 specimen of F. japonica from Ellacombe’s garden is now in the Grindon collection
(MANCH). Leo Grindon was a botanist and field naturalist interested in alien plants; he was an
avid collector, and maintained a wide circle of correspondents. In addition to Ellacombe’s,
specimens of Japanese Knotweed s./. in his collection come from Charles Bailey at Cleeve Hill
(Cheltenham), Mr. Garnett of Penketh (near Warrington), John Wood of Kirkstall (Leeds), J.
Cosmo Melvill at Prestwich, Mr Scott at Denzell (near Altrincham) and from Eastnor Castle
(Herefordshire). There were also specimens from Manchester Botanic Garden and the Garden of
the Royal Botanical Society, London (see Appendix 1).

Exchanges within and between these two groups could well have included one or more of the
Japanese knotweeds they were known to have grown. Further dissemination beyond these
immediate circles could have followed later.

CLUSTERS AND ASSOCIATIONS

In consideration of the subsequent history of the escape and naturalisation of these Fallopia taxa
(Conolly 1977; Bailey et al. 1996) we need to distinguish between F. japonica and the two taxa of
greater stature, F.sachalinensis and F. x bohemica. Since F. japonica var. japonica is now of
almost ubiquitous occurrence, it is in a sense too late to deduce anything from the absence or
presence of this taxon in any particular area. The other two taxa on the other hand have a much
less “mature” pattern of distribution, and are much more restricted in occurrence and _ habitat.
There are also important differences in the history of their introduction - F. sachalinensis having
been recommended as a forage plant. There is also evidence of its being used on large estates for
scenic plantings and cover for shoots. It is certainly true that F. sachalinensis is found more often
than F. japonica well off the beaten track on old estates, as if it had originally been deliberately
planted.

The case of the hybrid F. x bohemica is a little more complex; in addition to vegetative spread,
it can also arise from seed in locations where F. japonica occurs with male-fertile F. sachalinensis.
This is most likely to occur in nursery and botanic gardens and in private gardens extensive
enough to accommodate such large plants. The association of F. sachalinensis and F. x bohemica
with large estates and nursery gardens will be explored to see to what extent current stands of these
two taxa can be related to such estates either as residuals or escapes. We also examine in more
detail those areas where there are several adjacent spots on the B.R.C. 10-km recording scheme
map, which we refer to as clusters.

By far the largest cluster of F. x bohemica in the British Isles is in West Surrey (v.c. 17), where
there are more than 35 stands, some dozen of which are associated with F. sachalinensis. They
occur in the area bordered by Dorking, Guildford, Godalming and Haslemere - more specifically
the “Surrey Hills”. This is the heart of Gertrude Jekyll country, and it is tempting to see a direct
link with the fact that her house, ““Munstead Wood’, was in the centre of this area with its
numerous large estates and expensive country houses. Many of these houses were designed by
Lutyens, with Jekyll designing the gardens. Many of the present stands are close to such
properties, and the abundance of stands of both taxa in and around Holmbury St. Mary, close to a
group of Jekyll gardens, is surely significant. But proof of a direct recommendation by Jekyll and/
or of supply by, for example, Gauntlett’s at nearby Chiddingfold is more difficult to establish; and
in any case there were many other nursery gardens in the vicinity. Gauntlett’s did, however, offer
Polygonum sieboldii, P. sachalinense and the “mammoth” P. “Cookii” for many years. Moreover,
stands, possibly relics, of F. x bohemica grow today a short distance downstream from the former
Gauntlett site (Mrs J. Smith, pers.comm.).

In another cluster of this kind, in West Cornwall, there are a number of associations between
well known gardens and the occurrence of Fallopia taxa. Buryas Bridge, Penzance, where the
unusual F. japonica var. japonica x F. japonica var. compacta hybrid grows (Bailey & Conolly
1991), is near the Trewidden estate (formerly the property of T. Bolitho) where over 450 species

HISTORY OF JAPANESE KNOTWEED 105

of plants were grown at the end of the last century (Pett 1998). A former nursery garden is also
nearby. F. x bohemica persists at Lanarth on the Lizard peninsula, where P. D.Williams (1865—
1935), R.H.S. medallist and one of the greatest Cornish gardeners, had his estate. His relatives had
nursery gardens close to Redruth, where Gauntlett’s, prior to their 1906 move to Surrey, sold a
range of Japanese Knotweed taxa.

In South Wales, concentrated in Glamorgan , there is a further such cluster of F. x bohemica or
F. sachalinensis sites associated with estates of “landed gentry”. Clearly derived from or residual
are those at Black Pill near Clyne Castle (v.c. 41), former estate of the Vivians (Lord Swansea);
Tredegar Park (v.c. 35, Lord Tredegar); Cyfarthfa, Merthyr Tydfil (Crawshays estate); Craig-y-
nos, (once the home of Adeline Patti); Gabalfa House, Cardiff and Velindre, Whitchurch (now a
mental hospital).

In Scotland too, there are further such clusters of F. x bohemica sites, for example in the
Glasgow area, mainly along the River Kelvin, possibly relating to the Botanic Garden. Further
north, on the Morayshire coast (v.c. 95) around Forres is another Fallopia cluster. Here, in the
gardens at Kellas House to the east of Forres, both F. sachalinensis and the hybrid occcur, and
along the Muckleburn downstream of Dalvey House (west of Forres) F. sachalinensis is abundant.
Further downstream, where the Muckleburn approaches the river Findhorn up to a 100 ha of F. x
bohemica are found. The three taxa also occur at a number of other sites in the area. Kellas House
and Dalvey House are probably responsible for much of these introductions, but Brodie Castle,
home in the last century to the Brodie of Brodie - a noted horticulturist - is also a possibility (v.c.
95 information from I. Green, pers.comm.).

In England and Wales a high proportion of those stands which occur as solitary (non-clustered)
10-km entries are also associated with local estates and gardens. These include F. sachalinensis at
Edwinsford (Carms.); Falcondale (Cards.), by a lake; Nant-y-Frith, Flintshire (abandoned garden);
Easney, Ware (Herts.), a pheasant shoot cover; and at Leigh near Tonbridge (Kent) - all of which
are on or close to an estate. In the east of England F. sachalinensis has long out-lived the nursery
garden of R. Wallace at Kilnfield Gardens, Colchester, that once cultivated it (fide J. Heath) (Table
Zs).

There are, of course, many other isolated stands of these taxa, in quite other habitats, not
associated with estates but naturalised on roadsides, railway tracksides or banks, for which some
other explanation is called for.

THE FALL FROM GRACE

The “hype” of the early Siebold introductions, coupled with its high price and gold medal status,
leaves little doubt of the initial desirability of F. japonica to the Victorian gardener. It was
undoubtably a spectacular plant, reliable and hardy even in the far north of the country; and of
course there was a much more limited range of exotic garden plants available then.

John Wood of Kirkstall (1884, p. 208) heaps unqualified praise on F. japonica var. japonica for
its “..gracefully arching stems” and declares var. compactum to be a handsome bush. He
unaccountably declares F. japonica var. japonica to be “a capital plant for the small town garden”’.
He could, however, be accused of having a vested interest, since he was later to go into the nursery
business (Table 2), selling these selfsame plants at 6 pence to 9 pence each!

By the time of Robinson (1898) qualifications were beginning to creep in. Whereas F.
sachalinensis and var. compacta are still both recommended, var. japonica, we are warned “..1s
weedy, and in light soils springs up everywhere’. Similarly, Gertrude Jekyll (1899) cautions that
“P. compactum must be reduced” and “P. sieboldii planted with caution”. Walters (1887), in his
account of the flora of Alexandra Park in Oldham, noted that F. japonica var. japonica “..turns up
unexpectedly in nearly every piece of cultivated ground”.

Japanese Knotweed was not long content to remain in gardens and, from the beginning of the
century, increasing numbers of escapes were reported (Conolly 1977). By the 1930s F. japonica
was known in East Cornwall by the picturesque name of “Hancock’s Curse’, and its presence
reputedly reduced the price of a house there by £100 (Conolly op. cit.). It is tempting to link this
with Hancock’s nursery at Liskeard which is known to have been in business for at least the period
1897-1906 (Kelly 1897, 1902). In spite of its invasive proclivities now being well-known,
Japanese Knotweed was still being offered in the mid-1930s by nursery catalogues such as
Gauntlett’s of Chiddingfold (Table 2).

106 J.P. BAILEY AND A. P. CONOLLY

Today F. japonica var. japonica poses very serious problems for river authorities, amenity areas
and nature reserves alike, and a great deal of money is spent annually on surveying and control
measures. F’. sachalinensis is not such a problem, but can nonetheless occupy considerable areas
(e.g. Amroth v.c. 45); F x bohemica is a very invasive taxon, and its great stature makes its
occurrence even more intrusive. These taxa are a particular problem in South Wales; the 1998
survey of 400 km? of the Swansea city and county found that F. japonica var. japonica occupied
approximately 99 hectares, spread across a variety of habitats (S. Hathaway, pers. comm.). Its
frequent escape in Glamorganshire as early as 1907 was noted by Conolly (1977).

In 1981 the Wildlife and Countryside Act proscribed two land plants, Giant Hogweed and
Japanese Knotweed, making it an offence to introduce these species to the wild. This has
implications in terms of the redevelopment of sites infested with F. japonica since earth movement
and tipping are major factors in spreading the plant further. Duty of care obligations can add
considerably to the cost of site development, and contractors are becoming increasingly cautious
of taking on infected sites because of the costs of excavation and deep tipping of infested soil.
Hancock’s Curse is still with us!

FUTURE PROSPECTS

This paper leaves quite a few unresolved points and, although on first sight these might appear to
be insoluble, there are modern techniques that may be able to shed some light on these historic
matters. The main unresolved points are:

e Has our F. sachalinensis come from Sakhalin or Japan or both and on how many
occasions?

e How many times has F. x bohemica arisen in Britain?

e What are the relative roles of vegetative reproduction and seed production in the current

distribution of F. x bohemica ?
e Was P. “Cookii” really F. x bohemica?
e Where in Japan did the atypical British clone of F. japonica var. japonica originate?

The work of Hollingsworth & Bailey (2000) has indicated that it is possible to use DNA
techniques both to identify particular clones and to extract information on the native region of

origin of such plants. Work at Leicester University is currently underway on a number of these
problems, helped by B.S.B.I. funding.

POSTSCRIPT

Looking back at the original importers of Japanese Knotweed s./., little did Siebold realize what
was in store for the West, when he eventually got his precious Japanese plants back to Leiden, and
thus in 1850 despatched that fateful package to Kew Gardens. The role of Kew in subsequent
events may not be inconsiderable!

We admire rather than deplore the efforts of the early explorers from Russia who first collected
F. sachalinensis from Sakhalin and Japan: the naval surgeon Weyrich who “profited by every stay
of the vessel to botanize” (Bretschneidér 1898, p. 618), and later Maximovicz travelling back from
Japan with his enormous living collections. Their specimens were not collected primarily as
novelties for gardeners, but for their taxonomic value.

Nor should we blame the early English collectors of F. sachalinensis, such as Maries sent by
Veitch, and Oldham and Wilford from Kew, sent to gather potential horticultural jewels, who, in
spite of appalling conditions and risks, still managed to discover and bring back living new
species. For they could not have guessed that their collections might include potential invaders.
Indeed, this handsome Giant Knotweed has rarely if ever been a menace here. Nevertheless, with
the introduction of male-fertile plants came the potential for crossing with the male-sterile F.
japonica to produce our third taxon, the highly invasive F. x bohemica. A warning maybe of a
menace to come!

HISTORY OF JAPANESE KNOTWEED 107

At this point we recall the debt we owe to the efforts and encouragement of Duggie Kent and his
generosity in sharing his superior Knotweed knowledge, without which we would not now be
attempting to fit further pieces into the intriguing Japanese Knotweed jigsaw puzzle.

ACKNOWLEDGMENTS

The authors are indebted to David McClintock for generously sharing his years of Knotweed
experience. We thank the staff at Herbaria at the Natural History Museum (BM), Manchester
(MANCH), Kew (K) and Edinburgh (E) for assistance and loans. We are particularly grateful to
the Royal Botanic Gardens at Kew and Edinburgh and the National Botanic Gardens, Glasnevin
for access to their plant accession archives and Victorian nursery garden catalogue collections. We
gratefully acknowledge the assitance of The R.H.S. Lindley library, the Natural History Museum
botanical library, the Linnean Society’s library and in particular the Society’s librarian, Gina
Douglas. We also thank our many botanical collaborators and correspondents, Ann Boucher,
Arthur Chater, Lois Child, John Franks, Ian Green, W. Griffin, Shaun Hathaway, J. Heath, the late
H. H. Heine, George Hutchinson, Alan Leslie, Rose Murphy, Kevin Pyne, Joyce Smith, Clive
Stace and Roy Vickery. Finally, we acknowledge the assistance of the Herbaria at Uppsala (UPS),
Geneva (G), Leiden (L) and Leningrad (LE) for loan of herbarium specimens.

REFERENCES

ANDERSON, A. W. (1951). Dr. F. P. von Siebold. Gardeners’ Chronicle Feb. 17 1951: 52.

ANDRE, M. E. (1870). Un mois en Russie. Masson & Sons, Paris.

ANDRE, M. E. (1893). La Sacaline. Revue Horticole 64: 326-327.

ANON. (1943). Wisley in September. Journal of the Royal Horticultural Society 68:251-253.

BAILEY, J. P. (1989). Cytology and breeding behaviour of giant alien Polygonum species. Ph.D. Thesis,
University of Leicester.

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(Accepted October 1999)

HISTORY OF JAPANESE KNOTWEED 109

APPENDIX 1 HERBARIUM SPECIMENS CITED

F. JAPONICA VAR. JAPONICA

[Polygonum multiflorum «], Herb Thunberg 9705 UPS

Polygonum cuspidatum S. & Z. Legit v.S, Herb. Lugd. Batav. No. 908.174 826 L

. cuspidatum Sieb. Hort, Kew 1857, Herbarium Hookerianum K

. cuspidatum coll. R. Oldham, Japan No. 235 10/1861. Note saying “..up to 8ft high usually one
stemmed but sometimes branching near base”. Good seed set 2 sheets - male and female. K

. cuspidatum coll. R. Oldham, Japan No.337 with seed K

. cuspidatum coll. R. Oldham, Nagasaki Japan No 235 1862 female. K

. cuspidatum coll. Maximowicz Yokohama 1862. (Lvs crinkled like compacta.) K

. cuspidatum coll. Maximowicz iter Secundum, Yokahama 1862 (exserted stamens).

cuspidatum var. humilis rubiflora coll. Maximowicz Yokohama 1862 “cult Yedo” labelled
female K

P. cuspidatum coll. Maximowicz Nagasaki 1863, male and female (- compacta-like) K

[P. sibericum] cult. R.Bot. Soc. London Garden, 1867 (male-sterile) MANCH

P. cuspidatum Mr. Ellacombe’s, Bitton, Oct. 9 1882 (abundant good fruits) MANCH

pel ist as) ae) el ane) te}

F. JAPONICA VAR. COMPACTA

P. cuspidatum var. compactum, Mr. John Wood’s, Kirkstall, Leeds, July 16 1881 MANCH

P. cuspidatum var. compactum, Mr. Scott’s Denzel, (Altrincham), Aug 31 1883 only 3ft high
MANCH

[P.alpinum] Herb Charles Bailey, Cult. Cleeve Hill 13 Oct 1911 ex Smith’s nursery MANCH

[P.alpinum] Herb Charles Bailey, Cult. Cleeve Hill 30 July 1911 ex Smith’s nursery MANCH

P. cuspidatum var. compactum, Melrose Abbey 1915, coll. ILM. Haywood det. Thellung. E

P. cuspidatum var. compactum Bowles’ Garden Myddleton House, Bulls Cross. W.T. Stearn 3
September 1954. BM

F. SACHALINENSIS

Polygonum sachalinense F. Schmidt, Ins. Sachalin. Weyrich 1853 LE

[P. cuspidatum] Hakodadi Japan, 1859 coll. C. Wilford. K

P. sachalinense Sachalin Island, P. von Glehn 1861 K, BM

P. sachalinense Hakodate1861, Maximowicz iter Secundem. Ex Herb horti bot Petropolitani K

P. sachalinense coll. Maxim 2nd Japan, Hakodate Japan 1861 BM

P. sachalinense F. Sch. “Siberia” Hort. [cf. K] 23/9/1874 8-9ft fine foliage MANCH

[P. sachalinense] dated July 2 1878 Kew (F. japonica leaves on same sheet). 7 Sept. 1894 letter

from Dickson Brown & Tait attached. MANCH

P. sachalinense central mountains of Japan Aug1880, coll Mr. Maries and presented by Messrs
Veitch. Labelled male plant Hort Kew K

sachalinense Sent from Eastnor Castle, Herefordshire, Sept. 10 1886 MANCH

sachalinense Mr. Garnett’s Penketh, (nr Warrington) Sept. 1892 MANCH

sachalinense male-fertile specimen labelled from “Surgeon General ‘Cobbs’ (Cook’s) Garden
Lydney” Sept 1899 K

sachalinense Cult Kew Sept 1899. (In same hand as “Cobbs” (Cook’s) specimens, female) K

sachalinense Brook House, Prestwich, (Lancs.) J. Cosmo Melvill cult in hort herbaceo ex Insula
Saghalin, Aug. 1901 MANCH

We eras

F. x BOHEMICA

[P. sieboldii] Manchester Botanic Garden, Two specimens on one sheet; Oct. 1872 & Sept. 23
1876 MANCH

[P. sachalinense] Sent by Mr Cook, Lydney Gloucester 13.9.1901 K

[P. cuspidatum] Charles Bailey, Haymesgarth, Cleeve Hill, near Cheltenham 22/9/1911. (Male-
fertile) root from Smith’s nursery MANCH

[P. sachalinense] Marston, Oxford Sept 1922 coll. Claridge Druce (male-sterile) MANCH

110 J.P. BAILEY AND A. P. CONOLLY
FALLOPIA MULTIFLORA

Polygonum multiflorum Herb Thunberg 9706 UPS
[Polygonum (Helxine) Japonicum] Mihi. Vid Nat His. VHI Pl. 49 Fig 3, Herb Delessert collection
Burman unnumbered G

Watsonia 23: 111—121 (2000) tii

Hybridisation and clonal diversity in some introduced Fallopia
species (Polygonaceae)

M. L. HOLLINGSWORTH*

Royal Botanic Garden, Inverleith Row, Edinburgh, EH3 5LR

and
J. P. BAILEY

Department of Biology, University of Leicester, Leicester, LE] 7RH

ABSTRACT

Molecular markers have been used to examine the amounts of genotypic diversity and sample inter-
relationships among introduced invasive Fallopia taxa in Britain. The highly invasive Japanese Knotweed
(Fallopia japonica var. japonica) appears to be represented by a single clone in Britain. In contrast, the related
species Giant Knotweed (F. sachalinensis) and their hybrid (F. x bohemica) show much higher levels of
clonal diversity. The difficulties of distinguishing between recurrent origins and hybrid fertility are discussed
in relation to the variability of F. x bohemica. Finally, a very preliminary survey of native Asian F. japonica
has detected high levels of genetic diversity. This, coupled with some limited morphological and cytological
observations, suggests that a taxonomic revision of Fallopia section Reynoutria is needed to clarify the status
of British F. japonica.

INTRODUCTION

Many plant communities contain a large proportion of introduced species. Human activities have
vastly accelerated the movement of plants, carrying thousands of species, either deliberately or
accidentally, across natural barriers such as oceans and mountain ranges. Many of these introduced
species have become naturalised in the wild and some have become invasive weeds. In Britain,
alien plants form a significant component of the flora, with a recent account reporting 1391 aliens
out of the 2963 species recorded (Stace 1997).

Hybridisation is a frequent and important component of plant evolution and is widespread
amongst natural populations (Rieseberg & Ellstrand 1993). Occasionally, alien species hybridise
with native or other introduced species. In the New Flora of the British Isles, Stace (1997)
recognised 770 angiosperm hybrids; of these, 58 involve at least one non-native taxon and 12 are
hybrids between two alien species.

Hybridisation involving an introduced species, either with a native species or another introduced
taxon, can give rise to hybrid combinations that would be unlikely to occur naturally due to a lack
of sympatry of native ranges. The evolutionary implications of such post-introduction interactions
are well documented and provide some of the clearest and most unequivocal examples of
speciation such as the allopolyploid origins of Senecio cambrensis Rosser (Ashton & Abbott 1992;
Harris & Ingram 1992) and Spartina anglica C. E. Hubb. (Marchant 1967; Raybould et al. 1991).

Introduced plants make useful model systems for the study of evolutionary processes such as
hybridisation and speciation. Long term historical factors can be eliminated as confounding
variables if the introduction dates are known. We can study evolutionary events as they are
happening (or at least very recently after they have happened) rather than attempting to make
inferences about distant historical events. In addition, as some introduced plants show weedy
tendencies, it is also informative to study the amounts and partitioning of their genetic variation, as
this can provide information on the population genetic consequences of colonisation.

“Corresponding author: e-mail: M.Hollingsworth @rbge.org.uk

2 M. L. HOLLINGSWORTH AND J. P. BAILEY

INVASIVE FALLOPIA SPECIES IN THE BRITISH ISLES

FALLOPIA JAPONICA VAR. JAPONICA - JAPANESE KNOTWEED

Perhaps the most notorious invasive plant in the British flora is Japanese Knotweed, F. japonica
var. japonica. This species is native to Japan, Taiwan and Northern China where it grows along
roadsides, river banks and managed pasture, and it is one of the dominant colonists of lava fields
(.C.O.L.E. 1997). It is insect-pollinated and visitors to its flowers include flies, honey bees and
wasps (Tanaka 1966). F. japonica var. japonica, as recognised in Britain today, was introduced to
the British Isles in the 1850s (Bailey & Conolly 2000). Since then it has spread rapidly. The first
recorded escape from cultivation was in 1886 (Storrie 1886); by 1996 it had been recorded from
55% of the 2862 10-km squares of the Biological Records Centre mapping system of the British
Isles (B.R.C. unpublished data). Its spread has been documented by Conolly (1977) and Bailey &
Conolly (2000).

In Britain F. japonica var. japonica occurs in a variety of man-made habitats such as canal sides,
road verges, railway embankments and cemeteries, as well as stream and river banks. The
abundance of this plant in urban areas and the costs associated with its control have given it much
bad publicity. Its shoots are able to push up through asphalt and damage pavements, car parks and
other public facilities and the height to which the plants grow leads to reduced visibility along
roadsides and railways. Along water courses, decaying shoots can cause blockages and dense
stands on riverbanks can impede flow in high water, thus increasing the risk of flooding. Since
1981 it has been a criminal offence to knowingly introduce F. japonica var. japonica into the wild.

FALLOPIA SACHALINENSIS - GIANT KNOTWEED

A closely related species to F. japonica var. japonica is F. sachalinensis (Giant Knotweed).
Fallopia sachalinensis occurs in Britain, again stemming from a horticultural introduction
(Conolly 1977; Bailey & Conolly 2000). It is native to eastern Asia where it is known from the
southern part of Sakhalin Island, the southern Kurile Islands, the Japanese islands of Hokkaido and
Honshu, and Korea (Sukopp & Starfinger 1995). It occurs in disturbed habitats from sea level to c.
1050 m and can be found along roadsides, forest edges, cliff tops and river sides, and like F.
Japonica var. japonica, it 1s one of the pioneer colonists of the “lava deserts” caused by volcanic
eruptions (Sukopp & Starfinger 1995). Fallopia sachalinensis shows less invasive tendencies than
F. japonica var. japonica and is still a rather uncommon plant in much of the British Isles (Bailey
1997). The first account of this species growing outside cultivation was published in 1896 (Davies
1896).

OTHER INTRODUCED FALLOPIA SPECIES

In addition to F. sachalinensis and F. japonica var. japonica, other members of the genus which
have been introduced to the British Isles are F. japonica var. compacta and F. baldschuanica.
Fallopia japonica var. compacta is a dwarf variety of F. japonica which is only rarely found
outside of cultivation and is known from only 30 10-km squares of the Biological Records Centre
mapping system (B.R.C. unpublished data). Fallopia baldschuanica (Russian-vine) was
introduced from central Asia and is a common garden plant. It frequently occurs as discarded
material on waste ground, but is rarely well naturalised (Stace 1997). Although reference will be
made to these two taxa in this paper, we primarily concentrate on F. sachalinensis and F. japonica
var. Japonica.

REPRODUCTIVE BIOLOGY OF FALLOPIA JAPONICA AND F. SACHALINENSIS

Both F. japonica and F. sachalinensis are gynodioecious and can occur as either male-sterile
(female) or hermaphrodite plants. In Britain, both female and hermaphrodite forms of F.
sachalinensis and F. japonica var. compacta have been recorded, but only female plants of F.
Japonica var. japonica are known (Bailey 1994). Fallopia sachalinensis and F. japonica var.
compacta thus have the capacity to reproduce both sexually and asexually (via vegetative
propagation). However, despite the remarkable spread and abundance of F. japonica var. japonica,
as no male-fertile plants are known from Britain, it appears that seed dispersal is not responsible
for its success (no evidence for apomictic seed production has been recorded). Rhizome fragments

HYBRIDISATION AND CLONAL DIVERSITY IN FALLOPIA 113

as small as 0-7 g are capable of regenerating into new plants (Brock & Wade 1992) and the
transport and tipping of earth containing plant fragments and water-borne dispersal along river
systems are considered to be the major means of dispersal (Conolly 1977).

HYBRIDISATION

Morphological and cytological studies have provided convincing evidence that hybrids between F.
japonica s. 1. and F. sachalinensis occur in the British Isles (Bailey & Conolly 1985). The hybrid
between F. japonica and F. sachalinensis - F. x bohemica (Chrtek & Chrtkova) J. Bailey - was
first described in 1983 in the Czech Republic (Chrtek & Chrtkova 1983). Two taxonomic crosses
have been detected in Britain: plants with 2n = 66 stemming from a cross between F. japonica var.
Japonica (2n = 88) and F. sachalinensis (2n = 44); and plants with 2n = 44 stemming from a cross
between F. japonica var. compacta (2n = 44) and F. sachalinensis. Very rarely, hybrids have also
been reported between F. japonica var. japonica and F. japonica var. compacta; these plants have
2n = 66 (Bailey & Conolly 1991). Interestingly, the majority of seed on British F. japonica var.
Japonica appears to have F. baldschuanica as the pollen donor and these progeny are 2n = 54
(Bailey 1988). However, only one such hybrid is known to have become established in the wild in
Britain (Bailey 1988). In contrast the hybrid between F. japonica var. japonica and F.
sachalinensis is relatively common in Britain (Bailey et al. 1996). This hybrid shows some of the
invasive tendencies of F. japonica var. japonica and occurs as both female and hermaphrodite
plants.

Although morphological and cytological data have provided much useful information about this
complex in Britain, many questions regarding the biology of these plants require additional data.
For instance, while there is very good evidence that hybrids between some Fallopia taxa are
present in the wild, we have no information on how many times such hybrids may have arisen, and
indeed to what extent hybrid fertility allows for generations beyond the FI. In addition, little is
known about the amounts and partitioning of genetic diversity in the parental taxa. The lack of
male fertile plants of F. japonica var. japonica in Britain suggests that this taxon may have an
extremely narrow genetic base and, at an extreme, could consist of a single widespread clone. In F.
sachalinensis, which is represented in Britain as both hermaphrodite and female plants, the
possibility for both sexual and asexual reproduction exists, although the relative importance of
either is unknown. In this paper, we summarise our published and unpublished data which have
attempted to shed light on some of these issues. Particular attention is paid to the relative
importance of sexual versus asexual reproduction, and the dynamics of inter-specific hybridisation,
concentrating primarily on F. japonica var. japonica, F. sachalinensis and their hexaploid (2n =
66) hybrid F. x bohemica.

LEVELS OF CLONAL DIVERSITY IN THE PARENTAL TAXA

FALLOPIA JAPONICA VAR. JAPONICA

To assess the levels of clonal diversity of F. japonica var. japonica in the British Isles, we
collected 150 samples from localities ranging from Land’s End to Shetland (Hollingsworth &
Bailey, submitted). These samples were assayed for genetic variability using the arbitrary DNA
fingerprinting technique RAPDs: Randomly Amplified Polymorphic DNA (Williams ef al. 1990).
Using ten primers, we amplified a total of 108 reproducible bands. No genetic variation was
detected between any of the 150 samples collected from across the British Isles (Hollingsworth &
Bailey, submitted). These data, coupled with the absence of male fertile individuals in the British
Isles, provide strong evidence to support the hypothesis that the entire British population of F.
Japonica var. japonica consists of a single extraordinarily vigorous clone (Hollingsworth &
Bailey, submitted). Furthermore, preliminary studies on a few samples from Europe and North
America also showed the same genotype (genetic individual), which suggests that the distribution
of this genotype extends well beyond the British Isles (Hollingsworth & Bailey, submitted). It
seems possible that the enormous social and economic costs associated with the Japanese
Knotweed invasion can be attributed to a single, ill-advised introduction.

114 M. L. HOLLINGSWORTH AND J. P. BAILEY

FALLOPIA SACHALINENSIS

To provide a comparison with the data from F. japonica var. japonica, we carried out a more
limited study on the levels of genotypic diversity in F. sachalinensis. A total of 30 plants were
collected from twelve localities. Single samples were collected from most populations, but 17
plants were collected from the River Kelvin, Glasgow (Lanark, v.c. 77), and three samples were
collected from the River Clywedog, Caerynwch (Merioneth, v.c. 48). All samples were analysed
for genetic variation using 10 RAPD primers. In contrast to the data from F. japonica var.
Japonica, high levels of clonal diversity were detected. A total of 14 genotypes were recovered,
and no genotype was shared between sites (Hollingsworth 1998). Two genotypes were detected
from the 17 plants collected from the River Kelvin, with one of these genotypes restricted to a
single plant (Hollingsworth et al. 1998). Two genotypes were detected from the three plants
sampled along the River Clywedog. In the absence of additional well-sampled populations such as
the River Kelvin it is difficult to say much on the partitioning of genetic variability in this species
in Britain. However, in contrast to the simple situation of a single widespread clone of F. japonica
var. japonica, F. sachalinensis appears to be genetically variable in Britain. Bailey & Conolly
(2000) suggest that multiple introductions of F. sachalinensis to Britain may be the source of the
greater levels of genetic variability of F. sachalinensis compared to F. japonica var. japonica.
However, while M.L.H. agrees that this may be an important factor, she believes that there are
presently insufficient data to eliminate sexual reproduction as a source of at least some of the
variability. Fallopia sachalinensis has plants producing both functional ovules and pollen, and sets
viable seed. The potential for sexual reproduction in this taxon in Britain is clear, and the
correlation of this with genetic variability may well be meaningful. Progeny segregating from only
a single heterozygous individual can lead to a multitude of genetic combinations.

LEVELS OF GENETIC VARIABILITY IN 6x F. x BOHEMICA

Genetic diversity can occur among hybrid individuals due to multiple origins (defined here as
multiple gamete fusions) or hybrid fertility. Although meiosis is irregular in 6x F. x bohemica
(J. P. Bailey, unpublished) some seed set does occur, and the possibility of F2 (and beyond) hybrid
plants exists. To investigate the amounts and partitioning of genetic variability in 6x F. x
bohemica we have investigated two populations in detail, the River Kelvin, Glasgow (23 plants),
and the River Wnion system, Dolgellau (Merioneth, v.c. 48) (36 plants). In addition we also
collected a total of 16 samples from 14 other sites across Britain.

The River Kelvin site in Glasgow is of interest as, despite the fact that both F. japonica var.
Japonica and F. sachalinensis grow close to the hybrid, only female plants of the parental taxa are
present. In contrast, F. x bohemica occurs as both hermaphrodite and female plants along the river
(Hart et al. 1997). Their co-occurrence may be due to independent colonisation, rather than being
indicative of in situ hybridisation. This situation provides an opportunity to examine levels of
sexual reproduction within the hybrid population, with reduced probabilities of multiple origins as
a confounding variable.

Using arbitrary fingerprinting techniques (RAPDs and Inter-SSRs (inter simple sequence
repeats)), five genotypes were detected from 23 F. x bohemica plants (Hollingsworth et al. 1998).
Of the four female plants sampled, all had the same genotype. The 18 hermaphrodite plants
sampled had one of three different genotypes, with a final genotype restricted to a plant which did
not flower (and thus could not be sexed). Given that we detected five different genotypes, and
there are no male fertile individuals of the parental taxa at this site, it is possible that some of this
genetic variability is due to hybrid fertility, rather than recurrent origins of the hybrid
(Hollingsworth et al. 1998). Clearly, one cannot rule out the possibility that male fertile
individuals of the parental taxa previously occurred in the area (although a thorough search of
surrounding localities and checks of herbarium specimens found no evidence that this is the case
(Hart et al. 1997)), or that the different hybrid genotypes are independent colonists of the river.
However, given that the highest levels of genetic variation were detected in the only male fertile
taxon present along the river (see the above sections on F. japonica var. japonica and F.
sachalinensis), perhaps the most parsimonious explanation of the data is that at least some of the
genetic variation in F’. x bohemica is attributable to local sexual reproduction.

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HYBRIDISATION AND CLONAL DIVERSITY IN FALLOPIA

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116 M. L. HOLLINGSWORTH AND J. P. BAILEY

The second site we have examined in detail is a more complicated population along the River
Wnion system (Fig. 1). Fallopia japonica var. japonica is common at many sites along this river
system but there are only two known localities for F. x bohemica and one known locality for F.
sachalinensis. At Caerynwch Hall on the River Clywedog (Fig. 1) female F. japonica var.
japonica, hermaphrodite F. sachalinensis and both female and hermaphrodite F. x bohemica grow
together along the edge of the river. The production of Fallopia seedlings has been recorded at this
site (Bailey et al. 1996). The second area where F. x bohemica is found is 5 km downstream at
Dolgellau (Fig. 1). Both female F. japonica var. japonica and female and hermaphrodite hybrids
occur here, growing on the banks of the River Wnion and also in the surrounding field and
recreational area; F. sachalinensis is not present at this site. Two samples of F. japonica var.
Japonica, three samples of F. sachalinensis and 20 samples of F. x bohemica, including six
seedlings, were collected from Caerynwch Hall. Three samples of F. japonica var. japonica and
16 samples of F. x bohemica were collected from Dolgellau. All samples were examined for
genetic variation using ten RAPD primers.

The five samples of F. japonica var. japonica all gave the standard genotype for this taxon in
Britain, and are assumed to be ramets of the widespread genet. Two genotypes of F. sachalinensis
were detected from the three samples from this site (see above). A total of 14 genotypes were
detected from the 30 mature plants of F. x bohemica, with a further six genotypes detected from
the six seedlings (Fig. 2a & b). The genetic structure of F. x bohemica along the river system
suggests that both asexual and sexual reproduction have played a role in the history of this
population. At some locations the same genotype was found in 2—5 ramets, with these samples
being either adjacent to each other (i.e. genotype B5 Fig. 2b), or in two cases widely separated and
found at both Caerynwch and Dolgellau (i.e. genotype B2 & B3; Figs 2a & 2b). This suggests the
occurrence of both local (within site) and widespread (between site) dispersal via vegetative
propagation. All six seedlings we examined had unique genotypes; only a single RAPD primer
was required to discriminate between these samples, highlighting the sensitivity of the assay.

The presence of female F. japonica var. japonica with hermaphrodite F. sachalinensis at
Caerynwch Hall, coupled with the high number of genotypes of F. x bohemica, suggest multiple
hybridisation events as the source of this variability. However, other factors may be responsible.
The presence of a large number of F. x bohemica genotypes found only at Dolgellau, where F.
sachalinensis does not occur, implies the need for caution before multiple hybridisation events are
invoked as the causal explanation for the genetic variability of F. x bohemica in this river system.

The private genotypes of F. x bohemica at Dolgellau could be explained by water-borne
dispersal of seeds from Caerynwch Hall, or dispersal (vegetative or seed) from plants (unknown to
us) along parts of the river system not sampled in this study. However, if multiple independent
introductions to this site from outside of the river system (which seems unlikely given the large
number of genotypes detected) are excluded, there is one other explanation for this variability:
hybrid fertility.

Distinguishing between hybrid fertility and multiple origins as a source of genetic variability in
F. x bohemica is difficult. Some insights may be gained from examining the chromosome numbers
of the individuals in question. Although only a small sample, of the seven plants of F. x bohemica
from the River Wnion system from which cytological data are available, all are 2n = 66. Bailey
(1997) suggested that the irregular meiosis of F. x bohemica would tend to lead to subsequent
generations being aneuploids. Cytological analysis of seeds collected from isolated plants of F. x
bohemica have shown a diverse array of chromosome numbers (Bailey 1990; Bailey et al. 1996).
Thus one may suggest that if hybrid fertility was the source of the genetic variability in the F. x
bohemica population from the River Wnion system, then many of these plants would be expected
to be aneuploids. However, while the presence of aneuploids would presumably be good evidence
that the plants were the result of an irregular meiosis, it should be stressed that the converse is not
conclusive. Irregular meiosis could still give rise to 2n = 66 individuals, and this number is
presumably at least as likely as any other possible chromosome number from 2n = 66 plants. As
we have no information on the survival characteristics of aneuploid versus euploid F. x bohemica
it is premature to place too strong an interpretation on the sole recovery of 2n = 66 plants. It is
possible that there is a disproportionate survival of euploid plants over aneuploids (but see J.P.B.
opinion below).

From our limited survey of F. x bohemica from more widespread British localities (16 plants
from 14 sites), no genotypes were shared between localities (Hollingsworth 1998). The only
shared genotype was from Swansea, where, from our sample of three plants, two, growing 20 m
apart, gave identical RAPD profiles.

HYBRIDISATION AND CLONAL DIVERSITY IN FALLOPIA 117

B2&B11

B 15, B 16, B 17,
B 18,B 19 & B 20

River
Clywedog

0.1km

Caerynwch

FIGURE 2a. Distribution of Fallopia genotypes along the River Clywedog, Caerynwch.
J Fallopia japonica var. japonica; S n Fallopia sachalinensis, n refers to genotype code; B n Fallopia x
bohemica, n refers to genotype code

(
= a 7 Recreation ground ! N

Dolgellau
0.1km g :

—

FIGURE 2b. Distribution of Fallopia genotypes along the River Wnion, Dolgellau.
J Fallopia japonica var. japonica; B n Fallopia x bohemica, n refers to genotype code

118 M. L. HOLLINGSWORTH AND J. P. BAILEY

Although further work is required (with more intensive within-site sampling), 6x F. x bohemica
is a genetically variable taxon, and we conclude that sexual reproduction appears (in one way or
another) to have been an important determinant of population structure. There remains, however,
some question as to precisely how this has occurred, i.e. what is the relative importance of hybrid
fertility versus multiple origins as the source of the genetic variability? This is one area where we
as authors disagree. M.L.H. argues that the “jury is still out” and that the current data set
(Hollingsworth, 1998; Hollingsworth et al. 1998) is inadequate to distinguish between the two
phenomena. She does not believe the possibility of hybrid fertility giving rise to new genotypes
can yet be excluded. In contrast, J.P.B. believes that the lack of aneuploids among those plants
whose chromosome numbers have been counted is sufficient evidence against hybrid fertility
contributing to the observed genotypic diversity. Unlike many biosystematic problems, this issue
may be easily tractable. Fallopia japonica var. japonica appears to be represented in the British
Isles by a single clone. Our RAPD and inter-SSR data have shown that there are numerous
markers that distinguish this clone from our samples of F. sachalinensis. To distinguish between
hybrid fertility and multiple origins as determinants of the genetic variability in F. x bohemica the
following simple experiments need to be carried out. Controlled crosses should be made between
F. japonica var. japonica and F. sachalinensis. The parents and large numbers of progeny should
be examined for their RAPD and inter-SSR profiles (polyploid segregation tests require extensive
progeny arrays). Although RAPDs and inter-SSRs are dominant markers and it is not possible to
distinguish between homozygotes and heterozygotes for any given gene in individual samples,
examination of the progeny of controlled crosses will allow homozygotes to be identified. Bands
that are homozygous in F. japonica var. japonica should be present in all of its Fl hybrids. In
contrast, those bands that are heterozygous in F. japonica var. japonica and segregate, will be
present in some, but not all progeny. Following this line of reasoning, any wild-collected 6x F. x
bohemica which lacks bands that are homozygous in F. japonica var. japonica (present in all of its
controlled Fl progeny) may be considered to be of more complicated origin than Fl hybrids.

As a final point, however, it is important to note that there is one other potential confounding
variable which is not so easily tested for, and this is whether the variability was generated in situ at
all. It is not known to what extent material of F. x bohemica was originally distributed as seed for
horticultural purposes (see Bailey & Conolly 2000). Although we feel that the evidence presented
here for in situ sexual reproduction within Britain is virtually conclusive (as there are so many
different genotypes present), it remains possible that some of the variability in this taxon in Britain
could stem from horticulturally raised and distributed seed.

JAPANESE KNOTWEED IN ASIA

Although the results presented here provide insights into the reproductive biology and population
structure of these invasive Fallopia taxa in Britain, there is a conspicuous absence of reference to
native material. A detailed knowledge of the genotypic and genetic variability of these plants in
their native range would be desirable for comparative purposes and may also serve to identify the
geographic source of the introduced material. This is of value as F. japonica var. japonica is
represented by only a single genotype in Britain and it may be particularly susceptible to an
appropriate biological control. Identification of the source population may be a useful step in the
search for such controls. However, not only is there a need for molecular work on the native
material, there is also a need for more fundamental alpha taxonomic research. Whilst the
morphological and cytological variation in British Fallopia taxa has received considerable
attention (Bailey 1994; Bailey et al. 1996; Bailey et al. 1995; Bailey & Conolly, 1985; Bailey &
Stace, 1992), the variability in the native range has only begun to be appreciated. Plants of
Fallopia japonica collected from Japan and China show a high degree of intraspecific variability,
in both cytological and morphological characters (Bailey 1999). Material of F. japonica var.
japonica found in the British Isles today, although morphologically more similar to Japanese than
Chinese material, is not typical of either (Bailey 1999). Japanese F. japonica are predominantly
tetraploids (2n = 44), whilst the three Chinese and one Korean plant that have been counted so far
are octoploid or decaploid (2n = 88 and 2n = 110). With this in mind, in the following section we
refer to native material as F. japonica var. ‘japonica’ to distinguish it from British F. japonica var.
Japonica.

HYBRIDISATION AND CLONAL DIVERSITY IN FALLOPIA 119

F. japonica var. compacta

British Isles
F. japonica var. ‘japonica’
Peking
F. japonica var. japonica F. japonica var. ‘japonica’
Europe & USA China

F. japonica var. ‘japonica’

Tanagawa River, Tokyo,

Japan F. japonica var. ‘japonica’
Mt. Fujija,Japan

F. japonica var. ‘japonica’
Honshu, Japan
F. japonica var. ‘japonica’

Shinjuka, Tokyo, Japan : } - ee
F. japonica var. ‘japonica
Japan

F. japonica var. ‘japonica’
Mt. Tateyama Japan

FIGURE 3. Neighbour Joining tree of Fallopia japonica s.1.

We carried out RAPD analysis using 10 primers on 14 native samples of F. japonica var.
‘japonica’ and compared them to the British genotype of F. japonica var. japonica and three
British plants of F. japonica var. compacta (Table 1) (Hollingsworth 1998). The interrelationships
of the samples are shown in Fig. 3. Although most of the genetic distance between samples in this
data set is due to differences among individuals rather than groups of individuals (i.e. most of the
genetic distance in the figure is on the terminal, rather than internal branches) a few points stand
out. The three F. japonica var. compacta samples cluster together, and they together form a group
with the introduced genotype of F. japonica var. japonica. The two Chinese samples cluster
together, and there is a cluster containing the Japanese samples (although separated by only a very
short branch from the other samples). A key point is that there is considerable diversity among the
native samples, with much greater genetic distance between Japanese and Chinese samples of F.
Japonica var. ‘japonica’ than there is between the two different named varieties that occur in
Britain

This point is further emphasised by our studies on the chloroplast genome of these taxa. Using
either sequence data from the chloroplast trnL intron or PCR RFLPs of the trnK intron, we are
able to distinguish the chloroplast genomes of British material of F. japonica var. japonica, F.
japonica var. compacta and F. sachalinensis (Hollingsworth et al. 1999). When we compared the
trnL_ sequences of British F. japonica var. japonica and F. japonica var. compacta to five
accessions of native F. japonica var. ‘japonica’ we did not detect either of the British chloroplast
types among these native plants (Hollingsworth 1998). Instead, from the five native samples we
detected five additional chloroplast types (all different), although relationships among these were
poorly resolved in cladistic analyses, as most differences were autapomorphic (unique to
individual samples). However, although there is little taxonomic information in these sequence
data, the results corroborate those of morphology, cytology and RAPDs, i.e. that there is
considerable diversity among native plants named as F. japonica var. japonica, and this exceeds
the diversity between the British introduced varieties F. japonica var. japonica and var. compacta.
Furthermore, no material which precisely matches the introduced and highly invasive F. japonica
var. japonica is known to us from the native range.

120 M. L. HOLLINGSWORTH AND J. P. BAILEY

TABLE 1 COLLECTION DETAILS OF FALLOPIA TAXA

Taxon Locality Chromosome No.
F. japonica var. ‘japonica’ Peking Botanic Gardens, China 2n = 88
F. japonica var. ‘japonica’ China (no further details known) 2n = 116
F. japonica var. ‘japonica’ Honshu, Japan 2n = 44
F. japonica var. ‘japonica’ Japan (no further details known)

F. japonica var. ‘japonica’ Mt. Fujiya, Japan 2n = 44
F. japonica var. ‘japonica’ Mt. Tateyama, Midagahara, Japan 2n = 44
F. japonica var. ‘japonica’ Mt. Tateyama, Midagahara, Japan

F. japonica var. ‘japonica’ Mt. Tateyama, Midagahara, Japan

F. japonica var. ‘japonica’ Shinjuku, Tokyo, Japan

F.. japonica var. ‘japonica’ Shinjuku, Tokyo, Japan

F. japonica var. ‘japonica’ Tanagawa River, Tokyo, Japan

F. japonica var. ‘japonica’ Tanagawa River, Tokyo, Japan 2n = 44
F. japonica var. japonica Leicester (v.c. 55), England 2 =56
F. japonica var. compacta Brecon (v.c. 42), Wales 2n = 44
F. japonica var. compacta Bracken Hill, Surrey (v.c. 16), England 2n = 44
F. japonica vat. compacta Blarney Castle, Co. Cork (v.c. H4), Ireland 2n = 44

Thus there are different interpretations of what constitutes Japanese Knotweed (F. japonica var.
Japonica) in native (Asian) and introduced (Europe and North American) regions. To resolve and
clarify these differences, a detailed monograph of section Reynoutria (F. japonica s.l. and F.
sachalinensis s.1.) including morphological, cytological and molecular data is required.
Widespread collections should to be made from across the native range and the samples compared
using both phenotypic and genotypic data. Only when an understanding of material in both
introduced and native ranges is acquired will the most appropriate taxonomic designation for these
samples become apparent. One safe prediction, however, 1s that there will undoubtedly be a more
focused (narrow) consideration of what should be included under the name of F. japonica var.
Japonica.

ACKNOWLEDGMENTS

We are grateful to Pete Hollingsworth for his helpful comments on earlier drafts of this
manuscript. We are also grateful to Ann Conolly, Colin Ferris, Richard Gornall and Clive Stace
for many useful discussions regarding our Fallopia research, and to John Akeroyd and Daniel
Kelly for constructive review.

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(Accepted October 1999)

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Watsonia 23: 123-138 (2000) 123

Hybrid origin of the Oxford Ragwort, Senecio squalidus L.
R. J. ABBOTT, J. K. JAMES, J. A. IRWIN* and H. P. COMES’

Institute of Environmental & Evolutionary Biology, School of Biology, University of St Andrews,
St Andrews, Fife, KY16 9TH

ABSTRACT

A survey of morphometric and allozyme variation was conducted to test the hypothesis that Oxford ragwort,
Senecio squalidus L. (Asteraceae) is a diploid hybrid species which originated in cultivation from hybrid
material between S. aethnensis Jan. ex DC. and S. chrysanthemifolius Poiret that was introduced to the Oxford
Botanic Garden from Mt Etna, Sicily, in the 17th century. Morphometric analysis of material raised under the
same glasshouse conditions showed that S. squalidus closely resembled material derived from hybrid swarms
between S. aethnensis and S. chrysanthemifolius on Mt Etna, and produced a phenotype that was normally
intermediate in leaf shape and capitulum size to that of its two putative parent species. Allozyme analysis
further showed that populations of S. squalidus were very similar genetically to Sicilian material with some
populations clustering tightly with hybrid material and with S. chrysanthemifolius. Most notably, S. squalidus
and hybrid material were polymorphic and exhibited almost identical frequencies for two alleles (a and b) that
distinguish pure populations of the two parent species at the Acp-2 locus. Thus, in combination with previous
documentary evidence, the morphometric and allozyme results obtained from the present study provide strong
support for the hypothesis that S. squalidus is a new diploid hybrid species which originated allopatrically
through stabilization of hybrid material of S. aethnensis and S. chrysanthemifolius introduced to the British
Isles from the hybrid zone on Mt Etna, Sicily. Because allozyme diversity within S. sgualidus was found to be
much reduced to that present within Sicilian material, it is likely that only a small sample of hybrid material
(and allelic diversity) was introduced to the British Isles from Sicily.

KEYWORDS: hybrid speciation, morphometrics, allozymes, introduced plants, Asteraceae.

INTRODUCTION

In a series of articles published between 1955 and 1966, Douglas Kent chronicled the introduction,
escape from cultivation, and subsequent spread of the Oxford ragwort, Senecio squalidus L.
(Asteraceae), in the British Isles (Kent, 1955, 1956, 1957, 1960, 1963, 1964a,b,c, 1966). The
information contained in these reports is of considerable value for two reasons. First, there are very
few examples of an introduced invasive species for which such details of introduction and spread
have been rigorously documented. Second, a detailed knowledge of the spread of the Oxford
ragwort in the British Isles has been of crucial importance to the timing of two notable
evolutionary events that followed hybridization of the species with the native Groundsel, Senecio
vulgaris L., viz. the origin of the new allopolyploid species, S. cambrensis Rosser, independently
in Wales and Scotland, during the 20th century (Ashton & Abbott 1992a; Lowe & Abbott 1996),
and the origin of the radiate form of S. vulgaris (var. hibernicus Syme), a stabilised introgressant,
during the 19th century (Abbott 1992; Abbott et al. 1992a). Although Kent (1956) provided
information on the source of $. squalidus material introduced to the British Isles, no details were
available on how this material might have become modified during cultivation before it escaped
and became invasive. In this paper, we provide evidence that the material originally introduced to
the British Isles was hybrid in nature and that S. squalidus, as recognised in these islands, arose as
a derivative of this material most probably during the period of cultivation.

Present address:

*John Innes Centre, Norwich Research Park, Colney, Norwich NR4 7UH

TInstitut fiir Spezielle Botanik und Botanischer Garten, Johannes Gutenberg-Universitét Mainz,
D-55099 Mainz, Germany

124 R. J. ABBOTT, J. K. JAMES, J. A. IRWIN AND H. P. COMES

According to Kent (1956) and Druce (1927), S. squalidus (2n = 20) was collected from Mt Etna,
Sicily, and introduced to the Oxford Botanic Garden, England, soon after the Garden opened in
1621. The exact date of its introduction is unknown as early records from the Garden were lost in a
fire. Present records show that it has been in cultivation at Oxford since at least 1690 (Kent 1956).
Linnaeus described the species in 1753, from plants raised by him possibly from seed sent from
Oxford (Walker 1833), although evidence for the latter is disputed (Kent 1956). In 1792, the
species was reported growing on walls in the Oxford area (Druce 1927) and was described as a
garden escape. The subsequent spread of the species away from Oxford was aided initially by
exchange of seeds between botanic gardens but, in the late 19th century, following the
establishment of the railway system in the British Isles, S$. squalidus began spreading rapidly from
Oxford via the railway network. More recently, the species has spread along motorway verges in
Britain, and is now ubiquitous on wasteground especially in urban areas. Records show that the
species began to spread northwards from Oxford in the late 19th century and reached different
parts of the north of England between the early to mid-part of the 20th century (Kent 1964b.c).
The species became established in the wild in the Central Belt of Scotland in the mid-1950s (Kent
1955, 1966), and continues to spread northwards into Fife and north of the River Tay at the present
time (Abbott 1992, and pers. obs.).

The source and nature of S. sgualidus material now established in the British Isles has been
questioned in recent times from two standpoints. First, Walters (1963) emphasized the
morphological similarity between S. squalidus in the British Isles and S. rupestris Waldst. and Kit.
(2n = 20), a closely related species that occurs on mountains in central and southern Europe. This
raised the possibility that S$. rupestris was an alternative source of some, if not all, material of S.
squalidus established in Britain. Because of this morphological similarity, Alexander (1979) in a
taxonomic revision of Mediterranean Senecio, section Senecio, subsumed S. rupestris under S.
squalidus, although he considered British plants to be atypical of the taxon. Second, Crisp (1972),
in a biosystematic analysis of Senecio material growing on Mt Etna, was unable to recognise the
taxon S. squalidus found in Britain, and argued instead that British S. sqgualidus is a product of
hybridization between the diploid Sicilian species S$. aethnensis Jan. ex DC. (2n = 20) and S.
chrysanthemifolius Poiret (2n = 20), and that there is no evidence of material from elsewhere in
Europe having contributed to the British taxon. According to Crisp (1972), both Sicilian species
occur on Mt Etna with S. chrysanthemifolius present up to approximately 1000m altitude and S.
aethnensis from approximately 1600 to 2600 m altitude. Between these two species at 1300 m +/-
300 m, a series of hybrid swarms are present within which are found certain plants that bear a
close morphological resemblance to British S. sgualidus. Crisp (1972) advanced the view that
material introduced to the British Isles most likely was collected from these hybrid populations,
and following a century of cultivation in the Oxford Botanic Garden, stabilized derivatives of this
hybrid material escaped at the end of the 18th Century and have since spread rapidly to many parts
of the British Isles. He concluded that “British S. sqgualidus is of hybrid origin...” and “...can be
treated as a separate species as it is both geographically isolated from the parent species, and it has
evolved .... over many generations to a stage where it is morphologically distinct from either of
them, although still polymorphic and in general intermediate between them.” For the purposes of
this paper, and in keeping with the views of Crisp (1972), we treat as separate species S. squalidus
found in the British Isles and S. rupestris from central and southern Europe .

A survey of chloroplast (cp) DNA variation in S. sgualidus, S. aethnensis, S. chrysanthemifolius
and S. rupestris (Abbott et al. 1995) showed that all material of S. sqgualidus examined from the
British Isles (21 accessions from 20 populations) possessed the same cpDNA haplotype (haplotype
B) as found in material of S. chrysanthemifolius surveyed (11 accessions from seven populations
in Sicily). Haplotype B also occurred in three of five individuals of S. aethnensis examined, and
five of seven individuals surveyed from the hybrid swarms on Mt Etna. Other individuals from
these populations possessed haplotype A, which differed by a single restriction site mutation from
haplotype B. In contrast, only one accession surveyed of S. rupestris (from Abruzzi, central Italy)
possessed haplotype B, while other accessions of this species contained either haplotype A (from
the Italian Alps, Romania, N Bulgaria, and introductions to Germany), or haplotype C (from S$
Bulgaria and Greece). Haplotype C differed from haplotype A by a length mutation and three
restriction site mutations. The occurrence of haplotype B in all samples of S. squalidus from the
British Isles and in the two Sicilian species, S. aethnensis and S. chrysanthemifolius, and in their

HYBRID ORIGIN OF SENECIO SQUALIDUS Pes)

hybrid swarms, supported the hypothesis that S. squalidus is derived from hybrid material
introduced from Sicily. However, because haplotype B is also present in S. rupestris, albeit at low
frequency, it can not be ruled out that the British population of S. squalidus might be derived either
in part or entirely from introductions of S. rupestris.

Here we report the results of a comparison of morphometric and allozyme variation within
material of S$. squalidus from the British Isles with that within Senecio material sampled from Mt
Etna, Sicily, namely S. aethnensis, S. chrysanthemifolius, and hybrid populations between these
two species. These results provide further support for Crisp’s (1972) hypothesis of a hybrid origin
of S. squalidus in the British Isles.

MATERIALS AND METHODS

PLANT MATERIAL

Seed was collected separately from approximately 30 plants from each of the populations listed in
Table 1 and Fig. 1. This entailed sampling three populations of each of S. aethnensis, S.
chrysanthemifolius, and three of their hybrid swarms occurring on Mt Etna, Sicily, plus nine
populations of S. squalidus from the British Isles, and a single population of S. vernalis Waldst.
and Kit. from Germany. This material was analysed for allozyme variation, while subsamples were
used in a survey of morphometric variation.

MORPHOMETRIC ANALYSIS

Five plants (one offspring per mother plant) of each of S. aethnensis, S. chrysanthemifolius, and
also of hybrid material from Mt Etna (two plants from population 7, and three plants from
population 8), were raised with five plants from each of two populations of S. squalidus (from
Oxford and Edinburgh) (Table 1). The 25 plants were grown from seed to maturity as single
individuals in pots of 13 cm diameter containing compost. Pots were fully randomized within a
greenhouse. Illumination was provided by natural daylight supplemented by 400-W mercury
vapour lights to give a photoperiod of 16 h, while temperature was maintained at 20 °C +/- 2 °C.
On the day of full anthesis of the apical capitulum, each plant was harvested and measured for 32
characters. The character set recorded (Table 2) was modified from that used by Irwin and Abbott
(1992). Fifteen of the characters were descriptors of the capitulum and another 15 described leaf
size and shape. The remaining two characters were plant height and inflorescence length. Details
of measurement are given in James (1995). Data were subjected to univariate one-way ANOVA
and Duncan’s multiple range tests using the software package SPSS Base (version 7.5, SPSS Inc.

FIGURE |. (a) Locations of populations of Senecio squalidus ( @ ) sampled in the British Isles; (b) Locations
of populations of S. aethnensis ( & ), S. chrysanthemifolius ( ® ), and S. aethnensis x S. chrysanthemifolius
(™ ) sampled from Mt Etna, Sicily. Key to population numbers is given in Table 1.

126

R. J. ABBOTT, J. K. JAMES, J. A. IRWIN AND H. P. COMES

TABLE 1. GEOGRAPHIC LOCATIONS, SITE DESCRIPTIONS, COORDINATES, AND
ALTITUDE OF SAMPLED POPULATIONS OF SENECIO

Species No.

S. aethnensis 1

3%

S. chrysanthemifolius — 4*

S. aet. x S. chrys. Te

8*

S. squalidus

14

15
16
17

18

19

S. vernalis

Locality

Italy, Sicily, Mt Etna, 2 km E
Rifugio Sapienza, lava by
roadside

Italy, Sicily, Mt Etna, 3 km E
Rif. Sapienza, volcanic ash
Italy, Sicily, Mt Etna, Rif.
Citelli, volcanic ash

Italy, Sicily, Mt Etna, Bronte,
waste ground

Italy, Sicily, Mt Etna, 5 km W
of Nicolosi, roadside

Italy, Sicily, Mt Etna, 4 km
SE of Linguaglossa, roadside

Italy, Sicily, Mt Etna, 4 km
NNE of Ragalna Est, new lava
Italy, Sicily, Mt Etna, Paso
Cannelli, N of Pedara, iava
Italy, Sicily, Mt Etna, Villagio
Turistico Mareneve, roadside

British Isles, Scotland,
Kirkaldy, car park

British Isles, England, Oxford,
waste ground, Osney street
British Isles, Scotland,
Edinburgh, Leith, Mill Lane,
waste ground

British Isles, Ireland, Co.
Cork, S.of Kilbrittain

British Isles, England,
Norwich, waste ground at
railway station

British Isles, England, Bristol,
wasteground, Coronation Rd
British Isles, England,
Weymouth, waste ground
British Isles, Wales, Cardiff,
waste ground, Colington Rd
British Isles, Scotland,
Kirriemuir, waste ground

Germany, Schiisselacker
Weide, Eppelheim (nr.
Heidelberg)

*Populations included in morphometric analysis.

Collectors: RJA (R. J. Abbott), PAA (P. A. Ashton), JAI (J. A.

Lat. Long. Alt. (m)
37-42N 15:02E 1890
3741 N 15-03} Ee 1650
37-47N 15-04E 1750
37-45N 14:50E 760
37-:35N 14:59E 800
37-50 N’ 15:072E 750
37-37 N - 14:59.E) 19s
3740N 15-04E 1500
37-49N 15:04E 1400
56:08 N 03-10 W <50
51-45 N 01-09 W <50
55:58 N 03-10 W <50
51:37 N 08-35 W <50
52:-40N 01-20E <50
51:30 N 02:37 W <50
50:37 N 02-29 W <50
51-30N 03-10 W <50
56-43N 03:00W <I150
49-24N 08-38 E c.50

Irwin), JWK (J. W. Kadereit).

Collector (date)

RJA (13.06.88)

RJA (13.06.88)

RJA (16.06.88)

RJA (11.06.88)
RJA (12.06.88)

RJA (16.06.88)

RJA (12.06.88)
RJA (13.06.88)

RJA (16.06.88)

RJA (10.09.91)
RJA (22.05.91)

RJA & JAI
(01.06.90)

RJA (03.10.91)

PAA (08.07.90)

JAI (06.09.90)
JAI (03.09.90)
JAI (07.09.90)

JAI (07.07.91)

JWK (21.05.88)

HYBRID ORIGIN OF SENECIO SQUALIDUS 127

TABLE 2. CHARACTER MEANS OF SENECIO POPULATIONS

Populations
S. det. Schr. SEX AG S. squalidus P
3 4 oi) 8 11 (2

Height mm 606° 703° 585” 695° 593” 4833 #
Inflorescence length mm 28-8" IMS fo 21-6? 21-3" 31-3° DAA
Capitulum length mm Tiles? 9.98 10-4 10-5* 10-8 10-7” *
Pedicel length mm 173% 11-8” ited 17:3 20-5° 13-92" -*
No. of pedicel bracts 3-0° 3-4" 2-0? 5:32 3-2° 24% FR
Capitulum apical width mm 5-6° 4.23 4.9° 55" S-7" 4.9° ‘i
Capitulum basal width mm 5-8 4-7 5:8 5-3 5-9 5-7 ns
No. of phyllary bracts 21-8° 2ie?* 21-0? 21-0? 22-8° 21-6° a
No. of calyculus bracts 7-6 5-08 9-0° 6-7* 8-2° 9-0° ex
Calyculus bract length mm Sey 273° 2-93 30 3-0 2-83 5:
No. of florets per capitulum L16” 978 106° 119% 136° 106° ae
Ray floret length mm 132° 9.5" 9.4" 10-3*° 13-0° 2 a
Ray floret width mm Bk: 2-63 Bane 3-6° Bo7 3-6° cies
No. of disc florets LOS2% 84" O32 106° 123% O3” ear
Disc floret length mm 8-0° 6-8" 6-98 7-3” rl” 7-08 is
Disc floret corolla length mm 2-8° 2-48 a 2-8° Fe te 2-63 oon
Disc floret corolla width mm 0-99 0-9 0-89 0-94 0-92 0-87 ns
Longest leaf length mm 1334 128? [38° 144 IW 152” *
Midleaf length mm 107 108 116 124 125 126 ns
Midleaf lobe number 3-48 12-4° 13-0° 11-0° 16" 12-4° are
Midleaf maximum left width mm 18-7? 42-1° 42-7° 37:52 38-7" 41-7° ae
Midleaf maximum right width mm 19.48 46-0° 48-0° eee 45-1° 42-9» ee
Midleaf basal length (left) mm 61-7 51-9 46-5 72-0 64-9 72-6 ns
Midleaf basal length (right) mm 65-9° 44.8" 82-0° 64-0° TAeg? 71-6° i
Midleaf apical angle A‘ (deg) 109-0 62-78 69-0" 98-3 97.4% 83-6* nee
Midleaf apical angle B™ (deg) 41-48 58-6” FAS. 96:0" 104-2" 95-0% oe
Midleaf secondary vein angle? (deg) 20-48 66-64 61-0° 44.0 40-2° os ee i a
Square root of midleaf area mm 52 44 49 54 56 a ns
Midleaf perimeter mm 328 LOG2°* 1240° 797° 1186° 1240° 676%
Square root midleaf area/length mm 0-49 0-40 0-42 0-44 0-44 0-40 ns
Midleaf perimeter/length mm Sls 9-8° 10-7° 6-4° 9-4° 9-9° nee
Midleaf dissection™ 6-2? 24-1° 25-4° 14-6° Z lel? 24-8° iia

P = level of significant difference: * < 0-05; ** <0-01; *** <0-001.

Means followed by the same superscript letter are not significantly different at P = 0-05.

+ angle between the apex of the primary vein and the apices of the adjacent marginal tooth sinuses.
++ angle between the apex of the primary vein and the apices of the adjacent secondary veins.

+ angle between the midlobe secondary vein and the primary vein.

££ perimeter of the midleaf divided by the square root of its area.

1997) to detect differences between population means. Variables that were not normally
distributed were transformed into natural logarithms before analysis. Data were then standardized
by subtracting the character mean of the total sample from each individual value and then dividing
by the character standard deviation before a principal component analysis (PCA) was conducted
using NTSYSpc software (version 2.0, Rohlf 1998) on the derived trait correlation matrix.

ALLOZYME ANALYSIS

Allozyme variation was assayed for six enzymes by means of starch gel electrophoresis procedures
described previously (Ashton 1990; Abbott et al. 1992a; Glover & Abbott 1995). The enzymes
assayed were acid phosphatase (Acp), aspartate aminotransferase (Aat), B—esterase (B-Est),
glyceraldehyde-3-phosphate dehydrogenase (G3pd), phosphoglucoisomerase (Pgi), and
phosphoglucomutase (Pgm). Buffer systems and staining procedures for these enzymes were as
described by Comes & Abbott (1998).

128 R. J. ABBOTT, J. K. JAMES, J. A. IRWIN AND H. P. COMES

The genetics of electrophoretic variants of Aat, B—Est, and Pgi were deduced from previous
studies of their inheritance in S. squalidus and S. vulgaris L. (Abbott et al. 1992a,b; Ashton &
Abbott 1992b). The genetic interpretation of the other enzymes (Acp, G3pd, and Pgm) were based
on the generally conserved enzyme sub-structure, subcellular location, and isozyme number in
higher plants (Weeden & Wendel 1989). Three to four standards were run within each gel.
Isozymes and allozymes were designated numerically and alphabetically, respectively, according
to decreasing mobility rate.

Allozyme variation was estimated by computing the mean number of alleles per locus (A)
(including monomorphic loci), mean percentage of polymorphic loci (P), and mean unbiased
expected heterozygosities (H,) using the BIOSYS-1 software (vers. 1.7; Swofford and Selander
1989). Hardy-Weinberg equilibrium was tested across loci in all populations using the GENEPOP
software (vers. 3; Raymond & Rousset 1995) probability test option (Fisher’s method). Estimates
of genetic distance between populations were calculated (after Nei 1972) and subjected to an
unweighted pair group analysis (UPGMA) using BIOSYS-1.

F-statistics (Wright 1951) were estimated as outlined by Weir (1990) using the computer
program FSTAT (vers 1.2, Goudet 1995). Mean values of F, @ and f [Weir & Cockerham’s (1984)
estimators of Wright’s (1978) parameters of total inbreeding (Fit), subdivision among populations
(Fst), and inbreeding within populations (Fis)] were computed separately for S$. squalidus, and
their standard deviations were calculated by jackknifing over loci. An association between genetic
and geographical distance between populations of S. squalidus was examined in terms of the
correlation between @ and distance (km). The significance of the product-moment correlation was
evaluated by Mantel’s test (Mantel 1967) with 10,000 random permutations using GENEPOP.

RESULTS

MORPHOMETRIC ANALYSIS

Univariate analysis of variance detected significant differences between population means for all
but six of the 32 traits examined (Table 2). Duncan’s multiple range tests indicated that S.
aethnensis and S. chrysanthemifolius differed for 17 measured traits. S. aethnensis produced large
capitula and entire leaves, whereas S. chrysanthemifolius bore relatively small capitula and highly
dissected leaves. Comparisons between S. daethnensis and S. squalidus showed that the main
differences were in leaf shape. Senecio squalidus plants produced more dissected leaves than S.
aethnensis, but tended to have capitula of similar size. Differences between the two S. squalidus
populations and S. chrysanthemifolius were less straightforward. Whereas S. squalidus from
Oxford (population 11) produced larger capitula and leaves of a different shape to those of S.
chrysanthemifolius, S$. squalidus plants from Edinburgh (population 12) were more similar in
morphology to S. chrysanthemifolius. Interestingly, the means of the two S. squalidus samples did
not differ significantly from the means of one or other of the two hybrid samples examined
(populations 7 and 8) for nearly all traits examined, indicating a close resembance between S.
squalidus and hybrid material. However, the two hybrid samples did differ in mean for several
traits and it was evident that whereas the sample from population 7 tended towards S.
chrysanthemifolius in morphology, the other hybrid sample (population 8) was more similar in
morphology to S. aethnensis.

The first three components derived from principal component analysis (PCA) extracted 33-61,
19-93 and 7-45% of the total variance, respectively. A plot of principal component | (PC1) against
component 2 (PC2) separated plants in two-dimensions (Fig. 2). The two Sicilian taxa were widely
separated along the first axis with S. chrysanthemifolius plants exhibiting high scores and S.
aethnensis individuals having low scores. The majority of S. squalidus and hybrid plants (S.
aethnensis x S. chrysanthemifolius) exhibited intermediate scores along this axis, although one S.
squalidus plant (from Edinburgh) and a hybrid plant (from population 7) clustered with S.
chrysanthemifolius individuals. Examination of character coefficients (loadings) showed that PC1
was largely influenced by capitulum size and leaf width and leaf shape characters (total-sample
standardized coefficients >0-5; Table 3). Thus, $. chrysanthemifolius possessed small capitula and
highly dissected leaves in contrast to S. aethnensis which produced large capitula and almost entire
leaves (Fig. 2, Table 2). Senecio squalidus plants with capitula of intermediate size and less

HYBRID ORIGIN OF SENECIO SQUALIDUS 129

S. squalidus (11)
S. squalidus (12)
S. aethnensis

S. chrysanthemifolius

a + Pe @ O

S. aet. x S. chrys.

PC1

FIGURE 2. Plots for individuals of S. aethnensis, S. chrysanthemifolius, S. aethnensis x S. chrysanthemifolius
(populations 7 and 8), and S. squalidus (populations 11 and 12) on the first two principal components axes.

dissected leaves than S. chrysanthemifolius were also clearly separated from both Sicilian taxa
along PC2, but remained clustered with certain hybrid plants. PC2 was largely influenced by leaf
size and shape characters (Table 3).

ALLOZYME ANALYSIS

Three of the eight loci investigated (G3pd-/, Pgi-! and Pgm-1) were monomorphic in all 19
populations surveyed. The total number of alleles detected over the five polymorphic loci was 14
(range 2—3 per locus) (Appendix |). Of particular interest was the pattern of variation at the Acp-2
locus. All three populations of S. chrysanthemifolius were monomorphic for the Acp-2b allele
(Appendix 1). In contrast, all of the §$. aethnensis and hybrid (S. aethnensis x S.
chrysanthemifolius) populations investigated were polymorphic for the Acp-2a and Acp-2b alleles
as were five of the nine populations of S$. squalidus. The remaining four populations of S.
squalidus were monomorphic for the Acp-2b allele (Appendix 1). When data were pooled over
populations (Table 4) it was evident that the frequencies of the two Acp-2 alleles in S. squalidus
were very similar to those in the pooled hybrid material (S. aethnensis x S. chrysanthemifolius),
with the Acp-2b allele occurring at a higher frequency (approximately 0-8) than in S. aethnensis
material (approximatey 0-35).

At other polymorphic loci there were no major differences in allele frequencies between the two
Sicilian taxa, nor between them and their hybrid populations (Table 4). However, there was a
contrast between Sicilian material and S$. squalidus in that the latter was monomorphic at the
B-—Est-3 locus, and contained the Aat-3c allele at a much higher frequency.

UPGMA analysis of Nei’s genetic distances between populations based on allele frequencies at
all eight loci separated populations into two main clusters with S. vernalis positioned as a basal
group (Fig. 3). One cluster comprised seven populations of S. squalidus, while the other contained
the remaining populations. Within this second cluster, two subgroups were evident, one of which
contained three populations of S$. aethnensis and one of the hybrid populations (population 9),
while the other consisted of the three populations of S. chrysanthemifolius, two hybrid populations,
and two S. squalidus populations.

Levels of diversity varied across populations (Tables 5). Lower levels of diversity were present
in S. squalidus populations relative to populations of other taxa examined, based on measures of A
(mean number of alleles per locus), P (percentage of polymorphic loci) and H, (unbiased expected

130 R. J. ABBOTT, J. K. JAMES, J. A. IRWIN AND H. P. COMES

TABLE 3. TOTAL - SAMPLE STANDARDIZED COEFFICIENTS (LOADINGS) AND
EIGENVALUES FOR THE FIRST THREE PRINCIPAL COMPONENTS OF INDIVIDUAL
VALUES OF QUANTITATIVE TRAITS

Principal Components

1 Z 3
Height 0-131 0-241 0-172
Inflorescence length - 0-630 - 0-233 0-423
Capitulum length - 0-793 - 0-008 0-226
Pedicel length - 0-579 - 0-241 0-416
No. of pedicel bracts - 0-784 - 0-357 - 0-064
Capitulum apical width - 0-612 - 0-370 0-415
Capitulum basal width - 0-286 - 0-277 0-432
No. of phyllary bracts - 0-337 - 0-320 0-267
No. of calyculus bracts - 0-590 - 0-163 - 0-070
Calyculus bract length - 0-031 0-170 0-331
No. of florets per capitulum - 0-606 - 0-468 0-349
Ray floret length - 0-636 - 0-338 - 0-376
Ray floret width - 0-675 - 0-403 - 0-375
No. of disc florets - 0-629 - 0-448 0-344
Disc floret length - 0-693 0-290 0-207
Disc floret corolla length - 0-649 0-272 - 0-184
Disc floret corolla width - 0-317 0-202 - 0-410
Longest leaf length - 0-296 - 0-743 - 0-057
Midleaf length - 0-143 - 0-788 - 0-141
Midleaf lobe number 0-709 - 0-607 0-020
Midleaf maximum left width 0-663 - 0-643 0-018
Midleaf maximum right width 0-683 - 0-610 0-115
Midleaf basal length (left side) - 0-366 - 0-389 - 0-169
Midleaf basal length (right side) - 0-457 - 0-414 - 0-124
Midleaf apical angle A - 0-750 - 0-022 0-103
Midleaf apical angle B 0-020 - 0-778 0-104
Midleaf secondary vein angle 0-863 - 0-056 0-322
Square root of midleaf area - 0-494 - 0-575 - 0-398
Midleaf perimeter 0-582 - 0-759 - 0-226
Square root midleaf area/length - 0-557 0-075 - 0-397
Midleaf perimeter/length 0-727 - 0-573 - 0-177
Midleaf dissection 0-796 - 0-559 0-029
Eigenvalue 10-757 6-379 2-386
Variance (%) 33-616 19-935 7-455
Cumulative variance (%) 33-616 937501 61-006

heterozygosity). More diversity was present in S. aethnensis than in S. chrysanthemifolius and
their hybrid populations. Only in very few populations was the coefficient of inbreeding (f) high
and deviations from Hardy-Weinberg equilibrium significant. Thus in most populations random
mating was indicated.

The mean level of subdivision between S. squalidus populations (@ = F,,) over all loci (Table 6)
was significantly different from zero (as determined by bootstrap estimates of 95% CIs), indicating
significant population differentiation within this species in the British Isles. However, examination
of the UPGMA tree (Fig. 3) did not indicate any obvious geographical pattern to this structure, and
Mantel’s test of the correlation of @ and geographical distance between populations was not
significant (r = 0-106, P = 0-280).

Although the mean coefficient of inbreeding within S. squalidus populations (f = Fis) was not
significant, a high f value was calculated at the Acp-2 locus (Table 6). This might stem from the

HYBRID ORIGIN OF SENECIO SQUALIDUS 13:1

Ol. aet
09.axc S. aethnensis
Sicily
02. aet S. aethnensis x
S. chrysanthemifolius
03. aet
04. chr 4
OR axc
Sicily
S. chrysanthemifolius
05. chr
S. aethnensis x
06. chr S. chrysanthemifolius
08.axc S. squalidus
Sicily
nets British Isles
17. sq
11. sq
12. sq
14. sq
15. sq British Isles S. squalidus
10. sq
13 e2sq
16. sq
19. ver Germany S. vernalis
.30 .20 5/0) .0O

Genetic Distance

FIGURE 3. UPGMA tree constructed from genetic distances (after Nei, 1972) between Senecio populations.
Key to population numbers is given in Table 1.

132 R. J. ABBOTT, J. K. JAMES, J. A. IRWIN AND H. P. COMES

TABLE 4. ALLELE FREQUENCIES AT POLYMORPHIC LOCI IN SENECIO TAXA AFTER
POOLING DATA OVER POPULATIONS. (N = SAMPLE SIZE)

Taxon
Locus/allele S. aethnensis S. chrysanth. S. aet. x S. chrys. S. squalidus
Acp-2 N (62) (95) (76) (209)
a 0-645 ---- 0-204 0-170
b 0-355 1-000 0-796 0-830
Aat-3 N (62) (94) (76) (214)
b 0-911 0-925 0-960 0-327
c 0-089 0-075 0-040 0-673
B-Est-3 N (52) (87) (82) (215)
a 0-673 0-632 O-390 Pe ie re ===
b 0-327 0-368 0-409 1-000
Pgi-2 N (61) (88) (84) (211)
a OS 0
b 0-893 1-000 1-000 1-000
Pgm-2 N (59) (85) (81) (237)
Bn 0023 0 2 eee wee
b 0-881 0-906 0-920 0-996
C 0-119 0-071 0-080 0-004

occurrence of substructure within populations. However, previous studies (Abbott, unpublished)
have detected a null allele segregating at low frequency at the Acp-2 locus in some S. squalidus
populations. In the present study, null homozygotes were not recorded; however, it can not be
ruled out that a null allele was present at low frequency in some of these populations and affected
the correct identification of genotypes. The presence of a null allele for this dimeric enzyme would
lead to inflation of the number of homozygotes recorded due to masking of the null allele in
heterozygotes and, in turn, would result in inflation of the coefficient of inbreeding (/).

DISCUSSION

HYBRID ORIGIN OF SENECIO SQUALIDUS

The hypothesis that Senecio squalidus originated from hybrid material between S. aethnensis and
S. chrysanthemifolius introduced from Mt Etna, Sicily, to the British Isles in the 17th century
(Crisp 1972) is supported by the results of the surveys of morphometric and allozyme variation
reported here. Morphometric analysis showed that nine of ten S. squalidus plants raised under
glass bore a close morphological resemblance to hybrid plants derived from material sampled from
the hybrid zone on Mt Etna. Both types of plants produced phenotypes intermediate between S.
aethnensis and S. chrysanthemifolius. Thus, whereas S. aethnensis produced large capitula and
entire leaves, while S. chrysanthemofolius had smaller capitula and highly dissected leaves, S.
squalidus and the hybrid plants tended to produce capitula of intermediate size and leaves of an
intermediate degree of dissection. The only exceptions to this concerned one S. squalidus
individual and one hybrid plant which exhibited a phenotype similar to that of S.
chrysanthemifolius.

The survey of allozyme variation across populations confirmed a close similarity of certain
populations of S. sgualidus (populations 17 and 18) to S. aethnensis x S. chrysanthemifolius
hybrid material (populations 7 and 8), and also to S. chrysanthemifolius. These materials were
placed in a separate sub-cluster from S. aethnensis and one of the hybrid populations (9), in a
UPGMA tree constructed using Nei’s genetic distances between populations (Fig. 3). Other
populations of S. squalidus (populations 11-16) were positioned in a different cluster to Sicilian
material, although based on the genetic distances recorded they were closely related to this

HYBRID ORIGIN OF SENECIO SQUALIDUS 133

TABLE 5. ALLOZYME VARIABILITY ESTIMATES AND WEIR & COCKERHAM’S (1984)
COEFFICIENT OF INBREEDING (F = FIS) INSAMPLED POPULATIONS OF SENECIO.
TESTS OF HARDY-WEINBERG EQUILIBRIUM ACROSS LOCI WITHIN SAMPLES
(FISHER’S METHOD) ARE INDICATED.

Mean
Species Popn. sample size+ At P Hoy Baik)
S. aethnensis | 12-4 (1-2) 1-3 (0-2) 25:0 0-129 (0-084) -0-071
23-9 (0-1) 1-6 (0-2) 62:5 0-184 (0-068) 0-058
3 23-8 (0-6) 1-6 (0-2) 62:5. 0-208 (0-067) 0-209
Mean 1-5 50-0 = 0-174
S. chrysanthemifolius 4 41-0 (1-1) 1-5 (0-3) 37:5 0-107 (0-061) -0-071
>) 23°75 (1-1) 1-5 (0-3) 37:5 0-094 (0-065) 0-054
6 24-8 (0-2) 1-5 (0-3) 37:5 0-094 (0-062) 0-030
Mean 1-5 37:5 0-098
S. aet. x S. chrys. 7 37-8 (1-3) 1-5 (0-2) 50-0 0-076 (0-060) -0-319
8 27-6 (0-2) 1-5 (0-2) 50-0 0-138 (0-059) -0-159
9 15-6 (0-2) 1-5 (0-2) 50-0 0-138 (0-077) 0-365*
Mean 1-5 50:0 0-117
S. squalidus 10 28-4 (0°6) 1-3 (0-2) 25:0 0-022 (0-015) -0-031
11 24-8 (1-3) 1-3 (0-2) 25-0 0-091 (0-059) 0-045
12 26-4 (1-7) 1-1 (0-1) 12-5. 0-005 (0-005) 0-000
13 22-4 (2-1) 1-1 (0-1) 12-5. 0-049 (0-049) -0-139
14 25-4 (0-5) 1-3 (0-2) 25-0 0-113 (0-075) 0-467**
ifs) 25-5; Gl-5) 1-3 (0-2) 25-0 0-081 (0-060) 0-208
16 27-4 (0-3) 1-3 (0-2) 25-0 0-085 (0-063) 0-364*
17 18-8 (0-6) lel (0-1) 12-5 0-019 (0-019) -0-059
18 25-0 (0-7) 1-3 (0-2) 25-0 0-102 (0-069) 0-217
Mean 1-2 20:8 0-063
S. vernalis oy), 27-0 (0-0) 1-3 (0-2) 25-0 0-103 (0-068) 0-241

7 Standard errors in parentheses

++ Hardy-Weinberg equilibrium rejected at * P < 0-05 and **P < 0-01
A: mean no. of alleles per Jocus

P: percentage of polymorphic loci

H,.: mean expected heterozygosity

material. Strong evidence that S. squalidus is of hybrid origin, rather than a derivative of S.
chrysanthemifolius, was the presence of both Acp-2a and Acp-2b alleles in S. squalidus. Our
results showed that S. chrysanthemifolius is monomorphic for the Acp-2b allele and consequently
another taxon must have donated the Acp-2a allele to S. squalidus, which we propose is S.
aethnensis. It could be argued, of course, that S. squalidus might be a derivative of S$. aethnensis
rather than of hybrid origin, as all the allozyme alleles present in S. squalidus, including both
alleles at the Acp-2 locus, were also found in populations of S. aethnensis (Table 4). However,
there is good reason to believe that material of S$. aethnensis we sampled from Mt Etna was
introgressed. One of us (James 1999) has since found that populations of S. aethnensis that grow at
much higher altitudes on the mountain (>2,500 m) are monomorphic for the Acp-2a allele. We
propose, therefore, that S. sgualidus is derived from hybrids between S. aethnensis and S.
chrysanthemifolius rather than being derived directly from either parent.

A hybrid origin of S. squalidus assumes that the hybrid zone on Mt Etna formed following
secondary contact between S. aethnensis and S. chrysanthemifolius, and that material regarded as
hybrid on Mt Etna is not ancestral to S. aethnensis and S. chrysanthemifolius. If the latter were the
case, then S. squalidus would also be considered as directly derived from this ancestral taxon.

134 R. J. ABBOTT, J. K. JAMES, J. A. IRWIN AND H. P. COMES

TABLE 6. WEIR & COCKERHAWM’S (1984) ESTIMATES OF WRIGHT’S F-STATISTICS
CALCULATED SEPARATELY FOR EACH LOCUS FOR ALL BRITISH
POPULATIONS OF SENECIO SQUALIDUS

Locus F @ ei

Acp-2 0-659 (0-087) 0-239 (0-091) 0-546 (0-073)
Aat-3 0-443 (0-160) 0-426 (0-154) 0-028 (0-083)
Pgm-2 0-000 (0-000) 0-030 (0-014) -0-031 (0-015)
Mean (Std. Dev.) 0-478 (0-140) 0-385 (0-122) 0-219 (0-332)
95% confidence interval 0-000 - 0-655 0-016 - 0-420 -0-016 - 0-546

Means and standard deviations were obtained by jackknifing over loci. Confidence intervals were obtained by
bootstrapping over loci. Locus specific standard deviations were obtained by jackknifing over populations.
F = Fit (within total), @ = Fst (among populations), f = Fis (within populations)

Although this possibility can not be excluded, it is considered to be highly unlikely. In addition to
the fixation of different alleles at the Acp-2 locus in S. aethnensis and S. chrysanthemifolius, James
(1999) has found that populations of S. aethnensis and S. chrysanthemifolius from the extremes of
the altitudinal range on Mt Etna are monomorphic or almost monomorphic for many different
RAPD bands and also for cpDNA haplotype. Such nuclear and cytoplasmic differentiation
between these two species indicates that they are likely to have originated in isolation and recently
come into contact. Despite these molecular differences, $. aethnensis and S. chrysanthemifolius
produce fertile Fl and F2 hybrid populations, and are interfertile with S. squalidus (Abbott &
Forbes unpublished).

In another study (Abbott & Milne 1995) the Acp-2b allele, which occurs at high frequency in S.
squalidus and in hybrid material on Mt Etna, was shown to be absent from 16 populations of S.
rupestris sampled widely from the geographical distribution of this species. This shows that S.
squalidus could not be derived entirely from S$. rupestris material, although does not rule out the
possibility that it might be derived in part from this source. However, we propose that the
allozyme evidence combined with the absence of any known documentary evidence of an
introduction of S. rupestris to the British Isles, provide a strong argument against the possibility of
S. rupestris having contributed to the British taxon.

GENETIC STRUCTURE OF SENECIO SQUALIDUS
The level of allozyme diversity within S. sgualidus, measured in terms of expected heterozygosity,
was approximately one third of that recorded in S. aethnensis and approximately half that in the
hybrid material from which we propose S. sgualidus is derived (Table 5). This reduced level of
genetic diversity in S$. squalidus is probably due to a founder effect caused by the introduction to
the British Isles of a small sample of hybrid material (and allelic diversity) from the hybrid zone
on Mt Etna. Rieseberg (1997) has reviewed amounts of allozyme variation found in other
homoploid hybrid species relative to their parents, and shown them to differ between species. In
three Helianthus diploid hybrid species, levels of diversity were lower than expected, indicating a
small number of parental individuals involved in their origin. In contrast, in other homoploid
species, levels of diversity were approximately the same or slightly greater than those of parents.
Despite the low diversity in S. squalidus, significant population differentiation was found for
allozyme variation based on F statistics analysis (@ was significantly different from zero).
However, there was no obvious geographical pattern to this variation according to the UPGMA
tree constructed from genetic distances between populations (Fig. 3), and the non-significant
correlation of @ and geographical distance between populations. It is likely that population
differences in allozyme variation are the product of founder effects and genetic drift occurring
during the colonization of new sites, and that the recent massive range expansion of S. squalidus in
the British Isles has prevented a possible association of genetic and geographical distance from
arising between populations due to isolation by distance.

The mean coefficient of inbreeding (f) in S. squalidus was not significantly different from zero,
indicating that random mating is normal in the species - as noted by Abbott & Forbes (1993).

HYBRID ORIGIN OF SENECIO SQUALIDUS 135

However, a high f value was evident at the Acp-2 locus (across populations) and also in two
populations (estimated across loci). The high f value at the Acp-2 locus might stem from the
presence of a null allele at this locus which, if true, most likely occurs at low frequency, given that
null homozygotes were not recovered (see Results section). The high f values in two populations
could be due to population sub-structure (the Wahlund effect) which would promote biparental
inbreeding. Alternatively, some selfing might be occurring. Abbott & Forbes (1993) showed that
S. squalidus is strongly self-incompatible; however, the self-incompatibility reaction might be
weakened under certain conditions (Hodgkin et al. 1988) allowing selfing to take place.

Although S. squalidus was less variable than either of its parents on the basis of allozyme
variation, this finding does not tally with its morphological variation. In the field, it exhibits
considerable variation in leaf shape, with individuals at one extreme producing highly dissected
leaves similar to S. chrysanthemifolius, and plants at the other extreme that bear almost entire
leaves (var. subinteger, Druce 1927). However, most plants produce leaves intermediate between
these two extremes though tending towards S. chrysanthemifolius. Other characters known to vary
are capitulum size and seed type (Druce 1927). It would seem that morphologically the species
might be converging towards an intermediate phenotype, but is not yet completely stabilized.

ALLOPATRIC ORIGIN OF HOMOPLOID HYBRID SPECIES

The morphometric and allozyme evidence reported here, combined with previous documentary
evidence (Druce 1927; Kent 1956; Crisp 1972), provide support for the hypothesis that S.
squalidus is a new diploid hybrid species which originated allopatrically through stabilization of
hybrid material of S. aethnensis and S. chrysanthemifolius introduced to the British Isles from the
hybrid zone that occurs between the two species on Mt Etna, Sicily. Previous to this study, only
ten plant species had been indicated to be of homoploid hybrid origin based on rigorous molecular
analysis (Rieseberg 1997; Allan et al. 1997; Wolfe et al. 1998). Mechanisms of reproductive
isolation between homoploid hybrid species and their parents have been reviewed by Rieseberg
(1997) and Rieseberg & Carney (1998) and shown to involve postmating and/or premating
barriers. Two homoploid hybrid species, Pinus densata Masters and Encelia virginensis A. Nels,
are believed to be derived from parents that produce interspecific F, hybrids that are vigorous and
fertile (Wang & Szmidt 1994; Allan et al. 1997) and, in the case of E. virginensis, are known to be
interfertile with both parents. In these respects, therefore, they resemble S. squalidus, which is also
derived from parents which when intercrossed form vigorous fertile hybrid offspring, and whose
parents are interfertile with S. squalidus (Abbott & Forbes unpublished). Natural populations of P.
densata and E. virginensis are mainly allopatric with their parents and also ecologically isolated
from them. However, it has been argued that they are likely to have originated in sympatry with
gene flow between hybrid and parent species prevented by ecological isolation, and only later
became distributed allopatrically with their parents. Diploid hybrid species of Paeonia which are
also allopatric with their parents may have originated in the same way (Sang et al. 1995). In
contrast, S$. squalidus is an example of a diploid hybrid species which originated through
stabilization of hybrid material dispersed to locations geographically isolated from where
hybridization initially took place. Although the first step in this speciation process involved
hybridization between sympatric parents, the crucial process of stabilization resulting in the origin
of a new hybrid species occurred at an allopatric site where there was no possibility of hybrid
material backcrossing to either parent. During this process S. squalidus is likely to have also
become ecologically isolated from both parents, as neither parent species appears able to form
viable populations in the wild in the British Isles (Abbott, pers. obs.). Thus, the evolution of
premating barriers due, for example, to pollinator behaviour and/or ecological isolation (e.g. in
Penstemon, Wolfe et al. 1998; Pinus, Wang & Szmidt 1994; and Encelia, Allan et al. 1997), and
postmating barriers resulting from chromosomal or genic sterility factors (e.g. in Helianthus, Iris
and Stephanomeria, see Rieseberg (1997)), were not essential in the origin of S. squalidus, as
isolation was maintained by geographical separation.

Because of the possible ease with which homoploid hybrid speciation might occur at allopatric
sites, we may need to revise current views on the importance of this mode of plant speciation and
the frequency at which it occurs (Rieseberg 1997). That said, in the case of S. squalidus,
considerable assistance was required from man to aid the origin of this new species. Material from
Mt Etna was cultivated in the Oxford Botanic Garden for approximately 100 years before it
escaped to the surrounding area and started to spread beyond Oxford. It will be of interest in the

136 R. J. ABBOTT, J. K. JAMES, J. A. IRWIN AND H. P. COMES

future to determine what genetic changes might have occurred in this material, due to
recombination, segregation and mutation during this period, that could have led to the creation of a
plant type that is now well adapted to conditions in the British Isles.

ACKNOWLEDGMENTS

We thank David Forbes for assistance with the experimental work. Special thanks are due to
Professor Guiseppe Ronsisvalle (University of Catania) for his assistance and generous hospitality
during the collection of material in Sicily. The research was funded by grants to R.J.A. from the
NERC (GR3/7313) and The Russel Trust.

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(Accepted September 1999)

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Watsonia 23: 139-147 (2000) 139

A new hybrid between a European and a Chinese species of
Artemisia (Asteraceae)

C. M. JAMES

Department of Plant Breeding and Biotechnology, Horticultural Research International, East
Malling, Kent, ME19 6BJ

Bas. WURZEEL

47 Rostrevor Avenue, Tottenham, London, N15 6LA

and
C..A: STACE

Department of Biology, University of Leicester, Leicester, LE] 7RH

ABSTRACT

A new hybrid, Artemisia x wurzellii C. M. James & Stace hybr. nov., is described between A. vulgaris and
the Chinese A. verlotiorum. Fifteen localities are known in eastern London, and one in northern West Kent,
but there is no convincing evidence that the hybrid arose in situ at any of them. Characters distinguishing the
three taxa are summarized. Chromosome counts indicate that A. vulgaris is a hypodiploid and A. verlotiorum a
hypohexaploid. Artemisia x wurzellii has the expected chromosome number 2n = 34 (hypotetraploid).

KEYWORDS: Mugwort, Chinese Mugwort.

INTRODUCTION

The genus Artemisia L. (Asteraceae) is traditionally divided into three to six subgenera or sections,
one of which has more recently been recognised as the separate genus Seriphidium Fourr. (Bremer
& Humphries 1993). Various estimates put the number of species at 300-400, distributed in most
temperate parts of the world. In the former U.S.S.R. Poljakov (1961) recognised 116 species in
Artemisia sensu stricto, with a further 58 in Seriphidium. In China, Ling (1988) listed 186 species
in Artemisia sensu stricto. Tutin et al. (1976) recognized 54 native European species in Artemisia
sensu lato, of which only five reach the British Isles (four in Artemisia sensu stricto, one in
Seriphidium). In addition, 17 species of Artemisia and one of Seriphidium have been reported as
aliens in the British Isles (Clement & Foster 1994).

Hybrids are evidently rare in Artemisia, few having been reported. Of these, most are sporadic in
occurrence and several are dubious in identity. However, Kawano et al. (1995) reported triploid
hybrids (2n = 27) between A. capillaris Thunb. ex Murray (2n = 18) and A. japonica Thunb. ex
Murray (2n = 36) in Japan, and found evidence for the existence of backcrosses.

Of the two species relevant to the present paper, A. verlotiorum Lamotte is native to S.W. China
according to Brenan (1950), while A. vulgaris L. is almost circumboreal, being native to both the
British Isles and China. Ling (1988) placed the two species in adjacent series of subgenus
Artemisia section Artemisia. A. verlotiorum is treated as one of four species in series
Codonocephalae (Pamp.) Y. R. Ling, and A. vulgaris as one of 16 species in series Artemisianae
Y. R. Ling (but correctly to be known as series Artemisia). Diagnostic characters of the series were
not given. Ling (1988) also gave a much wider distribution for A. verlotiorum in southern and
south-eastern Asia.

140 C. M. JAMES, B. S. WURZELL AND C. A. STACE

Artemisia verlotiorum occurs as a naturalized alien in many parts of Europe as well as in
Australia and South America (Ryves 1984), presumably originating as an escape from herb-
gardens. Bini Maleci & Bagni Marchi (1983) reported on a morphological-anatomical comparison
of A. verlotiorum and A. vulgaris growing near Florence. The early occurrences in Britain were
listed by Brenan (1950), who also provided a detailed description and bibliography. The first
record was from Hounslow, Middlesex, made by C. G. Green in 1908 (OXF), and in the 1940s it
was recorded widely in metropolitan Surrey and Middlesex, many of the new sites being
discovered by D. H. Kent (Brenan 1950). By 1962 it was known as far north and west as W.
Norfolk and Dorset respectively (Perring & Walters 1962), and today it occurs north to Easterness
and Moray in northern Scotland (Vice-county Census Catalogue data in B.S.B.I. Database at
Leicester). It is still most common, however, in the Greater London area, where it occurs on all
kinds of rough ground, often with A. vulgaris very close by.

The putative hybrid between these two species was first found by B.S.W. in August 1987 on an
overgrown railway bank by Northumberland Park Station, Tottenham, Middlesex (Wurzell 1988).
Between then and 1995 B.S.W. discovered 13 additional localities for the putative hybrid in
eastern London, and one other has been reported there (see below). Although these localities occur
in three hectads (TQ/3.7, TQ/3.8 and TQ/3.9) and three vice-counties (Surrey, Middlesex, S.
Essex), and come within a few hundred metres of W. Kent, the total area of known distribution in
London covers only about 14 x 7 km. Elsewhere it has been recorded only in one outlying locality
in W. Kent (see below).

Living and herbarium material of the putative hybrid was sent by B.S.W. in 1987 to Leicester,
where it was studied by C.M.J. and C.A.S. in 1987 and 1988, and since then by C.A.S. This paper
presents the evidence for its putative parentage and describes the hybrid as A. x wurzellii.

MATERIALS

Living plants of the putative hybrid were obtained from the original locality at Northumberland
Park Station, Middlesex (TQ/34.91) and from near Walthamstow Marshes, S. Essex (TQ/35.87);
of A. verlotiorum from Tottenham, Middlesex (TQ/34.90), and Abersoch, Caerns. (SH/31.27); and
of A. vulgaris from Tottenham, Middlesex (TQ/34.90), near Bromsgrove, Worcs. (SO/98.68), and
central Leicester, Leics. (SK/59.04).

Herbarium material of the two species in LTR was also used for comparative purposes.

METHODS

Stem sections were cut on a freezing microtome at 25 um thick from a mid-region of the stem,
which was about 5 mm wide at that point.

Chromosome counts were obtained from root-tip mitoses using standard squash techniques. Pre-
treatment was with 0.002M 8-hydroxyquinoline at 4° C for 24h, fixation in ethanol : ethanoic acid
(3:1) for 2h, hydrolysis with 5M HCl for 6-10 min at room temperature, and staining with acetic-
orcein.

RESULTS

Brenan (1950) provided a full description of A. verlotiorum and a detailed table contrasting it with
A. vulgaris. All these differences are not repeated here, but the ones that were found to be most
useful in separating the two species and in recognizing intermediates (hybrids) are covered below
(see also Table 1).

VEGETATIVE MORPHOLOGY AND ANATOMY

The putative hybrids are generally intermediate between the parents in overall appearance,
although most of the characters of the leaves are difficult to quantify precisely. This is partly
because it is not easy to designate strictly comparable leaves on different plants, and partly
because of variation (mainly temporal and developmental, perhaps also environmental). We found
it essential to compare only the middle stem-leaves of mature flowering shoots. Such leaves, in

ARTEMISIA X WURZELLIT 14]

A B C

FIGURE |. Middle stem-leaves of mature flowering shoots of (A) Artemisia vulgaris, (B) A. x wurzellii and

(C) A. verlotiorum. The petiolar pectinations are delimited with *]

both parents and hybrid, conform to a basic pattern in which a single more or less trifid terminal
lobe decurrently connects with three pairs of variously incised lateral lobes which, in their turn,
lead downwards to a much smaller group of pectinate appendages (here called “petiolar
pectinations”) attached to that lowest section of the midrib that serves as the petiole (Fig. 1). The
leaves of A. verlotiorum have narrower primary lobes with fewer secondary lobes and serrations
than those of A. vulgaris, giving the impression that the former has longer leaf-lobes (although this
is often not so). There is, however, overlap, and the putative hybrids overlap with both parents as
well. Brenan (1950) found the same: “Segments of upper leaves usually comparatively short,
lanceolate to oblong or linear” (A. vulgaris); “Segments of upper leaves conspicuously elongate,
linear-lanceolate to linear” (A. verlotiorum) (our italics). Bini Maleci & Bagni Marchi (1983) also
expressed the differences in only comparative terms: “Foglie polimorfe; quelle tipiche pinnatofide
con lobi larghi, dentati ed auricole sviluppate” [leaves polymorphic, typically with wide, dentate
lobes] (A. vulgaris); “Foglie polimorfe; quelle tipiche pinnatofide con lobi stretti ed allungati, non
dentati ed auricole poco sviluppate” (A. verlotiorum) (our italics). We found the best
discriminators to be the length:breadth ratio of, and the number of serrations on, the main apical
segment of a middle stem-leaf of a mature flowering shoot (Table 1). Even so, there is some
overlap in ranges, and the putative hybrids are closer to A. vulgaris than to A. verlotiorum in the
leaf segment length:breadth ratio.

The petiolar pectinations (Fig. 1) also provide a useful field character for detecting the hybrid. In
A. verlotiorum they are narrow and few, but in each population of the hybrid so far examined they
are broader, shorter, more abruptly acute and more numerous (Table 1); thus they have served well
to distinguish these two rhizomatous taxa at a glance. The constancy of this feature in the hybrid
might well indicate that we are here dealing with a single clone. Sexually reproducing A. vulgaris,
in contrast, presents petiolar pectinations ranging between or even beyond those of the other two
taxa in shape and number.

Artemisia verlotiorum is more strongly aromatic than A. vulgaris, and Bini Maleci & Bagni
Marchi (1983) found that the former had 4—5 times as high a density of glands on the lower leaf-
surface (c. 20/mm? compared with 6-7/mm’). However, the very dense hair-covering present in
both species, and the difficulty of removing it, render this character of limited practical use in
identification.

142 C. M. JAMES, B.S. WURZELL AND C. A. STACE

According to the material we have examined, the nature of the stem pith (1.e. the tissue interior
to the ring of vascular bundles) differentiates the two species. In both, the pith region comprises a
central area of large, thin-walled, lignified cells (white in the fresh state) surrounded by a zone of
smaller, even thinner-walled, non-lignified, chloroplasted cells (green in the fresh state). In A.
vulgaris the white pith occupies a much greater proportion of the whole than in A. verlotiorum,
and in the putative hybrid the situation is almost exactly half-way between the two. We found this
to be the best single quantitative character separating the two species and the putative hybrid
(Table 1), but it should be remembered that we have not examined a large number of specimens.
Brenan (1950) also noted this character, but only in terms such as “relatively broad”. He also
claimed that the young stem of A. verlotiorum was more deeply furrowed than that of A. vulgaris,
but this was not confirmed by our observations.

The two species have an obviously different growth-habit, A. verlotiorum being extensively
rhizomatous and A. vulgaris clump-forming (with very short rhizomes). The putative hybrid more
closely resembles A. verlotiorum in this respect; both can, in favourable circumstances, become
quite invasive.

There are fairly reliable differences between the inflorescences of the two species. In A. vulgaris
the inflorescence is more branched, with longer, straighter branches, and leafy only at the base,
whereas in A. verlotiorum the branches are fewer and usually somewhat recurved, and the
inflorescence is conspicuously leafy almost to its apex. The putative hybrid is intermediate in these
characters, especially in inflorescence leafiness, although many more measurements would have to
be made before ranges could be allotted with confidence to the three taxa.

CAPITULUM CHARACTERS

The capitula of A. vulgaris are usually obovoid-campanulate (i.e. widest distally), whereas those of
A. verlotiorum are usually ovoid-ellipsoid (1.e. not wider distally than elsewhere), as pointed out
by Brenan (1950) but using a different terminology. The difference, however, is not marked and is

TABLE 1. CHARACTERS DIFFERENTIATING ARTEMISIA VULGARIS, A. VERLOTIORUM
AND THEIR HYBRID, A. x WURZELLII

A. vulgaris A. x wurzellii A. verlotiorum
Length:breadth ratio of apical 5-0-6-1 4-4—7-0 9-5-15
segment of mid-stem leaf
No. of serrations on apical segment 6-10 3-5 1-2

of mid stem-leaf

2—8 on each side,
lanceolate, acuminate,
5-6 x as long as wide

Petiolar pectinations (4)5—6(8) on each

side, ovate to

2—3 on each side,
lanceolate to

White pith as % diam. of total pith
Growth habit

No. of leaves in top 20 cm of infl.
Capitulum shape

Length & breadth of capitulum (mm)
Width of ‘bisexual’ florets (mm)
No. of stamens in ‘bisexual’ flowers
Receptacle shape

Flowering time

Seed-set in Britain

Chromosome no.

80-89
tufted
9-17
obovoid-campanulate
3-1-3-7 x 2-1-2:-5
0-25-0-3
2)
conical
July-September
yes
216
(? also 18)

obovate or narrowly
so, acute, 2-4(5) x as
long as wide
54-69
rhizomatous
18-24
ovoid-ellipsoid
1-8—2-4 x 1-6-1:8
0-2-0-3
0
+ spherical
October
no
2n = 34

narrowly so,
acuminate, 5—7 x as
long as wide

33-38
rhizomatous
25-32
ovoid-ellipsoid
3-5—4.2 x 2-4-3-0
0-4-0-51
5
low-domed
October-November
no
2mv= 505592
(? also 48, 54)

ARTEMISIA X WURZELLIIT 143

difficult to quantify. The putative hybrid is much closer to A. verlotiorum in this respect. Brenan’s
measurements for the two species broadly agree with the ranges seen by us, but the capitula of the
putative hybrid are smaller than in either species (mean 2-1 x 1-7 mm) (Table 1).

Differences in both the peripheral female and inner bisexual florets in the two species were also
recorded by Brenan (1950). There are usually some additional abortive bisexual florets right in the
centre of the capitulum. Brenan found (2) 5—22 and (0) 2-6 (11) bisexual florets for A. vulgaris
and A. verlotiorum respectively, but our figures (5—8 and 4-8) did not confirm these differences;
our range for the putative hybrid was 4—7. Our results do, however, confirm a difference in the
widths of the bisexual florets of the two species (Table 1); the putative hybrid has narrower florets
than in either parent.

Like Brenan (1950), we found no differences in the numbers of glands on the outside of the
corollas of the two species, although Bini Maleci & Bagni Marchi (1983) (and others earlier)
claimed that they were more abundant in A. verlotiorum (the more aromatic species).

We did find a slight difference in the shape of the receptacle in the two species. In A. vulgaris it
is elongated-conical whereas in A. verlotiorum it is low-domed; the putative hybrid is closer to the
latter but often higher-domed (sub-spherical).

PHENOLOGY AND FERTILITY

Artemisia vulgaris is a late-summer flowerer, reaching anthesis mainly from July to September in
southern England. A. verlotiorum flowers distinctly later, from October to November, and in many
years is damaged by frost before anthesis. The natural flowering periods of the two species do not
overlap, but plants of A. vulgaris that are cut down in summer will often produce late-flowering
stems, and, in the absence of frosts, undoubtedly both species could be in flower simultaneously.
The putative hybrid, when growing in the same general area as the two species, reaches flowering
at an intermediate time, mostly in October (two to four weeks before A. verlotiorum). Its flowering
stems are, however, very frost-susceptible and often do not reach anthesis in the wild.

Presumably due to its late flowering A. verlotiorum has not been found to produce ripe fruits in
this country. No fruits have been found on the putative hybrid. The florets of the putative hybrid in
the position of the bisexual florets of the parental species are much reduced, without the
conspicuously distally dilated corolla found in its two parents, and anthers are completely lacking;
only a reduced ovary, style and stigma in a very reduced corolla are present. Hence the putative
hybrids are probably totally male-sterile.

CHROMOSOMES

Clear chromosome counts (Fig. 2) were made from A. vulgaris from London, Birmingham and
Leicester (all 2n = 16), A. verlotiorum from London and Caernarvonshire (2n = 52 and 50
respectively), and the putative hybrid from both London sites (2n = 34) (Table 2). The numbers of
chromosomes possessing satellites in these three taxa were two, six and four respectively.

TABLE 2. CHROMOSOME NUMBERS OF ARTEMISIA TAXA IN THE PRESENT STUDY

Species

Locality Chromosome number
Artemisia vulgaris

Rough ground, Tottenham, London, Middlesex, v.c. 21 2n = 16

By Tardebigge Canal, east of Bromsgrove, Worcs., v.c. 37 2n = 16

On waste ground, central Leicester, Leics., v.c. 55 2n = 16

Artemisia verlotiorum
By public toilets, Abersoch, Lleyn, Caerns., v.c. 49 an = 50
Rough ground, Tottenham, London, Middlesex, v.c. 21 Zio

Artemisia x wurzellii
Railway bank, Tottenham, London, Middlesex, v.c. 21 (locality 14) 2n=34
Rough ground, near Walthamstow Marshes, London,
S. Essex, v.c. 18 (locality 11) 2 = 34

144 C. M. JAMES, B.S. WURZELL AND C. A. STACE

Cae
ee
‘ ee
wt
Sy
wy ©
e
i»
C ;

FIGURE 2. Chromosomes of: A. Artemisia vulgaris (2n = 16), B. A. x wurzellii (2n = 34) and C. A. verlotiorum
(2n = 52), from Tottenham, London, v.c. 21. Magnification x 1755.

DISCUSSION AND CONCLUSIONS

IDENTITY OF PUTATIVE HYBRIDS

Despite the paucity of quantitative data demonstrating the intermediacy of the putative hybrids,
there is strong circumstantial evidence that they are indeed hybrids between A. verlotiorum and A.
vulgaris (Table 1).

Their overall appearance is convincingly intermediate; this is particularly indicated by the width
and serration of the leaf-lobes, especially the apical leaf-segment of the middle stem-leaves, and
by the leafiness of the inflorescence. The pith anatomy and capitulum shapes are also intermediate
between those of the two putative parents, as is the flowering time in London. The putative hybrids
are totally male-sterile, no anthers being formed, a condition never observed in either parent.

The fact that other characters separating the two species are not intermediate in the putative
hybrid (e.g. rhizome growth, capitulum shape and size, petiolar pectinations) does not argue
against its hybrid origin, since this situation is common in nature. Hybrids are rarely exactly
intermediate in every diagnostic character, due to the varied genetic backgrounds of the observed
phenotypes.

Nevertheless, probably the most convincing evidence of hybridity comes from the chromosome
numbers. In our London material A. verlotiorum had 2n = 52, A. vulgaris had 2n = 16, and the
putative hybrids had the expected 2n = 34 (26 + 8).

In Asteraceae tribe Anthemideae, and in the genus Artemisia, the diploid number 2n = 18 is by
far the commonest reported chromosome count (Bolkhovskikh ef al. 1969, and later plant
chromosome number indices). Tetraploids with 2n = 36 and hexaploids with 2n = 54 are also well
distributed in Artemisia, but several aneuploid counts, such as 2n = 16, 14 and 34, exist in
addition. Hence our plants of A. vulgaris should probably be viewed as aneuploids, 2n = 2x — 2 =
16, where x (the base number) = 9. .

ARTEMISIA X WURZELLII 145

The majority of reported counts of A. vulgaris are 2n = 16 (widespread in Europe and in North
America, Russia, India and Japan), but there are a few 2n = 18 counts (India and Caucasus) as
well. The single previous British count, from Surrey, was 2n = 16 (Morton 1977). Most previous
counts of A. verlotiorum are hexaploids with 2n = 54 (from Italy, Germany, Japan and Russia), but
there are also two, presumably aneuploid, counts of 2n = 48 from China and Spain. Our counts of
2n = 50 and 52 for A. verlotiorum are new numbers for this species, and neatly complete the
reducing aneuploid series 54, 52, 50, 48.

It thus appears that both A. vulgaris and A. verlotiorum exist as both euploid and reducing
aneuploid cytotypes. On known evidence of the range of chromosome numbers of the two parental
species, hybrids could occur with chromosome counts of 2n = 32, 33, 34, 35 or 36. The number of
satellited chromosomes that we recorded in the three taxa confirm their suggested ploidy levels.

FORMAL DESCRIPTION OF HYBRID

The evidence presented above leads us to conclude that the intermediate plants are hybrids
between Artemisia vulgaris and A. verlotiorum. They are described below after their discoverer, B.
S.W.

Artemisia x wurzellii C. M. James & Stace, hybr. nov.

Hybrida inter A. vulgarem et A. verlotiorum intermedia sed ab ambabus floribus sterilibus
(anantheris) et capitulis minoribus (1-8—2-4 mm longis) differt. Caules medulla viridi et medulla
alba magis minusve aequicrassa. Folia caulina media segmentis apicalibus 4-4 — 7 x longioribus
quam latis.

HOLOTYPUS: Overgrown railway bank, Northumberland Park Station, Tottenham, Haringey,
London, Middlesex, map reference TQ/34.91. 2 October 1987, B. S. Wurzell s.n. (LTR).

ORIGIN AND DISTRIBUTION OF ARTEMISIA x WURZELLII
Since the original discovery in 1987, B.S.W. has found 13 additional sites for A. x wurzellii in
eastern and south-eastern London.

sumey, v.c. l7:

1 Roadside, near Stockwell Underground Station, Lambeth, TQ/30.76, 1988, still there 1990, not
visited since.

2 Roadside among bushes, near Ruskin Park, Camberwell, Southwark, TQ/32.75, 1990, not
visited since.

3 Grassland and scrub, off Neate Street, Burgess Park, Walworth, Southwark, TQ/32.77 &
TQ/33.77, 1995, still there 1998.

4 Shrubbery weed by road, Dog Kennel Hill, East Dulwich, Southwark, TQ/33.75, 1992, not
visited since.

5 Waste ground, off Tooley Street, by London Bridge Station, Southwark, TQ/33.80, 1988, not
visited since.

6 In rough grass, near Straker’s Road, Peckham Rye Common, Southwark, TQ/34.75, 1993, not
visited since.

7 Shrubbery weed in small park, near Chambers Wharf, Bermondsey, Southwark, TQ/34.79,
1988, not visited since.

8 Disturbed soil, King Stairs Gardens, off Brunel Tunnel Roundabout, Rotherhithe, Southwark,
TQ/34.79, 1992, not visited since.

9 By footpath, near Redriff Road, south end of Russia Dock Woodland, Rotherhithe, Southwark,
TQ/36.79, 1993, not visited since.

10 In newly landscaped shrubbery, near Salter Road, north end of Russia Dock Woodland,
Rotherhithe, Southwark, TQ/36.80, 1993, not visited since.

146 C. M. JAMES, B. S. WURZELL AND C. A. STACE
South Essex, v.c. 18:

11 Rough ground, near Coppermill Lane, off Walthamstow Marshes, Waltham Forest, TQ/35.87,
1987 & 1988, not visited since.

12 Stony waste ground, Bow Creek Ecological Park, Canning Town, Newham, TQ/39.81, 1994,
still there 1998.

Middlesex, v.c. 21:

13. Garden weed in new housing estate, Watermint Quay, Clapton, Hackney, TQ/34.88, 1994, still
there 1998.

14 Overgrown railway bank, Northumberland Park Station, Tottenham, Haringey, TQ/34.91,
August 1987, still there 1998.

In addition, two other records, neither of which we have any reason to doubt, have come to our
notice:

15 West Kent, v.c. 16: on rubbish tip, Swanscombe, TQ/60.74, G. Kitchener & M. Keene, 1994,
site cleared soon after (Bull. Kent Field Club 40: 35-36 (1995)).

16 Middlesex, v.c. 21: weed in shrubbery, Fair Street, St John’s Estate, Bermondsey, TQ/32.78, J.
R. Palmer, 1989, comm. J. E. Smith.

The first-discovered colony (no. 14 above) extends for at least 50 m along a railway bank
together with common coarse perennials of such habitats, including Armoracia rusticana,
Arrhenatherum elatius, Calystegia silvatica, Rumex obtusifolius and Urtica dioica. Artemisia
vulgaris occurs close by, but the nearest colony of A. verlotiorum is at least 400 m distant on
Tottenham Marshes. Moreover, the two parents do not normally overlap in flowering times, and A.
verlotiorum has not been seen to set seed in this country. A. vulgaris might also fail to set seed
from flowers opening as late as October. Hence, although the hybrid was at first thought to have
arisen spontaneously in sifu, it seems more likely that it arose elsewhere, perhaps not in Britain,
and was inadvertently dumped on the railway bank. All of the other localities are on rough or
disturbed ground or in recent habitats created by gardening, and it is very likely that the present
distribution is the result of soil and waste being transported by man. One need look no further than
Fallopia japonica to understand how effective this method of dispersal can be! It is, indeed,
possible that all 16 colonies represent a single clone which was introduced, perhaps as a medicinal
plant, from southern Europe, where A. verlotiorum is widespread and regularly forms fruits (and
where the hybrid should be sought), or even from China. On the other hand, the chromosome
number of the hybrid indicates that its A. verlotiorum parent had 2n = 52, a number not previously
reported but characteristic of the Tottenham clone. There are counts of only 2n = 48 and 54 (and
16) for this species from southern Europe (Italy and Spain); such plants could not have produced a
hybrid with 2n = 34 by crossing with plants of A. vulgaris with 2n = 16. Hence a London origin for
the hybrid is certainly possible.

ACKNOWLEDGMENTS

We dedicate this paper to Douglas H. Kent, who was the leading expert on the flora of Middlesex
(especially its introduced species), who considerably extended our knowledge of the distribution of
A. verlotiorum in London, and who was a very good friend to C.A.S. and B.S.W. over many years.
We thank Miss A. P. Conolly for providing living material of A. verlotiorum from Abersoch.

REFERENCES

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alcuni caratteri morfo-anatomici distintivi delle due specie. Webbia 37: 185-196.

BOLKHOVSKIKH, Z., GRIF, V., MATVEJEVA, T. & ZAKHARYEVA, O. (1969). Chromosome numbers of
flowering plants. Academy of Sciences of the U.S.S.R., Leningrad.

ARTEMISIA X WURZELLII 147

BREMER, K. & HUMPHRIES, C. J. (1993). Generic monograph of the Asteraceae-Anthemideae. Bulletin of the
Natural History Museum, Botany 23: 7-177.

BRENAN, J. P. M. (1950). Artemisia verlotiorum Lamotte and its occurrence in Britain. Watsonia 1: 209-223.

CLEMENT, E. J. & FOSTER, M. C. (1994). Alien plants of the British Isles. Botanical Society of the British
Isles, London.

KAWANO, S., NAGAI, Y. & HOSHTYA-USHIDA, S. (1995). A study on the natural hybrid swarms of two
Artemisia species, A. capillaris and A. japonica (Compositae) in central Honshu, Japan, with special
reference to its biological status. Journal of phytogeography and taxonomy 42: 133-153.

LING, Y. R. (1988). The Chinese Artemisia Linn. - the classification, distribution and application of Artemisia
Linn. in China. Bulletin of Botanical Research, Harbin 8: \-61.

MorTON, J. K. (1977). A cytological study of the Compositae (excluding Hieracium and Taraxacum) of the
British Isles. Watsonia 11: 211-223.

PERRING, F. H. & WALTERS, S. M. (1962). Atlas of the British flora. Thomas Nelson & Sons, London &
Edinburgh.

POLIAKOV, P. P. (1961). Artemisia, in SHISHKIN, B. K. & BOBROV, E. G., eds. Flora U.R.S.S. 26: 425-631.
Editio Academiae Scientiarum U.R.S.S., Moscow & Leningrad.

RYVES, T. B. (1984). Artemisia L. species in the British Isles. Watsonia 15: 109-112.

TUTIN, T. G., PERSSON, K. & GUTERMANN, W. (1976). Artemisia L., in TUTIN, T. G. et al., eds. Flora
Europaea 4: 178-186. Cambridge University Press, Cambridge.

WURZELL, B. S. (1988). Wurzell’s weeds. BSBI news 49: 53.

(Accepted September 1999)

a

.

Watsonia 23: 149-159 (2000) 149

Historical records of Lactuca serriola L. and L. virosa L. in
Britain, with special reference to Cambridgeshire (v.c. 29)

P. H. OSWALD
33 Panton Street, Cambridge, CB2 1HL

“Most of our botanical publications are taken one from the other; and thus if an eminent botanist has in the
course of his researches fallen into a mistake, the error has been propagated.” (Jacson 1797, pp. 229-230).

ABSTRACT

Thomas Johnson (1633) was the first British botanist to realise that three sorts of large wild lettuces occur in
England, though he apparently thought that what is now called forma integrifolia of Lactuca serriola L. was
related to L. virosa L. Records of L. virosa and L. serriola in Cambridgeshire (v.c. 29) are reviewed here. The
long-held belief that John Ray (1660) recorded both of them in the county is shown to be incorrect: he, in fact,
saw two forms of L. serriola. This paper casts doubt on some of the eighteenth and nineteenth century records
of L. virosa in Cambridgeshire. The present distribution of the Lactuca taxa considered in the paper is briefly
discussed.

KEYWORDS: Asteraceae, wild lettuces, distribution, Thomas Johnson, John Ray, Kent, Middlesex.

INTRODUCTION

Descriptions of Lactuca serriola and L. virosa in most British Floras are unsatisfactory because the
existence of two forms of each with differing leaf characters is ignored. The brief accounts in
Stace (1991, 1995, 1999) are accurate but lack many of the useful characters. The differences
between the four taxa are, however, satisfactorily described in Plant crib 1998 (Rich & Jermy
1998, pp. 292-293), with improvements in the corrigenda and addenda in the reprint of 1999, and
will be fully treated in Volume 4 of Peter Sell and Gina Murrell’s Flora of Great Britain and
Ireland (Cambridge University Press, Cambridge).

Both species are biennials or overwintering annuals, initially with a basal rosette and later with
leafy stems, branched, at least above, and usually with numerous capitula. Stunted plants occur,
but well-grown plants of L. virosa can exceed 2:5 m and L. serriola can attain 2 m. However,
because L. serriola often grows in recently disturbed sites, plants that have reached full size in the
absence of competition are frequent, whereas L. virosa grows mainly in post-disturbance grassy
vegetation, where it is reduced in size by competition (R. N. Carter, pers. comm.). The main
panicle of L. virosa is characteristically rhomboid, with little obvious leafage within it; that of
L. serriola, at least in taller plants, is less regular and usually less pointed or even flat-topped, with
the lower branches leafy. L. virosa begins to flower about a fortnight earlier than L. serriola,
normally in June, and the diameter of the open capitula is often double that of L. serriola (up to 20
mm: pace Clapham 1952 and later editions!); the capitula of both species close by late morning, so
these are often missed. Fruit characters are much more reliable than leaf characters: the fresh
achenes of L. virosa are strikingly purple or maroon and larger than the olive-grey achenes of
L. serriola. Another useful diagnostic character, which is unfortunately ignored in most British
Floras, is the smell when the plant is broken or bruised: L. virosa smells like Opium Poppy
(Papaver somniferum L.), L. serriola like Garden Lettuce (Lactuca sativa L.).

Two foliage forms of L. serriola occur, sometimes intermixed but often in separate populations,
the type with deeply runcinate-pinnatifid cauline leaves, the other with undivided or shallowly
lobed cauline leaves. There are also two forms of L. virosa, but here it seems that populations are
more often mixed and the difference between the two is less striking and has rarely been taken
note of: the type has unlobed, sinuate-dentate cauline leaves throughout; a form with the higher
stem leaves pinnatisect with wide lobes has been named var. /actucarii (Lamotte) Rouy, though
P. D. Sell proposes to reduce the name to forma level in Flora of Great Britain and Ireland.

150 P. H. OSWALD

Prince & Carter (1977) reviewed the characters distinguishing L. serriola, L. virosa and the
related but smaller L. saligna L. and revised the taxonomic treatment of L. serriola, giving to its
unlobed-leaved form the name forma integrifolia (S. F. Gray) S. D. Prince & R. N. Carter. Later,
Carter & Prince (1982) discussed the history of the taxonomic treatment of L. serriola and
L. virosa. They showed that, although Ray (1690) provided a “treatment of the British taxa ... as
good as any published since” and Linnaeus (1756) “laid the foundation for the taxonomy of
L. serriola and L. virosa as we know it’, by “the second half of the nineteenth century the
existence of an unlobed-leaved form of L. serriola was almost completely overlooked in Britain”,
the general practice being “to call all unlobed-leaved plants L. virosa without even according
varietal status to those that were really L. serriola”. They concluded that, as a result, “most pre-
1930 L. virosa records could as easily refer to L. serriola as to L. virosa”.

In this paper I show that the existence of three of the four taxa of large wild lettuces was clearly
understood by Thomas Johnson by 1633 but that John Ray (1627-1705) at first failed to recognise
them. As a combined result of this fact and the later confusion described by Carter & Prince
(1982), it has been wrongly believed since the time of John Martyn (1727), through that of C. C.
Babington (1860), and right up to the present day that L. virosa was first recorded in
Cambridgeshire by Ray (1660).

THE EARLY SEVENTEENTH CENTURY

In his first account of a botanical excursion into Kent, Thomas Johnson (1629) recorded “Lactuca
agrestis odore opi” on the Isle of Sheppey on 14 July 1629. This is the first localised British
record of L. virosa (Clarke 1900, p. 85), which is “still present here” (Gilmour 1972, p. 56). In a
list of 1632 of “the plants growing on Hampstead Heath and in the places visited ... on the way
there and back”, he recorded “Lactuca syl. odore viroso folijs dissectis, Lactuca sylv. prior, Trag.
sily. Matth.” and also “Lactuca syl. alter odore magis viroso folijs non dissectis, Lactuca agrestis
odore opij Lob. Ad. Endivia major & I. Trag. Thesion, Lugd.”. These names may be interpreted
thus: “Wild Lettuce with a fetid smell and divided leaves; the first wild Lettuce of Jerome Bock
(Tragus 1552, p. 259; Figure 1| in this paper); the wild Lettuce of P. A. Matthioli (Matthiolus 1558,
pp. 295-296)” and “The second wild Lettuce, with a more fetid smell and undivided leaves; the
field Lettuce with the smell of opium of Stirpium adversaria nova of Pierre Pena and Matthias de
l’ Obel (Pena & de Lobel 1571, p. 89); the greater, first Endive of Jerome Bock (Tragus 1552, p.
267); Thesion of Historia generalis plantarum by J. D’Aléchamps, published at Lyons
(Lugdunum) (Dalechampius 1587, pp. 564-565; Figure 2 in this paper)’. Clarke (1900, p. 85)
listed the former as the first British record of L. serriola, and Gilmour (1972, p. 128) identified the
two plants as L. serriola and L. virosa, while Kent (1975, p. 491) accepted the latter identification
but seems to have had doubts about the former. Both Clarke and Kent give “Hampstead Heath” as
the locality, but, for reasons that will soon become apparent, I believe that the two plants were
found on the way there or back, “betweene London and Pancridge [?Old St Pancras] Church’, and
were L. serriola forma serriola and forma integrifolia respectively, though Johnson had not yet
worked this out. If my theory is correct, these are the first British records for both forms of L.
serriola.

In the chapter of his Herball entitled “Of Lettuce”, John Gerarde (1597, p. 238) had stated that
there were sorts “both wilde and tame”, which he purposed “to laie down”; but in fact he dealt
only with the cultivated kinds. In his revision of Gerarde’s work, Johnson (1633) rectified this
omission, in Chapter 38 of Book 2 (p. 309), thus:

There are three sorts of wilde Lettuce growing wilde here with vs in England, yet I know not
any that haue mentioned more than two; yet I thinke all three of them haue beene written of,
though two of them be confounded together and made but one (a thing often happening in the
history of Plants) and vnlesse I had seene three distinct ones, I should my selfe haue beene of
the same opinion.

| The first and rarest of these hath long and broad leaues, not cut in, but only snipt about the
edges, and those leaues are they that are on the lower part of the stalke almost to the midle
thereof: then come leaues from thence to the top, which are deepely diuided with large
gashes: the stalke if it grow in good grounds exceeds the height of a man, (for I haue seene it
grow in a garden to the height of eight or nine foot) .... The whole plant is full of a clammy
milky iuice, which hath a very strong and grieuous smell of Opium [1.e. the latex of Papaver
somniferum].

HISTORICAL RECORDS OF LACTUCA SERRIOLA AND VIROSA 151

2 This hath broad leaues only cut about the edges, but not altogether so large as those of the
last described: the stalke, which commonly is two cubits or better high, is also smooth, and
diuided into many branches, bearing such floures as the last described; and this also hath a
milky iuice of the same smell as the last described, from which it differs only in magnitude,
and that this hath all the leaues whole, and not some whole and some diuided, as the former.
3 This in stalkes, floures and seedes is like to the last described, but the leaues are much
different, for they are all deeply gashed or cut in like as the leaues of Succory, or Dandelion.
This also is full of a milky iuice, but hath not altogether so strong a sent of Opium as the two
former, though it partake much thereof. The stalke of this is sometimes a little prickly, and so
also is the middle rib vpon the backeside of the leafe.

Johnson illustrated only the first and last of these plants (p. 309), labelling them respectively
“1 Lactuca syl. maior odore Opij. The greater wilde Lettuce smelling of Opium.” and “3 Lactuca
sylvestris folijs dissectis. The wilde Lettuce with the diuided Leafe.” The illustrations are poor
representations of the taxa; that of the last is a reduced and reversed version of Fuchsius’ (1542,
p. 301) full-page woodcut of “Lactuca syluestris”’, similar to that of Tragus (1552) (Figure 1 in this
paper) but single-stemmed.

In the section of Chapter 38 headed “The Names” (p. 310), Johnson wrote:

1 I take the first of these to be the Lactuca Sylvestris of Dioscorides and the Ancients, and
that which the Authours of the Adversaria [Pena & de Lobel 1571, p. 89] gaue vs vnder the
title of Lactuca agrestis scariole hortensis folio, Lactuce flore, Opij odore vehementi,
soporifero & viroso [field Lettuce with the leaf of garden chicory, the flower of Lettuce and a
strong, soporific and stinking / poisonous smell of Opium].

2 This is the Endiuia of Tragus [1552], pag. 268. and the Thesion of Daleschampius [1587],
pag. 564. Bauhine [Bauhinus 1623, p. 123] confounds this with the former.

3 This is the Lactuca Sylvestris prior, of Tragus [1552, p. 259]: the Lactuca Sylvestris of
Matthiolus [1558, pp. 295-296], Fuchsius [1542, pp. 298-303], Dodoneus, and others: it is
the Seris Domestica of Lobell [Pena & de Lobel 1571, p. 86].

There is no doubt that the first of these plants is Lactuca virosa (var. lactucarii from the
description of the upper leaves) and the last L. serriola forma serriola, to which Johnson allots the
two synonyms he gave to his first lettuce of 1632. The synonyms Johnson provides for his second
sort here are the last two names he gave to the “second wild Lettuce” of 1632, when it seems that
he too was still confounding two of them together and making but one, since he applied to one
taxon names that he later divided between two.’ Both Tragus (1552) and Dalechampius (1587)
(Figure 2 in this paper) show a plant with the undivided leaves of L. serriola forma integrifolia,
but it is odd that Johnson suggests that it too has a “smell of Opium’, which is true only of
L. virosa. However, it seems that he found it growing with L. serriola forma serriola, as he gives a
joint locality, “plentifully betweene London and Pancridge Church, about the ditches and highway
side”, whereas he records L. virosa only from John Parkinson’s garden, grown from seeds sent by
John Goodyer, who had found it growing in Hampshire (pp. 309-310). It is surprising that he does
not mention his locality of 1629 here, but perhaps, having realised that neither of the plants he had
found in 1632 was L. virosa, he was uncertain that he had really seen it in Kent either.

JOHN RAY’S ACCOUNTS

In his Catalogus plantarum circa Cantabrigiam nascentium, Ray (1660, Pp. 82-83) included two
kinds of large wild lettuces and a smaller one that he then thought was “not yet described’, i.e.
Lactuca Saligna, to which he gave the name “The least cut-leaved wild Lettuce’.” To the first large
lettuce, “Lactuca sylvestris costa spinosa C.B.”, he gave the name “Cut-leaved wild Lettuce” and
to the second “Endive-leaved wild Lettuce’. He described the former as growing “Ad agrorum
margines [By the margins of arable fields]. As by the way side from Ely to Cambridge, within a
mile or two of Ely.” The Latin synonyms given for it, which include “sylv. foliis dissectis Ger.
emac [i.e. in Johnson’s emaculated (or emended) edition of Gerarde of 1633] and “Seris
domestica Lob.” [Pena & de Lobel 1571, p. 86], make it clear that this was L. serriola forma
serriola.’ No locality or habitat is given for the second kind, but the first Latin name given is

152 P. H. OSWALD

“Lactuca sylvestris 2. Ger. emac. [1.e. Johnson’s second kind of wild Lettuce]. After this there
follow five synonyms, the first three including “viroso” and clearly relating to L. virosa; the last
two, however, are “Endivia Trag. p. 268.” and “Thesion Dalechampii Lugd. p. 564.”, which
Johnson (1633) gave as names for his second kind.

At the end of his account of his own second kind, Ray included a rather contorted note in Latin’,
which is poorly translated by Ewen & Prime” (1975, p. 77). This note is better rendered thus:

Johnson, in his edition of Gerarde, creates two kinds of wild lettuce with the smell of opium,
which he thinks are confused in the works of other writers; and he attributes to the first sort
earlier authors’ names which we think are more appropriate to his second sort, since we
believe that our plant, which is one with undivided leaves and is Gerarde’s second one, is the
only one described by botanical writers, rather than, as he suggests, Gerarde’s first.

This shows that, at this time, Ray believed there were only two kinds of wild lettuce other than
L. saligna. Moreover, he clearly thought that his own second kind fitted Johnson’s (1633) second
description rather than his first.

On this evidence alone, Martyn (1727), Babington (1860) and others might perhaps be excused
for having reached the conclusion that Ray’s second large Lactuca was L. virosa, but Ray’s
subsequent works make it certain that it was in fact L. serriola forma integrifolia.® Later writers,
including Ewen & Prime (1975), seem not to have found the entry in the appendix of 1685 to
Ray’s Cambridge catalogue about his second lettuce, “Lactuca sylvestris 2. Ger. emac.”. This may
be translated thus:

Delete all the synonyms. In their place add ‘Lactuce sylvestris sive Endivie multis dicte
folio laciniato dorso spinoso varietas [A variety of the wild Lettuce, or Endive as it is known
to many people, with a laciniate leaf with a spiny underside] J.B.’’ This does not differ from
Gaspard Bauhin’s wild Lettuce with a spiny midrib [Bauhinus 1623, p. 123] in any respect
other than in the leaves, which in it are not laciniate even on the stem. We now retract what
we propounded about this plant in the Catalogue and we assent to Johnson’s opinion.®

The last sentence had appeared already in Ray’s Catalogus plantarum Anglie (1670, 1677), in
which he included three taxa of large wild lettuces. In this work Ray indicated the plants that grew
wild around Cambridge (“Agri Catabrigiensis alumnas”’) by placing “C.” before their names, and
by this means he showed that he had recorded only two of the lettuces in Cambridgeshire and not
“The greater strong-scented wild Lettuce”. Even earlier, William How (1650) and Christopher
Merrett (1666) had listed four wild lettuces including “Lactuca sylvestris altera folio non laciniato,
odore minus vehementi / vehemente [The second wild Lettuce with the leaf not laciniate, with a
less powerful smell]”.

Presumably, by the time he wrote his retractions, Ray had seen Lactuca virosa for the first time,
somewhere outside Cambridgeshire, and had realised that his second taxon in the Cambridge
catalogue was not it. At any rate, in his later botanical works he distinguished three taxa of large
annual wild lettuces’. Thus, in Ray’s (1686) Historia plantarum, he listed “The greater strong-
sented wild-Lettuce”, “Cut-leaved wild Lettuce” and “Endive-leaved wild Lettuce”: each is
equated with one of Johnson’s three kinds, but they are reordered to put Johnson’s second kind
last. About this kind Ray commented:

Johnson, the emender of Gerarde, makes this species a variety of the first species
[L. virosa]'°, but J. Bauhin of the second [L. serriola forma serriola: see note 7], whom we
also support, since it is less tall than the former and has smaller leaves and also a less fetid
smell, at least to our nostrils. On the contrary, it does not differ in any character from the
second species except in the shape of the leaves, which in this species are not laciniate even
on the stalk. It grows in the same places as the preceding [i.e. second] species."

The three kinds are listed in the same order in the Synopsis (Ray 1690, p. 41), where the second
(i.e. L. serriola forma serriola) is called “Milder-scented, cut-leaved wild Lettuce’; the third,
which Ray here calls “Lactuca sylv. folio non laciniato. Sylv. 2. Ger. emac. [Wild Lettuce with the
leaf not laciniate;. the second wild one in the emended Gerarde.] Endive-leaved wild Lettuce’, 1s
described as growing “with the preceding one, of which it is a variety, but found more rarely”. &

HISTORICAL RECORDS OF LACTUCA SERRIOLA AND VIROSA 153

THES SON Dalechamp.
Endiuia Tragi.

FIGURE 1. Woodcut of “Lactuca siluestris” from FIGURE 2. Woodcut of “Thesion” from J. D’Aléchamps’
Book 1 of Jerome Bock’s Commentaries General History of Plants (Dalechampius 1587), with
(Tragus 1552), interpreted by Thomas Johnson uncut leaves which led Thomas Johnson (1633) to
(1633) in his edition of Gerarde’s Herball as regard it as a representation of Lactuca serriola forma
representing Lactuca serriola forma serriola. integrifolia. It is a redrawn and reversed image of

Tragus’ (1552) illustration of “Endiuia”.

EIGHTEENTH AND EARLY NINETEENTH CENTURY CAMBRIDGESHIRE RECORDS

John Martyn (1727, p. 21) listed “Lactuca sylvestris costa spinosa” and “‘Lactuca sylvestris major
odore Opii” in his account of the Cambridgeshire flora, equating the latter with “Lactuca sy!vestris
2. Ger. emac. Cat. Cant. [i.e. Ray 1660]. He was thus the first of a long line of Cambridgeshire
botanists to attribute a record of L. virosa to John Ray. In a working volume containing Ray’s
Catalogus and his own Methodus bound together, preserved in the Library of the Department of
Plant Sciences of the University of Cambridge, he wrote out a list headed “1727 Desiderata’”’,
among them the three Lactuca taxa of Ray’s Catalogus (including L. saligna); he later deleted
many of the taxa listed here, but none of the three Lactuca taxa, which suggests that he himself
never saw any of them in Cambridgeshire.

In his Plante Cantabrigienses (essentially a table providing the Linnaean names for the plants
recorded in Ray’s Catalogus and its appendices and in John Martyn’s Methodus), John’s son,
Thomas Martyn (1763a, p. 18), listed Lactuca virosa, its var. B and L. saligna. The table shows
that he too regarded Ray’s “Lactuca sylvestris 2.” as L. virosa, while equating Ray’s “Lactuca
sylvestris costa spinosa” (the cut-leaved form of L. serriola) with its var. B.'° Thomas Martyn also
annotated an interleaved copy of his father’s Methodus, which is also in the Library of the

154 P. H. OSWALD

Department of Plant Sciences at Cambridge. On a blank page opposite “Lactuca sylvestris major
odore Opii” he wrote “Near Clare hithe July 27 1759’, but later he added a line linking this to
“Lactuca sylvestris costa spinosa” instead. In a handwritten index of localities inserted at the end
of the volume, he entered “Lactuca sylvestris costa spinosa” under “Ely” and “Lactuca sylv. maj.
odore Opu” under “Clare hithe”; the latter has not been amended, but nevertheless it seems likely
that Thomas Martyn never saw L. virosa. It may be significant that neither species is listed in his
Herbationes Cantabrigienses (Martyn 1763b), which consists of lists of plants under localities
“comprehended in thirteen Botanical Excursions”.

Richard Relhan included only two large lettuces in his Flora Cantabrigiensis (1785, 1802,
i820) — L. virosa, “Strong-scented wild Lettuce”, and “L. Scariola” (a synonym for L. serriola),
“Prickly Lettuce”, the latter evidently forma serriola as it is equated with the second kind in Ray’s
Synopsis and with Johnson’s “L. sylvestris, foliis dissectis”. In 1785 the only locality given for the
former was “Burwell-pit’ and for the latter “Jn the Isle of Ely’. In the later editions “Jn the Road to
Cottenham, by the second Bridge from Histon” and “Ditch near Denny Abbey” were added for
L. virosa and “Hazenfield [Haslingfield], in the Lane leading into the Village from Cambridge’,
“Between Histon, and Rampton’, “In a Lane leading from Long Stanton towards Swavesey” and
“Burwell Pit’ were added for L. serriola. There seems little reason to doubt the localities for
L. serriola, but what about those for L. virosa? The fact that Burwell Pit was later given as a
locality for L. serriola suggests that the plant recorded there in 1785 may have been its forma
integrifolia. The specimen of “Lactuca Scariola” figured in English Botany (Smith & Sowerby
1795, no. 268) was sent to Sowerby from “near Denny Abbey” by the Rev. John Hemsted, so the
record from a ditch nearby may well have related to its unlobed-leaved form.'* Similarly, the
locality by the road to Cottenham is suspiciously close to that between Histon and Rampton. As
implied by Johnson (1633) and specifically noted by Ray (1690), the two forms of L. serriola
often grow together, whereas, at least until the last few decades, L. virosa seems rarely to have
occurred with L. serriola.’”

C. C. BABINGTON AND HIS CONTEMPORARIES

Despite Babington’s (1860) mistake in identifying Ray’s (1660) “Lactuca sylvestris 2.” as L.
virosa, there is no doubt that he himself knew the latter, at least at Cherry Hinton: the only new
locality for this species given in his Flora of Cambridgeshire is “Chalk-pit-close, Hinton”, on his
own authority, though the entry “Burwell Pit!; Relh.” signifies that he claimed to have seen a
specimen of Relhan’s from Burwell Pit. The Cherry Hinton locality is supported by three
specimens in CGE, the earliest collected by J. S. Henslow on 21 July 1825'° and the two others by
Babington himself on 10 June 1846 and 3 August 1855. Amendments (from “‘Scariola’ to
“vyirosa’) to the label of the earlier of the two specimens and to the record from “Chalk-pit close
Hinton” in the manuscript from which Babington prepared his 1860 Flora (bound in three large
volumes kept in the Library of the Cambridge Department of Plant Sciences) suggest that
Babington had some difficulty in distinguishing the larger Lactuca species. The only other
nineteenth century specimen from Cambridgeshire in CGE was collected by James Backhouse
(1825-1890) at “Elms n’. Wisbeach” (i.e. Elm, near Wisbech) in 1844."”

Babington (1860) gave three new localities for L. serriola.'* He also included a further entry of
“Burwell Pit!; Relh.” and a record from “Denny Abbey” based on Hemsted’s “specimen figured in
Eng. Bot. [Smith & Sowerby 1795, no. 268]” (as well as repeating Relhan’s record of L. virosa
there). Perring et al. (1964) suggested that in Babington’s time L. serriola was “as rare as L.
virosa’”.’” The four nineteenth century specimens from Cambridgeshire in CGE are all of forma
serriola.”

TWENTIETH CENTURY AUTHORS

Evans (1939) recorded L. virosa as “Very rare; probably extinct” in Cambridgeshire. After
quoting a supposed record of Ray’s “from Ely”*', Relhan’s records and Babington’s (1860) record
from “Hinton”, he added: “We have no later record.” He gave a number of localities for L.
serriola, “generally close to towns and villages”.

HISTORICAL RECORDS OF LACTUCA SERRIOLA AND VIROSA 155

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ine,

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2 3 a SS) 6 ih

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FIGURE 3. Map of Cambridgeshire (v.c. 29) showing post-1986 tetrad records for Lactuca virosa. Most of the
county is in TL (=52) but the northern part is in TF (=53).

156 P. H. OSWALD

Perring, Sell & Walters (1964) repeated the error that John Ray recorded L. virosa in
Cambridgeshire~ but they treated the species as extinct in the county, giving 1855 as the last
certain date for it.” However, Crompton & Whitehouse (1983) recorded that “it was certainly near
the gas works during the 1940—1950s, and it is still to be found on wasteland in the Newmarket
Road area and along nearby railway lines”. Perring et al. regarded L. serriola as “now a frequent
weed of roadsides and disturbed ground throughout the county” and suggested that a “rapid
spread” had taken place “between 1930 and 1945”, adding: “In recent years it may have declined
again.”°* They did not mention forma integrifolia, but one at least of the authors was well aware of
its existence (P. D. Sell, pers. comm.), despite their misleading comment that “the more nearly

simple leaves of L. virosa are usually sufficient to distinguish it”.

DISCUSSION

The confusion occasioned by the failure of most British Floras of the nineteenth century to
recognise that L. serriola can have simple leaves, so well described by Carter & Prince (1982), has
led to some uncertainties about the past status of this species and L. virosa in Cambridgeshire, as
elsewhere in Britain; but, in its case, the belief that L. virosa was known to Ray has until now
further clouded the picture. It is impossible to say with certainty when this species was in fact first
seen in the county, but it is not improbable that in the nineteenth century it occurred only at Cherry
Hinton (TL/48.56) and Elm (probably TF/46.06 or 47.06) and that it then disappeared until the
1940s. At any rate the first certain record for Cambridgeshire was made by the Rev. Leonard
Jenyns at Chalkpit Close, Cherry Hinton, on 12 August 1824 (see note 16). Previously more
characteristic of “naturally unstable habitats, such as sand-dunes and cliff-ledges” and “frequently
found on chalk and oolitic limestone” (Prince & Carter 1977, p. 336), L. virosa is becoming more
widespread in Britain in ruderal habitats like those long colonised by L. serriola.'> Since about the
1970s it has increased substantially in Cambridgeshire, especially on the margins of dual
carriageways (e.g. TL/3.4, 4.4 and 6.6) and on waste ground in Cambridge (TL/4.5). Figure 3
shows post-1986 records for 16 out of the 30 hectads that include a substantial area in
Cambridgeshire and 50 out of some 575 tetrads in the county.

L. serriola has increased even more markedly both in southern Britain and in Cambridgeshire,
though with an earlier start. Recording for Atlas 2000 shows its presence now in every hectad in
and around Cambridgeshire. Since 1986 records have been made in 201 tetrads in Cambridgeshire,
but these probably do not represent its complete distribution. What 1s particularly puzzling 1s why
its forma integrifolia, although already present at the time of Johnson and Ray, was formerly
apparently rarer than the nominate cut-leaved form, when now it is by far the commoner form. By
the 1950s Kent & Lousley (1954, p. 176) wrote of it (as L. serriola L. var. dubia (Jord.) Rouy):
“This is the aggressive entire-leaved form, possibly of adventive origin, which has spread so
rapidly in the London Area in recent years. Plants with runcinate leaves occur occasionally.” In
Cambridgeshire, P. D. Sell (pers. comm.) has suggested that forma serriola is native in the Fens
and forma integrifolia is a relatively recent immigrant, and Beckett & Bull (1999, p. 201)
apparently regard this as proven; though it is true that many localities for the former are in the
Fens (including those for all four nineteenth century specimens in CGE”), this theory seems too
simplistic, particularly since mixed populations often occur.

Prince & Carter (1977, p. 334) suggested that, “‘although there are no absolute barriers to natural
cross-pollination” between the two forms, “it is nevertheless an extremely rare occurrence”’.
Perhaps there are environmental factors that favour the predominance of one form or the other in
particular circumstances. Prince & Carter also wrote (p. 337): “Whereas the pinnatifid-leaved plant
is the less common variant in Britain (having a particularly restricted south-eastern distribution),
on the Continent it is the more abundant of the two.” My own observations suggest that this
generalisation is too sweeping: in much of western France the position is similar to that in south-
east England, and forma serriola certainly occurs in Wales. Could a more continental climate
favour the predominance of forma serriola and a more oceanic one that of forma integrifolia? This
might help to explain the former predominance of forma serriola in the Fens (with a more
continental climate than most of Britain), but the present predominance of forma integrifolia in
urban and roadside habitats is harder to account for: although the extremes of winter climate are
buffered by the warming effect of housing and traffic, these habitats are hotter and drier in
summer.

HISTORICAL RECORDS OF LACTUCA SERRIOLA AND VIROSA 157
ACKNOWLEDGEMENTS

My interest in Lactuca taxa was kindled by reading Prince & Carter (1977), when it suddenly
became clear why I| had experienced such difficulty in identifying these plants as a schoolboy in
the Thames Valley. In a nice piece of symmetry, their paper appeared in the issue of Watsonia
dedicated to the memory of J. Edward Lousley, in which Duggie Kent described Rumex x lousleyi
and provided a bibliography of Lousley’s published works. I am very grateful to the late Dr D. E.
Coombe for encouraging my interest in Lactuca, to Dr S. M. Walters for supplying the quotation at
the head of this paper, to Mrs G. Crompton for help in tracing Cambridgeshire records, to the staff
of Cambridge University Library’s Rare Books and Manuscripts Rooms for their courteous
service, to Dr D. Briggs. Mr P. D. Sell and Mrs G. Murrell for their kind assistance in Cambridge
University Herbarium (CGE), to Mr R. Savage for similar help in the Library of the Department
of Plant Sciences, to Dr C. D. Preston for reading a draft of this paper and making helpful
comments, and to Mr G. M. S. Easy and Mr D. A. Wells for supplying many of the records for
Figure 3. I also thank the Syndics of Cambridge University Library for the use of illustrations from
their books for Figures | and 2 and Mrs J. Gaunt for scanning them and cleaning them up for
printing.

REFERENCES

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BECKETT, G. & BULL, A. (1999). A Flora of Norfolk. Gillian Beckett, Stanhoe, Norfolk.

BAUHINUS, C. (1620, reissued 1671). Prodromus theatri botanici. Francofurti [Frankfurt].

BAUHINUS, C. (1623, reissued 1671). Pinax theatri botanici. Basileae [Basel].

BAUHINUS, J., CHERLER, J. H. & CHABRAEUS, D. (1651). Historia plantarum universalis. Tomus I. Ebroduni
[ Yverdon].

CARTER, R. N. & PRINCE, S. D. (1982). A history of the taxonomic treatment of unlobed-leaved prickly
lettuce, Lactuca serriola L., in Britain. Watsonia 14: 59-62.

CLAPHAM, A. R. (1952). Lactuca L., in CLAPHAM, A. R., TUTIN, T. G. & WARBURG, E. F., Flora of the
British Isles, pp. 1125-1127. Cambridge University Press, Cambridge.

CLARKE, W. A. (1900). First records of British flowering plants, 2nd ed. West, Newman & Co., London.

CROMPTON, G. (1997). Botanizing in Cambridgeshire in the 1820s. Nature in Cambridgeshire 39: 59-73.

CROMPTON, G. & WHITEHOUSE, H. L. K. (1983). A checklist of the flora of Cambridgeshire. Published by the
authors, Cambridge.

[DALECHAMPIUS, J.] (1587). Historia generalis plantarum. [Part 1]. G. Rovillius, Lugduni [Lyons].

EVANS, A. H. (1939). A Flora of Cambridgeshire. Gurney & Jackson, London & Edinburgh.

EWEN, A. H. & PRIME, C. T., trans. & ed. (1975). Ray’s Flora of Cambridgeshire (Catalogus plantarum circa
Cantabrigiam nascentium). Wheldon & Wesley Ltd, Hitchin.

Fucustus, L. (1542). De historia stirpium commentarii insignes. Basileae [Basel].

GERARDE, J. (1597). The herball or generall historie of plantes. J. Norton, London.

GiLMouR, J. S. L., ed. (1972). Thomas Johnson. Botanical journeys in Kent & Hampstead. The Hunt
Botanical Library, Pittsburgh, Pennsylvania.

[How, W.] (1650). Phytologia Britannica. Londini [London].

[JACSON, M. E.] (1797). Botanical dialogues, between Hortensia and her four children, ... designed for the
use of schools, by a lady. J. Johnson, London.

JOHNSON, T. [1629]. Iter plantarum investigationis ergo susceptum. Privately printed, [?London].

JOHNSON, T. (1632). Enumeratio plantarum in Ericeto Hampstediano locisque vicinis crescentium. T. Cotes,
[London].

JOHNSON, T. (1633). Of wilde lettuce, in GERARDE, J., The herball or generall historie of plantes, 2nd ed., pp.
309-310. A. Islip, J. Norton & R. Whitakers, London.

KENT, D. H. (1975). The historical Flora of Middlesex. The Ray Society, London.

KENT, D. H. & LOUSLEY, J. E., comp. (1954). A hand list of the plants of the London Area. [Part 4].
Supplement to London Naturalist 33.

LINNAEUS, C. (1753). Species plantarum. Holmiae [Stockholm].

LINNAEUS, C. (1756). Centuria II plantarum. Upsalae [Uppsala].

[MARTYN, J.] (1727). Methodus plantarum circa Cantabrigiam nascentium. Londini [London].

MARTYN, T. (1763a). Plantae Cantabrigienses. London.

MARTYN, T. (1763b). Herbationes Cantabrigienses. London.

MATTHIOLUS, P. A. (1558). Commentarii ... in libros sex ... Dioscoridis ... de medica materia. Venetiis
[Venice].

158 P. H. OSWALD

MERRETT, C. (1666, reprinted 1667). Pinax rerum naturalium Britannicarum. Londini [London].

PENA, P. & DE LOBEL, M. (1571, reissued 1605). Stirpium adversaria nova. Londini [London].

PERRING, F. H., SELL, P. D. & WALTERS, S. M. (1964). A Flora of Cambridgeshire. Cambridge University
Press, Cambridge.

PRESTON, C. D. & HILL, M. O. (1997). The geographical relationships of British and Irish vascular plants.
Botanical journal of the Linnean Society 124: 1-120.

PRINCE, S. D. & CARTER, R. N. (1977). Prickly Lettuce (Lactuca serriola L.) in Britain. Watsonia 11: 331-338.

[RAY, J.] (1660). Catalogus plantarum circa Cantabrigiam nascentium. Cantabrigiae [Cambridge] & Londini
[London].

RAY, J. (1670). Catalogus plantarum Angliae. Londini [London].

RAY, J. (1677). Catalogus plantarum Angliae. Editio secunda. Londini [London].

RAY, J. (1686). Historia plantarum. Tomus I. Londini [London].

RAY, J. (1690). Synopsis methodica stirpium Britannicarum. Londini [London].

[RAY, J. & DENT, P.] (1685). Appendix ad catalogum plantarum circa Cantabrigiam nascentium: continens
addenda & emendanda. Editio secunda, aucta plantis sexaginta. Cantabrigiae [Cambridge].

RELHAN, R. (1785). Flora Cantabrigiensis. Cantabrigiae [Cambridge].

RELHAN, R. (1802). Flora Cantabrigiensis. Editio altera. Cantabrigiae [Cambridge].

RELHAN, R. (1820). Flora Cantabrigiensis. Editio tertia. Cantabrigiae [Cambridge].

RICH, T. C. G. & JERMY, A. C. (1998). Plant crib 1998. Botanical Society of the British Isles, London.

SMITH, J. E. & SOWERBY, J. (1795). English Botany. Vol. IV. London.

STACE, C. (1991, 2nd ed. 1997). New Flora of the British Isles. Cambridge University Press, Cambridge.

STACE, C. (1999). Field Flora of the British Isles. Cambridge University Press, Cambridge.

TRAGUS, H. (1552). De stirpium ... usitatis nomenclaturis, propriisque differentiis, ... commentariorum libri
tres. [Book 1]. Argentorati [Strasbourg]. (Translation from the original German into Latin by D. Kyber).

FOOTNOTES

1 The second of the names that Johnson gave to the “second wild Lettuce” of 1632 was the name he had
given to the plant he found on the Isle of Sheppey in 1629 and a shortened version of the long “title”
from the Adversaria given here to the first sort (i.e. L. virosa).

“Lactuca sylvestris laciniata minima nondum descripta. The least cut-leaved wild Lettuce. This was found

on a bank and in a ditch by the side of a kind of drove or lane leading from London road to the river, just

at the water near a quarter of a mile beyond the spittle-house end |“‘near the southern end of the present

Coe fen” according to Ewen & Prime (1975, p. 78)].”” Ray later (1670, p. 37) realised that this plant had

already been described by Gaspard Bauhin (Bauhinus 1620, p. 68) as “Chondrilla viscosa humilis”.

3._L. serriola forma serriola now occurs with forma integrifolia by the roundabout where A142 (Witchford
Road) leaves A10 (the Ely bypass) (TL/527.794).

4 “Johnsonus apud Ger. duo facit genera lactuce sylvestris odore opii, que apud alios scriptores existimat
confundi; & nomina autorum que nos secunde ejus speciei magis propria putamus attribuit prime;
siquidem nostram que foliis est indivisis & 2 Ger. solam a botanographis descriptam arbitramur, non
prout ille vult, primam Gerardi.”

5 “Johnson after Gerard makes two kinds of lactuca sylvestris by the odour of the opium (= milky juice)
which he considers are confused by other writers; and he attributes the name given in his authors to the
first which we consider more appropriate to the second species. We think that our kind, which has
undivided leaves (i.e. the second of Gerard) is the only one described by botanical writers and not the
first as he wishes. [*This separation is not accepted today.]”

6 Babington (1860, p. 141) further confused matters by attributing Ray’s locality for his first plant (i.e.
L. serriola forma serriola) to this, his second plant, which Babington thought was L. virosa.

7 This is based on a sentence in Jean Bauhin’s account of L. serriola (Bauhinus et al. 1651, p. 1003):
“Huius Lactucee quedam mihi species obseruata folio non laciniato etiam per caules. [I have observed a
certain kind of this Lettuce with the leaf not laciniate even along the stems.]” On the next page Bauhin
states that at Lyons he had observed this species “with its earlier, fairly long leaves [in February and
March] not laciniate” (“Ibidem obseruaui foliis non laciniatis prioribus longiusculis.”’).

8 “Lactuca sylvestris 2. Ger. emac | Dele synonyma omnia. Horum loco adde Lactuce sylvestris sive
Endiviz multis dicte folio laciniato dorso spinoso varietas J.B. Hec non alia in re differt a Lactuca sylv.
costa spinosa C.B. quam foliis que ei ne in caule quidem laciniata sunt. Que de hac planta in Catalogo
tradidimus jam retractamus & in Johnsoni sententiam concedimus.” (Ray & Dent 1685, pp. 22-23).
Unlike Johnson (1633), Ray evidently did not regard “Endivia” and “Thesion Dalechampii” as reliable
synonyms for “Lactuca sylvestris 2.”.

9 In fact they are often biennial (see INTRODUCTION).

i)

10

ja]

12

19

20

21

22

Mee

24

25

HISTORICAL RECORDS OF LACTUCA SERRIOLA AND VIROSA 159

This is not actually stated by Johnson (1633) but is implied by the order in which he presented the taxa
and by his statement that his last taxon “hath not altogether so strong a sent of Opium as the two former”.
“Hance speciem Johnsonus Gerardi emaculator prime speciei varietatem facit; J. Bauhinus secunde, cui
& nos suffragamur; siquidem illa humilior est, & foliis minoribus, odore etiam mints gravi, nostris
saltem naribus: Non alia autem in re a secunda differt quam foliorum figura, que huic ne in caule quidem
laciniata sunt. Iisdem cum priore locis provenit.” (Tomus 1, Lib. V, Cap. I: p. 222).

“Cum priore, cujus varietas est, sed raritis.”” Nowadays it is by far the commoner form in Britain (see
DISCUSSION). Prince & Carter (1977, p. 337) are wrong in saying that Ray (1690) “described them as
altogether different plants”.

This should be L. serriola forma integrifolia, since Linnaeus classified this taxon as var. B of L. virosa in
1753; however, in 1756, when Linnaeus first described L. serriola, he made forma integrifolia its var. Y
(Carter & Prince 1982).

Sowerby’s engraving shows a plant with incised lower leaves but dentate upper cauline leaves and so
may have been drawn from specimens of both forma serriola and forma integrifolia.

Clapham (1952) described L. virosa as a “sub-Mediterranean species, associated in C. Europe with...
thermophilous plants’, but L. serriola as “associated with steppe species”. Preston & Hill (1997,
pp. 38-39, 102, 104) classified L. virosa as “Suboceanic Southern-temperate” and L. serriola as
“Eurosiberian Southern-temperate” but widely naturalised.

The earliest record from this locality appears to be that of the Rev. Leonard Jenyns in his journal for 12
August 1824 (Crompton 1997): “Walked this morning with Profess. Henslow to Hinton nr Cambridge.
.... In Chalkpit Close — ... The great lettuce — Lactuca virosa, .... Of the Lettuce we could not find more
than one specimen though we made the most diligent search.”

The sheet bears a printed label “EX HERB. JAMES BACKHOUSE, JUN.” and a later stamp “Ex Herb.
S. H. Bickham” (1841-1933), and the specimen was identified as “Lactuca scariola” and “Lactuca
Scariola” (amended in pencil in the hand of S. M. Walters to “L. virosa’). Despite mentioning this
specimen, Perring et al. (1964) did not include TF/4.0 among the grid squares for L. virosa; nor was this
square listed by Crompton & Whitehouse (1983). Recently Peter Sell has determined this specimen as
“forma lactucarii (Lamotte) P. D. Sell” (see INTRODUCTION). He has determined Henslow’s specimen
from Cherry Hinton as forma virosa, but both of Babington’s as forma lactucarii.

“Shuckburgh Castle, Newmarket Heath’, on his own authority, “Grunty Fen’, attributed to William
Marshall, and “Ely”, attributed to Henslow. An additional record, “Burrell’s Walk, Cambridge’, is
attributed to Relhan.

After giving Hemsted’s locality, J. E. Smith wrote: “It grows among rubbish and on waste ground in
other parts of that country, but rarely elsewhere, ....” (Smith & Sowerby 1795, p. 268). Leonard Jenyns
clearly regarded L. serriola as a rarity; in his journal for 1 August 1827 he wrote: “... today made an
expedition to Grunty Fen. .... In my return — observed, & gathered with much satisfaction for the first
time in my life, the prickly Lettuce (Lactuca scariola) which grew somewhat sparingly on a high ditch-
bank edging the fen very near its entrance from Ely thro’ the lanes.” Jenyns’ ‘Journal of Natural
History’, covering the years 1823-1846, is now among the Alfred Newton papers in Cambridge
University Library. This entry was not included by Crompton (1997).

They are labelled “Streatham Cambridgeshire” (C. M. Lemann, 22.07.1831), “edge of Grunty-fen
Cambridgeshire” (J. S. Henslow, 27.08.1833), “Ely, Cambs” (J. S. Henslow, 27.08.1833), perhaps from
the same site, and “Roswell Pits, Ely” (J. E. Little, August 1878). The date of “the oldest specimen in
CGE” given by Perring ef al. (1964) — “from Ely in 1853” — must be a mistake for 1833; but the Stretham
specimen is older. The earliest Cambridgeshire specimen of forma integrifolia in CGE was collected at
Barnwell in September 1937 by H. Gilbert Carter and identified by him as “Lactuca Scariola’’.

Evans was here maintaining an error of Babington’s; see note 6 above.

The “pre-1930” and “not since 1949” records for grid square TL/5.7, in Perring et al. (1964) and in
Crompton & Whitehouse (1983) respectively, suggest that these authors were also perpetuating
Babington’s incorrect application of Ray’s (1660) locality near Ely to this species.

This date was presumably based on Babington’s specimen in CGE labelled “Chalk-pit, Cherry Hinton,
Cambridgeshire.”

Even if this was ever true, today it is frequent in Cambridgeshire. Crompton & Whitehouse (1983) wrote:
“Many new records yet it is reported to be declining.”

Not only does forma integrifolia of L. serriola have nearly simple leaves, but var. lactucarii of L. virosa
has pinnatisect cauline leaves (see INTRODUCTION).

(Accepted December 1999)

7?

cil

Watsonia 23: 161—165 (2000) 161

Calystegia (Convolvulaceae) hybrids in West Wales
R. K. BRUMMITT

The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AE

and
A. O. CHATER

Windover, Penyrangor, Aberystwyth, Ceredigion SY23 1BJ

ABSTRACT

Intensive studies of populations of Calystegia (Convolvulaceae) in Cardiganshire, v.c. 46, and Merioneth, v.c.
48, have recognised C. sepium subsp. sepium and subsp. roseata as native taxa there while C. silvatica subsp.
silvatica and subsp. disjuncta and C. pulchra are introduced and naturalised. Numerous probable hybrids
between these taxa are postulated, some as single populations and others apparently having spread locally
probably by vegetative means. Even back-crosses between adjacent populations may be postulated. In
particular, one putative hybrid between C. pulchra and C. sepium was recognised over an area of several
kilometres in Cardiganshire, resembling C. pulchra in general appearance but differing in having a paler pink
corolla and less markedly square leaf sinus.

KEYWORDS: Calystegia, Convolvulaceae, hybridisation, Wales.

INTRODUCTION

Hybrids among the native and naturalised species of Calystegia in the British Isles have long been
known (Stace 1961, 1975, 1997, Brummitt 1963, 1980, 1998). The whole genus, in which some 25
species world-wide may conveniently be recognised, is taxonomically difficult, and few if any of
the species are morphologically clear-cut. They mostly vary considerably over their ranges and
merge geographically one into another, and division into species and subspecies is of necessity
somewhat arbitrary. In the British Isles, apart from C. soldanella (L.) R.Br. which is not
considered in this note, there are one native and two naturalised species.

The native C. sepium (L.) R. Br. is represented by the widespread typical subsp. sepium which is
white-flowered (rarely pink-flowered, f. colorata (Lange) Dorfl.) and glabrous, and the
predominantly western coastal subsp. roseata Brummitt which is pink-flowered and usually hairy
at the shoot apices and leaf sinuses. In addition, one further subspecies, subsp. spectabilis
Brummitt, has been recorded once as an introduction. The commoner of the two naturalised
species, C. silvatica (Kit.) Griseb., is represented here by the typical subsp. si/vatica from the east
Mediterranean, with broad, saccate bracteoles rounded or emarginate at the apex, and subsp.
disjuncta Brummitt from the west Mediterranean, with narrower, less saccate bracteoles, obtusely
pointed at the apex (Brummitt 1996, 1998). Rather less common, though widely distributed, is C.
pulchra Brummitt & Heywood, of unknown origin, with large pink flowers, hairy stems, petioles
and pedicels, and often winged peduncles.

The following notes are based on fieldwork and specimens collected by both of us in two
periods of several days in Cardiganshire, v.c. 46, and Merioneth, v.c. 48, in July 1996 and August
1998, and on various collections by A.O.C. in the former county over several years. Nearly all the
recorded British taxa occur in this area, and much of the variation we found seemed likely to be
the result of hybridisation. Although in the absence of experimental evidence we are unable to
confirm most of our interpretations, we feel that it is worth putting our observations on record in
the hope that they may be a stimulus to comparable observations being made in other parts of the
country so that a more complete picture of the state of the genus in Britain can be built up. Details

of the relevant specimens are given. Collections of all taxa made by us are deposited at NMW or
K or both.

162 R. K. BRUMMITT AND A. O CHATER

FIELD OBSERVATIONS

C. sepium subsp. sepium was widespread in the area, and the chief variation noted was that in
several coastal localities plants occurred with distinctly hairy stem apices. This is unusual for this
taxon, and the possibility that the hairiness is due to introgression from subsp. roseata must be
borne in mind. Subsp. roseata occurred in several coastal and estuarine sites, appearing completely
native in reedbeds in both the Dyfi and Mawddach estuaries. It also occurred in coastal hedges,
and at one site 18 km inland in Salix carr by the disused railway at Cors Caron where it may have
been an accidental introduction. A specimen collected in 1961 by P. M. Benoit from the railings of
a waste garden at Bron Fegla near Arthog in Merioneth, now at LIV, was identified by R.K.B. as
subsp. spectabilis, a taxon widely naturalised in Scandinavia, and is the basis for the inclusion of
this subspecies in Stace (1997). The pink flowers and coastal habitat would fit well with that of
subsp. roseata, but the large size of the flowers and the broad leaf sinus suggest that it is more
likely to have been a garden introduction. Searches for the plant in 1998 by both Benoit and
ourselves were unsuccessful, and we assume that it is now extinct at the locality.

Among the populations of C. silvatica examined, 30 were considered to be clearly subsp.
disjuncta, 23 subsp. silvatica and 5 were intermediate. 13 plants were assumed to be hybrids
between C. sepium and C. silvatica. The name C. x lucana (Ten.) G. Don is available for this
interspecific cross, but since there is a wide range of morphological forms resulting from the cross,
R.K.B. prefers not to apply a binomial (see Brummitt 1998 and further notes below). The putative
hybrid Cardiganshire populations varied considerably in morphology of the bracteoles, and in a
few cases this was extreme enough to suggest whether it was subsp. si/vatica or subsp. disjuncta
that was involved. A preponderance of hybrids among collections made late in the season suggests
that they may continue flowering later into the autumn than the parents.

C. pulchra appeared fairly uniform in West Wales, with deep pink flowers, notably square leaf
sinuses and large rounded bracteoles. However, in the lower Teifi valley, in an area bounded by
Cardigan, Penpare and Cenarth, and particularly conspicuous at intervals along the A484 road, we
found plants (some of which had been previously recorded as C. pulchra) with paler pink flowers,
more rounded leaf sinuses and somewhat irregularly shaped bracteoles. Morphologically they
appeared to be intermediate between C. pulchra and C. sepium subsp. sepium, which was frequent
in the same area, and we assumed that they were a hybrid between these two taxa. The name C. x
scanica Brummitt is available for this interspecific cross, but the plant to which this name was
originally applied, naturalised in several places in southern Sweden, is very different, having white
flowers, bracteoles less strongly overlapping, and large leaves with a very characteristic shape.
The differences between these nothomorphs is at least as great as the difference between the parent
species, and again R.K.B. prefers not to refer to “the hybrid” with a binomial but regards these
Cardiganshire populations as “a hybrid” presumably between C. pulchra and C. sepium. At one
roadside hedge and scrub site near Llandygwydd where both this hybrid (Brummitt 20058 &
Chater) and C. sepium subsp. sepium (Brummitt 20057A & Chater) occurred, we found a white-
flowered plant similar to C. silvatica subsp. disjuncta, but with very pointed bracteoles and
narrowly pointed leaves (Brummitt 20057B & Chater); morphologically this looked very likely to
be a back cross between the putative C. pulchra/sepium hybrid and C. sepium subsp. sepium.

Near Aber-porth we found a single plant or small population in a roadside hedge (Brummitt
19513, 20054 & Chater) rather like Brummitt 20057B & Chater. The corolla was white or very
slightly flushed pink inside, and with pink mid-petaline bands outside like those often found on C.
silvatica subsp. disjuncta, with the dull green, matt leaves of C. pulchra and some stem apices
glabrous and others hairy. It too may have been a back cross between the C. pulchra/sepium
hybrid and C. sepium subsp. sepium, or possibly a hairy form of C. silvatica subsp. disjuncta
showing some introgression from C. pulchra.

Near Newcastle Emlyn and at Llandysul, 8 and 15 km further up the Teifi valley respectively,
plants with white flowers and grotesquely misshapen bracteoles occurred (Brummitt 20064A,
20064B, 20065 & Chater) which appeared to be a white-flowered nothomorph also resulting from
hybridisation between C. pulchra and C. sepium. At Llandysul these plants grew in close
association with the putative parents. Further north in Cardiganshire, in a roadside hedge at Ffos-y-
ffin, was a mixed population of C. pulchra (Brummitt 20066B & Chater) and white-flowered
plants. Some of the latter were glabrous and some had pubescent stem apices, while the bracteoles

CALYSTEGIA HYBRIDS IN WEST WALES 163

were very variable though not grotesquely enlarged and distorted as in the Llandysul and
Newcastle Emlyn plants. They appeared to us to be a mixture of C. sepium subsp. sepium
(Brummitt 20067A & Chater) and a white-flowered hybrid between C. pulchra and C. sepium
(Brummitt 20067B & Chater).

Close to Morfa Mawddach Station, near Arthog in Merioneth, we noted three populations in
subcoastal habitats typical of where C. sepium subsp. roseata grows. One was white-flowered and
was Clearly introduced C. silvatica subsp. disjuncta (Brummitt 20051 & Chater). The others were
pink-flowered, one (20050A) growing among brambles together with the white flowered 20051,
and the other 50 m away climbing up Phragmites (20050B). The latter in particular was in typical
habitat of subsp. roseata and looked very like it, but our suspicions were raised by the unusually
broad bracteoles and flowers rather large for C. sepium. There is a possibility that both the pink-
flowered plants were hybrids of C. sepium subsp. roseata with C. silvatica subsp. disjuncta. On
the other hand, the leaf sinus is narrowly V-shaped, and rather large bracteoles like these may be
found in subsp. roseata more often than in subsp. sepium, and hybridisation is by no means
proven.

In the genus as a whole, abnormal bracteoles appear often to be an indication of hybridity, and
perhaps a quarter of all putative hybrids have the bracteoles of some flowers variously unequal in
size or shape, more or less leaflike, not opposite, or puckered or distorted.

DISCUSSION

The pattern emerging from our fairly intensive survey of the genus in one area of Wales probably
reflects the situation over much of northern Europe. Different taxa have been introduced, probably
originally as garden plants, and have become established in the wild through their vigorous
vegetative reproduction. They have then hybridised with native taxa and other introduced taxa. In
some cases these hybrids may be restricted to single populations, but in a few cases they appear to
have spread - probably by vegetative means, although little is really known about how the
rhizomes are moved from one locality to another - and one clone may be recognised in numerous
populations over one area. The putative hybrid between C. pulchra and C. sepium which we saw
frequently in different colonies over an area of several kilometres in Cardiganshire is probably as
morphologically distinct and almost as widely distributed as the putative hybrid between C.
silvatica and C. pulchra which is well established at different localities in eastern Nottinghamshire
and adjacent Lincolnshire and which was named as C. howittiorum by Brummitt (1980). There
should probably be a limit, however, to the number of such clones which are given formal names,
and we decline to describe this Cardiganshire hybrid as a new taxon. The uncertainty about the
actual parentage of such plants is an additional reason for not applying formal binomial
nomenclature to putative hybrids, at least until much more is known about their origins and
reproductive behaviour.

SELECTED HERBARIUM SPECIMENS (ALL DET. R. K. BRUMMITT).

C. SEPIUM SUBSP. SEPIUM x C. SILVATICA

v.c. 46, Cards., among Phragmites at N end of boardwalk just E of A467 bypass, Teifi Marshes
Reserve, Cardigan, altitude 4 m (SN/182.458), 22 September 1995, A. O. Chater (NMW).

v.c. 46, Cards., among Fallopia japonica on waste ground, NW corner of Glanyrafon Industrial
Estate, Llanbadarn Fawr, 7 m (SN/608.803), 7 July 1995, A. O. Chater (NMW). C. sepium
subsp. sepium x C. silvatica subsp. silvatica.

v.c. 46, Cards., upper part of salt marsh 100 m S of Isolation Flats, Ystwyth estuary, 4 m
(SN/580.805), 7 September 1995, A. O. Chater (NMW).

v.c. 46, Cards., abundant on shingle beach opposite mouth of Aberystwyth harbour, 3 m
(SN/580.803), 11 September 1995, A. O. Chater (LTR, NMW). 22 July 1996, R. K. Brummitt
19505 & A. O. Chater (K). C. sepium subsp. sepium x C. silvatica subsp. disjuncta.

v.c. 46, Cards., scrub by road bridge E of Rhoscellan-fawr, 1-5 km N of Llangorwen, 70 m
(SN/599.854), 12 September 1995, A. O. Chater (NMW).

v.c. 46, Cards., N hedge of A487 at Penffynnon, Penparc, 135 m (SN/217.481), 13 September
1995, A.O. Chater (NMW).

164 R. K. BRUMMITT AND A. O CHATER

v.c. 46, Cards., NW hedge of road 500 m WSW of Brynllynan,Y Ferwig, 95 m (SN/185.484), 13
September 1995, A. O. Chater (NMW).

v.c. 46, Cards., forming dense growth over rank grass by road and ditch adjacent to refuse tip, 300
m S of Borth church, 4 m (SN/611.894), 25 July 1996, A. O. Chater (NMW); and ibid. 12
August 1998, R. K. Brummitt 20049B & C & A. O. Chater (K, NMW).

v.c. 46, Cards., abundant by track in conifer plantation, Long Wood, 1-4 km S of Llangybi, 210 m
(SN/610.515), 1 August 1996, A. O. Chater (NMW). C. sepium subsp. sepium x C. silvatica
subsp. disjuncta.

v.c. 46, Cards., in scrub on sandy soil 300 m from coast, 100 m W of Ty-gwyn, Mwnt, 30 m
(SN/197.518), 15 August 1996, A. O. Chater (NMW).

v.c. 46, Cards., roadside hedgebank 600 m S of Temple Bar crossroads, 170 m (SN/533.537), 21
August 1996, A. O. Chater (NMW).

v.c. 46, Cards., in hedge and rank marsh 100 m NW of Pentre, Llundain-fach, Abermeurig, 75 m
(SN/559.567), 8 September 1996, A. O. Chater (NMW).

v.c. 46, Cards., scrub at edge of wood 200 m S of Aberaeron church, 35 m (SN/456.626), 2 July
1997, A. O. Chater (NMW).

v.c. 46, Cards., hedge on N side of garden, Wallog house, 10 m (SN/590.857), 3 October 1998, A.
O. Chater (KK).

POSSIBLY C. SEPIUM SUBSP. ROSEATA x C. SILVATICA SUBSP. DISJUNCTA

v.c. 48, Merioneth., among brambles 150 m SE of Morfa Mawddach Station, Arthog, 5m
(SN/6297.1401), 12 August 1998, R.K.Brummitt 20050A & A.O. Chater (K, NMW).

v.c. 48, Merioneth.,On Phragmites 100m SSE of Morfa Mawddach Station, 5m (SN/6290.1405),
12 August 1998, R.K. Brummitt 20050B & A.O. Chater (K, NMW).

C. PULCHRA x SEPIUM, PALE PINK NOTHOMORPH

v.c. 46, Cards., hedge and scrub on N side of B4570/A484 road junction, 800 m E of Cardigan
church, 5 m (SN/189.460),13 August 1998, R. K. Brummitt 20055 & A. O. Chater (K, NMW).

v.c. 46, Cards., hedge on S side of A484 road by Llechryd waterworks, 10 m (SN/230.435), 13
August 1998, R. K. Brummitt 20056 & A. O. Chater (K, NMW),.

v.c. 46, Cards., scrub on S side of A484 road 800 m SE of Llwynduris, 12 m (SN/244.427), 13
August 1998, R. K. Brummitt 20058 & A. O. Chater (K, NMW).

v.c. 46, Cards., S hedge of A487 road 150 m WSW of crossroads in Penparc village, 130 m
(SN/211.479), 19 August 1998, A. O. Chater (K, NMW).

C. PULCHRA x SEPIUM, WHITE NOTHOMORPHS

v.c. 46, Cards., hedgebank on SW side of road T-junction 600 m SW of Aber-porth church, 111 m
(SN/250.507), 23 July 1996, R. K. Brummitt 19513 & A. O. Chater (K, NMW). 13 August
1998, R. K. Brummitt 20053 & A. O. Chater (K, NMW). Possibly a back cross between C.
pulchra x sepium and C. sepium subsp. sepium.

v.c. 46, Cards., scrub by lay-by at bottom of wooded slope on N side of A484 road 800 m SE of
Llwynduris, 12 m (SN/244.427), 13 August 1998, R. K. Brummitt 20057B & A. O. Chater (K,
NMW). Probably a back cross between C. pulchra x sepium and C. sepium subsp. sepium.

v.c. 46, Cards., scrub at NE corner of Rugby Club car park, 500 m E of Newcastle Emlyn bridge,
25 m (SN/314.409), 13 August 1998, R. K. Brummitt 20061 & A. O. Chater (K, NMW).

v.c. 46, Cards., rank vegetation and wall between S part of Llandysul churchyard and Teifi river,
60 m (SN/419.406), 13 July 1998, R. K. Brummitt 20064B & A. O. Chater (K, NMW).

v.c. 46, Cards., scrub by gardens 150 m N of Llandysul church, 70 m (SN/418.408), 13 August
1998, R. K. Brummitt 20065 & A. O. Chater (K, NMW).

v.c. 46, Cards., W hedge of A487 road 350 m SSE of Henfynyw church, Ffos-y-ffin, 115 m
(SN/449.609), 13 August 1998, R. K. Brummitt 20067B & A. O. Chater (K, NMW).

REFERENCES

BRUMMITT, R. K. (1963). A taxonomic revision of the genus Calystegia. Ph. D. thesis, University of
Liverpool.

BRUMMITT, R. K. (1980). Further new names in the genus Calystegia (Convolvulaceae). Kew Bull. 35: 327—
334.

CALYSTEGIA HYBRIDS IN WEST WALES 165

BRUMMITT, R. K. (1996). Two subspecies of Calystegia silvatica (Kit.) Griseb. (Convolvulaceae) in the
Mediterranean region. Lagascalia 18: 338-340.

BRUMMITT, R. K. (1998). Calystegia. In RICH, T. C. G. & JERMY, A. C., eds. Plant crib 1998. Botanical
Society of the British Isles, London.

STACE, C. A. (1961). Some studies in Calystegia: compatibility and hybridisation in C. sepium and C.
silvatica. Watsonia 5: 88-105.

STACE, C. A. (1975). Hybridisation and the flora of the British Isles. Academic Press, London etc.

STACE, C. A. (1997). New Flora of the British Isles, 2nd ed. Cambridge University Press, Cambridge.

(Accepted October 1999)

ate

é

Watsonia 23: 167—172 (2000) 167

Ludwigia x kentiana E. J. Clement: a new hybrid aquatic
E. J. CLEMENT

54 Anglesey Road, Alverstoke, Gosport, Hampshire, PO12 2EQ

ABSTRACT

A hybrid Ludwigia L. (Onagraceae) found wild in England is described and illustrated: its morphology
suggests a parentage of L. palustris (L.) Elliott x L. repens J. R. Forst., and it probably equates to a plant
known to aquaria enthusiasts as L. x mullertii hort.

KEYWORDS: hybridisation, vegetative spread, water weed, Onagraceae, Hampshire-purslane.

INTRODUCTION

In September 1995 U. Sutcliffe & Ms A. Sutcliffe found in a round pond on Putney Heath (GR
TQ/237.733), Surrey, v.c. 17, England, a waterweed that was identified as Ludwigia palustris.
Subsequently when Mrs E. Norman revisited the locality, material was sent to me for confirmation
and I realised that it differed appreciably from our native species. It was noticeably large and
vigorous, and was probably equatable to a plant known by aquarists as L. x mullertii hort. (with
orthographic variants ranging from L. x mulerttii to L. X muellertii), with a probable parentage of
L. palustris (L.) Elliott x L. repens J. R. Forst. A request for further information contained in a
preliminary report (Clement 1997) yielded nothing. I have still not found a botanical description of
this taxon nor a validation of the name, nor any other matching taxon, hence my decision to
publish herein a new and unambiguous epithet.

This hybrid has not yet been definitely confirmed from a second locality, but further records are
expected. Any vigorous Ludwigia colony should be sampled after permission to collect has been
obtained. The lack of preservation of good, herbarium vouchers has hampered this research paper
and it is becoming a recurrent problem in alien plant studies in Britain. The conservation
movement has discouraged the collection of all natural history material: photographs, alone, being
acceptable - hence there is a dramatic decrease in the collection of herbarium material.

THE PROBABLE PARENT SPECIES

The taxonomy of the two probable parents presents no problems, but their nomenclature calls for
some explanation, as considerable confusion exists in the literature.

L. palustris (L.) Elliott (published in 1817) Usnardia palustris L.) has, fortunately, never been a
serious nomenclatural or taxonomic problem. At least two varieties have been recognised - e.g.
Munz (1961) - but none of them seems worthy of higher status. The statement therein (Munz op.
cit. p. 215) that in contradistinction to the American forms “the green bands of the hypanthium
terminate well below the summit’in the European form seems untrue (see Fig. 1). The word
“hypanthium” has, incidentally, different definitions in different books: Stace (1997) says that the
hypanthium is absent in this genus, but his use of the term “hypanthium” (of an epigynous flower)
apparently refers solely to that part of the extension of the receptacle beyond the summit of the
ovary. (His glossary fails to make this fact clear).

L. palustris is a widespread native in the northern hemisphere, occurring in W., C. and S. Europe
(north as far as New Forest, England), N. Africa, W. Asia, N. and C. America and W. Indies. It is
naturalised in southern Africa, S. America, the Pacific area, Hawaii, S.E. Australia and New
Zealand. Illustrations may be found in Fig. 1, and in many British and foreign Floras.

L. repens J. R. Forst. (Isnardia repens (Sw.) DC.) presents more problems. It was published in
Forster (1771) with a query (?) before the specific name - and some authors appear to have

168 E. J/CLEMEN®

rejected the epithet as being “not accepted by the author” (and hence an invalid name), but Article
34.1 in Greuter et al. (1994) makes it very clear that this is not so. The description seems adequate
to define the taxon, the question mark indicating merely that Forster was unsure whether his plant
was a new species or not. I have not searched for a type specimen: the statement by Stafleu &
Cowan (1976) “it is not possible to state where the original Forster herbarium (father and son) is
preserved” acted as a deterrent. The entry in Jackson (1893) has also mislead many of the unwary:
it falsely equates this species to L. palustris as well as misquoting the page number in Forster’s
work: it should be p. 6 (not 22). This is especially annoying, since this book contains no index of
page numbers even for generic names! The homonym L. repens Sw. (published in 1797) is
synonymous, whereas Jussiaea repens L. (1753) is only distantly related (see synonyms below
after the key). Finally, the name L. natans Elliott (published in 1821) does indeed refer to this
plant, and is unambiguous, but is, alas, a later synonym that should not be used, in spite of the
recent choice of it by e.g. Garve & Meijden (1997) and Clement (1997).

The species is restricted to S. and S.E. United States, Mexico and the West Indies. It is
naturalised in S. Asia and Japan, also recently in Europe e.g. in Spain (Nieto Feliner 1997).
Illustrations may be found in, e.g. Britton (1918), Fawcett & Rendle (1926) and Radford et al.
(1968).

DESCRIPTION OF THE HYBRID

To validate the new epithet proposed herein, a formal diagnosis is required:

Ludwigia x kentiana E. J. Clement, hybr. nov.

Hybrida probabiliter inter Ludwigiam palustrem (L.) Elliott et L. repentem J. R. Forst., ab eis
speciebus fructibus sterilibus, petalis circa 0-5 mm longis differt.

HOLOTYPUS: England, Surrey (v.c. 17), Putney Heath, Round Pond, 6 October 1997. Mrs E.
Norman, s.n. (BM); isotypus in Herb. EJC. A glabrous perennial with stems 20-80 cm, prostrate
or weakly ascending, often largely or entirely submerged in water, abundantly rooting at lower
nodes. Leaves 20-50 x 5-25 mm, opposite, the blade narrowly rhombic-obovate, gradually
narrowed to a long petiole, with the margin entire. Flowers solitary in leaf axils, + sessile.
Bracteoles linear, c. | mm. Calyx-tube + cylindrical, 4-sided, uniformly pale green; calyx teeth 4,
c. 2 mm, deltoid. Petals 4, minute (c. 0-5 mm) cream, + spathulate, fugacious (usually dropping off
as soon as the buds open); stamens 4, 0-8 mm long; style 0-8 mm long with subglobose stigma
divided into 4 indistinct lobes. Fruit a capsule, partially developing and then dropping off, the
ovules not expanding.

The epithet kentiana is for D. H. Kent in appreciation of his great assistance to me for over 30
years during my studies on the adventive flora of Britain. The aquarium trade may be disappointed
that I have not upheld the name that they currently use. The epithet that I have chosen is intended
to cover all variations of the hybrid, some undoubtedly not worthy of any aquarium tank. The best
clone (preferably the original, correct one!) can be more accurately named as L. x kentiana
‘Mullertii’, which may be legitimately abbreviated to L. “Mullertii’ - the simple quotes being the
current convention to indicate a cultivar (i.e. equal to cv. Muellertii in older literature). At present,
I refrain from allocating the clone described here as true L. ‘Mullertii’. An English name is also
called for - Kent’s Hampshire-Purslane seems appropriate.

TABLE 1. COMPARISON OF LUDWIGIA PALUSTRIS, L x KENTIANA AND L. REPENS

L. palustris L. x kentiana L. repens
Leaf blade Widest near its middle Widest in uppermost third Widest close to its apex
Bracteoles 0-0-5 mm, free c. 1mm 1—2 mm, shortly adnate to ovary
Sepal lobes — About as long as broad Longer than broad Much longer than broad
Petals Absent 0-5 mm, cream 3-5 mm, yellow
Fruit Subrotund and persistent; +Cylindrical and caducous; — Cylindrical and persistent;

with 4 dark green bands of | uniformly pale green uniformly greenish yellow

LUDWIGIA x KENTIANA 169

= Imm

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=
©
2
=
©
3)
vn

FIGURE 1. Ludwigia x kentiana (A-G) and L. palustris (a—g) del. D. J. P. Smith © 1998.
A,a. habit of plant; B,b. leaf from middle of stem; C,c. side view of immature fruit with persistent calyx teeth;

D,d. transverse section of immature fruit showing ovules; E,e. view of flower from above; F,f. stamen; G,g
stigma and style.

170 E. J“ CLEMENT

L. x kentiana can be distinguished from its probable parents by the characters shown in Table 1.

Several North American Floras e.g. Fernald (1987) claim that L. repens has no petals - the
virtually invariable state in herbaria since the petals drop so quickly - but there seems to be no
truth in this statement (Godfrey & Wooten 1981). I notice that Radford er al. (1968) claim “petals
absent” in their description, but their wiser key says “petals present”. Clement (1997) claimed (1—)
4 petals for the hybrid, but a more careful study of fresh specimens revealed an invariable number
of just 4. The illustration by D. J. P. Smith (Fig. 1) clearly shows these differences between L.
palustris and L. x kentiana; in addition other minutiae may be noted, though less easily put into
words, including the shape of the anthers. The provenance of the plants drawn are, respectively,
near Hatchet Pond, New Forest, (v.c. 11, S. Hants) and Putney Heath.

D. J. P. Smith also drew my attention to the fact that two of the opposing anthers, in both taxa,
are noticeably larger than the other two - two of them may be better described as staminodes, a fact
that I have not seen mentioned elsewhere.

KEY TO LUDWIGIA SPECIES OCCURRING IN EUROPE

The following key, partly based on Raven (1963), is provided to alert readers to the possibility of
other species occurring in Britain in the future. All are typically perennial aquatics, although
annuals, shrubs, and even a tree occur within the genus. Only L. palustris is native in Europe and it
shows a markedly southern distribution.

l. Leaves opposite; flowers + sessile; petals 4, 0-4 mim........2......cossncseateneeatee serene eet: «a8 2.
I. Leaves alternate; flowers pedicelled; petals 5(—6), 7—30 mmm ....:5. cc.steecncee ee ee +
2. POtals aS Gti assis casas dcactevscccdistheacchcccaduidcasadennssigudelecealuatee een epee. eee eee L. palustris
Dis Petals present; DUt LUSACIOUS 2..isscs.vecceeosenessnncectsnennsbonsicncedael copeagetecBicce sete eee eeeeeet ee eee: 2
3: Petals Cy 0:5 dy CHEB IN 5a0u5iedeaasspnwsvnvaaessvauewatinntende chaveeiees oReece cutee caer eee L. x kentiana
Zi Petals: 35 titi; VOUOW sccscssseccapcassabssonncswcesuedaseiea doesaes Uegmodouinsndessec eee tea emene tment L. repens
4. Petals white or pale yellow, with a dark yellow claw ..............cccseescecceeeeeeeeeeees L. adscendens
4. Petals bright yellow throughout .....:::scccccssesssslececeseseachiesaseeeseccnasasoarteeteaneae seemeeeeeenen eee meee ese 5
De Calyx-tube finely hairy to glabrous; petals 7—16 mm ...............sss,ccsssa.deteneeeereres L. peploides
oj Calyx-tube WitsUte icsccsaccccssecsinsasxcaacueavenxassiadandedusiudaccatineccisvsteeesgd eee ee eee 6
6. Stem_and leaves pubescent; petals 15—24 mom ....2.c.006.....46.0s0sedected tee eee L. grandiflora
6. Stem and leaves = glabrous; petals 7—15 Mh cssccrcesnccssas iced L. hexapetala

The four additional species not discussed above are:

Ludwigia adscendens (L.) Hara (Jussiaea repens L.)

S. and S.E. Asia and tropical Australia. (a casual? in The Netherlands). Illustrations may be found
in, e.g. Soerjani (1987).

L. peploides (Kunth) Raven

Tropical and subtropical N. and S. America, and perhaps native in Australia. Naturalised in S.W.
France and New Zealand. A variable plant, sometimes divided into four subspecies. Illustrations
may be found in, e.g. Godfrey & Waoten (1981) and Hoch (1993).

L. grandiflora (Michx) W. Greuter & Burdet

(L. uruguayensis (Camb.) Hara; Jussiaea repens sensu Coste, non L.)

Pantropical and subtropical. Naturalised in S. France, Spain, The Netherlands and Belgium.
Illustrations may be found in, e.g. Coste (1903) and Godfrey & Wooten (1981).

L. hexapetala (Hook. & Arn.) Zardini, Gu & Raven

Tropical America. Records exist for S. France and E. Spain, but they are probably referable to the
closely allied L. grandiflora. Some authors, such as Hoch (1993), provide an illustration but
clearly do not separate these two species even as varieties.

Other potential escapees are also grown in aquaria - e.g. L. arcuata Walter x L. repens
(Kasselmann 1985). The first parent has flowers on long pedicels 15-35 mm long, and this
influence would presumably be evident in any hybrid that it forms.

LUDWIGIA X KENTIANA 171
DISTRIBUTION OF THE HYBRID

The hybrid L. x kentiana is probably widespread in aquaria in Britain and elsewhere, but no
previous record of its escape into a wild location has come to my notice: it may well have arisen in
an aquarium in Europe. Indeed, since both parents appear to be invariably self-pollinating in the
wild, with the anthers and stigmas in juxtaposition and the consequent 100% set of fruits, the
hybrid may have been deliberately made by human manipulation. The parents overlap in their
native range only in S. and S.E. United States and the West Indies, but I can trace no mention of a
hybrid in the N. American literature.

It is possible, however, that one (or more) of the current records for L. repens in Europe are
better incorporated within L. x kentiana, even though a different clone may be involved and so not
perfectly matching the description given herein.

In England, I have seen and confirmed only the Putney Heath record, but the following are likely
to be either the same taxon or possibly pure L. repens. The voucher for v.c. 59 in BM possesses
only flower buds: it is certainly not pure L. palustris, as labelled. The earliest British record
appears to be in 1927 (from v.c. 59).

v.c. 11 (S. Hants). Southampton, Sholing, Miller's Pond, GR SU/45.10, 1958-1965, R. P.
Bowman. Probably an aquarist’s introduction; pond now infilled.

v.c. 14 (E. Sussex). Seaford Head, South Hill Barn, GR SZ/505.980, 31 August 1991, P. D. L.
Maurice. In a dew pond.

v.c. 16 (W. Kent). Tonbridge, GR TQ/5.4, 1989-1990, S. Melville (MNE). In a garden pond, but
not deliberately planted.

v.c. 18 (S. Essex). Earls Path Pond, Epping Forest, GR TQ/415.967, 3 August 1976, K. J. Adams
& J. O. Mountford. In an acidic pond.

v.c. 59 (S. Lancs). Manchester Docks, GR SJ/8.9, 1927, L. Adams.(?LTN). By Failsworth Canal,
GR SD/8.0, 14 May 1960, Miss V. Gordon (LIV); ibid., 26 July 1960, J. E. Lousley & Rev. C.
E. Shaw (BM). Plentiful for several years.

A map showing all Ludwigia records for the British Isles is given by Preston & Croft (1997), but
all the above colonies, except Seaford Head, are probably now extinct.

Surprisingly, the v.c. 9 (Dorset) record predicted by Clement (1997) to be this alien has since
been determined by D. Pearman & myself as pure (and presumably native) L. palustris: it has been
seen in 1996-1998 in a newly dug-out old pond at Sutton Holms, Edmonsham (Pearman 1999),

DISCUSSION

Conservationists may well be alarmed to hear that yet another vigorous aquatic plant is threatening
to oust our native plants from almost any pond, particularly if acidic. The history of this hybrid, to
date, suggests that it does not jump from site to site as readily as Crassula helmsii (Kirk)
Cockayne and also that extreme cold weather appears to exterminate it completely - but a wary eye
on its progress is necessary. Indeed, according to Salisbury (1972), even our native L. palustris
“probably ...often behaves as an annual and ‘perennates’ perhaps only when the winter is not
severe’. The ability of the hybrid Ludwigia to climb up and over other low vegetation in ponds is a
little worrying, while the tendency for warmer summers will certainly favour its spread.

More than one clone of the hybrid may have occurred in Britain, each colony spreading solely
by vegetative propagation, and with a probable variation in hardiness (and hence its potential
threat). A glance at Clement & Foster (1994) reveals that this is the first alien Ludwigia to be
found in the British Isles, but others seem likely to follow: worldwide, 82 species are currently
recognised (Raven 1963) and many of them have weedy tendencies.

ACKNOWLEDGMENTS

I am grateful to Prof. C. A. Stace for advice and for providing me with the Latin diagnosis. Dr C.
D. Preston kindly provided me with information on all Ludwigia alien records in Britain.

172 E.32.CLEMENT

Mrs E. Norman and P. Stanley enabled D. J. P. Smith to observe material in peak, fresh
condition for his beautiful artwork presented here. Additional help has been freely given by R. P.
Bowman, Miss V. Gordon, D. Pearman, Miss P. Miles, M. Mullin, A. Underhill and B. Wurzell.

REFERENCES

BRITTON, N. L. (1918). Flora of Bermuda. Charles Scribner’s Sons, New York:

CLEMENT, E. J. & FOSTER, M. C. (1994). Alien plants of the British Isles. Botanical Society of the British
Isles, London.

CLEMENT, E. J. (1997). Ludwigia x muellertii Hort. new to Britain. BSBI News 77; 54.

CosTE, H. (1903). Flore de la France 2: 83. Paul Klincksieck, Paris.

FERNALD, M. L. (1987). Gray’s Manual of Botany 2: 1055. Dioscorides Press, Portland, Oregon.

FORSTER, J. R. (1771). Flora Americae septentrionalis. B. White & T. Davies, London.

GARVE, E. & MEIJDEN, R. VAN DE (1997). Ludwigia natans Elliott bei Hannover sowie hinweise zur
bestimmung adventiver Ludwigia-arten. Floristische Rundbriefe 31(1): 9-12.

GODFREY, R. K. & WOOTEN, J. W. (1981). Aguatic and wetland plants of southeastern United States.
Dicotyledons. University of Georgia Press, Athens.

GREUTER, W. et al. (1994). International code of botanical nomenclature (Tokyo code). Regnum Vegetabile
131.

Hoc, P. C. (1993). Ludwigia in HICKMAN, J. C., ed. The Jepson Manual, pp. 800 & 807. University of
California Press, Berkeley.

JACKSON, B. D. (1893). Index kewensis 2: 123. Clarendon Press, Oxford.

KASSELMANN, C. (1985). Viele jahre in Kultur, erst jetzt erkannt: Ludwigia repens Forster x arcuata Walter.
Aguarien - und Terrarien - Zeitschrift (DATZ) 38(8): 377-380 (Not seen).

Munz, P. A. (1961). Ludwigia L. in LUNDELL, C. L. et al. Flora of Texas 3: 214-220. Texas Research
Foundation, Renner.

NIETO FELINER, G., ed. (1997). Ludwigia L. in CASTROVIEJO S. et al. Flora iberica 8: 87-90. Real Jardin
Botanico, Madrid.

PEARMAN, D. (1999). Ludwigia palustris in Dorset. BSBI News 80: 17-18.

PRESTON, C. D. & CROFT, J. M. (1997). Aquatic plants in Britain and Ireland, pp.107—109. Harley Books,
Colchester.

RADFORD, A. E. et al. (1968). Manual of the vascular flora of the Carolinas. University of North Carolina
Press, Chapel Hill.

RAVEN, P. H. (1963). The Old World species of Ludwigia (including Jussiaea), with a synopsis of the genus
(Onagraceae). Reinwardtia 6: 327-427.

RAVEN, P. H. (1977). Onagraceae. Flora Malesiana series I, 8(2): 98-113.

SALISBURY, E. J. (1972). Ludwigia palustris (L.) Ell. in England with special reference to its dispersal and
germination. Watsonia 9: 33-37.

SOERJANI, M. ef al., eds. (1987). Weeds of rice in Indonesia. Balai Pustaka, Jakarta.

STACE, C. A. (1997). New Flora of the British Isles, 2nd ed., p. 449. Cambridge University Press, Cambridge.

STAFLEU, F. A. & COWAN, R. S. (1976). Taxonomic literature, 2nd ed., 1: 857. Bohn, Scheltema & Holkema,
Utrecht.

(Accepted November 1999)

Watsonia 23: 173-177 (2000) 3

A new section and species of Festuca (Poaceae)
naturalized in England

R. FLETCHER and C. A. STACE

Department of Biology, University of Leicester, Leicester, LE1 7RH

ABSTRACT

An introduced species of Festuca, well naturalised in a limestone quarry in N.W. Yorkshire for at least 70
years, is identified as F. gautieri (Hack.) K. Richt. subsp. scoparia (A. Kern. & Hack.) Kerguélen. The same
taxon occurred on a roadside in S. Devon. A description and chromosome counts are given, and the origin of
the plants is discussed.

KEYWORDS: Fescue, grass.

INTRODUCTION

British species of Festuca L. are usually placed in three sections: Bovinae Hack. (F. pratensis
Huds., F. arundinacea Schreb. and F. gigantea (L.) Vill.), Montanae Hack. (F. altissima All.) and
Festuca (Ovinae Hack.; the F. rubra L. and F. ovina L. aggregates). Species of section Eskia
Willk. (Variae Hack.) resemble those of sect. Festuca in being relatively small plants with fine
leaves, but as natives they do not approach Britain closer than the Jura, Alps and Pyrenees.

Herbarium material (well past flowering) of an unknown Festuca was sent from N.W. Yorkshire
to C.A.S. in 1993 by Mr R. M. Payne, and living material as well as fully flowering herbarium
material of the same plant by Mrs D. Millward in 1994. This was studied at Leicester in 1994 and
1995 by R.F. and C.A.S., and identified as F. gautieri (Hack.) K. Richt. subsp. scoparia (A. Kern.
& Hack.) Kerguélen, a member of section Eskia from the Pyrenees. An introduced Festuca
discovered in S. Devon in 1990 by Mr R. Takagi-Arigho had been confirmed by Dr T. A. Cope as
F. gautieri, and herbarium material (well before flowering) was sent to Leicester in 1994. Living
material was sent to C.A.S. in 1994 by Mr L. J. Margetts. This was also identified as F. gautieri
subsp. scoparia. The Yorkshire site was also visited by R.F.

MATERIAL

As well as the living and herbarium material from the two English localities, herbarium material of
F. gautieri subsp. gautieri and subsp. scoparia and of related species in BM, K and LTR was used
for comparative purposes.

METHODS

Leaf-sections were cut on a freezing microtome either from fresh leaves or from dried leaves
resuscitated by boiling in water for 5 min.

Chromosome counts were obtained from root-tip mitoses using standard squash techniques. Pre-
treatment was with saturated a-bromonaphthalene at 4°C for 24 hr, fixation in ethanol: ethanoic
acid (3 : 1) for 30 min, hydrolysis with 5M HCl for 10 min at room temperature, and staining with
aceto-orcein. Giemsa C-banding techniques were those used by Bailey & Stace (1992), and DAPI
staining techniques were those used by Bailey et al. (1993).

174 R. FLETCHER AND:.C.%>. STACE

RESULTS

CYTOLOGY

Good squashes were obtained of the S. Devon and N.W. Yorks plants and the number 2n = 14
found in both. Mitotic C-banding of the chromosomes with Giemsa staining, and staining with
DAPI, showed most of the chromosomes with one or two narrow interstitial bands. This contrasts
strongly with species of section Festuca (F. ovina agg. and F. rubra agg.), which have thick
terminal bands following both of these techniques (Bailey & Stace 1992, Bailey et al. 1993).
These bands are considered to represent AT-rich heterochromatin (Bailey et al. 1993).

IDENTIFICATION

Although superficially resembling a segregate of F. ovina agg., the then unknown Yorkshire
Festuca was quickly recognized as a member of section Eskia rather than of section Festuca by the
following two characters:

1.Lemmas scarious for most part, with only a narrow central part green or purplish-green;
2.Caryopsis not adherent to the lemma or palea.

The following additional characters also distinguish the Yorkshire plant from any British taxa of
the F. ovina aggregate:

1.Growth habit a spreading, springy cushion, not a firm dense tuft;

2.Leaf-blades pungent (sharply pointed) at the apex. The combination of the sharp leaf-tips and
the springy cushions give the growing plant the feel of a live hedgehog;

3.Ligules of culm-leaves at least 0-5 mm long;
4.Anthers at least 3 mm long;

5.Ovary and caryopsis pubescent at apex (in the British taxa of section Festuca this character is
found only in F. heterophylla Lam., a member of the F. rubra aggregate).

Use of the keys in Markgraf-Dannenberg (1980) and Kerguélen & Plonka (1989) unequivocally
identified the plant as F. gautieri. The combination of ligule less than 2 mm, panicle less than 7
cm, leaves less than 0-7 mm wide and with 5(—7) (not more) vascular bundles, spikelets often >9
mm, and leaves pungent at the apex distinguish it from other European species of section Eskia.

Markgraf-Dannenberg (1980) did not recognise any infraspecific taxa in F. gautieri, but “F.
scoparia A. Kerner ex Nyman” was cited in the index as a synonym of F. gautieri. Kerguélen &
Plonka (1989) recognized two subspecies under F. gautieri: subsp. gautieri and subsp. scoparia
(A. Kern. & Hack.) Kerguélen. The differences between those concern chromosome numbers,
habitat preferences, and rather minor quantitative features. Table 1 lists the differentiating
characters given by Kerguélen & Plonka for the two subspecies, and our measurements of these
characters for herbarium material of the two subspecies from the Pyrenees and for plants from the
two English localities.

The diploid chromosome number of the English plants clearly identifies them as subsp.
scoparia. With regard to the morphological characters said by Kerguélen & Plonka (1989) to
distinguish the two subspecies, the majority indicate that the Yorkshire plant is subsp. scoparia.
According to our measurements of herbarium material, lemma length (up to 6-2 mm) appears to be
the best discriminant (see Discussion). Unfortunately the flowering material of the Devon plant
was too immature to provide comparative measurements.

HABITAT

Kerguélen & Plonka (1989) described subsp. scoparia as “often calcicole in the Pyrenees”, and
said that G. Claustres claimed that it is a strict calcicole in Dép. Ariége (central French Pyrenees).
On the other hand they state that subsp. gautieri occurs on siliceous soils. Both grow in short turf
and scree in the Pyrenees and eastern Spanish mountain ranges south to the Sierra Nevada and
Morocco (Kiipfer 1974).

FESTUCA GAUTIERI 175

TABLE 1. CHARACTERS OF THE TWO SUBSPECIES OF FESTUCA GAUTIERI
ACCORDING TO KERGUELEN & PLONKA (1989) AND THE PRESENT AUTHORS

Subsp. Subsp. Subsp. Subsp. Subsp. Subsp.
gautieri fide —_ scoparia fide gautieri scoparia scoparia — scoparia
Kerguélen& Kerguélen & (Pyrenees) (Pyrenees) ex ex Devon
Plonka (1989) Plonka fide Fletcher fide Fletcher Yorkshire
Ligule of culm leaves 0-5-1 Ce 0-5—1-2 0-4-1 0-7-1-2 0-6-1-1
(mm)
Panicle length (cm) 4-5-7 7-9 3-7-4-7 5-2-6-7 5-6-7:3 -

Spikelet length to tip (9-5) 10-12 up to 9 (10) 9-5—12-5 8-6-10-6 9-5—11-5 -
of 4th lemma (mm)

Upper glume length 4-]-4-5 4-5-5 3-54-6 4-1-5-6 3-4-4:3 -
(mm)

Lemma length (mm) 6-7-7 5-2-6-2 5-5-7-1 4-9-6-2 4-3-6-2 -
Anther length (mm) 2-9-3-5 (2-1) 2-2-3 (4) 1-7-3-3 3-4-3-6 2-9-4-] -
Chromosome number P= 29 2n= 14 - - 2n = 14 2n= 14

In Yorkshire the plant grows in Seata Quarry, Aysgarth, N.W. Yorks., colonizing a flat
limestone slab at the edge of the quarry, which must to some degree mimic the Pyrenean habit of
subsp. scoparia. There is a single patch measuring about 4-5 x 1-5 m.

In Devon the plant grew on a road embankment near Exeter, S. Devon.

ORIGIN AND HISTORY IN ENGLAND
The S. Devon plant was “fairly obviously an introduction” (Margetts 1993), and presumably a
recent one. We have not been able to ascertain whether it still exists there.

Although F. gautieri makes an excellent and decorative rock-garden plant it is not mentioned in
The European Garden Flora (Walters et al. 1984); however, its relative (also with pungent leaf
apices and from the Pyrenees), F. eskia DC., is and it is possible that F. gautieri could be supplied
under that name. In addition, F. gautieri is now listed in The Plant Finder (Philip, 1999) as being
available from several nurseries.

The N.W. Yorks. plant occurs in Seata Quarry with two other notable aliens, Erinus alpinus L.
and Hypericum nummularium L., which also are native in the Pyrenees. The long exserted anthers
of F. gautieri indicate that it is an outbreeder, since most such festucoids are self-incompatible
(Barker & Stace 1982). Hence the lack of seed-set in the Yorkshire plant is what would be
expected from a single clone introduction. If that is the case, a patch now measuring 4-5 m across
will be far from a recent arrival.

Mrs D. Millward has informed us that the quarry ceased to be worked around the early 1920s,
and that as far as local knowledge goes the “aliens” (Erinus and Hypericum) were there then. It is
possible that they were introduced deliberately by the Backhouses, who were planting “The
Rockery” (now a listed building despite being a rock-garden!) not far away in the middle of the
19th century.

NOMENCLATURE

This species was first described as F. varia Haenke subsp. scoparia A. Kern. & Hack. in 1881 at a
time when only two species of section Eskia were recognized in Europe (Hackel 1882). Hackel
(1882) recognized three varieties of subsp. scoparia: var. genuina, the new var. gautieri Hack.,
and var. lutea Hack. The last (from calcareous areas in the Romanian Carpathians) was completely
ignored by Markgraf-Dannenberg (1980), but probably represents a distinct species. At the species
level F. scoparia (A. Kern. & Hack.) Nyman was created also in 1882, but this name is pre-dated
by F. scoparia Hook. f. (1844). Festuca gautieri (Hack.) K. Richt. was created in 1890, and is
therefore the correct name at species level for the two taxa together; Kerguélen (1983) combined
scoparia as a subspecies of F. gautieri.

176 R. FLETCHER AND CC. A. STACE

DESCRIPTION OF ENGLISH PLANTS
F. gautieri (Hack.) K. Richt. subsp. scoparia (A. Kern. & Hack.) Kerguélen

Stems spreading, + stoloniferous and forming dense springy cushions or mats up to 30 cm high;
rhizomes 0; innovations all intravaginal. Leaf-blades up to 9 cm, 0-4—0-6 mm diameter, glabrous
abaxially, often curved, sharply and stiffly pointed at apex, setaceous, distinctly 5—7-angled, with a
discrete abaxial sclerenchyma girder in each angle, with 5 (—7) vascular bundles, with 2 grooves;
ligule 0-6—1-2 mm; leaf-sheaths glabrous, with free overlapping edges in distal half, with distinct
rounded auricles. Culms up to 45 cm, erect, glabrous; panicles 5—7-5 cm, with rather few (up to
15) spikelets, rather compact, with scabridulous branches and pedicels. Spikelets 5-9—11-5 mm (to
tip of 4th lemma), with 3-6 florets, glabrous; lower glume 2-9-3-5 mm, with | vein, hyaline for
most part; upper glume 3-4—4-3 mm, with 3 veins, hyaline for most part; lemma 4-3—5-5 mm (excl.
awn), with 5 veins, rounded on back, hyaline for most part, with awn 0-25—0-6 mm; palea shorter
than lemma, 2-keeled and 2-veined, bifid, with scabrid apex and ciliate keels; lodicules 2, bifid;
stamens 3, with blackish-purple, long-exserted anthers 2-9-4-1 mm; ovary obovoid, pubescent in
apical part. 2n = 14.

DISCUSSION

The distinction between the two subspecies of F. gautieri 1s not easy to make, and probably it
would not merit subspecific rank if the differences in chromosome number and habitat preferences
did not exist. The main differences noted by Hackel (1882) were the thinner, green (not glaucous-
green), 3-ribbed (not 1-ribbed) leaves with 5 (not 7) veins, a more angular (not more rounded)
abaxial surface and discrete bundles (not a continuous band) of abaxial sclerenchyma, and
spikelets 9-10 mm (not 10-12 mm) in subsp. scoparia compared with subsp. gautieri. These
discriminants are partly corroborated (e.g. spikelet length, leaf angularity in section) but partly
contradicted (e.g. leaf colour and sclerenchyma distribution) by Kerguélen & Plonka (1989), and
our observations do not all coincide with either (Table 1). It seems clear that all three sets of
measurements are based on an insufficient number of specimens, and firm conclusions cannot yet
be drawn as to the best diagnostic characters that can be used to separate the two subspecies.
However, there is agreement between Kerguélen & Plonka and ourselves that subsp. scoparia has
a longer panicle and shorter spikelets, upper glumes and lemmas than subsp. gautieri, although the
actual measurements do not agree. Our measurements of the Yorkshire plant agree with our
measurements of Spanish herbarium material of subsp. scoparia more than with those of subsp.
gautieri with respect to panicle and lemma length, but not with respect to spikelet and upper glume
length. The different habitat of the Yorkshire plant might at least partly account for this anomaly.

The Yorkshire plant almost certainly originated from a deliberate planting at least 75 and
probably well over 100 years ago. With the increased use of foreign grass-seed mixtures on
roadsides, etc., and the introduction of ever more exotic and esoteric garden ornamentals, it seems
likely that further records of F. gautieri will be made in the wild in Britain.

ACKNOWLEDGMENTS

We are grateful to R. M. Payne, D. Millward, L. J. Margetts and R. Takagi-Arigho for sending us
material and supplying us with information, and to Dr J. P. Bailey for help with the cytological
work.

REFERENCES

BAILEY, J. P., BENNETT, S. T., BENNETT, M. D. & STACE, C. A. (1993). Genomic in situ hybridization
identifies parental chromosomes in the wild grass hybrid x Festulpia hubbardii. Heredity 71: 413-420.

BAILEY, J. P. & STACE, C. A. (1992). Chromosome banding and pairing behaviour in Festuca and Vulpia
(Poaceae, Pooideae). Plant systematics and evolution 182: 21—28.

BARKER, C. M. & STACE, C. A. (1982). Hybridization in the genera Festuca and Vulpia: the production of
artificial F; plants. Nordic journal of botany 2: 435-444.

FESTUCA GAUTIERI La

HACKEL, E. (1882). Monographia Festucarum europaearum. Theodor Fischer, Kassel & Berlin.

KERGUELEN, M. (1983). Les Graminées de France au travers de ‘Flora Europaea’ et de la ‘Flore’ du C.N.R:S.
Lejeunia, n. s., 110: 1-79.

KERGUELEN, M. & PLONKA, F. (1989). Les Festuca de la flore de France. Société Botanique du Centre-Ouest,
Dignac.

KUPFER, P. (1974). Recherches sur les liens de parenté entre la flore orophile des Alpes et celle des Pyrénées.
Boissiera 23: 11-322.

MARGETTS, L. J. (1993). 85th report on botany. Vascular plants. Report and transactions of the Devonshire
Association for the Advancement of Science, Literature and the Arts 125: 227-235.

MARKGRAF-DANNENBERG, I. (1980). Festuca, in TUTIN, T. G. ef al., eds. Flora Europaea 5: 125-153.
Cambridge University Press, Cambridge.

PHILIP, C., ed. (1999). The RHS Plant finder 1999-2000. Royal Horticultural Society/Dorling Kindersley,
London.

WALTERS, S. M. et al., eds. (1984). The European garden flora 2. Cambridge University Press, Cambridge.

(Accepted April 1999)

fee

:
Febe.

an

Watsonia 23: 179-196 (2000) 179

Assessing the status of Stratiotes aloides L. (Water-soldier) in
Co. Fermanagh, Northern Ireland (v.c. H33)

R. S. FORBES

Institute of Continuing Education, The Queen’s University of Belfast, Belfast, Northern Ireland,
BT7 INN

ABSTRACT

A case is argued for considering Stratiotes aloides L. (Water-soldier) (Hydrocharitaceae) of native status in
Co. Fermanagh (v.c. H33), and possibly elsewhere in Ireland. The case is circumstantial in the absence of
fossil evidence from the current interglacial, and consists of early 19th century discovery, long persistence in
natural water bodies, rare and scattered occurrence, and lack of the “boom and bust” dynamics of many
introduced aquatic species. No new stations have been found for over 100 years which strongly suggests that
either S$. aloides is not much used in Irish water gardens, or it does not readily escape into the wild. The
species complement of two vegetation communities involving S. aloides in Upper Lough Erne, Co.
Fermanagh is strikingly similar to communities described in the Norfolk Broads, where the species has long
been considered native. Conservation arguments for sites frequently revolve around the occurrence of rare
indigenous species, making the question of status more than a purely academic matter.

KEYWORDS: Hydrocharitaceae, aquatic macrophyte, native status, garden history.

INTRODUCTION

Botanists for too long have uncritically accepted notions of indigenous or introduced plant status
largely derived from early observations and old studies made in southern England, and have
continued to apply them to situations elsewhere without a great deal of reconsideration (Watson
1883; Bentham & Hooker 1900; Hyde & Wade 1957; Clapham et al. 1962; Webb 1985). Progress
in botanical knowledge in Ireland has always lagged behind that in Great Britain, originally by up
to a century or more, but it has tended to tramp the same path. However, it needs to be said that
what is true and self-evident for the south of England does not necessarily apply in the different
biological, historical and human circumstances operating in Ireland, or in other parts of these
islands. Wealth, leisure and education combine in promoting both natural history and garden
pursuits, and in timing and intensity these endeavours developed differently in the two major
islands. In particular, the passion for gardening and the fashions in it did not in Ireland slavishly
follow the English scene, but had a regional, climatically-governed identity, frequently constrained
by limited resources rather than by any lack of imagination (Nelson & Brady 1979; Lamb & Bowe
neo).

The status of Stratiotes aloides L. (Water-soldier) is a case in point, a species whose native
distribution throughout its whole European range is problematic, since it has been grown as an
aquatic ornamental for nearly three centuries and its native occurrence has been obscured by
numerous escapes from cultivation (Cook & Urmi-K6nig 1983; Preston & Croft 1997).

S. aloides is described by Stace (1997) as native, its distribution in the British Isles summarised
(abbreviations expanded here) as, “now very local in E Anglia, N and S Lincolnshire, SE
Yorkshire and Denbighshire, formerly locally frequent in central Great Britain, introduced in
scattered places in Great Britain, north to central Scotland.” He makes no mention whatsoever of
S. aloides occurring in Ireland. The species is described as being a dioecious perennial, and Stace
adds the significant remark, “only female plants occur in Britain” (but see below). Perring &
Walters (1976) in the Atlas of the British Flora are less dogmatic with regard to the status of the
plant, their map distinguishing between “probably native” and “probable or certain introductions”.
On the basis of the 17th century records, Preston & Croft (1997) regard it as likely that S. aloides
is native in eastern England “and perhaps also in Cheshire” (v.c. 58) (but see below).

180 R. S. FORBES

With regard to S. aloides in Ireland, Perring & Walters (1976) indicate five widely scattered
stations of “probable or certain introduction” status: three around Belfast Lough, Co. Antrim, one
on Upper Lough Erne, Co. Fermanagh, and one in Co. Cork. Webb et al. (1996) state that the plant
is naturalised in Upper Lough Erne and near Cork. The Census Catalogue of the Flora of Ireland
(Scannell & Synnott, 1987), lists $. aloides as an introduction in three vice-counties: West Cork,
Fermanagh and Antrim.

R. D. Meikle, on the other hand, takes a totally different view of the status of S. aloides in his
unpublished (1957) typescript Flora of Co. Fermanagh, based on collaborative field work with E.
N.Carrothers, R. C. Davidson & J. McK. Moon. In this Meikle states his opinion that, “Stratiotes
aloides has as good a claim to be considered indigenous in Fermanagh as Hydrocharis morsus-
ranae (Frogbit) or any other aquatic”’.

The present paper considers the status of S. aloides in Fermanagh in the light of definitions and
criteria for such deliberation which were carefully delineated by Webb (1985), and supplemented
by Preston (1986). Basic to Webb’s definition of native or alien plant status is the consideration of
human involvement (i.e., Neolithic man onwards). Fossil evidence from the period before
Neolithic farming activity is the only concrete proof of native status that Webb accepts; all other
evidence is circumstantial and, as such, for each species it must be subjected to careful, objective
scrutiny and the balance of probabilities weighed.

I will begin by reviewing the distribution and history of the species in relation to human
activities in Europe and the British Isles, in order to determine what patterns emerge, how they
might apply in Ireland, and how the status of S. aloides in Fermanagh appears in the light of such
investigation.

FOSSIL EVIDENCE

S. aloides can be recognised to species level both from macro-fossils and from fossil pollen
(Andrew 1970). Dickson (1970) describes how the species can be distinguished from the extinct S.
intermedius Hartz by its tough resistant leaf spines, which have a main pointed cell and subsidiary
basal cells. Both species are illustrated by Katz et al. (1965). The species is also recognised from
its fossil seeds which are often abundant, and the history of the genus has been traced back by
means of these seeds into the Tertiary Era for some 48 million years, through as many as 15 fossil
species (Chandler 1923; Cook & Urmi-Konig 1983).

In the South of England, fossil pollen has been discovered in early Pleistocene interglacial sands
and clays of the Pastonian (West 1961). This interglacial pre-dates both the Anglian (the first East
Anglian glacial stage) and the preceding Cold Beestonian stage. S$. aloides also occurs as fossil
pollen in the shelly sands of the Corton Beds, the name given to warm interstadial deposits within
the Anglian glacial in East Anglia, which are of Middle Pleistocene age (Godwin 1975). In the
Late Pleistocene it is found as seed in the Hoxnian interglacial, and again as macroscopic remains
in the Ipswichian interglacial at Wretton and Selsey. Fossil pollen from the Ipswichian has also
been recorded from Bobbitshole and Selsay (Godwin 1975). Other aquatics associated with S.
aloides in the Selsay fossil record included Hydrocharis morsus-ranae, Sagittaria sagittifolia
(Arrowhead) and Butomus umbellatus (Flowering-rush). Godwin obliquely poses an unanswered
question about S. aloides’ means of dispersal when he comments that, “It is of interest that whilst
the fossil record provides no evidence whatever of this thermophilous plant in the glacial stages, it
occurs nevertheless in every interglacial (my emphasis) from the Pastonian onwards” (Godwin
1975). Despite the implication contained in this quote, no specific reference is given here to a
fossil presence of S. aloides in the early part of the current interglacial, the Flandrian, and I am not
aware of any such record.

The above plentiful fossil occurrences, read in conjunction with early first records, help support
the native status of S. aloides in at least south-eastern England, yet fail to prove it conclusively in
Webb’s terms. Fossil material has been noted in only one Irish deposit so far, at Shortalstown, Co.
Wexford, where a single grain of Stratiotes pollen, thought to be of Ipswichian (Late Weichselian)
age, was discovered by Colhoun & Mitchell (1971). Although this appears to be the only Irish
fossil of Stratiotes, the reason for this may well be that no one has been looking for it; in addition,
to date no palynological studies of Holocene lake sediments have ever been made in Fermanagh or
Cavan, the region of S. aloides’ main past and current distribution (Dr Valerie Hall, pers. comm.,
1998).

STATUS OF STRATIOTES ALOIDES IN CO. FERMANAGH 181

GEOGRAPHICAL DISTRIBUTION AND HISTORICAL EVIDENCE

EUROPE
The map produced by Cook & Urmi-K6nig (1983) shows S. aloides occurring in the British Isles
and in Europe south to Spain, Italy, Bulgaria and European Turkey, north to Finland, and east to
Siberia (Fig. 1). Throughout the range Cook & Urmi-KGnig discovered it difficult to determine its
status, but they reviewed the scientific opinion in relation to the fossil occurrences, floristic
literature (Sometimes documenting definite introductions), and herbarium records. They showed
that the distribution of S. aloides today agrees very closely with the distribution of the 15 fossil
species. The single exception was the find of S. tuberculatus E. M. Reid from the Pliocene in
Portugal, which alone supports the two unconfirmed literature records of S. aloides in Spain from
the 18th and 19th century (Willkomm & Lange 1861). This is a particularly interesting disjunction
in the species range, as it is generally accepted that S. aloides is not native to France on grounds of
its late first mention in the relevant literature (1810), a series of documented introductions, and the
lack of any fossil finds. A similar puzzle exists with respect to Italy (where it is considered native
in the eastern part of the Po Plain) and Switzerland (“certainly not native’) (Cook & Urmi-K6nig
(1983)).

Cook revised his opinion of the native range of S. aloides two years later, asserting that it is
indigenous only in Central Europe and is introduced and established in Western Europe (Cook

TA

: t
: L
.) L
— e t} ' L
a ? tte i a
0%
4 ee @
elon 4 343 nai ie
@ 42 6
4 C4 00 rode) e@ us L L
dae L 2
4 ° e
=i L L L a
L L L L
4
4 ry L
a e eA
¥ @ “oe int: @,. 2en L
@ a6 c
hg ta a L be :
aK L L & L 1S
L
L L Lada
e O
N ® L
> L L

FIGURE 1. Distribution of Stratiotes aloides L. in Europe, with sexes determined from herbarium material
(after Cook & Urmi-K6nig 1983), and with the British Isles updated by the author. @ female plant, A male
plant, O plant of unknown gender, L record from the literature only.

182 R. S. FORBES

1985). No explanation accompanies this reassessment, but in the light of his earlier Stratiotes
review (Cook & Urmi-Konig (1983)) I believe it may involve the distribution of a dragonfly
associate (See section on insect associates below).

GREAT BRITAIN

In the British Isles, on the basis of early records, the species is regarded by most naturalists as
indigenous in the east of England (Geldart 1906; Stewart et al. 1994; Preston & Croft 1997) (Fig.
2). It was first reported by Thomas Johnson in the East Riding of Yorkshire in 1626 (Crackles
1990), followed by Lincolnshire in 1633 by the same finder (Gerarde 1633; Gibbons 1975). Old
records from the 17th and early 18th centuries must be accepted with caution, as the name
Stratiotes was applied by different authors to a number of quite unrelated plants, including
Achillea millefolium, Hydrocharis, Hottonia palustris and even some Umbelliferae (Geldart 1906).
Geldart also lists the English common names given by Johnson in the second edition of Gerard’s
Herball (Gerarde 1633) as, “Water houseleek, Knights’ pondwort, Knights’ water sengreen,
Freshwater soldier, Wading pondweed, Soldier’s yarrow and Knights’ water woundwort’. The fact
that seven English common names for the plant were in use by 1633 suggests that it was already
widely known by local people when the first plant record was published.

The native range of Stratiotes has contracted sharply in the last 150 years (Fig. 2). It is now
extinct in its previous stronghold, the Cambridgeshire fens (Preston & Croft 1997), and it has
largely disappeared from Lincolnshire and Yorkshire also (Stewart ef al. 1994). The greatest
concentration of surviving Stratiotes sites regarded as native in England occur in East Norfolk (v.
c. 27), where it is currently found in 43 tetrads (nine 10 km squares), in ditches (or dykes) in
grazing marshes around the River Waveney. Even in this corner of England, however, there are ten
to twelve more-or-less isolated sites where the species is suspected of being introduced.

Apart from S.E. England and East Yorkshire, where the plant has a long history, there is
justifiable doubt and uncertainty with regard to its status in the rest of Great Britain due to a
combination of late first records, documented introductions, colonies in man-made or heavily
disturbed habitats, and a lack of local fossil evidence, very much paralleling the situation in France
(Watson 1883; Cook & Urmi-K6nig 1983; Stewart et al. 1994; Preston & Croft 1997).

Preston & Croft (1997) suggested that S. aloides might be native in Cheshire (v.c. 58). It
transpires that this assertion was made on the same basis as that used by Stewart et al. (1994)
while working on the Scarce Plants in Britain project, 1.e., that when required to separate native
from alien records, unless there are good grounds for a difference, the treatment of the species
follows that given by the Atlas of the British Flora (Perring & Walters 1962 and subsequent
reprints with minor corrections and additions). “We think that this (1.e., The Atlas) presents a
realistic assessment of the status of most species, and also recognise that there is an arbitrary
element to the decisions which have to be made and there is no point in presenting alternative
opinions if the evidence to support them is weak.” (Stewart et al. 1994). Perring & Walters (1962)
accepted the view of De Tabley (1899) who stated in his Flora of Cheshire that S. aloides seemed
to him as native in central Cheshire as anywhere in Britain, although he gave no reasons for
making this assertion. In the same work De Tabley also cited several known introductions of the
species in Cheshire (G. McKay, pers. comm., 1998). The more recent Flora of Cheshire (Newton
1971) does not claim native status for S. aloides in the county, and C. D. Preston (pers. comm.,
1998) now agrees that in retrospect, “the case for native status in Cheshire is weak and we should
perhaps have shown less deference to the Atlas map.”

S. aloides occurs in four 10 km squares in Denbighshire, and Ellis (1983) claimed them native
(J. Green, pers. comm., 1998). The first Denbighshire record was made by the Rev. W. E. Jones in
1938 and was included in the 2nd edition of Welsh Flowering Plants by Hyde & Wade (1957),
who assumed it to be native. Ellis updated the records from the same area for his Flowering Plants
of Wales, and repeated the assumption of native occurrence since he had no reason to question it
(Ellis 1983). In the light of the Stratiotes map published in Stewart et al. (1994), Ellis has revised
his opinion and has written, “without any firm evidence to the contrary, I would probably class the
v.c. 50 populations as introduced.” (G. Ellis, pers. comm., 1998).

It is not at all surprising that the older Cheshire and Denbighshire records were simply assumed
to be native by the botanists of the time. While describing the range of circumstantial evidence that
needs to be assembled and weighed to enable a realistic consideration of a plant’s status, Webb

STATUS OF STRATIOTES ALOIDES IN CO. FERMANAGH 183

FIGURE 2. Distribution of Stratiotes aloides L. in the British Isles. Symbols represent at least one record in a
10 km square. O pre-1975 putative native records, ® putative native records made in or after 1975, + records
of introduced population made before 1950, x records of introduced populatons made in or after 1950. Based
on Preston & Croft (1997) and updated by the Biological Records Centre as requested by the author.

(1985) commented that in assigning native status there is often an element of wishful thinking,
sometimes amounting to local patriotism, or an impression that the amount present in an otherwise
undisturbed “natural looking” habitat was sufficient evidence in itself to claim a find as being
native. Webb also gave telling examples of why this level of assessment is unreliable.

IRELAND

Co. Antrim (H39)

The first Co. Antrim record was made in 1815 by John Templeton, a well-known and respected
amateur naturalist. He described the site at Stranmillis on the (then) outskirts of Belfast as, “In the
back drain of the Belfast watercourse”. Praeger (1938) and Hackney (1992) both regarded this and

184 R. S. FORBES

the other Co. Antrim records as introduced, noting that three out of four stations were extinct by
1938. Hackney also records that the plant has not been seen since 1930 at the last remaining Co.
Antrim station, the mill-dam at Woodburn, where it was reported as “occupying two acres” (Fisher
1930). Praeger (1938) commented in the second edition of The Flora of the North-East of Ireland
that Stratiotes, “should be looked on as planted in our district; in Ireland it seems fully naturalised
in Lough Erne alone” (my emphasis). In Co. Antrim the arrival, spread and departure of S. aloides
fits the “boom and bust” behaviour pattern shown by many plant introductions (Simpson 1984;
Clement & Foster 1994) and, apart from the early first discovery in a natural water course, there is
no evidence with which to argue the contrary. I would prefer to consider Templeton’s Lagan site
as of “possibly native” status but, mindful of Webb’s warning of vision clouded by local
patriotism and recognising that the evidence really is insufficient, it must remain a (now extinct)
“probable introduction”.

Co. Fermanagh (H33)

S. aloides was first found in Fermanagh in 1805, placing it amongst the very first plants recorded
in the county. The discoverer of S. aloides, the Rev. Robert Scott, M.D., was Professor of Botany
at Trinity College, Dublin from 1785 until 1808, and is regarded as the father of Fermanagh
botany. A native of the county from Scottsborough House, Magheraveely, while primarily a
bryologist, Scott evidently had a good eye for flowering plants (especially aquatics), and he was
responsible for adding Utricularia intermedia Hayne (Intermediate Bladderwort), to the flora of
the British Isles (Praeger, 1949).

The Fermanagh records of Dr. Scott were published along with his finds in other parts of Ireland
in the first edition of J. T. Mackay’s Catalogue of Rare Plants found in Ireland (1806). Mackay
himself visited Fermanagh and the adjoining part of Cavan (H30Q) shortly after Scott, for in the
same work he records his own find of S. aloides, “in drains on the roadside near Castle
Saunderson, in the vicinity of Lough Erne’. Scott’s original S. aloides site is described as, “
growing in a drain near Crum Castle, on the banks of Lough Erne, county of Fermanagh, in
August last.” Crum is currently spelt, “Crom”, and this locality near the Fermanagh-Cavan
boundary where the Erne connects southwards with Belturbet, is still the main area of distribution
for the plant today. S. aloides has recently been recorded again from the Cavan side of the border;
the species was refound in 1996 at two loughs near Belturbet (Round Lough, H15.39, and
Anoneen Lough, H22.35) (P. Reilly, pers. comm., 1998). This represents a gap of 128 years since
the species was last listed in the same area by David Moore (Moore & More 1866). Robert
Northridge and I, as joint B.S.B.I. vice-county recorders for Fermanagh (H33), have assembled a
total of 44 S. aloides records in 15 tetrads (Fig. 3). Fourteen tetrads contain post-1975 records. The
species was first recorded at the S.E. boundary of the county and subsequent finds extended its
known occurrence northwards along Upper Lough Erne rather slowly. It was not found at
Belleisle, about eight km. south of Enniskillen, until 1900. In 1990 a major survey of aquatic
vegetation, commissioned by the Department of the Environment for Northern Ireland, extended
the known distribution as far north as Lough Yoan, within 2 km of the town. In reality, however,
the evidence is much too scant to conclude that the species is spreading in Fermanagh. Botanical
recording in the county (as elsewhere in Ireland) has been extremely occasional, sporadic and
unsystematic until the 1980s, and it has involved only a handful of people, few of whom resided
locally. One can say with confidence, however, that S. aloides has not exhibited an explosive
spread in the manner characteristic of some definitely introduced aquatic species, such as Elodea
canadensis (Canadian Waterweed), or E. nuttallii (Nuttall’s Waterweed). Neither has it declined or
disappeared on Upper Lough Erne as it has in its Co. Antrim stations, where its occurrence on
man-made or disturbed aquatic habitats always tended to support the suspicion it might be an
introduction.

The headquarters of S. aloides in Fermanagh remains very much where it was first found almost
two centuries ago, the southernmost fen-fringed lakes and inlets of Upper Lough Erne, where it
sometimes dominates open water. In mid-August 1994, at the southern of two lakes in the district,
both called “Corracoash”, Robert Northridge and I came across a striking example of its growth
potential; a dense population covered the lake surface, approximately 3 ha in extent. In 1998 the
communicating Mill Lough, approximately twice the size of Corracoash Lough, was so overgrown
with S. aloides that it was difficult to row a boat through it (J. Bullock, pers. comm., 1998).

STATUS OF STRATIOTES ALOIDES IN CO. FERMANAGH 185

a
/
.

a
a

FIGURE 3. Tetrad distribution of Stratiotes aloides L. (Water-soldier) in Co. Fermanagh (H33). Hatched
symbols represent pre-1975 records, black symbols post-1975 records.

Other stations in Ireland

Following the Fermanagh, Cavan and Antrim finds already discussed, Mackay published a further
site for S. aloides, “on the banks of the Shannon, near Portumna” (Co. Tipperary, H10) in his
Flora Hibernica (1836). Mackay’s listing for this site showed S. aloides occurring on the major
river of central Ireland with Sagittaria sagittifolia and Eleocharis acicularis (Needle Spike-rush).
The Tipperary S. aloides had gone by 1866 when Moore & More published their Cybele
Hibernica.

In addition to its earlier northern sites, Moore & More (1866) also list S. aloides occurring,
“naturalized (my emphasis) in Ballyphehane Bog” (Cork city; H4), a site which Mackay (1825)
previously mentioned in passing, and which was near the Cork Botanic Garden. Significantly,
Mackay (1825) does not list S. aloides for this site, although twelve other species from
Ballyphehane Bog are attributed to his major Cork contributor, Mr Drummond. The Ballyphehane
station is again ignored by Mackay in his Flora Hibernica (1836), which also suggests he knew it
was not a native occurrence of the species. Drummond was the curator of the short-lived Cork
Botanic Gardens, which was situated on a low limestone hillock in the city. It appears that
Drummond simply used the nearby marsh (it was never a bog), as a convenient out-station for
growing aquatic plant species for the Botanic Garden (M. Scannell, pers. comm., 1998). This is the
only unequivocally introduced site for Stratiotes in Ireland.

Colgan & Scully (1898) mention S. aloides as having occurred fleetingly (in 1829) in a canal
near Drogheda, Co. Louth (H31). The lack of persistence and artificial nature of the waterway both
suggest that a garden origin 1s likely in this case.

186 R. S. FORBES

Moore & More (1866) considered S. aloides, “probably not indigenous” in southern Ireland, but
suggested it might possibly be native in the north of Ireland. They recognised that the species was
a rapidly spreading introduction elsewhere in the British Isles, and compared it with Elodea
canadensis, (a plant which made its first appearance in these islands in Co. Down around 1836).

As was the case in Co. Antrim, the S. aloides plants in these three widely scattered locations in
southern Ireland between 1829 and 1836 appear to have become extinct soon after their discovery,
except at Ballyphehane Bog where the species survived until some date after 1900. The apparently
short-lived nature of the other two south of Ireland stations is typical of most introductions; the
persistence of the species at Ballyphehane may be the result of deliberate management.

The fact that there have been so few sightings of S. aloides in Ireland over the 192 years since its
first discovery suggests that it is either truly rare, or so hard to detect that it is seldom found. As is
the case with many Irish lakes, the waters of Upper Lough Erne are an opaque dark brown colour
due to peat staining, so that a grapnel is an essential item of equipment to drag for submerged
macrophytes. Even if one is actively looking for S. aloides, it can be difficult to see the
characteristic “pineapple-like” leaf rosettes in deep murky lake water, and there could be very
many plants invisible at depth. Under conditions of low illumination the leaves do not develop
sufficient gas in their air-canals and intercellular spaces to raise the plant rosettes to the surface,
and it is quite normal to find it as a bottom-dwelling plant in water between | and 5 m deep (Cook
& Urmi-Konig 1983). In more shallow waters, particularly after a prolonged spell of warm
weather, very rapid growth can produce the sort of Stratiotes “explosion” that occurred at
Corracoash and Mill Loughs.

HABITAT

The typical habitat of Stratiotes is still or slow-flowing, meso-eutrophic water (Preston & Croft
1997). Upper Lough Erne consists of a multitude of small, shallow, linked lakes and intervening
islands - a drowned drumlin landscape - and although there is a channel navigable for motor
launches, the great majority of its shores are undisturbed. Drainage ditches are cleared from time
to time by local farmers and, although fringing emergents encroach, open water is plentiful, as is
rainfall throughout the year. Agriculture is entirely pastoral and low intensity on the surrounding
rushy meadows and, while slurry is spread, eutrophication from run-off has not become a
widespread problem. In recent years 60% of Fermanagh farmland has been covered by a
government-funded Environmentally Sensitive Area farm subsidy scheme.

In England S. aloides was formerly found in lakes and slow-moving rivers but now is virtually
confined to drainage ditches (Preston & Croft 1997). In the Broads of south-eastern England S.
aloides occurs in three habitats, all of them man-made. The Broads are flooded peat excavations,
the fen dykes were created to allow boat access into the fens, and the network of ditches in the
reclaimed marshes were created for land drainage (R. J. Driscoll, pers. comm., 1998). S. aloides
appears to prefer the still water of grazing marsh ditches, particularly where the water is
moderately rich in nutrients and calcareous, and it grows where regular cleaning of the ditches or
ponds, (often grazed on their margins), suppresses the growth of shading reeds and other tall
emergent species. S. aloides is even more intolerant of fluctuating water levels than its relative and
frequent associate, Hydrocharis morsus-ranae, and unlike it does not survive on mud to form
turions (Cock & Lii6nd 1982).

Eutrophication, and habitat disturbance and destruction due to agricultural changes, are readily
identified as the major reasons for the rapid decline in distribution and decreasing abundance in the
20th century. In east Norfolk, the main area of surviving populations considered indigenous,
habitat loss continues due to a change from pasture to arable agriculture, subsequent pollution and,
since the area is low-lying and near the coast, sporadic tidal inundation (A. Bull, pers. comm.,
1998). On the other hand, Wheeler & Giller (1982) suggested that S. aloides, in the dyke system of
the Catfield and Irstead fens in the Norfolk Broads, showed abundant growth at the fen margins,
possibly in response to a degree of agricultural nutrient run-off, whilst the species has become rare
in the central areas of these fens during the last 30 years. On the basis of water chemistry they
considered it at least possible that this decline might be due to nutrient depletion associated with
oligotrophication and acidification.

A similar catastrophic decline of the species in. the Netherlands from the 1960s onwards
followed increased drainage associated with modern agricultural practices, which diminished the

STATUS OF STRATIOTES ALOIDES IN CO. FERMANAGH 187

influence of ground water. To prevent the low-lying peat soil from drying out, sulphate-enriched
alkaline water from the Rhine was admitted, which produced a complex of environmental stresses
involving iron limitation, sulphide toxicity, a strong increase of non-rooted species due to internal
and external eutrophication, and ammonium toxicity (Smolders ef al. 1996).

The habitat requirements and tolerances of S. aloides are thus very specific and narrow (see
community ecology section below).

REPRODUCTIVE PATTERN

Phytogeographically, Stratiotes is regarded as a Continental species by Matthews (1955), a
position refined by Preston & Hill (1997), who described it as Eurosiberian Boreo-temperate.
When trying to relate a species’ distribution to geography and climate, however, one must take
particular account of the fact that different optimum and minimum conditions may occur for life
processes such as seed set, germination and vegetative growth, and that biotypic variation may
modify these physiological characteristics within the total range of a species. This is certainly the
case with regard to growth and sexual reproduction of Stratiotes.

Currently, reproduction of S. aloides in the British Isles and in most of Europe is entirely
vegetative and is carried on by the production of new open leaf rosettes (or closed turions
(Kornatowski 1979)), budded off on monopodial lateral shoots (Cook & Urmi-K6nig 1983).
Renman (1989) studied the life histories of two clonal populations of S. aloides in northern
Sweden and found that emergent forms of the plant produced more daughter turions than did
submerged forms.

There is the added complication of the dioecious nature of the plant and the present-day
distribution of the sexes. The dioecy of Stratiotes is not absolute. Dioecy is secondary in origin
and in S. aloides flowers of both sexes contain whorls of staminodes, which in the female flowers
secrete nectar at their base (Daumann 1931). Although the staminodes are normally sterile, rarely
the female flowers contain a few apparently fertile stamens on their outer whorl (Geldart 1906;
Cook & Urmi-K6nig 1983; Preston & Croft 1997). Hutchinson (1948) went so far as to describe S.
aloides as “subdioecious’’. The existence of hermaphrodite plants (even at low frequency) casts a
different light on the possible reproductive history of the species in the post-glacial period.

Full dioecy is represented when plants (or genets - defined as genetically distinct individuals),
are totally male or totally female. According to Richards (1986), who was writing on dioecy in
general terms, when sex is phenotypically unstable, hermaphrodites that are capable of selfing may
occur. When only some of the flowers on a female genet become hermaphrodite (as in the case of
S. aloides), they are termed subgynoecious, and this is considered the result of unstable sex
expression in a genetically dioecious plant (Richards 1986). (When all the flowers of a female
plant or genet become hermaphrodite, the species is said to show gynodioecy (i.e. female and
hermaphrodites occurring together), but this does not apply in the case of Stratiotes). Plants that
are fickle in their sex expression are often subject to environmental influence and, in the case of
thermophilous species such as S. aloides, temperature is clearly the most likely significant factor -
possibly linked with daylength. Sex expression in plants and its physiological control is a
fascinating area of study (Meagher 1988), and touching on it ought to remind us that concepts such
as “incompatible” and “dioecy”’, are decidedly “leaky”, and that our description of the natural
world is frequently a convenient oversimplification of the variation that really exists (Proctor et al.
1996).

Cook & Urmi-Konig (1983) reviewed the current distribution of the sexes in S. aloides, and
found that while female plants strongly predominate north of a line drawn from Cherbourg to
Leningrad (St Petersburg), isolated male plants have been found in Denmark, Sweden and Finland
(Fig 1). North of the Cherbourg/Leningrad line, however, they did not find any evidence of S.
aloides bearing seed and, although fruits may form, no viable seed has been found in the British
Isles either, since the Pleistocene (Crackles 1982). This suggests three non-exclusive possibilities;
(a) that for successful seed production, S. aloides requires a warmer summer than that currently
found north of the Cherbourg/Leningrad line; (b) that male S. aloides plants are more susceptible
to cold and have become extinct in what is now a scattered, relict population; or (c) male flowers
are never or only rarely triggered to occur by the longer summer daylength or other environmental
factors to the north.

188 R. S. FORBES

The fossil assemblage found with S. aloides in the Corton Beds in East Anglia of Middle
Pleistocene age indicated to West & Wilson (1968) a strong northerly aspect to the vegetation,
with some persistence of Arctic-alpine species which occurred, “together with a number of
aquatics, including S. aloides and Potamogeton crispus, with a mainly southern distribution”, (my
emphasis). Considering the occurrence of fossil Stratiotes along with Cladium mariscus,
Hydrocharis morsus-ranae and Lemna minor, West (1961) suggested that their joint presence
indicated a considerable summer warmth in the Cromerian and Pastonian interglacials of the
Norfolk coast.

Evidence like this supports the notion that the prevailing temperature in the north and west of S.
aloides’ current range is too cool for successful sexual reproduction, as even in regions where both
sexes occur today (i.e. Sweden, Finland and Denmark), no viable seed appears to be set. Further
south, in a warmer climate, seed is successfully produced, e.g. in the Netherlands (Smolders et al.
1993). Palynologists tell us that we are past the climatic optimum in the current interglacial period
(the “Littletonian” in Ireland ( Mitchell & Ryan 1997)), so it is arguable that S. aloides flourished,
reproduced and spread by seed in an ancient warmer climatic phase, and when cooler conditions
developed it declined to its current disjunct relict areas, persisting by vegetative reproduction. To
explain the occurrence of single sex populations in the British Isles, we have to postulate
differential survival capacities of the sexes. A similar explanation involving differential vigour,
ecology and survival between the sexes is used to account for the restricted occurrence of female
plants of Petasites hybridus in the British Isles (one site only in Ireland), while males are
widespread throughout both islands (Valentine 1946, 1947; Perring & Sell 1968).

DISPERSAL ABILITY

As Preston & Croft (1997) point out, without seed S. aloides appears to lack a means of long-range
dispersal over land. The current lack of fruiting ability may have arisen, however, after small
founder populations of seed successfully established in both England and Ireland, some plants
perhaps temporarily retaining a degree of fertility through the possession of a subgynoecious habit.

“Baker’s Law” (Baker 1955; Stebbins 1957) suggests that dioecious species appear to be at a
major disadvantage when colonising islands compared to self-compatible hermaphrodite plants.
Bawa (1982), however, has suggested that dioecious taxa may have been disproportionately
successful in colonising islands. Analyses of numerous oceanic island floras by Baker & Cox
(1984) indicated that dioecious taxa do not do better in long-distance dispersal than self-
compatible hermaphrodites, but neither do they fare worse, a result that was unexpected and was
not predicted by extensive numerical simulations (Cox 1985). While the distances and the
timescales involved in colonisation of oceanic islands are far greater than those associated with
continental islands such as the British Isles, the evidence remains that jump dispersal is effective
and plants do colonise islands. The fact that in S. aloides dioecy is not absolute but “leaky” (Cox
1985), and that it is capable of producing subgynoecious self-compatible hermaphodites from time
to time, overcomes some of the conceptual difficulties we face in imagining jump dispersal of the
species to Britain and Ireland.

Like two aquatics often associated with Stratiotes, Ceratophyllum demersum (Rigid Hornwort),
of which Scannell (1976) found that it flowered but failed to ripen fruit in Ireland, and
Hydrocharis morsus-ranae, which flowers erratically and seldom fruits (Cook & Liiénd 1982;
Preston & March 1996), S. aloides possesses a successful means of vegetative multiplication. It is
possible that a single founder or a single aggressive clone may have survived many years of
asexual competition and a slowly changing, cooling environment, giving rise to high levels of
sterility, even in species or regions where both sexes are currently sympatric (Richards 1986).
Throughout north-west Europe S. aloides appears to fit this pattern closely, and on offshore islands
the probability of a restricted genome ts clearly increased.

The occurrence of fertile seed-producing plants across the North Sea in the Netherlands, and the
small but possibly highly significant seed set which self-compatible subgynoecious plants could
produce, provide an ancient seed source for colonisation of the British Isles, most likely through
transportation by birds. Jump-dispersal of turions is possible, but diaspores adhering securely tc
the dispersal agent externally (i.e. exozoic transport) is somewhat more difficult to envisage thar
endozoic transport of ingested seed.

STATUS OF STRATIOTES ALOIDES IN CO. FERMANAGH 189

Fruit of S$. aloides ripens late, (end of September to end of October), at a time when birds are
migrating. Fresh seeds of S. aloides are heavily clothed with mucilaginous hairs and are liberated
in a gelatinous mass by longitudinal splitting of the fruit (Cook & Urmi-K6nig 1983), which
suggests the possibility that they might stick to feet or feathers of birds feeding among fertile
fruiting plants. The exotesta which bears the mucilaginous hairs is however shed, shortly after the
seed is released from the fruit (Cook 1987), and in any case the mucilage tends to be sticky when
dry, or almost so, but non-sticky when wet (Sculthorpe 1967). However, while S. aloides is said to
attract a range of herbivores including water snails, beetles, insects and “several
vertebrates” (Cook & Urmi-Ko6nig 1983), birds are not specifically mentioned as feeding on the
plant. This may simply reflect the fact that, at present, seed is rare on Stratiotes plants anywhere in
Europe.

Similar dispersal circumstances attend other non- or seldom-fruiting aquatic macrophytes,
including several that are commonly associated with Stratiotes in Britain and Ireland and generally
considered native. Other native aquatic species of the same community freely set seed, yet appear
to remain poorly adapted for long-range jump dispersal as their seeds are totally digested when
eaten by birds and fish, e.g. Nymphaea alba (White Water-lily) and Nuphar lutea (Yellow Water-
lily) (Preston & Croft 1997).

All long-range or jump-dispersal events are rare and unlikely and as such are difficult to
envisage (Carlquist 1974; Cook 1985). Birds, particularly waterfowl, are often cited as agents of
dispersal. Because of their rarity, natural transport events are almost impossible to observe and, for
the same reason, they are not readily open to experiment. In the great majority of cases we do not
know how or when the dispersal event(s) happen, but we have indisputable evidence that they do
occur.

GENETIC DIVERSITY

As noted by Webb (1985), this is one area where data for most plants are very rarely available. I
doubt that there are any relevant data on S. aloides apart from comments made by Cook & Urmi-
Konig (1983). These authors undertook in their revision of S. aloides to rid the accumulated
literature on it of contradictory observations and views, by reviewing the species on the basis of
their own direct experience of fresh and fixed material. After their study they concluded that they
had found no genetic or ecotypic variability worthy of formal taxonomic recognition. Three
features they mentioned were:

a. minor differences existed with respect to seed sizes between Swiss and Czechoslovakian
plants,

b. British specimens were somewhat stouter than Continental and Scandinavian ones, yet fell
within the overall pattern of variation for the species,

c. totally submerged plants with more flaccid, stomata-less leaves up to 100 cm long were more
common in the north of the range of the species.

The reproductive pattern and current behaviour of S. aloides suggest that a small or very small
founder population might well be all that the species succeeded in dispersing across the North Sea
and the Irish Sea, so that plants on each island may represent one clone or a small range of clones.
Comparative genetic investigation using isozymes is required to answer the questions posed by
this clonal hypothesis.

COMMUNITY ECOLOGY

The Middle Pleistocene fossil assemblage in East Anglian sediments described by West & Wilson
(1968) showed S. aloides occurring with Potamogeton crispus (Curled Pondweed), Cladium
mariscus (Sword Sedge), Hydrocharis morsus-ranae and Lemna minor (Common Duckweed). All
these fossil allies occur today with S. aloides in Upper Lough Erne, Co. Fermanagh, together with
numerous other frequently associated aquatics, e.g., Lemna trisulca (Ivy-leaved Duckweed),

190 R. S: FORBES

Sagittaria sagittifolia (Arrowhead), and several narrow-leaved Potomogeton species typical of
more shallow waters, such as P. pusillus (Lesser Pondweed), and P. pectinatus (Fennel
Pondweed). In phytosociology, communities are drawn floristically by species presence, plus a
particular set of frequency and abundance values of all the species found in the samples or relevés.
A sample may be pigeon-holed into a previously described community even if the name-giving
species, or some of the other characteristic “constant species”, are absent. The information
available from Upper Lough Erne forms part of the substantial Fermanagh Flora database and
consists, not of standardised quadrats or relevé samples, but of site specific species lists which do
not incorporate frequency and abundance values. It is possible, however, to glean an idea of
species frequency from the overall species statistics of records and tetrads occupied, and I present
these estimates of species presence here (Table 1).

The National Vegetation Classification survey of aquatic vegetation in Great Britain (Rodwell
1995) shows S. aloides present in two plant associations, prominently as a characteristic constant
species in the Hydrocharis morsus-ranae—Stratiotes aloides community (A4) (Table 1), and as a
rare companion species in the Potamogeton pectinatus—Myriophyllum spicatum community (A11).
The Aydrocharis morsus-ranae—Stratiotes aloides community is equivalent to the continental

TABLE 1: FLORISTIC TABLE FOR THE HYDROCHARIS MORSUS-RANAE-STRATIOTES
ALOIDES COMMUNITY (A4) OF RODWELL (1995)

Norfolk Broads Fermanagh
Species Frequency Cover value No. recs. No. of tetrads
Hydrocharis morsus-ranae V 1-4 78 19
Stratiotes aloides V 1-7 4- 12
Lemna minor V 1-3 720 50+
L. trisulca V 1-5 487 50+
Callitriche platycarpa V 1-3 13 5
Polygonum amphibium IV 1-3 370 47+
Utricularia vulgaris IV 1-7 98 30+
Ceratophyllum demersum IV 1-7 0 0
Myriophyllum verticillatum Il 1-3 26 6
Nuphar lutea Il 1-2 619 50+
Potamogeton obtusifolius Il 1-6 240 50+
Elodea canadensis Il 1-4 604 50+
Potentilla palustris II 1-2 674 50+
Hottonia palustris I 1-2 0 0
Berula erecta I ] 23 6
Ceratophyllum submersum U 1-6 0 0
Oenanthe aquatica Ul 1-3 126 2a,
Sparganium erectum I 1-3 613 50+
Nasturtium officinale I 1-3 432 35+
Sium latifolium I 1-3 Sed, 50+
Alisma plantago-aquatica I 1-3 708 50+
Nymphaea alba i 1-3 185 18
Ranunculus lingua I 1-3 181 35+
Sagittaria sagittifolia I 1-3 87 28
Potamogeton crispus I 1 98 18
Sparganium emersum I 3-4 267 50+
Number of samples 12
Number of species/sample 14 (11-19)

Based on data (columns | & 2) from the Norfolk Broads (Wheeler & Giller 1982), compared with
data from the Co. Fermanagh Flora database (unpub.). Column 3 lists the total number of records
for the species in the database, and column 4 indicates the number of tetrads in around Upper
Lough Erne in which it occurs.

STATUS OF STRATIOTES ALOIDES IN CO. FERMANAGH 191

Hydrochario—Stratiotetum (Westhoff & den Held 1969) or the Hydrocharitetum of Oberdorfer
(1977). Rodwell admits that he has characterised this community in Great Britain almost entirely
on what he refers to as, “the small data set of Wheeler & Giller (1982)” from the Norfolk Broads,
and he recognises that more comprehensive sampling is needed to arrive at a satisfactory definition
of this kind of assemblage in Britain. Keeping this reservation in mind, examination of the
published floristic table for the Hydrocharis morsus-ranae-Stratiotes aloides community
demonstrates that in addition to the two eponymous aquatics, there are 24 additional species listed,
of which 21 occur in Upper Lough Erne (Table 1). The only plants missing from Upper Lough
Erne are the two Hornworts, Ceratophyllum demersum and C. submersum (Soft Hornwort), plus
Hottonia palustris (Water Violet). Of these three, C. demersum alone has a prominent role as a
“constant species” in the plant community, but the other seven “constant” species are all present.

The fact that S. aloides occurs in Fermanagh backed up by a very nearly complete complement
of the characteristic species of the Ceratophyllum—Stratiotes nodum (Wheeler & Giller 1982), or
the Hydrocharis morsus-ranae—Stratiotes aloides community of Rodwell (1995) - and this despite
the depauperate flora of Ireland - must lend very strong circumstantial support to the claim for
Water-soldier’s native status in Upper Lough Erne.

With regard to the second community, in which Stratiotes plays a much more minor role, the
Potamogeton pectinatus—Myriophyllum spicatum community (All) of Rodwell (1995), Upper
Lough Erne supports 47 of the 56 characteristic species listed by Rodwell for the three sub-
communities he recognises (neglecting the moss Fontinalis antipyretica). The nine missing species
include five which are nationally scarce in Great Britain, two that are strictly coastal, and two
which have never been recorded anywhere in Ireland.

Thus we can say with confidence that Stratiotes occurs in Upper Lough Erne with very much the
same vascular plant species complement that accompanies it in Great Britain and, importantly, that
in turn these match Continental equivalents.

INSECT ASSOCIATES

Preston (1986) suggested an additional criterion for assessing native status of plants to the eight
discussed by Webb (1985), namely, the relationship of the plant species in question to
oligophagous herbivorous insects. Entomological evidence, Preston indicated, needs to be used
with the same degree of caution as Webb’s criteria, for introduced plant species may attract insect
feeders from related long recognised native species, as is known in the case of Impatiens noli-
tangere (Touch-me-not Balsam), which supplies oligophagous insects to the definitely introduced
garden escape /. parviflora (Small Balsam) when the two species are adjacent.

With regard to S. aloides, the absence of the dragonfly Aeshna viridis Eversmann, which is
described as dependent on the macrophyte, led Cook & Urmi-K6nig (1983) to argue that S. aloides
might not be native in Great Britain and other areas of Europe from which the insect is missing.
These authors report the findings of Miinchberg (1956), who researched the habits and life history
of the insect: it is carnivorous and, although the larvae live on S. aloides rosettes for two or three
years until they reach maturity, they do not appear to harm the leaves of the host plant at all.
Knowing this, it is not at all clear why the dragonfly should be specifically associated with S.
aloides. Higler (1977) found over 200 invertebrate species associated with S. aloides in the Dutch
broads, but apart from a few Lepidopteran caterpillars, which did insignificant damage, none of the
other species appeared to feed on the living plant tissues - they merely used the plant as a habitat
surface or shelter. The argument linking Aeshna viridis to the native status of S. aloides is further
weakened by the fact that the distribution of the dragonfly only partially overlaps that of S.
aloides, and the dragonfly is absent from N.E. Italy and the mouth of the Danube where Cook &
Urmi-K6nig believe S. aloides is likely to be native (Cook & Urmi-K6nig 1983).

Another dragonfly, Anaciaeshna isosceles (Miiller) (Norfolk Hawker), a protected, endangered
species, is restricted in Great Britain to the Broadlands and there heavily dependent on S. aloides,
again apparently purely for shelter, although it is also found to a lesser extent on Glyceria maxima
(Reed Sweet-grass), Juncus effusus (Soft-rush) and Iris pseudacorus (Yellow Iris) (Leyshon &
Moore 1993). This dragonfly was found in the examined Suffolk Broadland dykes only where a
dense population of S. aloides occurred.

192 R..S. FORBES

EASE AND SOURCE OF KNOWN NATURALIZATION

GREAT BRITAIN

S. aloides was recommended by Philip Miller as a suitable plant for water gardens in his famous
Gardener’s dictionary, first published in 1724 and with many subsequent revisions and abridged
editions. Until Loudon’s An encyclopaedia of gardening appeared in 1822, Miller’s publication
was the most widely used and influential garden handbook and his ideas became fashionable.
William Robinson’s The English flower garden (first published in 1883 and which also ran to
many subsequent editions), continued the fashion, although he was not impressed with Stratiotes,
“spike of unattractive blossoms”. A recently published manual of water gardening sponsored by
the Royal Horticultural Society (Robinson 1997) continues to recommend S. aloides as a hardy
submerged aquatic species for garden pool decoration.

The map of the British Isles distribution of S. aloides in Preston & Croft (1997) and the revised
version published here (Fig. 2) provide a convenient summary of current knowledge of the
distribution of the species, the extent to which it has become naturalised in Great Britain, and its
decline this century. (The coverage of Preston & Croft’s map gives a very much more complete
and detailed picture of the status of S. aloides than the Atlas map (Perring & Walters 1976), having
information plotted for 216 squares compared with just 93). It requires only a cursory glance to
appreciate that only a small minority (16%) of the post-1969 symbols represent native
occurrences, the balance being records of introduced populations. The revised map (Fig. 2)
contains four classes of record symbols, 25% representing post-1969 native occurrences. The
introductions extend north to Brechin (v.c. 90), south to Portsmouth (v.c. 11), and west to Bideford
(v.c. 4) but are, as one might expect, centred on the major conurbations of the English midlands.

The other impression quickly gained from Fig. 2 1s the extent to which S. aloides as a species
(irrespective of the status of the particular station) is declining in the Britain Isles. Even after
revising the status of the Fermanagh and Cavan records, the number of native symbols on the map
has dropped from a total of 75 10-km squares pre-1950 to just 23 today, while in areas where it
was introduced, the decline has only been from 149 squares pre-1950 to 69 squares after that date.

IRELAND

In Co. Fermanagh the main garden activities in the late 18th and early 19th centuries were still
landscaping with trees, and the less formal, more natural Rococo style of gardening initiated by
Alexander Pope, his friend Dean Jonathan Swift and his friend, the Dean of Down, Patrick
Delaney. The Rococo style involved vistas of trees grown in a natural manner, rather than the
clipped dress of the formal baroque garden, and it had subsidiary vistas in the form of meanders or
angles, the whole effect being intricate, whimsical and natural (Lamb & Bowe 1995). It is known
that Delaney advised on garden improvements in Co. Fermanagh, certainly at Belleisle near
Enniskillen, and perhaps also at Manor Waterhouse (Malins & the Knight of Glin 1976). The
Manor Waterhouse estate (H3730), a few km south of Lisnaskea and five km north of the Crom
estate, belonged to an Englishman, John Madden (1652-1703). Madden and his son Samuel were
interested in plants (Nelson 1981), and gardening (including the use of water features); however,
there are no records of S. aloides from Manor Waterhouse and no evidence suggesting that the
Maddens knew the species, or that they had any hand in introducing it to the county.

The grounds of Crom Castle, where (or close to where) S. aloides was first discovered in 1805,
at that particular time were probably extremely neglected from the gardening point of view, since
Crom Castle had been destroyed by a major fire in 1764, and was not rebuilt until 1834. The
grounds around the castle were then laid out in the new classical Landscape style, complete with
false ruins and a picturesque cottage or garden house on nearby islands in Upper Lough Erne.

Miller’s Gardener’s dictionary (1724 and subsequent editions) had many Irish subscribers, and
three Dublin editions were published, in 1732, 1741 and 1764 (Nelson 1980). Although Miller
recommended S. aloides for use in garden water features, | cannot unearth any evidence that the
plant was ever fashionable or widely available in Ireland. The Garden History Society has
published The availability of hardy plants of the late eighteenth century (Harvey 1988), in which
there is no mention of either S. aloides, or its often associated relative, Hydrocharis morsus-ranae,
indicating that S. aloides was not available for purchase in the horticulture trade immediately prior
to its discovery in Ireland.

STATUS OF STRATIOTES ALOIDES IN CO. FERMANAGH 193

S. aloides was first found associated with or close to landed estates e.g., Crom and Castle
Saunderson. Does this not suggest a cultivated origin? Such estates were sited in places that were
visually interesting and strategically important and, being large enough, often contained relatively
undisturbed habitats that were likely to contain interesting plants. Educated people lived in these
surroundings and they used the lakes for sailing and pleasure boating and were likely to notice
unusual plants. S. aloides has an unusual life-cycle, rising and sinking as the year progresses, and
we can surmise that estate residents would comment on it to each other and to interested visitors.
Botanists like Scott and Mackay, while not directly involved in the life of the grand houses,
through their knowledge of plants and gardens, had access to the estates, their owners and
gardeners, a tradition of hospitality amongst plantspeople which happily continues to the present
day. The botanists would thus gain knowledge of local wild plants from the residents of the large
country houses, and very many of the older plant records in the Fermanagh Flora database are
associated with the names of old estates, e.g., Florencecourt, Belleisle, Crom and Castle Coole (R.
H. Northridge pers. comm., 1998).

If, contrary to published information (Harvey 1988), sources of S. aloides were widely available
and it was used for decorative effect in water gardens throughout Ireland for a period, why, if it
readily escapes (as it appears to do in Great Britain), have there been no further reports of it in the
wild in Ireland during the last 160 years?

CONCLUSIONS

The only incontrovertible proof of native status for S$. aloides in Co. Fermanagh, pre-Neolithic
macrofossil material, is absent so far - here and elsewhere in the British Isles. Therefore we must
weigh up the circumstantial evidence offered here to decide if it is sufficient to allow us to say, as
Meikle did in 1957 (unpub. ms.), that S$. aloides is as indigenous to Co. Fermanagh as Hydrocharis
morsus-ranae, or any other aquatic plant species.

The major points in favour of an indigenous origin for S. aloides in Fermanagh are:

1 Early discovery (in an Irish context at least), by a reputable and very experienced botanist who
knew the locality intimately and could therefore have judged the likelihood of garden origin,
yet did not argue for it, or comment on it as a possibility.

2 The rare and scattered occurrence of records and the lack of spread of the species when
compared to other known aquatic introductions.

3. The total lack of new stations for the species for well over 100 years, despite the recording
efforts of a growing body of botanical recorders, suggests either that S. aloides is not widely
grown in Ireland, or if it is, it does not escape or spread.

4 §. aloides was not available in the horticulture trade in the late 18th century and no
information suggests that it ever became a popular or fashionable garden plant in Ireland.

5 Persistence for almost 200 years in a natural watercourse (i.e. Upper Lough Erne) compared
with its relatively rapid disappearance at sites where introduction is either known (ie.
Ballyphehane Bog), or possible.

6 The remarkable degree of similarity in the range of associated species when compared to the
vegetation described from the Norfolk Broads, the most likely native station in Great Britain.

7 The existence in Upper Lough Erne of accepted native aquatic plant species, some rare, some
with disjunct distributions, and some which also currently lack seed dispersal.

8 The habitat in Upper Lough Erne fits the habitat typical for S. aloides everywhere in Europe,
namely slow-flowing meso-eutrophic water.

194 R. S. FORBES

The main evidence suggesting S. aloides could be an introduction is:

1 Miller’s Dictionary of Gardening (1724 & subsequent editions) recommended S. aloides to
gardeners creating water features. Early 19th century occurrences of the species in a few
widely scattered stations in Ireland, including canals, flax dams and other artificial waterways,
might imply some level of cultivation and subsequent escape.

2 Some of these possibly escaped S. aloides colonies persisted for up to 120 years before dying
out and could therefore appear naturalised for a time.

3S. aloides appears to lack an obvious means of long-range “jump” dispersal over land, and it is
thus difficult to account for its presence anywhere in the British Isles, and more so on the
offshore and more ancient island of Ireland.

4 The restricted distribution of its extant Irish populations, especially its absence from other
watercourses in drumlin regions (and even from the lower parts of the River Erne catchment).
However, as discussed earlier, the habitat requirements and tolerances of S. aloides are very
specific and narrow, and locations that /ook appropriate may not provide suitable sites (e.g., the
physiography and water chemistry of Lower Lough Erne differs considerably from the Upper
Lough).

I believe that, on balance, the circumstantial evidence assembled here strongly supports Meikle’s
assertion that S$. aloides has as much right to indigenous status in Ireland as Hydrocharis or any
other aquatic species. Like Webb (1985), I feel sure that there are other cases where the current
status of a plant species is heavily dependent on received wisdom rather than reasoned debate over
assembled facts. Conservation arguments for sites frequently revolve around the occurrence of rare
and declining indigenous species, making the question of status much more than a purely academic
matter.

ACKNOWLEDGMENTS

I especially thank Robert Northridge, my partner as joint Recorder for Co. Fermanagh (H33), for
over 20 years of enjoyment during field work, and for persistence in the creation of the Fermanagh
Flora database which forms the basis of this work. Drs Damian McFerran and Julia Nunn at the
Centre for Environmental Data and Recording (CEDaR), Ulster Museum, Belfast assisted with the
creation and maintenance of the database. Prof. Christopher Cook provided a copy of his published
map of European distribution and kindly permitted me to modify it (Fig. 1). Drs Chris Preston and
Jane Croft at BRC provided the revised British Isles distribution (Fig. 2). The Co. Fermanagh
distribution (Fig. 3) was drawn using Dr Stuart Ball’s RECORDER and Dr Alan Morton’s
DMAPW software. I also thank colleagues who offered information or responded to enquiries:
Joan Bullock, R. J. Driscoll, Gwynn Ellis, Dr Charles Nelson, Michael Walpole, Shaun Wolfe-
Murphy and B.S.B.I. v.c. Recorders, Maura Scannell (H3), Patrick Reilly (H30), Alec Bull (27),
Jean Green (50) and Graeme Kay (58). Dr Valerie Hall (Queen’s University, Belfast) provided
helpful information and assistance. Finally, my wife Deirdre read the paper and encouraged its
submission.

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(Accepted March 1999)

Watsonia 23: 197—208 (2000) 197

The distribution, ecology and conservation of Arenaria norvegica
subsp. anglica Halliday (Caryophyllaceae)

K. J. WALKER

LT.E., Monks Wood, Abbots Ripton, Huntingdon, Cambridgeshire, PE17 2LS

ABSTRACT

Arenaria norvegica subsp. anglica is endemic to the Craven area of Mid-west Yorkshire (v.c. 64) where it
grows exclusively amongst limestone fragments (clitter) and in shallow solution hollows with a thin covering
of soil on level or slightly inclined limestone exposures. Since its discovery at Ribblehead in 1889 over 20
populations have been found, all but one of which occurs on the eastern slopes of Ingleborough. Over 70% of
the entire population, of around 2000-3000 plants, occurs within two adjacent pastures where numbers are
apparently stable. However, subsp. anglica has declined on tracks and bridleways where it was first recorded
over 100 years ago. Although colonies within pastures appear to have fared better, their frequent small size
(less than 50 plants) makes populations susceptible to drought and disturbance. The current rarity of subsp.
anglica owes more to climate change (contraction of habitat) and aspects of its autecology (limited seed
production and dispersal) rather than human activities. However, grazing and recreational disturbance still
threaten some populations. As a result conservation measures are being introduced in order to reduce grazing
levels and protect sensitive trackside populations. In addition, the creation of “safesites” and collection of
seeds for genebanking will ensure its ultimate survival.

KEYWORDS: taxonomy, population size, rarity, habitat management, Yorkshire.

INTRODUCTION

Arenaria norvegica Gunn., Arctic Sandwort (Caryophyllaceae, Subfamily Alsinoideae), is an
arctic-montane therophyte which occurs in Iceland, Norway, Sweden, Finland, Shetland and north-
west Scotland. Two subspecies occur in the British Isles (Fig. 1): A. norvegica subsp. norvegica' is
a plant of base-rich screes, river shingle and fell-field, confined to a small number of sites in
Scotland (Beinn Iadain, v.c. 97; Beinn Sgulaird, v.c. 98; Rum and Eigg, v.c. 104; Inchnadamph, v.
c. 108; Unst, v.c. 112) and one somewhat problematical site in the west of Ireland (Gleninagh
Mountain, v.c. H9), where it may be extinct (Webb & Scannell 1983). A. norvegica subsp. anglica
Halliday, English Sandwort, (Arenaria gothica auct., non Fries hereafter referred to as subsp.
anglica) appears to be endemic to England occurring in only two 10 km squares in the Craven
region of Mid-west Yorkshire (v.c. 64). This subspecies can be distinguished from subsp.
norvegica by its often annual (rarely biennial) habit, somewhat larger flowers, laxer habit, greater
ciliation (especially on the base of the sepals) and narrower, more lanceolate leaves (Halliday
1960b).

Subsp. anglica is an undoubted Yorkshire speciality which has attracted many generations of
botanists to the area, not least because it shares its habitat with a number of other rarities (e.g.
Minuartia verna and Sedum villosum). The population in Craven, which is in the order of (1000-)
2000—3000(—4000) plants, is confined to the Carboniferous limestone outcrops on the eastern
slopes of Ingleborough (Walker 1995). Remarkably there is only one very small population
outside a restricted (5 km square) area, despite the occurrence of suitable habitat elsehwere. As a
result it is now recognised as “endangered” (Hodgetts et al. 1996) and afforded full protection
under Schedule 8 of the Wildlife and Countryside Act (1981).

This paper describes the history of the plant in Britain, in terms of its taxonomy, distribution and
Status. Information on aspects of its ecology are also presented, and are discussed in relation to

rarity as well as conservation measures proposed within a recent action plan (Walker & Corkhill
1996).

' Nomenclature follows Kent (1992) for vascular plants and Smith (1978) for bryophytes.

198 K. J. WALKER
TAXONOMY

Arenaria norvegica is included within the Arenaria ciliata L. complex, a polymorphic taxon
incorporating a number of closely related montane plants with highly disjunct distributions
stretching some 40° from northern Spain to 82° north in the Arctic. The taxonomic history of this
complex is very confused, with up to four species and seven geographically distinct subspecies
being recognised (Chater & Halliday 1993).

Subsp. anglica, in particular, formerly posed a number of taxonomic problems for British
botanists. On discovery in July of 1889 it was initially thought to be a form of A. ciliata, although
it was later confirmed by F. A. Lees and Professor C. C. Babington as A. norvegica Gunnerus
(Rotheray 1889b). At the time this species was known from only two sites in the British Isles; on
Unst where it had been known of since 1837 and in West Sutherland on the Scottish mainland
where it had been recently discovered (Scott & Palmer 1987). However, a number of botanists
who were familiar with the Scottish plants, including J. Gilbert Baker of Kew, commented on the
laxer habit, narrower leaves and differing shoot structure of the “Yorkshire sandwort” (Baker
1889). After visiting the Ribblehead populations in September of 1889 both Baker and Lees began
to doubt these earlier determinations, particularly Lees who suggested that the Yorkshire plants
may represent an “altered form” of A. norvegica (Rotheray 1889b). As a result he sent specimens
to William Whitwell and Arthur Bennett who had a small collection of European Arenaria
material sent to him by Nilsson in 1882.

After comparing these with Lees’ plants and Fries’ original descriptions Bennett concluded that
the Ingleborough plant was in fact A. gothica Fries (Rotheray 1889b; Bennett 1891), a very rare
plant of dry limestone habitats and lake shores in southern Sweden (V4stergétland), Gotland and
the Swiss Jura (Lac de Joux) (Williams 1898; Albertson 1946). The Ribblehead population
represented a notable extension of its curiously disjunct European distribution although both
Bennett and Baker thought that its true place was with A. norvegica either as a subspecies of A.
ciliata or as a variety (Baker 1889; Bennett 1891).

Despite these misgivings botanists remained faithful to Bennett’s determination until Halliday’ s
study of the A. ciliata complex in the late 1950s (Halliday 1960a). Using a variety of techniques
Halliday showed the Yorkshire plants to have a greater affinities to A. norvegica than either the
Swedish or Jura A. gothica and remarked that “‘apart from differences in habit, the British plant is
closer to A. norvegica in most other respects, in particular in leaf shape, colour, ciliation and
inflorescence size” (Halliday 1960a). Furthermore, British plants were diploid (2n = 80; Swedish
plants 2n = 100), could interbreed freely and produce fertile seeds whereas seed production ©
declined in Swedish x English hybrids (Halliday 1958, 1960a, b). On this basis Halliday
concluded that the English plant was better placed under A. norvegica and proposed two
geographical subspecies; A. norvegica subsp. norvegica and A. norvegica subsp. anglica, “taxa
sufficiently distinct morphologically and geographically separated but capable of interbreeding
freely and with a common origin” (Halliday 1960a). Subsequent taxonomic work on the A. ciliata
complex has supported this division (Wyse-Jackson & Parnell 1987). In this study the two
subspecies were shown to be closely related and distinct from all the other European members of
the complex, including both A. ciliata s.s. and A. gothica.

DISTRIBUTION

HISTORICAL RECORDS: 1889-1980

Subsp. anglica was originally discovered by Lister Rotheray at Ribblehead Station, near
Ingleborough, on 12 July 1889 where it was growing “in some profusion” amongst limestone
“staging” on a recently constructed track (Rotheray undated, 1889a, b; Whitwell 1889). F. A. Lees
who visited the site soon after located hundreds of plants, including a new population growing
“300 and 400 yards [350 m] away” from the original colony on the opposite side of the railway
line (Whitwell 1889). Its occurrence on a man-made track initially raised doubts as to whether the
plant was native (Baker 1889; Whitwell 1889; Bennett 1892). However, the use of local stone for
track-building led Lees and J. G. Baker to predict its presence on “the neighbouring hills” (Baker
1889; Whitwell 1889). Indeed a small population was discovered on a track at Selside in the

ARENARIA NORVEGICA SUBSP. ANGLICA 199

FIGURE 1. The past and present distribution of Arenaria norvegica subsp. norvegica (MM 1970 onwards; A pre-
1970) and subsp. anglica (@ 1970 onwards) in the British Isles. Sites where undetermined fossil seeds of A.

ciliata agg. have been found are indicated as follows: A late Devensian/Flandrian, 0) mid Devensian,O early
Devensian.

following August and as a result botanists began to search more widely (Whitwell 1890). Then in
September of 1894 Reginald J. Farrer, the famous plant-collector of Ingleborough Hall, reported
its presence at “Sulber Nook” [Nick], in a spot so remote as to confirm its native status (Farrer
1894; Rotheray 1895). Several more populations were found by him in the following year, all of
which occurred on or adjacent to the Clapham Lane, a popular bridleway connecting Selside and
Crummack Dale (Whitwell 1890, 1895; Rotheray 1895). Farrer communicated these new finds to
Rotheray who promptly visited the following August. His account of this trip gives a good
indication of the numbers and location of plants as well as describing a number of populations
which were undoubtedly new to Farrer (Rotheray undated).

An intriguing record was the discovery by Farrer of a small population within the grounds of his
own residence, Ingleborough Hall, some 9 km from Ribblehead (Whitwell 1895). Given the
numbers of alien plants which have been found in the vicinity in recent years (Abbott 1996) one
wonders whether such a keen plant-collector could have moved it there?

200 K. J. WALKER

Although Rotheray and Farrer undertook extensive searches of the Craven limestones over
subsequent decades no new populations were found until the 1950s when Halliday discovered a
small colony close to Farrer’s original Sulber Nick site (Halliday 1960a). A number of other
populations were discovered by Bartley & Clark (1978) who were the first to describe the
extensive flush population within Sulber pasture (refound in 1993), and the two outlying colonies
to the south-east; at Foredale, in a disused quarry, and close by on pavement at Juniper Gill.

To date the only verified record away from the eastern slopes of Ingleborough is the small
trackside colony at Dawson Close which was discovered by Ruth Kilby during a Yorkshire
Naturalists’ Union field excursion in 1952 (Sledge 1952). This site on the northern slopes of
Fountains Fell was a notable extension in range, some 6 km to the east of the nearest Ingleborough
population. Further afield Lees’ (1937) unconfirmed report of having “picked up two or three
specimens from a limestone-metalled road near Healuagh” (Swaledale) in 1906 is conceivable
given the transport of Ingleborough limestone for track-building, although this population has
never been relocated. Most unlikely is Arthur Bennett’s claim of having found a fragment of A.
gothica whilst collecting Potamogeton obtusifolius from Grasmere (Cumbria) given the absence of
suitable habitat within the vicinity (Halliday 1960a).

CURRENT DISTRIBUTION AND STATUS

Since 1981 staff of the Nature Conservancy Council (and latterly English Nature) have carried out
a number of detailed population censuses of all extant populations (Table 1). In 1995 all these sites
were visited by the author and extensive searches made of the neighbouring limestone pastures
(Foley 1995; Walker 1995). The results of these surveys have given a much clearer indication of
the plant’s distribution and abundance. However, given the discovery of at least eight new
populations over the last half century, one of which is situated some 6 km to the east of Sulber, it
is not inconceivable that subsp. anglica may well await discovery elsewhere in Craven.

In 1995 over 2700 plants were recorded in approximately 20 colonies within eleven | km
squares. Over 70% of the total population (Table 1; populations 2—7 & 8—14) occurred within two
adjacent pastures at Sulber on the eastern slopes of the Ingleborough massif (Walker 1995). Here a
series of loosely connected populations are confined to shallow soils on limestone exposures,
flushes and rough tracks all of which occur on or within 500 m of the Claphman Lane. Elsewhere
small populations occur by tracks, at Dawson Close and Selside, in a disused quarry at Foredale
and on limestone exposures at Juniper Gill and Thieves’ Moss.

Although overall populations sizes are small (mean 143 +43; n = 18) these vary consistently by
habitat. Trackside populations are by far the smallest (mean 16 +4; n = 6) and only account for 3%
of the total population. As a result these populations are vulnerable to trampling and drought
which presumably caused the localised extinction of subsp. anglica at Selside and Sulber Nick in
1992. Populations on gently inclining pavement, which account for 53% of the entire population,
are much larger (mean 161 +54; n = 9) and apparently less prone to fluctuations in overall numbers
(Table 1). In contrast the two flush colonies at Sulber, which alone account for over 40% of the
entire population, show marked fluctuations in numbers from year to year presumably due to the
ephemeral water-supply within spring-fed flushes.

CHANGE SINCE 1889

The distribution of subsp. anglica has changed little since it was discovered in 1889 and it can still
be found in the localities described by Rotheray and Farrer over a hundred years ago. Extinction
has occurred in only two sites; in Spring Valley at Ingleborough Hall (where it may have been
introduced) and at Ribblehead, whete over-collecting led to its demise within two years of
discovery (Whitwell 1890). Rotheray’s accounts and the results of recent surveys suggest that it
has also been lost, or declined, from numerous sections of the Clapham Lane, particularly those
which traverse Sulber and the adjacent Borrin’s pasture (Rotheray undated; Whitwell 1895).

The most likely cause of decline is presumably the increased recreational usage of tracks within
the Sulber area over the past century. Since the opening of the National Park in 1953 visitor
numbers have increased dramatically, particularly at Ingleborough where there are now around
250,000 visitors annually (S. Rogers, pers. comm., 1995). This is undoubtedly due to the
increasing popularity of the Three Peaks footpath which passes directly through Sulber Nick (and
a number of populations of subsp. anglica). This figure is likely to increase further when the
Clapham Lane is incorporated into the “Pennine Bridleway’, a new long-distance routeway
designed to cater for walkers, horse-riders and cycling enthusiasts.

201

ARENARIA NORVEGICA SUBSP. ANGLICA

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FIGURE 2. Changes in the mean numbers of plants of Arenaria norvegica subsp. anglica on: a. tracksides (@)
(populations 1, 9a, 10, 11, 12 & 16) and b. limestone pavement/flush habitats (MI) (populations 3a, 7, 9b—d, 13
& 14).

In contrast, populations on “primary” habitats, such as limestone pavements and flushes appear
to be steadily increasing (Fig. 2). The extent to which this is a result of more information or
relaxation in grazing pressure is unclear. In the past overgrazing has presumably reduced plant
numbers, particularly in the Sulber pastures where census returns have shown a tripling in sheep
numbers this century (from around 13,000 to just over 35,000 in 1989; P. Corkhill, pers. comm.,
1995). Although stock are important in maintaining open conditions and transporting seed,
excessive defoliation and trampling by stock has been observed to cause some mortality,
especially around the edge of flushes where grazing is particularly intense (Walker 1995).

ECOLOGY

HABITAT, SOILS AND CLIMATE
Subsp. anglica is a plant of shallow skeletal soils (lithomorphic protorendzinas) on sparsely-
vegetated level or slightly inclined limestone exposures. Here plants grow amongst limestone
fragments (clitter) and in shallow solution hollows with a thin covering of soil. At one site it also
grows directly on tufaceous “crusts” (hydromorphic gleys) which have developed on flat or gently
sloping limestone slabs around the edge of flushes and seepage lines. In addition, it has been long
established within “ruts” of trackways and amongst shattered limestone on a quarry floor. Recently
it has also been observed on disturbed rabbit scrapes (Walker 1995).

In all of these habitats soils are shallow (usually <7 cm), circumneutral (mean pH of 7-59 + 0-06
n = 37), with both high levels of calcium (+30 mg Ca 100 g'') and organic material (mean 67% +
2). In most habitats moisture supply is limited and unpredictable and as a result populations are
small and sparsely distributed. In contrast, where water-supply is more predictable, i.e. around the
edges of flushes and springs, populations are larger and numbers less variable from year to year.

The altitudinal range of populations is between 310-390 m (see Appendix). Here the climate is
distinctly oceanic with rainfall being uniformly high (average of 1590 mm yr’; 1981-95) tending
towards a spring minimum and autumn maximum. At this altitude the growing seasor
(accumulated temperatures >6°C) lasts for 5-6 months (April to September) and the distribution o
the plant appears to be limited to areas where the oceanic influence ensures that the summe
temperatures do not exceed 23°C (Dahl 1951).

ARENARIA NORVEGICA SUBSP. ANGLICA

203

TABLE 2. SPECIES ASSOCIATED WITH ARENARIA NORVEGICA SUBSP. ANGLICA IN
NON-FLUSHED, FLUSHED AND DISTURBED SITES.

Species Frequency Mean Non-flushedt Flushed (n Disturbed
(n = 43) abundance (n= 2) = 19) (n:=:3)
Arenaria norvegica subsp. anglica 43 2-2 V(1-3) V(1-4) V(1-3)
Festuca ovina agg. 42 5-1 V(3-7) V(2-8) V (5-7)
Linum catharticum 42 1-3 IV(i-3) V(i-3) IV(2)
Sesleria caerulea 34 2:5 V(i-5) V(1i-7) (3)
Minuartia verna 33 1-8 ~ V(i-4) IV(i-3) (2)
Ctenidium moliuscum p24 | 1-4 IV(1-5) Nidi-4) IV(1)
Thymus polytrichus subsp. britannicus 26 1-3 IV(i-4) lidi+) V(5)
Carex panicea 25 7 Ii(1-2) V(i-5) V(4-6)
Briza media 22. 1-0 Ili(i—3} Tii¢di-3) V(i-4)
Tortella tortuosa Ze jb IV(1-7) Ii(¥i-4) Ii(1)
Agrostis capillaris 16 1-1 Til (i-4) iii) H(A)
Koeleria macrantha 15 1-0 Wd) Hi(3-5)
Euphrasia officinalis s.1. 14 <1-0 I(1-3) (1) TV (i-2)
Carex caryophyllea 10 <1-0 Iii (1-4) I1(6)
Plantago lanceolata 10 <1-0 I(1) Ii(1-2) V (2-4)
Cerastium fontanum 9 <1-0 II(1-3) I(i-2) li(1)
Schistidium apocarpum 9 <1-0 Il(i-4) IiI(i-4)
Juncus articulatus 9 <1-0 I(3) II(i—2)
Prunella vulgaris 8 <1-0 1h) I(1) V(2-3)
Taraxacum officinale s.1. 8 <1-0 (1) IV(1)
Hypnum cupressiforme i, <1-0 Iq)
Sedum villosum q| <1-0 I(t) I(1-2) II(1)
Potentilla erecta 6 <1-0 II(i-3) I(1) I(5)
Sagina nodosa 6 <1-0 I(1-3) (1) (2)
Viola riviniana 6 <1-0 I(1) 1(4) I(t)
Bellis perennis 5) <1-0 I(1) I(1) V(i-2)
Carex viridula subsp. oedocarpa 5 <1-0 II(2-5)
Carex flacca 5 <1-0 I(1-3) I(5) IV(3)
Carex pulicaris 4 <1-0 I(1-2) I(1) II(1)
Cratoneuron commuiatum 4 <1-0 Ii¢i-7)
Ranunculus bulbosus + <1-0 I(1) V(i-2)
Agrostis canina 3 <1-0 i(2-3)
Poa annua 3 <1-0 I(1) I(3) IV(i-2)
Bare soil 39 5-6 V(1-8) V(4-9) V(5)
Bare limestone 36 4.9 V(1-9) IV(i-8) (5)
Limestone gravel (<10cm) os 3-0 IV (3-8) IV(i-5) IV(1)
Limestone gravel (>10cm) 24 2-4 IV(i-7) IV(1-9)
Faeces (rabbit/sheep) 11 0-3 Ii(i—3) I(1) IV(1)

Notes: + Frequency classes: I = 0—20%; II = 20-40%; III = 40-60%; IV = 60-80%, V = 80-100%.
Additional species in <2 quadrats: Non-flushed; Antennaria dioica, Cirsium sp., Dicranum scoparium,
Drepanocladus_ revolvens, Erophila verna, Galium sterneri, Lotus corniculatus, Polytrichum
juniperinum, Pseudoscleropodium purum, Vulpia bromoides. Flushed; Bryum argenteum, Plantago
media, Primula farinosa, Racomitrium lanuginosum. Disturbed; Achillea millefolium, Ajuga reptans,
Cardamine pratensis, Hypericum montanum, Polygala vulgaris.

204 K. J. WALKER

However, drought can have a devastating effect on plant numbers as in 1992 where low spring
rainfall (83 mm for May and June as opposed to an average of 163-5 + 16-6 mm between 1981-
1995) caused a number of localised extinctions (Fig 2.). As a result subsp. anglica is most
frequently found within sheltered microtopographichal niches (e.g. rock fissures, moss cushions,
solution cups etc.) where water-loss and exposure to extreme temperatures are reduced (Halliday
1960a).

VEGETATION

The habitat of subsp. anglica typically occurs as scattered “islands” of open ground, usually only a
few metres in extent, within upland limestone pastures dominated by Sesleria caerulea (CG9;
Rodwell 1992). Within this localised habitat subsp. anglica occurs within three closely related
assemblages with differing microtopographic, edaphic and disturbance regimes (Table 2).

On non-flushed soils overlying gently inclining limestone exposures subsp. anglica occurs with
a number of other diminutive species (e.g. Minuartia verna, Sagina nodosa and Erophila verna)
on bare patches of soil/gravel amongst scattered clumps of Festuca ovina and Sesleria caerulea.
Here vegetative cover rarely exceeds 30% and is characterised by the presence of stress-tolerants
such as Thymus polytrichus and Carex caryophyliea.

On flushed soils and gravels vegetative cover is less fragmentary and more species-rich. Here
the constants, Sesleria caerulea, Festuca ovina, Minuartia verna and Linum catharticum are joined
by a number of species of moister soils most notably Primula farinosa, Carex viridula subsp.
oedocarpa and C. panicea. Presumably this is the Arenaria norvegica subsp. anglica — Sedum
villosum heath described by Bartley & Clark (1978) which they considered a transitional variant
between the more species-rich flushes and the surrounding calcareous grasslands.

Subsp. anglica also occurs within a species-poor “disturbance” assemblage on tracks and rabbit
scrapes. Here it is associated with ruderals of bare soils, such as Prunella vulgaris and Poa annua,
and rosette-forming hemicryptophytes such as Bellis perennis, Plantago lanceolata, Taraxacum
officinale and Ranunculus bulbosus which can tolerate heavy trampling pressure.

LIFE-CYCLE

Classified as an annual or biennial (e.g. Clapham et al. 1987), the majority of plants germinate in
the autumn, over-winter as leafy shoots and flower and set-seed in the following spring or summer
(winter annual) (Halliday 1960a; Walker 1995). Under exceptional circumstances plants have also
been observed to flower and set seed within one season (summer annual) (Walker 1995), or
survive two winters (biennial) (Halliday 1960a).

Throughout its life subsp. anglica is exposed to extreme variations in both temperature and .
moisture supply; as a consequence there is no general relationship between the time of flowering
and season. This is achieved by the production of polycarpic shoots (cymes) which enable the
plant to flower continuously (indeterminately) over an extended period. Thus although flowering
typically takes place between early May and September, it is not uncommon to find plants
flowering as late as December if conditions are favourable (Whitwell 1895). This trait (sometimes
termed uniseasonal iteroparity) allows annuals such as subsp. anglica to avoid the potential
dangers of synchronous flowering within an unpredictable habitat, and ensure that some seed is
dispersed under favourable conditions (Harper 1977; Symonides 1988).

On average plants produce 2—3 hermaphrodite white flowers. As with many other members of
the Alsinoideae, reproduction may be entirely autogamous (i.e. within the same flower) due to the
automatic transfer of pollen between the homogamous reproductive surfaces. On maturity both the
inner and outer whorl of anthers are forced into contact with the maturing stigmas ensuring that if
they have not already been pollinated any remaining pollen grains are picked up (G. Halliday,
pers. comm., 1998). In addition, plants which are artificially self-pollinated, and isolated from
insect visitors with a fine muslin, are able to reproduce and set seed (Walker 1995). However, the
production of obvious signals (simple white receptive flower, sweet scent) and rewards (nectaries)
suggest that insects do visit the plant and that cross-pollination may take place.

Subsp. anglica produces relatively large seeds (0.8 x 0.9 mm), the tough black testa having
numerous raised tubercles which presumably serve to reduce desiccation. The viability of fresh
seed is usually very high, with over 90% germinating readily when placed on moist filter paper (S.
Terry, pers. comm., 1995; Halliday 1960a). However, experimental studies carried out by the
author have shown that germination declines with increased illumination and temperature, and at
low moisture levels (Walker 1995). This suggests that optimal conditions for germination are in
sheltered micro-sites where moisture supply is not limiting.

ARENARIA NORVEGICA SUBSP. ANGLICA 205

Capsules contain on average 3—4 seeds (mean 3-59 per capsule, n = 291) although under
cultivation this can be much higher (Walker 1995; G. Halliday, pers. comm., 1998). After
fertilisation it can take up to three weeks for the pericarp to split and reveal the seeds inside.
Although some seeds are released at this stage the majority remain firmly attached by placentae to
the capsule until detachment and decomposition of the pericarp on the surrounding soils. As a
result dispersal is extremely limited and usually not more than a few centimetres from the parent
plant. However, the presence of subsp. anglica on tracks, in particular the Clapham Lane and the
outlying colony at Dawson Close, suggests that seed may be transported over greater distances,
presumably within soil attached to humans, vehicles or sheep which are often “away-wintered”
from the Sulber pastures (P. Corkhill, pers. comm., 1995; Sledge 1952). In addition, like Thlaspi
perfoliatum it may have been widely dispersed within limestone ballast used in the construction of
road and railways in the past (Rich et al. 1989). However, although both species are now well
established on artificial sites (such as quarry floors, tracks and railway embankments) the lack of
any specialised seed dispersal mechanisms means there is little tendency for further spread to
occur.

Although experiments suggest that relatively few seeds are incorporated into the seed-bank, the
sporadic reappearance of the plant within a number of trackside sites suggest that some seed may
persist for short periods.

CONSERVATION

It seems likely that subsp. anglica may well have evolved, through isolation, from a more widely
distributed and variable population (Pigott & Walters 1954; Raven & Walters 1956). The
occurrence of fossil seeds of Arenaria cf. ciliata, to which subsp. anglica undoubtedly belongs,
within Mid- and Late Devensian deposits in the south and east of England and the east of Ireland
suggests a much wider glacial distribution (Fig. 1). These species presumably survived the height
of glaciation within periglacial refuges free of ice (i.e. unglaciated massifs, coastal areas) or on the
periglacial “park-tundra” to the south of the ice limits (Godwin 1975). However, as a result of
climatic changes during the post-glacial this once continuous lowland distribution has been
restricted to a number of disjunct montane and coastal refugia where immature soils still persist
(Godwin 1953; Rose 1957; West 1988).

Today the rarity of these habitats, as well as poor dispersal ability has presumably compounded
this restriction in range. As such subsp. anglica is likely to remain extremely rare being restricted
by biological and ecological factors rather than as a direct result of human activities.

CURRENT THREATS

Although there is no evidence for an overall decline in numbers this century many trackside
populations have been lost or severely degraded as a result of the increased usage by walkers, “‘off-
road” vehicles, scramble and mountain bikers. This has led to severe soil erosion and a decline in
plant numbers at numerous sites along the Clapham Lane. In particular track-widening and
Shortcuts created by walkers threaten large colonies at Sulber Gate and Crummack Dale. At
Selside, where the plant grows on the edge of a popular track leading to Alum Pot, car-parking has
reduced numbers to the verge of extinction.

Many of the pastures in which the plant occurs were formerly overgrazed (sheep numbers tripled
between 1940 and 1980). As a result sheep numbers have been reduced through management
agreements since designation as a S.S.S.I. in 1986. However, many populations are still highly
susceptible to disturbance caused by off-road vehicles.

There are no 20th Century reports of collecting of subsp. anglica and disturbance caused by
visiting botanists is unlikely to pose a threat to many of the larger populations. However, smaller,
more accessible colonies (some with less than ten plants) are extremely vulnerable to trampling,
particularly at Selside which is frequently visited by botanists.

CONSERVATION MANAGEMENT

Subsp. anglica has been included within English Nature’s Recovery Programme as a species
considered to be in danger of extinction and requiring special conservation measures (Whitten
1990). As a result an action plan has recently been proposed in order to ensure its long term self-
sustained survival in the wild (Walker & Corkhill 1996).

206 K. J. WALKER

By the early 1990s all the sites in which subsp. anglica occurred received statutory protection
either as part of the Ingleborough National Nature Reserve (N.N.R.) or as Sites of Special
Scientific Interest (S.S.S.L) (Ingleborough, Pen-y-Gent and Foredale Quarry). As a consequence
grazing levels are now agreed with farmers (in some cases voluntarily under the Wildlife
Enhancement Scheme), and ensure that sheep numbers do not exceed two ewes (and followers) per
hectare throughout the year. In addition, one pavement on which the plant occurs (Juniper Gill) has
been recently stock-proofed (P. Corkhill, pers. comm., 1995). The steady increase in plant
numbers recorded within these pastures (Fig. 2) suggests that these measures are having
favourable results.

In order to arrest the decline of subsp. anglica on vulnerable track-side localities a number of
measures have been proposed. At Sulber Gate on the Clapham Lane a diversion of the Pennine
Bridleway has been agreed with the Countryside Commission in order to protect a large colony. In
addition, better way-marking of routes will reduce the numbers of “unofficial” paths which have
appeared in recent years. At Selside limestone boulders have been placed around sensitive areas in
order to discourage car-parking.

In the long term ex-situ measures, such as seed-banking and the establishment of populations in
“safe-sites”’, will safeguard the plant from ultimate extinction as well as providing material for
further research and public viewing. In 1992, 500 seeds were collected from two extant
populations and are now held at the Royal Botanic Gardens Kew Seed Bank at Wakehurst Place.
In the same year seed was also artificially introduced to Scar Close N.N.R. where it has apparently
failed to establish. Introduction into further “safe-sites” is currently being discussed with a view to
creating a public viewing facility in order to reduce pressures on extant populations (P. Corkhill,
pers. comm., 1995).

In order to assess the success (or otherwise) of these management activities a sample of
populations are being monitored every three years. The results of recent surveys suggest that the
efforts of conservationists and landowners have ensured the survival of this endemic plant for
many years to come.

ACKNOWLEDGMENTS

I am especially grateful to Peter Corkhill at Ingleborough without whose support and invaluable
assistance this project would not have been possible. I would also like to thank Mike Foley, Barrie
Goldsmith, Adrian Soames, Janet Terry and Peter Welsh for information and advice on various
aspects of this work. Chris Preston, Geoffrey Halliday and Owen Mountford all made valuable ~
comments on an earlier draft of this paper and Dave Roy at Monks Wood produced Fig. 1.
Fieldwork was supported by grants from English Nature as part of the C-E-L Scheme and Species
Recovery Programme.

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(Accepted March 1999)

K. J. WALKER

208

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Watsonia 23: 209-212 (2000) 209

Annual variations in the size of a population of
Cardamine impatiens L.

L.R. WILLIAMS

Brent Parks Service, c/o Brent Town Hall, Forty Lane, Wembley, Middlesex, HA9 9HD

ABSTRACT

A population of Cardamine impatiens Narrow-leaved Bittercress was monitored by annual counts of the
number of plants during the decade 1988-1998 at Fryent Country Park, Middlesex (v.c. 21). There were large
annual variations in the number of individuals, both in total and at each colony. Variations appeared to be due
to the cyclic nature of populations at each colony, to unexplained variations possibly related to the weather,
and to ground disturbance. The species increased in response to occasional ground disturbance. The
reappearance of plants where it had not been observed for several years, suggested the role of a buried seed
bank. The average spread, in a south-easterly direction, was 17 ma year over the decade.

KEYWORDS: Narrow-leaved Bittercress, Brassicaceae, monitoring, Middlesex, hedgerows.

INTRODUCTION

Cardamine impatiens, Narrow-leaved Bittercress is a scarce plant in Britain with scattered
populations nationally (Stewart et al. 1994). It is a plant of varied habitats including shady
woodland, rocks and screes, shady banks of streams and rivers and of damp roadsides. It occurs on
both calcareous and neutral soils. In London and in Middlesex (v.c. 21) it was considered extinct
(Kent 1975; Burton 1983) until a population was rediscovered in 1985 at Fryent Country Park,
(TQ/198.881), in the former parishes of Kingsbury and Wembley. Prior to that the most recent
record had been ‘near Harrow’ in 1901. Wembley was part of the Harrow parish until 1894.
Monitoring of C. impatiens commenced in 1988 with the aim of providing information on how
best to conserve the species locally.

METHOD

The investigation was undertaken at Fryent Country Park, approximately 15 km north-west of
central London and in the London Borough of Brent. The Country Park covers an area of
approximately 100 hectares of lowland countryside surrounded by suburbia and bisected by the
A4140 road. The landscape is of hay meadows cut annually, with hedgerows, woodlands, ponds,
and areas of scrub, rough grassland, amenity grassland and horse grazing. The park is on London
Clay, with a pH of about 6-5, though Barn Hill, to the south, is capped with gravel and more acid
soils. The altitude varies between 45-83 m O.D. The flora of the park was described by Williams
(1996) and the landscape by Williams & Northcroft (1992).

Counts were made by the author to estimate the number of individual plants. Each more-or-less
continuous group of plants was mapped as a “colony”. Obviously there were limitations to this
approach, since some of the colonies merged during the investigation. Counts were made towards
the end of May (to early June) in each of the years 1988 to 1998.

RESULTS

The number of individuals at each colony are given in Table 1. The species was found in the
northern, central part of the Country Park. Records were from both the Kingsbury and the
Wembley parish areas, and from the park on both sides of the A4140 road. There was considerable

210 L. R. WILLIAMS

variation in the population size from year to year (Table 1 and Fig. 1). The highest count was in
1997 (9977) and the lowest in 1995 (504), though the 1994 count (1076) would probably have
been lower if it had not been for two new colonies, on disturbed ground, totalling 900 plants.

There was an increase in the number of colonies during the investigation, mainly within the
original range. However the linear spread, in a south-easterly direction was approximately 170 m,
or an average of 17 m a year. At the end of the investigation the range of the species covered about
25 ha, or about a quarter, of the park.

Cardamine impatiens was predominately associated with hedgerow edge habitats. In almost all
other cases it could be found within 10 m of a hedgerow, for example, on ditch and streamsides, in
nearby scrub and on ground that had been disturbed near to hedgerows. C. impatiens preferred the
herbaceous hedgerow edge, but was occasionally found under the hedgerow canopy in the centre
of the hedgerows. The flora and history of the hedgerows has been described by Williams &
Cunnington (1985), and in Williams et al. (1987).

A few plants were found in the adjacent hay meadows, probably as a result of ground
disturbance or poaching of the ground. It was however, unable to survive in hay meadows beyond
the first year. A colony established on an adjacent railway cutting. It survived as a weed in the
garden of one property backing onto a hedgerow. Several plants were found in the drying mud of
seasonal ponds.

In addition to the annual variation in the total population there was considerable variation in the
counts of each colony. The relative variation in the numbers at individual colonies did not
necessarily reflect the overall changes. Three main factors appeared to be responsible for the year
to year variations in the population at each colony. These factors were cyclic variation, the weather
and ground disturbance. In addition, the total population may have increased as a result of the
establishment of new colonies.

At each colony there appeared to be a cycle as the population increased and then decreased. The
duration of these cycles was difficult to estimate due to the other factors involved, but two to four
cycies per decade appeared to be the norm. To some extent the cyclic nature would be expected
from a biennial species, but the cycies could aiso be a facet of the weather-related variation.

The weather was possibly a significant factor affecting the population, particularly when the
populations at most colonies increased or decreased in synchrony. Thus 1994 and 1995 were years
of low populations, while the highest number of plants was recorded in 1997. It is not known
which weather factors were responsible for these effects and at which stage/s in the life cycle. In
1990 many of the plants were smaller than usual, possibly as a result of the dry conditions of that

year. Nevertheless, many plants were flowering even though they were less than 20 cm tall. In |

1995 many of the young rosettes were affected by a late frost.

1888 . 1989 1896 1982 1983 1984 1995 1996 1987 1988

FIGURE 1. Changes in the population of Cardamine impatiens at Fryent Country Park, Middlesex, 1988-1998.

211

VARIATION IN A CARDAMINE IMPATIENS POPULATION

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212 L. R. WILLIAMS

Ground disturbance was responsible for some major fluctuations in populations. During the
investigation ground disturbance was often a result of mechanical works on the park. The effect
was also noted after work by conservation volunteers on hedgerow edges, ponds and streamsides.
This suggested germination of seeds from a buried seed bank and/or from seeds released by
adjacent plants. The evidence for a buried seed bank gains support from the observation that in
some years no plants could be found at some colonies, and yet the colonies would revive in later
years (Table 1).

DISCUSSION
It is accepted that counting errors could affect the results, though the results probably give a

(1991) suggested a number of recording problems during counts of plant populations. Possible
easons for error include reductions in the number of seedlings surviving through the course of the
growing season. Whilst the survey was undertaken in late May (to early June) each year, it is
possible that in a ‘late’ season, more seedlings would have appeared subsequently. Hutchings
(1991) emphasised that the growing plants do not necessarily reflect the size of the population of
any dormant viable seeds in the soil. Indeed, it was obvious from the resuits that the seed bank
olayed an important role in the reappearance of C. impatiens at sites where it had not been
observed for one or more years.

The effect of ground disturbance on the maintenance of populations of C. impatiens was noted
by Stewart et al. (1994) and by Briggs (1990). Where C. impatiens is present in southern
ashwoods, seedlings appear in cleared sites within the year following felling. After three years,
very few plants survive against the competing vegetation. The dormant seed buried in the soil,
however, remains viable for many years. Corner (1988) described the habitat of a population of C.
impatiens in Scotland where it occurred on scree in old woodland and noted that there was some
ground disturbance at that site as a result of cattle and sheep grazing. Sinker et al. (1985) also
noted that C. impatiens was intolerant of competition, but invasive in recently disturbed habitats.
At Fryent Country Park, C. impatiens was recorded on recently re-dug garden soils adjacent to a
hedgerow colony of the plant.

Whilst the species spreads mainly by the explosive release of seeds, it is possible that some of
the new colonies, assuming that they are new, were established by seeds carried by other means.

In the London investigation, C. impatiens was largely associated with the hedgerow edge; a
habitat additional to those listed by Rich (1991), but consistent with the shade preference of this
species. The results suggest that locally, the species is best able to survive in light herbaceous
vegetation, and on recently disturbed or exposed ground. It prefers moderate shade and limited
competition from other plants.

~

REFERENCES

BriGGs, M. (1990). Sussex plant Atlas supplement. Booth Museum of Natural History, Brighton.

BURTON, R. M. (1983). Flora of the London area. London Natural History Society, London.

CORNER, R. W. M. (1988). Cardamine impatiens L.: A native Scottish locality. Watsonia, 17: 91-92.

HUTCHINGS, M. J. (1991). Monitoring plant populations: census as an aid to conservation, in GOLDSMITH, F.
B., ed. Monitoring for conservation and ecology, pp. 61-76. Chapman and Hall, London.

KENT, D. H. (1975). The historical Flora of Middlesex. The Ray Society, London.

RICH, T. C. G. (1991). Crucifers of Great Britain and Ireland. Botanical Society of the British Isles, London.

SINKER, C. A. et al. (1985). Ecological Flora of the Shropshire region. Shropshire Trust for Nature
Conservation, Shrewsbury.

STEWART, A., PEARMAN, D. A. & PRESTON, C. D. (1994). Scarce plants in Britain. J.N.C.C., Peterborough.

WILLIAMS, L. R. (1996). The flora of Fryent Country Park, Middlesex. The London naturalist, 75: 45-66.

WILLIAMS, L. R. & CUNNINGTON, W. (1985). Dating a hedgerow landscape in Middlesex: Fryent Country
Park. The London naturalist, 64: 7-22.

WILLIAMS, L. R., MCLAUCHLIN, J. & HARRISON, T. G. (1987). Hedgerows surviving in suburban Kingsbury.
The London naturalist, 66: 35-39.

WILLIAMS, L. R. & NORTHCROFT, K. (1992). Heritage land: the case for Fryent Country Park. The London
naturalist, 71: 33-42.

(Accepted March I 999 )

Watsonia 23: 213—220 (2000) 218

Notes

UPDATE ON BIRDSEED ALIENS (1985-1998)

In the 13 years since our joint paper (Hanson & Mason 1985), which listed a total of 425
species, of which 318 were actually cultivated by the authors from various birdseed mixtures, I
have continued to examine packets purchased in supermarkets, pet shops, garden centres and street
markets. The less common seeds have been extracted for cultivation and the remainder mixed with
Sunday-roast fat for the garden bird table. Many of the resulting exotic weeds have turned out to
be the usual familiar aliens but several new, upgraded and overlooked records can now be added to
the list of definite birdseed aliens as the sources of birdseed constantly alter due to changing world
commerce.

The number of such additions amounts to a further 44 taxa while the local sewage works has
produced another six “probables”’, i.e. species growing together with known birdseed aliens.

In addition to the above, Clement & Foster (1994) and Ryves et ai. (1995) include another 34
species as possible birdseed aliens although some of these are much more likely than others thus
making a grand total of just over 500 taxa. Continued cultivation experiments will certainly
enlarge the proportion of definite birdseed aliens to more than the 70% of this grand total which
will itself also certainly marginally increase. Any further additions on a major scale to the total
number of taxa now seems most unlikely.

DETAILS OF ADDITIONAL BIRDSEED ALIENS BETWEEN 1985-1997

Cult. cultivated by

CGH C. G. Hanson

FH Mrs F. Houseman

Miiller recorded in Miiller (1950)

gardens recorded in gardens in association with other birdseed aliens
tips recorded on rubbish tips with other birdseed aliens
occasional a few plants seen most years

sporadic single plants appearing irregularly

The following abbreviations are used:
The nomenclature and sequence of families follow Flora Europaea. The genera and species are
listed alphabetically.

Polygonaceae
Persicaria glabra (Willd.) M. G6mez (Polygonum glabrum Willd.): Tips and parks; sporadic.
P. pensylvanica (L.) M. Gémez: Tips; sporadic.

Chenopodiaceae

Atriplex prostrata Boucher ex DC.: Cult. CGH.

A. sagittata Borkh.: Cult. CGH.

Bassia scoparia (L.) Voss (Kochia scoparia (L.) Schrader): Sewage works and tips; sporadic.

Amaranthaceae
Amaranthus bouchonii Thell.: Cult. CGH; tips; sporadic, probably overlooked as A. hybridus.
A. capensis Thell. subsp. uncinatus (Thell.) Brenan: Cult. CGH.

Ranunculaceae
Nigella arvensis L.: Tips; sporadic.
N. sativa L.: Tips; sporadic. Cult. CGH.

Papaveraceae
Papaver dubium subsp. lecogii (Lamotte) Syme: Cult. CGH.

214 NOTES

Crucifereae

Brassica rapa L. subsp. oleifera (DC.) Metzger: Cult. CGH; tips; sporadic.

B. tournefortii Gouan: Cult FH; also observed as a birdseed alien in Belgium (F. Verloove pers.
comm.)

Lepidium campestre (L.) R. Br.: Cult. CGH.

Sisymbrium loeselii L.: Sewage works and tips; sporadic.

Caesalpiniaceae
Senna obtusifolia (L.) Irwin & Barneby: Sewage works; sporadic.

Fabaceae

Glycine soja Siebold & Zucc.: Cult. CGH.

Lotus glaber Miller: Cult CGH.

Lupinus sp.: Cult. CGH from niger seed (Guizotia) but not yet identified.
Onobrychis viciifolia Scop.: Cult. CGH.

Sesbania exaltata (Raf.) Cory: Sewage works and tips; sporadic.

Vicia hirsuta (L.) Gray: Cult. CGH.

V. sepium L. var. eriocalyx Ceiak.: Cult. CGH.

Vigna angularis (Wilid.) Chwi & Ohasmi (Adzuki bean): Cult. CGH.

V. radiata (L.) Wilczek: Cuit. CGH; tips; sporadic.

Umbelliferae
Apium graveolens L.: Cult. CGH.
Conium maculatum L.: Cult. CGH.

Boraginaceae

Amsinckia micrantha Suksd.: Cult. CGH; tips; sporadic. My own specimen at least is this and not
A. intermedia as listed in Hanson & Mason (1985).

Borago officinalis L.: Cult. CGH; tips; sporadic.

Verbenaceae
Verbena officinalis L.: Cult. CGH.

Lamiaceae
Ballota nigra L.: Cult. CGH.

Solanaceae

Lycopersicon esculentum Miller: Cult. CGH; very frequent on tips and sewage farms but not as a
birdseed alien.

Solanum nigrum L. subsp. schultesii (Opiz) Wessely: Cult. CGH; tips; sporadic, probably
overlooked in the past.

S. physalifolium Rusby: Cuit. CGH; tips; sporadic. My own specimen at least is this and not S.
sarrachoides as listed in Hanson & Mason (1985).

Polemoniaceae

Navarretia squarrosa (Eschsch.) Hook. & Arn.: Gardens; sporadic. Reported as a birdseed alien in
Holland.

Sesamum capense Burm. f.: This is probably the species which occurs sporadically at our local
sewage works.

Asteraceae

Anthemis altissima L.: Recorded from a Kentish field with other birdseed aliens after a dressing of
sewage sludge.

A. arvensis L.: Cult. CGH.

Cichorium pumilum Jacq.: Sewage works and tips; sporadic.

Tripleurospermum inodorum (L.) Schultz-Bip.: Cult. CGH.

Xanthium strumarium L.: Miller; gardens; tips; sporadic; cult. CGH

q
i
r

NOTES 215

Poaceae

Bromus diandrus Roth.: Cult. CGH and FH. -

Echinochloa crus-galli (L.) P.Beauv. var. oryzoides: Cult. CGH.

Eleusine coracana (1...) Gaertn.: Cult. CGH; recorded also at Ghent (Belgium) as a birdseed alien
by F. Verloove (pers. comm.). A cultivated millet in tropical Africa.

Lolium x boucheanum Kunth (L. multiflorum x L. perenne): Cult. CGH.

Oryzopsis miliacea (L.) Benth. & Hook.: Cult. CGH and FH; tips; sporadic.

Panicum subalbidum Kunth: Cult. CGH; tips; sporadic.

Paspalum dilatatum Poir.: Recorded from several urban areas with other birdseed aliens.

Phleum exaratum Griseb.: Miiller; reported as a rare birdseed alien at Roeselvare, Belgium 1994
by F. Verloove (pers. comm.).

Rostraria cristata (L.) Tzvelev: Gardens; tips; sporadic; reported as a rare birdseed alien at Ghent,
Belgium in 1994.

Triticum durum Destf.: Miller: cult. CGH; tips; sporadic

T. turgidum L.: Miller; tips and waste ground; sporadic.

Note 1 Suggestions are strong that the following species could well be included as birdseed aliens
but definite proof is still awaited. Confirmation from readers would be most welcome.

Aegilops neglecta Lolium persicum
Amaranthus spinosus Medicago scutellata
Crepis suberostis M. truncatula
Eleusine muitiflora ~ Nigella hispida
Galactites tomentosa Phleum paniculatum
Lathyrus inconspicuus Picris cupuligera
L. ochrus Polygonum bellardii
Leptochloa uninervia Silene conoidea

Note 2 The following aliens are claimed as of possible birdseed vector in the references below
but I have been unable to corroborate them in my studies to date. E. Clement (pers. comm.)
agrees that a significant number appear to be distinctly unlikely and also that several
identification errors are suspected. Confirmation is again welcome.

Agrostis gigantea Cucumis myriocarpus Oenothera canovirens
Anthemis austriaca Euphorbia falcata Paspalum distichum
Anthoxanthum ovatum E. segetalis Physalis ixocarpa
Artemisia annua Gilia capitata Salvia viridis

Bowlesia incana Ayoscyamus albus Schkuhria pinnata
Brachiaria eruciformis Juncus pallidus Scolymus maculata
Carduus thoermeri Lathyrus tingitanus Tetragonolobus purpureus
Centaurea diffusa L. tuberosus Triticum monococcum
Chenopodium missouriense Lepidium virginicum Verbascum phoenicium
Clarkia quadrivulnera Linum grandiflorum Vulpia geniculata
Convolvulus sabatius Malope trifida

ACKNOWLEDGMENTS

Once again I am greatly indebted to Eric Clement for checking the identity of most of the taxa and
to Filip Verloove who has kept me informed of his birdseed discoveries in Belgium.

REFERENCES

CLEMENT, E. J. & FOSTER, M. C. (1994). Alien plants of the British Isles. Botanical Society of the British
Isles, London.

HANSON, C. G. & MASON, J. L. (1985). Bird seed aliens in Britain. Watsonia 15: 237-252.

MULLER, K. (1950). Die Vogelfutterpflanzen. Mitt. Ver. Math. Naturw, Ulm 23: 3-33.

RYVES, T. B., CLEMENT, E. J. & FOSTER, M. C. (1995). Alien grasses of the British Isles. Botanical Society of
the British Isles, London.

C. G. HANSON
I Coltsfoot Road, Ware, Herts.

216 NOTES
HIERACIUM KENTII SP. NOV. (ASTERACEAE)

During our long spell of working on the British hawkweeds, Cyril West and I had to lump together
some species into groups which we were unable to deal with at the time. The largest of these
groups was what we referred to as the Hieracium exotericum aggregate. The two botanists who
have done the most work on this group are Alex Jordan, whose final account was published by A.
Boreau (1857), and N. Hylander (1943). The plant here described as H. kentii belongs to this group
and is particularly distinct in the stellate hairiness of the underside of the cauline leaves. We could
not match it with any of the species in Boreau (1857) or Hylander (1943).

Hieracium kentii P. D. Sell, sp. nov.
HOLOTYPE: Busbridge, Godalming, Surrey, v.c. 17, 28 May 1896, E. S. Marshall in Set of British
Hieracia by E. F. & W. R. Linton, no. 37, as H. murorum var. pellucidum (CGE).

Ab H. grandidenti Dahist. foliis latioribus, pilis stellatis densis in pagina inferiore foliorum
caulinorum, involucri squamis pallidioribus, pilis glanduliferis in involucris pedunculisque
luteonigrioribus distinguitur.

Herba perennis phyllopoda, caudice ramoso. Caulis 40-70 cm altus, pallide luteoviridis, ad basin
saepe purpurascens, stricte erectus, striatus, in parte inferiore pilis simplicibus eglandulosis
numerosis, sursum gradatim paucioribus, mediocribus vel longis pallidis et in parte superiore pilis
stellatis numerosis pilisque glanduliferis numerosis brevissimis vel brevibus luteonigris vestitus.
Folia in pagina superiore mediocriter luteoviridia, in pagina inferiore pallidiora neque
purpurascentia, in paginis ambabus pilis simplicibus eglandulosis numerosis brevibus_ vel
mediocribus, et in marginibus costaque infra longioribus, subrigidis pallidis vestita, caulina pilis
stellatis numerosis vel densis infra vestita; folia basalia pauca vel satis numerosa, exteriora 4—10
cm longa, 2-5-6 cm lata, subrotunda vel late ovata vel late elliptica, ad apicem late rotundata
mucronuiataque vel obtusa, dentibus magis minusve mammiformibus undulatodentata, ad basin
truncata dentium pari basali saepe deflexo, interiora (quando adsunt) 6-10 cm longa, 2-5-5 cm
lata, elliptica vel oblongoelliptica, ad apicem subobtusa vel magis minusve acuta, exterioribus
similiter dentata, petiolis saepe brevibus, sed interdum ad 8 cm longis et pilis simplicibus
eglandulosis numerosis longis pallidis vestitis; folia caulina 0O—1(—2), lanceolata vel ovata, ad
apicem acuta vel acuminata, plerumque acute sed interdum dentibus mammiformibus dentata, ad
basin truncata, sessilia vel breviter petiolata. Inflorescentia corymbosa, saepe ramo inferiore longo
praedita, acladio breviusculo; pedunculi plerumque aliquantum breves gracilesque, pilis stellatis
densis pilisque glanduliferis numerosis brevissimis vel brevibus inaequalibus luteonigris pilisque
simplicibus eglandulosis sparsis brevibus pallidis vestiti. Capitula 30-35 mm diametro, ad basin
rotundata. Involucri squamae ante anthesin incumbentes, 4-9 mm longae, 1-0-1-2 mm latae,
olivaceae, interiores marginibus pallidis, anguste linearilanceolatae, ad apicem magis minusve
acutum gradatim decrescentes, pilis glanduliferis densis brevissimis vel brevibus maxime
inaequalibus luteonigris et secus margines adque basin pilis stellatis numerosis vestitae, sine pilis
simplicibus eglandulosis. Ligulae flavae glabrae. Styli paulo obscuri. Receptaculi alveoli margine
breviter dentati. Cypselae 3-0—3-5 mm longae, rubronigrae.

Phyllopodous perennial herb with a branched stock. Stem 40-70 cm, pale yellowish-green, often
purplish towards the base, strictly erect, striate, with numerous, medium to long, pale simple
eglandular hairs below, which become gradually fewer up the stem, and numerous stellate and
numerous, very short to short, yellowish-black glandular hairs in the upper part. Leaves medium
yellowish-green on the upper surface, paler beneath and not purplish, with numerous, short to
medium, subrigid, pale simple eglandular hairs on both surfaces and longer ones on the margins
and midrib beneath, the cauline with numerous to dense stellate hairs beneath; basal few to fairly
numerous, the outer 4-10 x 2-5-6 cm, subrotund, broadly ovate or broadly elliptical, broadly
rounded and mucronulate to obtuse at apex, undulate-dentate, the teeth more or less mammiform,
and truncate at base with the basal pair of teeth often deflexed, the inner, when present, 6-10 x
2-5-5 cm, elliptical to oblong-elliptical, subobtuse to more-or-less acute at apex, toothed like the
outer, the petioles often short, but sometimes up to 8 cm and with numerous, long, pale simple
eglandular hairs; cauline leaves 0—1(—2), lanceolate to ovate, acute to acuminate at apex, dentate,

NOTES ONT,

usually sharply but sometimes with mammiform teeth, truncate at base, sessile or shortly petiolate.
Inflorescence corymbose, often with a long lower branch, with the acladium fairly short; peduncles
mostly rather short and slender, with dense stellate hairs, numerous, very short to short, unequal,
yellowish-black glandular hairs and occasional short pale simple eglandular hairs. Capitula 30-35
mm in diameter, rounded at base. Jnvolucral bracts incumbent in bud, 4-9 x 1-0—1-2 mm, olive
green, the inner with pale margins, narrow linear-lanceolate, gradually narrowed to a more-or-less
acute apex, with dense, very short to short, very unequal, yellowish-black glandular hairs and
numerous stellate hairs along the margins and at the base, without simple eglandular hairs. Ligules
yellow, glabrous. Styles slightly discoloured. Receptacle pits with margin shortly dentate. Achenes
3-0-3-5 mm, reddish-black. Flowering May and June.

V.C. 13. W. SUSSEX
Shady path near Pitts Hill, August 1918, W. C. Barton (BM).
Bexley Hill, near Lodsworth, 22 June 1957, F. R. Browning & C. West (CGE).

V.C. 14. E. SUSSEX

Bexley Heath above Eastbourn, 10 August 1912, C. C. Lacaita (BM).

Mickleham, 8 June 1938, H. W. Pugsley (BM).

Roadside east of Holtye Common, near East Grinstead, 30 June 1958, B. A. Miles (CGE).

V.C. 16. KENT

Tunbridge Wells, 1855, A. G. More (BM, CGE).

Wood near Shoreham, 29 May 1922, L. B. Hall (BM).

Near Ightham, 11 June 1938, F. A. Browning (BM).

Ashurst Park, 30 August 1942, H. W. Pugsley (BM); June 1951, C. West (CGE).
Shadwell Wood, near Sheldhurst, 6 June 1943, H. W. Pugsley (BM).

V.C. 17. SURREY

Banks ascending to Hindhead, 4 June 1889, E. S$. Marshall 479 (BM, CGE).

Church Lane, Whitley, 7 June 1890, F. J. Hanbury (BM); 28 June 1958, B. A. Miles (CGE).
Whitley, 23 May 1896, E. S. Marshall (BM, CGE).

Busbridge, Godalming, 28 May 1896, E. S. Marshall (BM, CGE).

Under trees, White Hill, near Caterham, 21 June 1906, C. E. Salmon (BM).

Wotton, 12 July 1916, C. E. Britton (BM).

Margery Wood, Colley Hill, 8 June 1919, C. E. Salmon (BM).

Pilgrims Way, Caterham, 28 June 1932, C. E. Britton (BM).

Roadside on Gravelly Hill towards White Hill, Caterham, June 1948, C. D. Pigott (CGE).

I am very pleased to name this plant after Duggie Kent. Hieracium was not a genus he was
particularly interested in, but he spent a large amount of time searching out original publications
for me so that we could get their nomenclature straight. His contribution to British botany was
prolific over many years and I am personally particularly indebted to him for meticulously reading
the proofs of volume 5 of Flora of Great Britain and Ireland. | am indebted to Philip Oswald for
supplying the Latin description and diagnosis and to David McCosh and Chris Preston for sorting
out and arranging the loan of specimens from the BM. A complete revision of the British Hieracia
will appear in volume 4 of Flora of Great Britain and Ireland and is almost finished.

REFERENCES

BOREAU, A. (1857). Flore du centre de la France, 3rd ed. Roret, Paris.
HYLANDER, N. (1943). Die Grassameneink6mmlinge Schwedischer Parke mit Besonderer Beriicksichtigung
der Hieracia silvaticiformia. Symbolae Botanicae Upsalienses 7: 106-274.

PD SEL
Dept. of Plant Sciences, University of Cambridge, Downing St, Cambridge CB2 3EA

218 NOTES

CHARA BALTICA BRUZ. (CHARACEAE), BALTIC STONEWORT, REDISCOVERED
IN HICKLING BROAD, NORFOLK 5

The stoneworts are a small group of unusually complex aquatic green algae which make up the
family Characeae. Baltic Stonewort, Chara baltica Bruz., is one of 16 species on the British Red
List (Stewart 1996), and was once widely distributed around the coast of Britain, with records
from 14 sites in Cornwall, Devon, Kent, Norfolk, Anglesey and the Western and Northern Isles of
Scotland. The species has declined in recent years, and there are only six modern records, some of
which are doubtful and need confirmation (N. Stewart, pers. comm.). Its sole site in Norfolk is
Hickling Broad, which is the richest site for stoneworts in the UK.

The earliest documented record of C. baltica at Hickling was in 1898 by G. R. Bullock-Webster,
one of the pioneer workers on charophytes of the British Isles. Bullock-Webster recorded the
species again in 1899 and 1904 (specimens at NMW) and in 1902 (specimen in BM), and noted
that the plant had “occurred freely for a number of years in company with Chara hispida and
Chara aculeolata (now Chara pedunculata) in the northern area of Hickling Broad” (Groves &
Bullock-Webster 1924a). Although it was abundant around this time, C. baltica was not observed
in subsequent visits to Hickling Broad prior to 1924 (Groves & Bullock-Webster 1924b). There
are sporadic reports of C. baltica for the middle of the century; one is attributed to E. A. Ellis in
the mid-1940s (Ellis 1965), and G. H. Rocke recorded it in 1955 and 1956 (Wallace 1956, 1957).
A sample of the material collected in 1956 is also in BM. The last record of this species at
Hickling was made by Rocke in 1965 (Petch & Swann 1968). All historical records are for the
variety rigida (originally described by Bullock-Webster) of which Hickling is the type and only
known locality. The distinctive characters are supposed to be its firm, rigid habit, comparatively
long internodes and patent (standing out at right angles) spine cells (N. Stewart, pers. comm.).

Hickling Broad was known to be dominated by stoneworts for the first 60 years of the 20th
century, with 15 species being recorded. Water quality subsequently deteriorated due to
eutrophication, increasing salinity and accelerating rates of organic sedimentation. The main
causes were nutrient input from a large winter roost of Black-headed Gulls in the late 1960s, and
deep drainage of the surrounding arable land which breached the saline water table (George 1992).
The changes in water quality resulted initially in a periodic explosion in growth of filamentous
algae, especially Cladophora sauteri (Nees von Eisenbeck ex Kiitzing) Ktitzing, and the
replacement of the charophyte-dominated communities by waterweeds typical of the more fertile
conditions.

The 1965 record of C. baltica in Hickling Broad was prior to the near total disappearance of
aquatic macrophytes and the shift to turbid, phytoplankton-dominated water which accompanied
further nutrient enrichment. Charophytes were rarely recorded from then until the late 1970s, when
they were an occasional component of the submerged plant communities (Phillips & Moss 1978).
By the 1980s, nutrient levels were decreasing due to the decline in the size of the gull roost and
progressive washing-out of the residual gull guano from the Broad’s sediment.

It is uncertain whether C. baltica continued to survive at very low densities in Hickling after
1965, but the species was not detected by the Broads Authority’s annual macrophyte surveys
which commenced in 1982. It may, however, have been overlooked, since these surveys sample
only selected transects across the Broad.

In 1995 the Broads Authority implemented a weedcutting regime for Hickling to remove Spiked
Water-milfoil Myriophyllum spicatum, which was impeding movement of boats across the open
water. Intensive sub-aquatic monitoring of the aquatic macrophytes (including charophytes) is an
integral part of the weedcutting programme and has yielded much detailed information about
species distributions, growth characteristics, physical structure and composition of the plant
communities. In June 1998, C. baltica was rediscovered in the northern arm of the Broad which
had not been monitored since 1994. This location is consistent with the historical records which
were also for the “northern area” (Groves & Bullock-Webster 1924a, b) and suggests that oospores
of this species may have survived in the sediment from the 1960s. C. baltica was found to be co-
dominant with the Intermediate Stonewort Chara intermedia, forming dense, closed lawns up to
0-8 m tall. The species was only associated with soft, organic sediments and was growing at depths
of 0-8 to 1-2 m. Oogonia and antheridia were abundant in June and July on branchlet nodes at the
surface of the charophyte lawn which experienced the highest light levels. This species is almost
certainly perennial, overwintering with Chara intermedia. Other occasional associates were Chara
connivens, Nitellopsis obtusa, Potamogeton pectinatus, Najas marina and Myriophyllum spicatum.

NOTES 2119

C. baltica has been recorded predominantly from standing water bodies up to 7 m deep and is
usually associated with sandy substrates (Stewart & Church 1992). It is restricted to sites near the
coast, which suggests a requirement for salt, and in the Baltic Sea it is known to tolerate salinities
up to 1-8% (approximately 10, 000mgCl 1') (Olsen 1944). Salinity levels in Hickling Broad ranged
between 1,000 and 2 400mgCl l' for the period 1978 to 1998. Total phosphorus concentrations
were approximately 0-1mgP I’ from 1980 to 1990, but showed a steady fall to about 0-05mgP I’!
by summer 1998 (Schutten & Davy 1998). There has also been a gradual increase in water clarity
in Hickling Broad during the 1990s, culminating with a switch to clear water in summer 1998.

Further surveys are needed to determine whether C. baltica maintains its current pattern of
abundance and distribution in the northern area of Hickling Broad, or whether it appears only
sporadically. Experience has shown that it is sometimes difficult to distinguish sterile material of
this species from long-spined plants of C. intermedia. The two species are, however, readily
separated by the larger size of the antheridia of C. baltica, which are over 0-5 mm in diameter.

ACKNOWLEDGMENTS

I am most grateful to Nick Stewart for confirmation of C. baltica, historical records and much
helpful advice. Thanks are also due to Dr Martin George for providing historical information about
Hickling Broad.

REFERENCES

ELLIS, E. A. (1965). The Broads. Collins, London.

GEORGE, M. (1992). The land use, ecology and conservation of Broadland. Packard, Chichester.

GROVES, J. & BULLOCK-WEBSTER, G. R. (1924a). Notes on the British Charophyta. Journal of botany 62: 33-
3i5

GROVES, J. & BULLOCK-WEBSTER, G. R. (1924b). The British Charophyta, Vol. 2. Ray Society, London

OLSEN, S. (1944). Danish Charophyta. Chorological, ecological and biological investigations. Det
Konggelige Danske Videnskabernes Selskab. Biologiske Skrifter 3(1): 1-240.

PETCH, C. P. & SWANN, E. L. (1968). The Flora of Norfolk. Jarcold, Norwich.

PHILLPS, G. L. & Moss, B. (1978). The distribution, biomass and productivity of submerged aquatic
macrophytes in the Thurne Broads, Norfolk, 1975-1977. Report to the Nature Conservancy Council.
SCUTTEN, J. & DAvy, A. J. (1998). Hickling clear; can we understand why? Technical Research Report

98/02. University of East Anglia, Norwich.
STEWART, N. F. (1996). Stoneworts — connoisseurs of clean water. British wildlife 8: 93-99.
STEWART, N. F. & CHURCH, J. M. (1992). Red Data Books of Britain and Ireland: Stoneworts. J.N.C.C.,
Peterborough.
WALLACE, E. C. (1956). Plant records. Proceedings of the Botanical Society of the British Isles 2: 26-44.
WALLACE, E. C. (1957). Plant records. Proceedings of the Botanical Society of the British Isles 2: 245-268.

J. E. HARRIS
Kepwick Cottage, Wymondham Road, East Carleton, Norfolk, NR14 SJB

220 NOTES
THLASPI PERFOLIATUM L. NEW TO WORCESTERSHIRE (V.C. 37)

On 13 April 1999 whilst recording near the Gloucestershire border for the forthcoming Flora of
Worcestershire, KB found a large population of a Thlaspi species on the edge of a remote arable
field, which was tentatively identified as T. perfoliatum. The identification was subsequently
confirmed by TCGR and two specimens have been deposited in the National Museum & Gallery
of Wales herbarium (NMW). This is the first confirmed record for Worcestershire (v.c. 37). The
only previous record for Worcestershire sensu lato is an undated one from Evenlode (c. SP/22.29)
by W. Cheshire (Lees 1867), which at the time appears administratively to have been a detached
parish of Worcestershire in East Gloucestershire (v.c. 33).

This is a remarkable discovery of a rare, declining and statutorily protected species, hitherto
surviving in only nine native and three introduced sites in Britain (Rich et al. 1998). The locality is
about 20 km from the nearest sites in the Cotswoids, situated in an otherwise unremarkable area
with many arable fields and straight, hawthorn-dominated hedges on a low ridge of Blue Lias. The
soil was derived from the Blue Lias bedrock with loss of the upper horizons due to ploughing and
soil creep, and formed a shallow, stony, greyish clay soil pH 7-3 (measured in a 50:50 slurry with
distilled water using a Hanna pHep2 meter). Rich et al. (1989) noted T. perfoliatum was generally
only persistent for long periods of time on Oolite Limestone soils, though it had persisted on
similar formations (including Blue Lias) elsewhere.

The arable habitat is unusual in Britain, the species only having been recorded in arable sites
four times previously (Rich et al. 1989). The plants were exceptionally tall and robust, probably
resulting from fertiliser. They occurred both in the crop and on the adjacent bank, which
presumably acts as a refuge when unsuitable crops are grown. The bank was 1-1-5 m tall
indicating a long history of ploughing and soil creep at the field margin. Associated species in the
arable crop were Convolvulus arvensis, Polygonum aviculare, Tripleurospermum inodorum and,
on the bank, Anisantha sterilis, Arrhenatherum elatius, Bromus hordeaceus, Knautia arvensis,
Lamium purpureum, Malva moschata and Senecio vulgaris.

The population was estimated to be about 5000 plants, most of which occurred in an area of c.
77 m x 3 m on the edge of the arable field, but significant numbers were also present along an
adjacent bank. This is thus the third or fourth largest population in Britain (Rich et al. 1998).

The information available to date gives no obvious reason to suspect it is an introduction, and
the site is accepted as native. It is close to the known native range on similar geology and soils,
and the landscape context suggests it could have survived in situ from the Enclosure of this part of
Worcestershire. There were no obvious alien species present, and the large population indicated it
had been present for some time. The location has been notified to Plantlife, English Nature and the
Worcestershire Trust for Nature Conservation, and efforts are being made to conserve the site.

In 1998, a few plants of 7. perfoliatum were also reported from the Gloucestershire Trust for
Nature Conservation Reserve of Cutsdean Quarry. They are rumoured to have been sown (pers.
comm. C. Studholme to M. and C. Kitchen 1998), but the person responsible is unknown as is the
origin of the seed. Plants were last recorded here in 1967 for certain and possibly in 1971, and
were lost when Buckle Street was realigned along the west side of the quarry (S. C. Holland, pers.
comm., 1988). No plants were seen in repeated searches in recent years by TCGR and co-workers
(1983, 1986, 1992, 1994 and 1996). In 1999, there were six plants scattered in the middle of
deliberately created bare open banks.

REFERENCES

LEES, E. (1867). The botany of Worcestershire. Worcester Naturalists’ Club, Worcester.

RIcH, T. C. G., KITCHEN, C. & KITCHEN, M. A. R. (1989). Thlaspi perfoliatum L. in the British Isles:
distribution. Watsonia 17: 401407.

RICH, T. C. G., LAMBRICK, C. R., KITCHEN, C. & KITCHEN, M. A. R. (1998). Conserving Britain’s
biodiversity: Thlaspi perfoliatum L. (Brassicaceae), Cotswold Pennycress. Biodiversity and conservation
7: 915-926.

K. BARNETT
49 Hastings Road, Malvern, Worcestershire, WR14 2XE

| T. C. G. RICH
Dept. of Biodiversity and Systematic Biology, National Museum & Gallery, Cardiff, CF 10 3NP

Watsonia 23: 221—230 (2000) 271

Book Reviews

A Flora of Norfolk. G. Beckett and A. Bull pp. 320. G. Beckett 1999. Price £38.00. ISBN 0-
9534999-0-1

Just when recent county Floras, such as Hampshire and Cumbria, had set new standards in content,
mapping and presentation that seemed unlikely to be easily surpassed, along comes this new Flora,
covering all political and Watsonian areas of Norfolk and incorporating bryophytes as well as
vascular plants and ferns. It is a superbly integrated work, produced over a relatively short period
at a price I consider quite acceptable.

The introductory chapters have a particularly valuable section on man’s influence on the various
habitats, showing a deep and welcome understanding of the subject. This is supplemented by
chapters on habitats using the National Vegetation Classification (NVC) and on soils, reminding
readers of the wealth and diversity of habitats, particularly wetlands, created by drift deposits,
calcareous and acid, overlaying chalk. There is an excellent section on Norfolk botanists which
manages to place predecessors into context rather than just be a recital of names.

These introductory chapters end with an excellent and welcome map, an unaccountable rarity in
modern Floras. Because of this there is no gazetteer, which would have been useful if only to
locate the places named in the illustrations. It would also have been useful to have had an
indication of where political and vice-county boundaries differ.

The species accounts cover the bulk of the book; vascular plants and ferns, 215 pages, and
bryophytes 22 pages. These are supplemented by maps on a tetrad scale for the plants and on a 10
km scale for the bryophytes, showing only records made during the survey (1985 to, I assume,
1998). This latter is my main reservation over the book — the complete lack of older records on the
maps, although declines are often noted in the text. By and large I feel that “snapshot” floras,
mapping only those species and sites seen during the survey, omit an important dimension by not
illustrating change and it is this that is a weak point in recent Floras, of, say, Kent and Wiltshire.
Purists might argue that there is a degree of difficulty in allocating tetrads to old records. I think
this problem is overstated. In this flora historical maps of species such as Lycopodiella inundata,
Lathyrus palustris, Trifolium ochroleucon and many others would have added much to the text. A
minor point would have been a wish that the authors had resisted temptation and stuck to their
original 1987 start date — to tie in with the BSBI Monitoring Scheme and the new Atlas 2000.

A further concern is the extent of incorporation of published records. I noted, following a query
on some aquatic plants, that records on Potamogeton compressus, in Scarce Plants in Britain
(Stewart et al. 1994) and on Potamogeton X cooperi, in Aquatic Plants in Britain and Ireland
(Preston & Croft 1997) were not included. Also, it did not seem that extensive surveys by C.
Doarkes (on aquatic plants) or R. J. Driscoll (on Luronium and other subjects) had been used.

The text of the accounts is first rate, and is one of the few I would use when writing ecological
accounts on a countrywide scale for the new Atlas 2000. There is also a very welcome injection of
horticultural knowledge, particularly in the accounts of aliens. The germination details are also
very useful and rarely encountered elsewhere. Relatively unimportant criticisms here would
include a frequent lack of any comment on native or alien status, numerous errors and omissions
(over 25) in the national status of rare plants, and occasional difficulties in tying up tetrad
frequencies with the text for the unmapped species. The coverage of Brambles is absolutely first-
class, and not at all too long.

The maps and the superimposing of the tetrad distribution over coloured soil types where
relevant, or over rivers or roads for aquatic or wayside species, are really elucidating and pleasing
to the eye. They add explanation to the distribution of all but a very few species. I found it
difficult, by eye, to assign a tetrad dot to a precise grid reference and have made myself a little
overlay. I wish there were more maps of aliens, but it may be that the authors considered those
they mapped told a story, whereas those they omitted did not. Since tetrad totals are given for the
species not mapped it would have been useful to have had totals for those mapped. There are many
more records, particularly in East Norfolk, of species that were mapped in Scarce plants in Britain

222 BOOK REVIEWS

(Stewart et al. 1994). Finally the colour illustrations are first-rate, and incorporated into the text
with very little loss of quality — a most pleasing arrangement, and a great improvement over the
traditional and obligatory block.

[t is not often that my non-botanist friends enthuse over the county Floras that they see strewn
around our dining room. A Flora of Norfolk is a rare exception, with the species pages setting new
standards in presentation and integration of maps, plates and text. It is instantly accessible, with
the coloured underlays of soil types and rivers and roads a real bonus. Just as its predecessor was
the first to carry colour (Flora of Norfolk, C. P. Petch and E. L. Swann, 1968), so this takes us into
the computer age in new ways that must arouse admiration and trepidation in equal measure in
those who would emulate in other counties.

D. A. PEARMAN

Jilustrations of the British and Irish orchids. D. M. Turner Ettlinger. Pp. 214. Published privately
by the author (Royden Cottage, Cliftonville, Dorking, Surrey). 1998. Price £24.50 + £1.10 p & p.
ISBN 0—9530380-1-7.

My recent review in Watsonia (22: 201-202) of what I termed “an iilustration-free iconograph” by
Derek Turner Ettlinger (Notes on the British and Irish Orchids, 1997) has, thanks to the fiscal
generosity of an insightful uncle, now been joined by an illustration-rich, genuinely iconographic
companion. This welcome A5-sized volume consists of 519 well reproduced photographs
organised in 100 colour plates — the images, drawn from a vast reservoir, are excellent. Their
primary role is to document the wide range of morphological variation observed within the 49
bona fide species recognised as native by the author; coverage varies from one plate for the
cisappoitingly uniform to five for notoriously variable species such as Dactylorhiza incarnata
and D. fuchsii, plus E. helleborine if considered to include the genetically undifferentiable E.
“youngiana.” Colour morphs, achlorophyllose plants and, where possible, pollinating insects are
included, supplemented with a few “accidental” and extinct species.

Opposite every plate is a page of information relating to specific photographs; each carries
approximate locality, habitat, date, and magnification for close-ups. Here, much space is wasted
and the opportunity to highlight diagnostic characters of the taxa is largely eschewed. Rather,
brief, footnote-style commentaries are occasionally provided, most relating to conservation sensu
lato; examples include under- and over-grazing, de- and re-afforestation, hoof-poaching, draining,
vandalism, landfill, development, over-collection and re-introduction.

Most taxonomic insights from recent DNA studies (e.g. Pridgeon et al., Lindleyana 12: 89-109,
112-141, 1997) are supported in principle, though in practice the classification of Ettlinger (1997)
is followed for consistency (if not brevity, given that it includes 72 often-obscure varieties). The
taxonomy is especially brave when dealing with critical genera such as Epipactis and
Dactylorhiza. One exception to the uniformity between the two volumes is the upgrading of
Stace’s (New Flora 2nd ed., 1997) three “varieties” of G. conopsea to full species on the basis of
extensive (albeit mostly unpublished) molecular data, though they remain notional subspecies in
the associated account of hybrids. Flicking between selected plates can be instantly informative;
for example, it usefully reinforced my prejudices regarding the synonymy of the orchidological
“Celtic fringe” represented by the Scottish Dactylorhiza majalis “ssp. majaliformis” and the
Welsh “ssp. cambrensis.”

The sparse treatment of hybrids (focusing on those between distantly related species) and
omission of the many wonderful teratologicai “hopeful monsters”, each a potential species in its
own right (Bateman and DiMichele, in D. S. Ingram & A. Hudson, eds., Shape and form in plants
and fungi, 63-102, 1994), lead this reviewer to hope that a third volume may one day emerge. In
summary, like its predecessor, this book could be viewed as targeted primarily at committed
orchidophiles. However, I trust that its extensive use of colour will encourage the wider readership
that it merits.

R. M. BATEMAN

BOOK REVIEWS 223

The Box Hill Book of Box. Lalage Grundy. Pp 334. Friends of Box Hill, Dorking. 1998. Price
£3.00*. ISBN 0—9534430—0-0

Box Hill in Surrey was one of the earliest properties of the National Trust, acquired at the turn of
the century. In spite of its enormous popularity, it retains a wildness and a rich and varied flora and
fauna, including many rarities. Of these the eponymous box (Buxus sempervirens) is locally so
plentiful that its special interest deserves to be drawn to the attention of visitors. This attractive,
well illustrated and informative booklet with a superb colour photograph on its front cover will
surely fulfil this purpose.

The booklet is, however, much more than a glossy advertisement. It is written in an engaging
style, which should make it accessible to a wide range of people, and it is full of facts about the
botanical interest of box: its relatives world-wide, its toxicity to mammals and the specialised
invertebrates which nevertheless feed upon it, its extraordinary smell, its geography, ecology,
history and its uses to man. The wood of box is valuable and from time to time the trees on Box
Hill have been cropped and there is evidence that some areas on Box Hill are old plantations. For
this reason it has often been questioned whether a sub-Mediterranean species is truly native so far
north, even though a place-name, Boxlands, shows that it was a significant feature in the
fourteenth century. There is, however, a remarkable feature of Box Hill which makes the native
status much more plausible: the river Mole undercuts the south-western slope to produce a very
steep curved amphitheatre facing the afternoon sun. Here, as the author points out, temperatures
can rise to 40°C! On this slope, known as the Whites because of its long-established stripes of
eroded chalk, box thrives and is dominant over several hectares and forms, with yew and privet, a
sombre dark green scrub which is unique in Britain. One of the beautiful photographs shows this
special place with its two sentinel whitebeams.

This type of booklet has great value, both educationally and for adding to the visitor’s
enjoyment, and, as a credit to the National Trust who protect Box Hill, it mirrors the handsome
guide-books which are provided for historic houses. Box is of course unusual because it is a rarity
that is not endangered by advertisement but the Friends of Box Hill and the author are to be
congratulated on setting a precedent which should surely be followed with attractive guides to the
geology, vegetation and animals of other open-space properties.

C. D. PIGOTT
(* obtainable from the Friends of Box Hill, Pixham Mill, Pixham Lane, Dorking, Surrey, RH4 1PQ at £3.50 to
include postage and packing.)

Three-language list of botanical name components. A. Radcliffe-Smith. Pp. vi+ 143. Royal Botanic
Gardens, Kew. 1998. Price £9.99. ISBN 1—900347-50-4.

In 1675 John Ray produced his Dictionariolum trilingue, a vocabulary of physical, biological and
other terms under 32 headings in English, Latin and Greek, mainly for the use of schools, best
known today through the Ray Society’s facsimile edition of 1981. This might well have been the
inspiration for the present list, but Mr Radcliffe-Smith mentions in his introduction that he heard
of it only in 1984, the year after he embarked on a 16-part list of Greek “roots or elements”, with
their Latin and English equivalents, published in Kew’s Nomenclatural forum. This formed the
basis of his new list (or rather lists, for, thanks to “the amazing alphabetizing ability of the
Spreadsheet’, the list is repeated with the Latin roots first and then with the English meanings
first).

The original list was apparently compiled “in order that mixtures of Latin and Greek in new
compound-coinages might be avoided”, but “an additional purpose, namely to help its users
understand the meanings of Latin and Greek compound-coinages already in the literature”, seems
of wider relevance, at least for the ‘Greek-first’ and ‘Latin-first’ sections. The main lists, it should
be noted, are of ‘building-blocks’ for compound words. Thus “poly- multi- many” and “-
phylla -folia -leaved” allow the creation of the specific epithets polyphylla and multifolia but not
multiphylla or polyfolia. But it is nowhere explained that in adjectival terminal elements the lists
usually give precedence to the feminine form: Lupinus (masculine) must be polyphyllus. There is a
strong bias towards initial elements: for example, I looked in vain for “-prasum’” (as in the specific
epithets of three Allium species) and ‘“-scordum’” (as in the names of the related genera
Nectaroscordum and Nothoscordum).

224 BOOK REVIEWS

There follow plant-name and numerical supplements, each again in three sections. It would have
been easier to find one’s way about the book if the resulting nine sections had carried running
heads. The first supplement contains 297 Ancient Greek plant-names for which plausible Latin and
English equivalents can be found, not always according with modern botanical usage (e.g. “daphne
laurus laurel”). The second is a mixture of indispensable roots such as “proto- primi- first” and
“amphi- bi- two’, useful extras (e.g. “didymo- gemini- twin’), and impossibly complex
monstrosities like “diplasiepitetrapempto- duo et quattuor partes [not in fact an element usable in
a botanical name, even supposing one wanted to!] two and four-fifths”.

It was wise to transliterate the Greek (unlike Ray) “to keep costs down’, but surely also to be
accessible to present-day readers, but crazy to do it so literally, with “k” for kappa, “ai” for alpha
iota, and “gch” for gamma chi (usually but not consistently). The Romans invented a consistent
system of transliteration, which is generally used in botany: thus “chamaikissos” (groundivy)
would be better written as “chamaecissos”’, “orobagche” (broomrape) as “orobanche”, and
“sogchos” (sowthistle) as “sonchos”’.

Another mistake was to present the second and third sections of each part as exact replicas of the
first (apart from their word order). This has produced five successive entries for “cup” followed by
four for “curl”, with much repetition in the Latin column also, because of the richness of the Greek
language. (The Greeks often had several words for it!) Again, because “tip” appears in the ‘Greek-
first’ section only as a synonym for “point” (under “-acme’’), it cannot be found alphabetically in
the ‘English-first’ section, while there are ten entries there for “point”. So much for that “amazing
ability of the Spreadsheet’! The moral is surely not to trust a computer to operate without close
supervision.

One final grumble: in the plant-name supplement, 13 Latin names have abbreviated (but
unexplained) generic names. Especially odd are “p. cerasus” and “pr. cerasus”, with different
English and Greek equivalents and separate “alphabetized” places in the “Latin-first’ section.

But these are all things which, with a little human interference, could easily be set right in a
second edition. Mr Radcliffe-Smith deserves warm congratulations on his stupendous labour and
the Royal Botanic Gardens on a tidily presented book at a very modest price.

P. H. OSWALD

The Plants of Nottingham, A City Flora. P. Shepherd. Pp. v + 76. Wildtrack Publishing, Sheffield.
Price hbk. £15.00, pbk. £8.00. ISSN 1354 0270.

This is a source of much interest and information. Accounts are given of Nottingham’s very varied
geology and of the impact of the urban environment on plants. There are descriptions of the City’s
habitats and vegetation, following, as far as possible, the National Vegetation Classification. The
final chapter lists, in Kent order, the 700 plant taxa recorded since 1985. Localities are indicated
but no grid references given.

There are, however, numerous mis-spellings, (including the rather apt “ravishes” of Dutch elm
disease), ambiguities and inaccuracies in the text, for example, Saxifraga granulata is said to be
“Restricted to Church Cemetery and Forest Recreation Ground .....” and then four more localities
are given; Cochlearia danica 1s said to be alien but some known non-natives are not so designated;
Sagittaria sagittifolia is called Marsh arrowgrass — and are all the city’s railways really laid on
clinker and ash instead of the more usual ballast?

At least two cited references are missing from the list. Two sketch maps have no scale and the
rivers are represented differently on-all three maps. There is repetition; “Calcicole” is defined
twice, though not on first appearance, (but we are never told what a forb is, nor what is meant
when Fumaria officinalis is described as a “widespread segetal species of flower beds, cultivated
ground and recently landscaped areas”) and I wonder if it is necessary to give both English and
Latin names each time a plant is mentioned. I would have preferred to have the parentage of
hybrids given in the text and to have the major roads shown on the maps.

This very useful publication would have been greatly improved by more rigorous editing but,
nonetheless, I look forward with eagerness to the promised historical account of the flora of
Nottingham.

A. BURNS

BOOK REVIEWS 225

John Lindley 1799-1865 gardener-botanist and pioneer orchidologist. Edited by William T.
Stearn. Pp. 232. Antique Collectors’ Club in association with the Royal Horticultural Society,
Woodbridge. 1999. Hardback. Price £29.50. ISBN 1—85149-296-8.

John Lindley is probably most familiar through his association with the Royal Horticultural
Society. The Lindley Library has his own library at its core, while the Lindley Medal is awarded
for exhibits of special scientific or educational interest at RHS shows. His name is also linked to a
few of our native plants, including the eponymous Rubus lindleianus, and many more garden
plants — he was responsible for such generic names as Chaenomeles, Photinia, and Victoria.
However it is in the world of sumptuous, exotic orchids that he is most notable.

John Lindley was born at Catton near Norwich on 5 February 1799, the son of a nurseryman and
pomologist — this symposium volume is published to mark that bicentenary. He was a
contemporary of William Jackson Hooker, and his life overlapped with that of Sir James Edward
Smith. This Norwich-born triumvirate dominated English botany through the early nineteenth
century. While Smith’s The English flora (1824-1828) stuck doggedly to the obsolescent,
artificial, Linnaean system of classes, Hooker in his Flora Scotica (1821), with Lindley’s help, and
later Lindley himself, championed natural orders (families) as a better method for classifying
flowering plants. Lindley, appointed Professor of Botany at the University of London in May
1828, went so far as to forbid his students from using any of Smith’s botanical texts because of the
archaic classification system, and so he published A synopsis of the British flora; arranged
according to the natural orders ... in 1829 (new editions appeared in 1835 and 1841).

This celebratory book contains six original essays. The first is an invaluable biography by
Professor William T. Stearn, incorporating information from the hitherto obscure archive of family
papers now in British Columbia (these papers are the subject of a brief essay by Kathryn Bridge).
Lindley the botanist, perpetually writing and publishing, is interwoven with Lindley the family
man, administrator and inter alia defender of Kew Gardens. Dr Phillip Cribb discusses Lindley’s
contribution to orchidology, concluding that he really was the founding father of orchid taxonomy.
To illuminate this, the book is illustrated with 35 colour plates of orchids — there are also six
colour plates showing plants from other families. Chris Brickell expands on Lindley’s career as a
horticulturist, and Professor W. G. Chaloner discusses The Fossil flora of Great Britain (1831-—
1837) that was jointly produced by Lindley and William Hutton. Dr Brent Elliot, the present
Librarian of the Lindley Library, describes the history of the library and John Lindley’s own
books. A history of the Lindley Medal by William L. Tjaden is reprinted from Archives of natural
history. Miss J. M. Allford’s bibliography of John Lindley, compiled in 1953, forms the tenth
chapter and, to quote Stearn, “graphically demonstrates [his] astonishing industry”. Yet this
bibliography of 238 works is unlikely to be complete as it takes no account of Lindley’s 20 years
as editor of Gardeners’ chronicle which he co-founded. Two of Lindley’s lectures, delivered at the
University of London in 1824 and 1834, are also reprinted.

All told, this is a scholarly, handsomely produced tribute to a botanist whose studies and
publications become even more astonishing when it is remembered that, as a boy, he lost the sight
of one eye.

E. C. NELSON

Alternative Agriculture. A History. From the Black Death to the Present Day. Joan Thirsk. Pp.
365. Oxford University Press. 1997. Hardback £25.00, ISBN 0—19-820662-3.

This book can be taken as precisely what its title suggests, a scholarly examination of a rather
obscure aspect of social history. The author, now President of the British Agricultural History
Society, was formerly a Reader in History at Oxford. It came to me highly recommended by a
fellow botanist, and I looked forward to it as a useful reference book, a help with questions about
the historic status of plants. We all now accept that our precious and declining arable weeds must
largely have arrived from the continent as crop seed impurities, and that plants from monkish
gardens and apothecaries’ satchels become long-term denizens, so I certainly hoped that
Alternative Agriculture would be useful. I had not anticipated a real delight, a book so rich in
interest and information that the writer has joined Pennington, Rackham and Salisbury as one of
the authors who has changed my life.

226 BOOK REVIEWS

Dr Thirsk writes with a sparse donnish elegance. Each chapter is concise, factual, lucid, and
immaculately referenced. Her general argument is that agriculture follows historical cycles.
Mainstream agriculture, concentrating on producing beef and corn, reaches a stage when supply
exceeds demand, prices drop, and farmers then have to diversify. This thesis is extremely topical,
as since the 1980s Britain has been entering an alternative phase, and it is both fascinating and
reassuring to read that the current “ruin” moans of farmers have been uttered “at least three times
before in our documented history’. The phases of alternative diversity have been roughly 1350-—
1500, 1650-1750, 1879-1939, and we are now in the fourth phase. A glance at the popular press
can confirm this current trend, as crops such as borage, linseed, evening primrose and calendula
have been attracting notice.

For instance oilseed rape, castigated since the 1970s as a modern blight on the landscape, and
currently even more of a bugbear as part of the GMO controversy, was actually introduced in the
1560s, and was grown for some 300 years before going out of fashion in the 1880s.

Some of the species involved, such as woad, have direct relevance to county recording, as they
survive as rare denizens, and some fascinating possible lines of spread (and countries of origin)
can be postulated from this book. Others such as madder and saffron are gone, but remain in place
names and folk lore. (Saffron caused my only complaint: Latin names are not always given, and I
had a panicky little search of other references before I was sure that Crocus sativus was the plant
in question, not Colchicum autumnale). The only direct botanical reference in the bibliography is
Syme’s English Botany 1863-86, which isn’t perhaps quite familiar to most of us, but the
botanical (and in particular horticultural) information is accurate and to the point. Both native
plants, such as teasels or weld Reseda luteola, grown commercially for a time to supply changing
industries or trade shortages, and more exotic species such as safflower or vines are discussed. The
accompanying details are one of the great pleasures of the book, and often illuminate historical
county records.

Not only is this a clear and readable account of the characteristic crop plants in the British
landscape for more than six centuries, but the material for the book is enriched by a multitude of
ripping yarns. Diversification produces entrepreneurs, people of vision, imagination, daring, and
the new crops they tried, the deals, journeys, tithe wars, quirks of fate make wonderful reading.
The larger patterns of wars, peace, international trade, disease and immigration are clearly
presented, but within this matrix individual stories have entrancing life. For instance early in the
nineteenth century in Lincolnshire, a Mr Cartwright improved his father’s business based on
movable woad mills by working a fixed 200 acres by the New Forty Foot Drain near Boston, and
the labourers were housed in a specially built hamlet called “Isatica’! Treasures like this abound,
with much “serious” information which is directly relevant to field botany, such as the use of
Anthyllis vulneraria to fodder rabbit warrens, or the novelty of crop legumes in the seventeenth
century (the treatise Sainfoin Improved was printed in 1671). My many unsuccessful searches for
plants such as Mentha pulegium or Damasonium alisma were explained when I read that William
Cobbet saw 10,000 geese on one Surrey common in the 1820s, where now there may be only dogs
on leads. I was also delighted to read of the feminist sensibility of a poultry commissioner in 1896
acknowledging the superiority of French egg production where women are in charge and “men
there do not hinder as in some cases here”’.

Enthusiasm makes for an unbalanced review, but this really is a book with relevance for most of
the botanical community. Tales of individual species, and fashions in land use, and the origins of
imported plants, all give clues to current plant distributions. For conservationists, reassurance is
perhaps the most welcome lesson. A paradox affecting many of us is that we work to conserve, but
believe that it’s already too late. Dr Thirsk’s calm and elegant exposition of “things coming around
and around” is most comforting, especially against the current background of millennium hype.
And nobody with any curiosity about the oddities of nature, or with a liking for a good story,
should resist this book.

R. FITZGERALD

BOOK REVIEWS 227

Flora of the Russian Arctic, Vol. I, Polypodiaceae — Gramineae, ed. J. G. Packer, transl. G. C. D.
Griffiths. Pp. xxxviii + 368. University of Alberta Press. 1995. Hardback $65, ISBN 0-88864—
2695. (A translation of the original Russian edition ed. A. I. Tolmachev.)

The ten volumes of the original Russian work were published in softback between 1960 and 198
and cover the vascular flora of the Russian arctic islands from Franz Joseph Land to Wrangel
Island and the generally 200-300 km wide mainland from the Murmansk peninsula to the western
shores of the Bering Sea. The Flora contains a wealth of information of interest to western
botanists but this has remained largely untapped on account of language difficulties. It was,
therefore, a bold and imaginative initiative on the part of the University of Alberta Press to publish
in hardback an English edition in six volumes under the editorship of Professor J. G. Packer. The
first volume covers the first two Russian volumes and deals with the Pteridophyta, Gymnosperms
and monocots from Sparganiaceae to Gramineae. In his preface Packer gives projected publication
dates for the remaining five volumes concluding in 1998. However, this laudabie and longed-for
project terminated prematurely with the publication of volume 2, Cyperaceae — Orchidaceae, in
1996. At least these volumes cover the two largest families in the arctic: Cyperaceae and
Gramineae; the latter includes such exclusively arctic genera as Arctophila, Dupontia and Phippsia
and is represented in the Russian arctic by at least 162 species.

A welcome feature are the detailed keys, sometimes even unduly detailed, and the expansive
remarks on many critical species, there being no attempt to produce accounts of uniform length. OF
especial interest for western botanists are the long accounts of the critical genera Calamagrostis
(including all the British taxa) and Puccinellia. The remarks under Lycopodium annotinum and
Huperzia selago should help to focus western attention on their neglected subspecies. There are
useful habitat and ecological notes and data on distribution outside Russia. About half the species
are provided with dot maps and the distributions of all the species in the 16 districts are
summarised by tables in the Appendix.

Since nearly 40 years have elapsed since the original Russian edition it is not surprising that
several statements are now outdated. The northern limit for Potamogeton (P. filiformis in
Greenland) is now 77°N, Puccinellia tenella occurs in Spitsbergen, Poa hartzii in Spitsbergen and
Canada, Alopecurus borealis in England, and Poa pseudoabbreviata is not a Russian endemic
since it occurs in Alaska. There are few other errors: Hierochloe odorata and Alopecurus aequalis
occur in Greenland, Cystopteris dickeana is stated to occur in the mountains of Scotland and for
some reason Puccinellia capillaris is said to occur in Britain “only as an introduction”.

The species concept is generally similar to that in the west. A glaring exception to this is the
inclusion of Festuca baffinensis in F. brachyphylla. No chromosome numbers are cited and there
is no indication that only tetraploid Polypodium vulgare occurs in the arctic. The nomenclature has
in many cases been superseded. Dryopteris species are erroneously called shield ferns and I was
blissfully unaware that Roegneria was the “rhizomeless wheat grass” genus.

The format is pleasing and clear although the margins are excessively wide and the running
heads are in smaller type than the text.

While applauding the appearance of this fine volume one must deeply regret the demise of such
an imaginative project. One can only guess at the disappointment experienced by Professor Packer
and his able translator and hope that an altruistic publisher will come to their rescue.

G. HALLIDAY

Carmarthenshire rare plant register. R. D. Pryce. Pp. xvi + 88. Published by the author and
available from him at Trevithin School Road, Llaneli, Carmarthenshire, SA15 4AL. 1999. Price
£15.00. No ISBN number.

Just as the Atlas of the British Flora encouraged the production of many county local plant atlases
after its publication in 1962, so too has the publication of national Red Data Books encouraged the
production of local equivalents. In some cases these have been published books dealing with rare
species in many different plant and animal groups, such as the Red Data Books for
Cambridgeshire and Cornwall reviewed in Watsonia 22: 289-290 (1999). Other counties have
decided not to follow this route, preferring instead to produce “rare plant registers”, working
documents listing the nationally and locally rare species in a county and detailing the sites in
which they occur.

228 BOOK REVIEWS

The current volume covers the vice-county of Carmarthenshire. It has its origins in a rare plant
list produced by Richard Pryce in 1984, “the product of the early days of home computing”. It has
been revised and produced as a spirally bound report which makes full use of current computer
facilities. It includes internationally rare species, nationally rare and nationally scarce species and
species which are rare, scarce or declining in the vice-county. Red text is used to labei
international and national rarities and green for the local rarities. Thus the criteria for inclusion are
set out clearly, and species such as Liparis loeselii are marked by a plethora of conservation
designations, detailed in red. The register is richly illustrated by colour photographs and also
includes tetrad distribution maps of the more frequent species.

The criteria for inclusion are said to follow those recommended by Perring & Farrell in BSB/
news 71: 10-11 (1996), though they have been interpreted very liberally. One example is that
species are included if they qualify under the criteria regardless of whether or not they are native
or alien in the county, so that Galanthus nivalis is included as an internationally protected species.
even though only native populations are protected and the species occurs in the county as an alien,
and Arenaria balearica, Erinus alpinus and Lagarosiphon major are listed as locally rare species.
A particularly interesting inclusion is Yucca recurvifolia, thought to have become established on
sand dunes from seeds or vegetative fragments which have drifted from North America. If so this
would establish the Agavaceae as a family native to Europe! Subspecies, varieties and hybrids are
included where appropriate. Occasionally the criteria appear to have been “bent”, as for the
orchids Anacamptis pyramidalis and Gymnadenia conopsea, which are frequent on the coast but
included as they are rare inland.

The accounts of individual species include a list of records made since 1950, detailing the grid
reference, native status, date of last record, recorder, site protection details, site name and habitat
for each record. The species accounts are a model of clarity, an exception being the baffling
treatment of Asparagus officinalis subsp. officinalis, which identifies several native populations of
this introduced subspecies and applies to it an English name (“Wild Asparagus”) and a
Biodiversity Action Plan priority rating which presumably refer to subsp. prostratus. Many of the
accounts include a brief paragraph describing trends in the distribution and current threats to the
plant. These provide a valuable insight into the changes taking place in the flora of this area. The
individual accounts could have been usefully supplemented by an introductory chapter drawing
together these themes, and explaining the various acronyms and initiatives mentioned in the tex:
(CADW, Tir Cymen scheme, Millennium Coastal Park scheme) which might be familiar to the
residents of the area but less well known outside Wales.

This register is designed to be comprehensible to both botanists and others such as county .
planners who might need it, and it will undoubtedly provide a useful tool for conservationists in
the vice-county. It will be interesting to see how effective it will be in promoting the conservation
of the threatened species which are identified. For Lemna trisulca, for instance, will the “urgent
remedial measures required by local authority in order to maintain this species in the vice-county”
be undertaken in time? In addition, the register will certainly be of considerable interest. to
botanists outside the vice-county. Most of the records in this register come from Carmarthenshire
Flora Database, built up by recorders working towards a Flora of Carmarthenshire. This
publication increases the anticipation with which we await this important Flora.

C.D PRESTON

Field Flora of the British Isles. C. A. Stace. Pp. xiii + 736. Cambridge University Press,
Cambridge. 1999. Price £17.95. ISBN 0-521-—653150-0.

The New Flora of the British Isles by C. A. Stace was first published in 1991 and appeared as a
second edition in 1997. It has, as predicted, now become the standard British Flora. Only in
Ireland, where many botanists remain loyal to Webb’s An Irish Flora, does it have a serious rival.
The aim of the new Field Flora is, in the words of the author, “to present the essential data
contained in my New Flora...in a more compact and portable form suitable for taking into the field
or on holiday”. Thus it includes “the means of identification of all the species and subspecies
contained in the’ New Flora” (although I note that no attempt is made to separate the three
subspecies of Dryopteris affinis recognised in the latter!).

The Field Flora is a light and compact book with a flexible plastic cover. It weighs one pound,
less than a third of the weight and occupying less than a third the volume of its predecessor. This

BOOK REVIEWS 229

condensation has been achieved by reducing the keys and descriptions which formed the bulk of
that book to brief descriptions of the families and genera and to annotated keys. In addition to the
details required to key out a species, the keys include a brief indication of the size of the plant as
well as ecological and distributional information about the species. Thus the dichotomy which
separates two Helianthemum species appears as:

4 Petals white; leaves grey-tomentose on upperside. Stems to 50cm. Native; dry limestone
grassland; N Somerset and S Devon
White Rock-rose - H. appeninum (L.) Mill.

4 Petals yellow; leaves green-pubescent on upperside. Stems to 50 cm. Native; base-rich grassland;
common in suitable places over most of Br, E Donegal

Common Rock-rose - H. nummularium (L.) Mill.

The hybrid between these species is mentioned in a brief note after the key. Where there is only

a single species in a genus the data are presented as a numbered “monachotomy” rather than
immediately after the generic description. Thus for Tuberaria:

1 Stems + procumbent to erect, to 30cm but often <10 cm; Native; dry barish ground near sea; W
& SW Ir, NW Wa, Jersey and Alderney
Spotted Rock-rose - T. guttata (L.) Fourr.

The line drawings of individual (often alien) species which appear in the New Flora are omitted,
but the comparative illustrations of features such as the leaves of Alchemilla and Sorbus, the
utricles of Carex, the fruits of Rumex and the labella of Dactylorhiza are included.

Although the format of this Flora is designed for use in the field, the content remains similar to
that of the larger Flora. Rare native and alien species are given the same treatment as commoner
plants. This is a wise decision, as although these species are less likely to be encountered while
undertaking routine recording, they are just the species one might need to check if one thinks one
has come across them. As Stace points out, this is a major difference from Clapham, Tutin &
Warburg’s Excursion Flora of the British Isies, which orovides descriptions of commoner species
but simply keys out the rarer species. Perhaps more debatable is Professor Stace’s decision to
retain the same characters to identify the species as are used in the New Flora; no attempt is made
to concentrate on characters which are easy to observe in the field. Many of the keys do make
good use of vegetative characters, but sometimes (as with Trifolium) I feel that more use could
have been made of vegetative characters at the early stages of the key to species. The paradox that
sO many species in our flora are instantly identifiable to the experienced botanist on sight but can
only be keyed out in a Flora by a series of difficult technical questions will continue to puzzle and
frustrate beginners.

I would also have liked to see, in addition to the keys which are included in the book, some extra
aids such as the vegetative key to commoner grasses included in C. E. Hubbard’s British Grasses.
Most ecologists would expect some help with the identification of vegetative grasses in a Field
Flora, but little is provided here. All one is told about vegetative characters in the account of
Arrhenatherum is that A. elatius is a loosely tufted perennial.

It is always easy to suggest additional features which a book might contain. I must therefore
make it clear that, having carried the book in the field for several weeks, I already regard it as
totally indispensable. Not only does it help identify unknown plants, it is an ideal aide-mémoire for
dim-witted botanists like me who can never remember whether Viola reichenbachiana or V.
riviniana has the darker spur, and what are the differences between the fruits of Rumex
conglomeratus and R. sanguineus. Even though botanists who have bought the first two editions of
the New Flora may initially feel they have already paid twice for the information in the Field
Flora, it is sure to prove a worthwhile purchase. It is in some ways a more satisfactory book than
the New Flora itself, for whereas the absence of extensive descriptions in the larger Flora is
disappointing, the format of the Field Flora is perfectly suited to its intended purpose. Whereas I
have retained my copy of Clapham, Tutin & Warburg’s Flora to use alongside the New Flora, |
am sure that I can now discard my copy of the CTW Excursion Flora. (Indeed, I have already tried
to throw it away, but I succumbed at the last minute to sentimental affection for this battered and
water-stained companion of 18 years fieldwork.) Professor Stace has again placed us greatly in his
debt.

C. D. PRESTON

230 BOOK REVIEWS

Urban Flora of Belfast. S. Beesley and J. Wilde. A Project of the Belfast Naturalists’ Field Club.
Pp. viii + 196. The Institute of Irish Studies and The Queen’s University, Belfast 1997. Price
£8.50. ISBN 0 85389 695 X.

Ever since writing the Flora of Inner Dublin (Wyse Jackson & Sheehy Skeffington 1984) I find it
hard to walk in any city without examining what is emerging from the pavement cracks or
glancing at a promising bit of waste ground. I was therefore delighted to be able to review the
Urban Flora of Belfast. Belfast is, in many ways, similar to Dublin, as both are large cities on the
east coast of Ireland with a large commercial port and therefore comparisons are inevitable.

The Flora is very systematically surveyed. The city, which has been divided into 76 one
kilometre squares, covers the whole of urban Beliast, much more than the Dublin inner city flora,
which covers an area of approximately 9 km’ and is defined along “natural” lines. The Belfast
Flora also includes suburban and more rural areas. As with the Dublin city flora, many aliens and
garden escapes are present. The criteria for their inclusion are well set out and a list of garden
escapes is also included. The Flora inventory was carried out over a short time interval so that it
could act as a baseline for future study of this ever-changing environment. Each square was visited
at least twice over the period 1993-1995, with seasonal differences taken into account where
possible.

The total number of species is 592, 63% of which are native. The proportion of non-native
species is high, as in inner Dublin, where the number of casuals and aliens was also recorded as
approximately 35%. The Dublin city Flora has less records (358), but this is not proportionally less
in relation to the area covered (c.9 km? vs 76 km’), which suggests the existence of an intrinsic
urban flora, regardless of the area in question. Twenty-five species were found in every square, 4!
very predictable (e.g. Taraxacum, Plantago spp., Sonchus spp. and several grasses) and fence 19
one tree, the elder, Sambucus nigra. Though an opportunist, it seems quite an achievement for it to
turn up in all 76 squares. Buddleja is also common, as in Dublin, though it is not so common in
other Irish cities.

There is an impressive list of records new to the counties of Antrim and Down, but aiso many
are listed as new to the Census Catalogue of Ireland (Scannell & Synnott 1987). However the
majority of these last are adventives, mostly garden escapes ranging from potato Soianum
tuberosum to ornamentals such as Mahonia aquifolia and nasturtium Tropaeolum majus. | would
not have expected these to be included in the Irish census, which includes native species or
“established aliens”. However, it does bring into question whether some, such as the Mahonia,
merit inclusion in the Census. I would not consider potato or nasturtium as established, though
they probably do get renewed from cultivation very regularly. Other species, such as Sanguisorba.
minor are native to Ireland, but are still considered adventive to the area - as is the surprising
record for Euphorbia hyberna - almost certainly a garden escape.

The introductory chapters deal with the methodology and summarise the results in terms of
records per square and new records. A most valuable section discusses each square in turn, listing
its main habitats and species of note. This is not only very useful for the current botanist, but also
for future comparison of changes. It also gives some insight into the problems of urban recorders,
permission being necessary to enter some grounds. In the case of the Balmoral Golf Club,
recording involved getting up at 5a.m., presumably to avoid early golfers!

Conducting a botanical survey of any inner city area requires a certain amount of courage and
great credit is due to the Belfast Naturalists’ Field Club for this survey of a not entirely untroubled
spot. There are some interesting asides like the reference to the Malone Road residents who
(sadly!) keep their gardens and pavements weed-free! But it is clear too that everyday life,
including that of botanists, continued throughout a very rough time, hopefully now relegated to the
past.

The Flora is very accessible to the amateur or professional, whether resident or not in Belfast
and readers should derive much pleasure in dipping into this book. It is not much bigger than An
Irish Flora (Webb et al. 1996) and therefore can also be carried about the streets of Belfast. It is
also illustrated by some attractive line drawings of plants in a Belfast setting —which are very
similar in style to those of the Flora of Inner Dublin. I am therefore disappointed that they do not
refer at all to this sister Flora, even as suggested further reading.

M. S. SKEFFINGTON

RITISH ISLES
(B.S.B.1.)

ES

Ve
Bobl

The B.S.B.I. traces its origin to the Botanical Society of London founded in 1836 and has a membership
of 2,850. It is the major source of information on the status and distribution of British and Irish
flowering plants and ferns. This information, which is gathered through a network of county recorders,
is the basis for plant atlases and of publications on rare and scarce species and is vital to botanical
conservation. The Society published Atlas of the British flora in 1962 and revised editions and is
working towards a new atlas, to be completed in 2000. It organises plant distribution surveys, publishes
handbooks on difficult groups of plants and has a panel of referees available to members to name
problematic specimens. The B.S.B.I. arranges conferences and field meetings throughout the British
Isles and, occasionally, abroad. It welcomes as members all botanists, professional and amateur alike.

Details of membership and any other information about the Society may be obtained from:

The Hon. General Secretary,
Botanical Society of the British Isles,
c/o Department of Botany,

The Natural History Museum,
Cromwell Road,

London, 5W7 5BD.

BSBI Handbooks

Each Handbook deals in depth with one or more difficult groups of British and Irish plants.

No.1 Sedges of the British Isles
A. C. Jermy, A. O. Chater & R. W. David. Revised edition, 1982. 272 pp., with descriptions,

line drawings and distribution maps for all 73 species of Carex. Paperback. ISBN 0 901158 05
4.

No.2 Umbellifers of the British Isles

T. G. Tutin. 1980. 200 pp., with descriptions and line drawings of 73 species of Apiaceae
(Umbelliferae). Paperback. ISBN 0 901158 02 X.

No.3 Docks and knotweeds of the British Isles
J. E. Lousley & D. H. Kent. 1981. 208 pp., with descriptions and line drawings of about 80
native and alien taxa of Polygonaceae. Paperback. Out of print. New edition with distribution
maps in preparation; orders recorded.

No.4 Willows and poplars of Great Britain and Ireland
R. D. Meikle. 1984. 200 pp., with descriptions and line drawings of 65 species, subspecies,
varieties and hybrids of Salix and Populus. Paperback. ISBN 0 901158 07 0.

No.5 Charophytes of Great Britain and Ireland

J. A. Moore. 1986. 144 pp., with descriptions and line drawings of 39 species and varieties of
Characeae and 17 distribution maps. Paperback. ISBN 0 901158 16 X.

No.6 Crucifers of Great Britain and Ireland
T. C. G. Rich. 1991. 344 pp., with descriptions of 148 taxa of Brassicaceae (Cruciferae), 129 of
them with line drawings, and 60 distribution maps. Paperback. ISBN 0 901158 20 8.

No.7 Roses of Great Britain and Ireland
G. G. Graham & A. L. Primavesi. 1993. 208 pp., with descriptions and line drawings of 13

native and nine introduced taxa of Rosa, descriptions of 76 hybrids, and 33 maps. Paperback.
ISBN 0 901158 22 4.

No.8 Pondweeds of Great Britain and Ireland
C. D. Preston. 1995. 352 pp., with descriptions and line drawings of all 50 species and hybrids
of Potamogeton, Groenlandia and Ruppia, most of them with distribution maps; detailed
introductory material and bibliography. Paperback. ISBN 0 901158 24 0.

No.9 Dandelions of Great Britain and Ireland
A. A. Dudman & A. J. Richards. 1997. 344 pp., with descriptions of 235 species of Taraxacum,
most of them illustrated by silhouettes of herbarium specimens; drawings of bud involucres of
139 species and 178 distribution maps. Paperback. ISBN 0 901158 25 9.

Other publications

English names of wildflowers
J. G. Dony, S. L. Jury & F. H. Perring. 1986 (2nd ed.). 126 pp. Recommended English names
for British and Irish vascular plants (Latin-English and English-Latin). Paperback.
ISBN 0 901158 15 1.

List of vascular plants of the British Isles
D. H. Kent. 1992. 400 pp. The same nomenclature and sequence as in Clive Stace’s New Flora
of the British Isles (1991, 1997), with selected synonyms. Paperback. ISBN 0 901158 21 6.
Supplied with five errata lists and Supplement I (December 1996) of 36 pp.

Alien plants of the British Isles
E. J. Clement & M. C. Foster. 1994. 616 pp. Lists 3,586 recorded non-native species (of which
885 are established), with English names, frequency of occurrence, status, areas of origin,
location of voucher specimens, references to published descriptions and illustrations, and
selected synonyms. Paperback. ISBN 0 901158 23 2.

Alien grasses of the British Isles
T. B. Ryves, E. J. Clement & M. C. Foster. 1996. 234 pp. A companion volume to the last,
listing over 700 non-native grasses. Includes keys to bamboos and eight of the larger and more
difficult genera and 29 pp. of illustrations. Paperback. ISBN 0 901158 27 5.

Plant crib 1998
T. C. G. Rich & A. C. Jermy. 1998. 400 pp. An identification guide for some 325 ‘difficult’

taxonomic groups, with explanations, keys and illustrations of plant details. A4 paperback.
ISBN 0 901158 28 3.

British Red Data Books I Vascular plants
M. J. Wigginton, ed. 1999 (3rd ed.). 468 pp. Up-to-date information and maps for 408 taxa
(including 118 microspecies) regarded as threatened in Great Britain, with details of their
habitats and associated species, maps showing their pre-1970, 1970—1987 and post-1987 records
by 10-km squares, and 1-km square frequency maps for 63 taxa. Hardback, published by JNCC,
Peterborough. ISBN | 86107 451 4.

Scarce plants in Britain
A. Stewart, D. A. Pearman & C. D. Preston, comp. & ed. 1994. 518 pp. Accounts of 254
nationally scarce taxa (occurring in 16-100 10-km squares in Great Britain) and of 71 taxa
formerly thought to be so, with details of their habitats and associated species, reproductive
biology, changing British distribution and world range; updated distribution maps. Hardback,
published by JNCC, Peterborough. ISBN 1 873701 667.

Aquatic plants in Britain and Ireland
C. D. Preston & J. M. Croft. 1997. 365 pp. Summarises the distribution, habitat and
reproductive biology of 200 aquatic plants in 72 genera, with 200 distribution maps and 72 line
drawings. Hardback, published by Harley Books, Colchester. ISBN 0 946589 55 0.

Available from the official agents for BSBI Publications, Summerfield Books, (John & Sue Atkins)
Summerfield House, High Street, Brough, Kirkby Stephen, Cumbria CA17 4BX.
Fax & Tel: 017683 41577. E-mail: summerfield @iclweb.com

INSTRUCTIONS TO CONTRIBUTORS

Scope. Authors are invited to submit Papers and Notes concerning British and Irish vascular
plants, their taxonomy, biosystematics, ecology, distribution and conservation, as well as topics of
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Watsonia

February 2000 Volume twenty three Part one

Douglas H. Kent Memorial Issue

Contents

PRESTON, C. D. & SANFORD, M. N. Douglas H. Kent Memorial Issue of
OBITUARY Douglas Henry Kent (1920-1998)

ALLEN, D. E. & STACE, C. A. Botanical publications of Deusten. Fees Kent
ALLEN, D. E. A selective supplement to British and Irish Herbaria

CROMPTON, G. & NELSON, E. C. The herbarium of William Skrimshire (1766-
1829) of Wisbech

BOWLT, E. M. & BOWLT, C. Tobi Bieta (713- 1753) a wader apothecary
and botanist of his time

NELSON, E. C. A history, mainly noneleoian of St ahene S Heath

PRESTON, C. D. Engulfed by suburbia or destroyed by the er the ecology of
extinction in Middlesex and Cambridgeshire

HOLYOAK, D. T. Hybridisation between Rumex heart ie Gall (Polygonacese)
and other docks .

BAILEY, J. P. & CONNOLLY, A. P. Prize- -winners to ae
Knotweed s.l. (Polygonaceae) in the British Isles

Q Biseory of j apanese

HOLLINGSWORTH, M. L. & BAILEY, J. P. Hybridisation and clonal diversity in

some introduced Fallopia species (Polygonaceae)

ABBOTT, R. J., JAMES, J. K., IRWIN, J. A. & COMES, H. P. Hybrid origin of the
Oxford Ragwort, Senecio squalidus L.

JAMES, C. M., WURZELL, B. S. & STACE, C. A. A new or eee a European
and a Chinese species of Artemisia (Asteraceae) sae

OSWALD, P. H. Historical records of Lactuca serriola L. and L. virosa L. in Baltain
with special reference to Cambridgeshire (v.c. 29)

BRUMMITT, R. K. & CHATER, A. O. ues amie tic. hybrid in West
Wales wee

CLEMENT, E. J. a eee x ae E. Je clement a new fataril saves.

FLETCHER, R & STACE, C. A. A new section and species of Festuca (Poaceae)
naturalized in England ee :

OTHER PAPERS

FORBES, R. S. Assessing the status of Stratiotes aloides L. Silene ic: in Co.
Fermanagh, Northern Ireland (v.c. H33) :

WALKER, K. J. The distribution, ecology and conservation of ee norvegica
subsp. anglica Halliday (Caryophyllaceae)

WILLIAMS, L. R. Annual variations in the size of a Asepelaiion of Coe
impatiens L. i sone > “ime

NOTES
Hanson, C. G. Update on birdseed aliens (1985-1998)
Sell, P. D. Hieracium kentii sp. nov. (Asteraceae)

Harris, J. Chara baltica Bruz. (Characeae), Baltic Stonewort, valves in
Hickling Broad, Norfolk

Barnett, K. & Rich, T. C. G. Thlaspi ee L. new to Worcseteichiie e ce 37)
BOOK REVIEWS
Published by the Botanical Society of the British Isles

Typeset by D. K. & M. N. SANFORD
Printed in Great Britain by THE BOOK COMPANY, PO BOX 243, IPSWICH, SUFFOLK

139-147

(49-150 14

161-165
167-172

173-177

179-196
197-208

209-212

213-215
216-217

218-219
220
221-230

met

WUT

NN! A

Botanical Society of the British Isles

Volume 23 Part 2 August 2000
Editors: M. Briggs, D. L. Kelly, D. R. McKean,
D. A. Pearman, M. N. Sanford, D. A. simpson

= Botanical Society of the British Isles

Patron: Her Majesty Queen Elizabeth the Queen Mother

Applications for membership should be addressed to the Hon. General Secretary,
c/o Department of Botany, The Natural History Museum, Cromwell Road, London,
SW7 5BD, from whom copies of the Society’s Prospectus may be obtained.

Officers for 2000—2001

President, Dr G. Halliday
Vice-Presidents, Mr M. Walpole, Mr D. J. McCosh, Prof. C. A. Stace
Acting Honorary General Secretary, Miss A. Burns

Honorary Treasurer, Mr M. E. Braithwaite

Editors of Watsonta

Papers and Notes, D. L. Kelly, D. R. McKean, M. N. Sanford*, D. A. Simpson
Book Reviews, D. A. Pearman
Obituaries, M. Briggs

*Receiving editor, to whom all MSS should be sent (see inside back cover).

© 2000 Botanical Society of the British Isles
The Society takes no responsibility for the views expressed by authors of Papers,
Notes, Book Reviews or Obituaries.

The cover illustration of Meconopsis cambrica (L.) Viguier (Welsh Poppy) was
drawn by Rosemary Wise.

Watsonia 23: 231-235 (2000) Zo

Presidential Address, 1999

MARY BRIGGS

LIBRARIES

THE STUDY OF A LOCAL FLORA

An appreciation of the wild plants of the British Isles

The theme of this Address is to show how the study of a local flora can lead one to a greater
understanding of the larger themes of botany, and how a local botanist can contribute directly to
these larger themes from his or her own local knowledge and investigations. It is based on my own
experience as a B.S.B.I. member and as County Recorder for West Sussex.

The B.S.B.I. entered the technological age in our Atlas of the British Flora (Perring & Walters
1962). For this, records collected by members from all corners of the British Isles were edited by
Max Walters and Frank Perring, and punched onto cards to produce maps. There was immediate
excitement as patterns of distribution were clearly demonstrated, often for the first time, as the
maps appeared. The Atlas led directly to the first legislation for the protection of wild plants in
Great Britain in the Wild Creatures and Wild Plants Act 1975, followed by the Wildlife and
Countryside Act 1981, with the schedules of protected plants reviewed every five years since. The
Society was deeply involved with both these Acts of Parliament and the subsequent Quinquennial
Reviews. It is difficult for us now to realise that before the first Atlas, there was no quick reference
to the distribution of individual species across the British Isles. Reference to the distribution of
each species gave the first general indication of rarity.

However, in the conservation field, as far back as the “Battle for Teesdale” in the 1960s, when
we attempted - without success - to save the rare plants at Cow Green from inundation by a new
reservoir (Lousley et al. 1965; Tutin et al. 1965), we learned then to our disadvantage that learned
opinions based on experience were insufficient as legal evidence for Planning Inquiries. In the
thirty years since we have steadily endeavoured to assemble our information on plants into a more
rigid form, with the statistics required for legislation. This has progressed through the recording
methodology used for our Monitoring Scheme sample survey 1987-1988 organised by Tim Rich
and reported in Rich & Woodruff (1990). Currently we have another leap forward in recording,
record storage and the availability of records planned for the future through our present Atlas 2000
project, and on into the next century with the Threatened Plants Database Project, in which the B.
S.B.I. is heavily committed. Inevitably this standardisation of recording format could well lead to
the recording of plant distribution becoming largely a number-crunching exercise.

A sub-theme of this Address is to show that, in spite of advanced modern technology, there can
be more to the study of a local flora than just the accumulation of records. My sub-title proposes
that there is as much a place for the wider study of the natural history of our plants, and for the
enjoyment of the plants, at the turn of the millennium as ever before.

For a deeper perspective and understanding of a local flora, an obvious start is with the geology.
Taking Sussex in the context of the Weald, we have a number of habitats: chalk downland, coastal
plain, narrow bands of upper and lower greensand and extensive heavy weald clay. In small local
areas the clay is lightened by bands of Paludina limestone and Horsham stone (Gallois 1965). In
Mesozoic times the Weald was a lake, in which a snail, Viviparus sussexiensis flourished along
with dinosaurs. The snails sank on death into the sediment at the water’s edge, and their shells can
still be seen now embedded in the local “winkle stone”, which is sufficiently firm to cut into
blocks for building. Some local houses were built from this stone, for example Hungerhill House
built in 1911 from winkle stone quarried from a nearby field on the farm and the wall blocks
clearly show the rounded shells. Winkle stone can also be polished, with the shell outlines giving a
marbled marking; it is known locally as Sussex, Petworth or shelly marble. In local churches
memorials are framed with polished local “marble”, and many churches have baptismal fonts
carved from winkle stone. At the time of the Wealden Lake the vegetation was largely of conifers,
cycads, ferns and horsetails; now we look in the bands of winkle stone for plants not common on

(Aue 17 wo |

232 M. BRIGGS

the surrounding clay, e.g. Adoxa moschatellina, Lathraea squamaria and Asperula cynanchica. A
similar local sandstone is Horsham stone, a thinly bedded ripple-marked calcareous sandstone.
which was also used in building, as roof tiles for houses built with oak rafters sufficiently stout to
carry the weight of the stone. In long-drained Horsham stone quarries the occasional Ophrys
apifera has been recorded, far from the usual chalk downland localities for this orchid. Adoxa
moschatellina has also been recorded in fields marked “brickfield” on old maps at the Records
Office. In the north-west of Sussex there is a narrow wedge of Bargate beds, a sandgate bed in the
lower greensand; the map for Fallopia dumetorum in Sussex Plant Atlas (Hall 1980) can be
exactly superimposed on the Bargate beds on a geological map of Sussex.

The heavy weald clay supported large areas of deciduous forest, and historically Sussex oaks
were used to build the naval sailing ships at the time of the Armada. Still today Sussex has a
higher percentage of deciduous woodland than most English counties. For many centuries the
sticky clay was impassable in the winter months to men, horses and horse-drawn vehicles, and a
Domesday Book map of mid-Sussex shows the absence of human settlements along the Forest
Ridge. Ifield and Worth are the only place names mapped in the mid-north Weald in 1086 (Morris
1976). A recently discovered link with geological distribution is in Tilia platyphyllos - nationally
scarce as a native tree, and until 1987 considered to have been only planted in Sussex. In 1987
Francis Rose realised that a succession of copses of T. platyphyllos discovered along north scarps
of the South Downs in northwest Sussex all had features of ancient woodland (Rose 1991). Donald
Pigott accepted these as native trees and there is a paper in preparation.

Another feature of the local flora is the Mediterranean element. Many Mediterranean species are
recorded that are dependent on hot summers for good seed production, this seed in turn being
dependent on early spring rainfall for germination. When recording population size of these
species from year to year the variation, from a handful of individuals to some thousands in
occasional years, is very evident. The unpredictability of orchid populations is well known. This
year at a meadow reserve (not in Sussex) a local recorder was apologising for the Orchis morio
pictured on a notice listing the plants to be seen, saying that this had now not been seen for many
years at this site, when in the meadow that day 20,000 plants of Orchis morio were counted.
Similar variations in population sizes in different years are shown by Gentianella anglica in East
Sussex, but even more dramatic are figures from Dorset, where in 1994 numbers of this species
were estimated at 500,000, but in spite of searches at the same sites in 1996 only 50,000 were
recorded. This was referred to by my predecessor, David Pearman, in his Presidential Address
(Pearman 1997). To some extent these variations in population size cast doubt on the validity of
“snapshot” recording as a monitoring tool.

In the New Naturalist monograph on Lords and Ladies by Cecil Prime, the pattern of Arum
italicum subsp. neglectum in southern Britain is explained by a map correlating the distribution
with the area in which snow lies for less than five days in the year (Prime 1960). Since then a
possible weather pattern change due to global warming could extend the range of this plant further
north following a retreating snowline, but the further restrictions of habitat requirements to either
the springlines between chalk and greensand, or the coastal plain on brick-earth and gravels, could
limit any dramatic change. A recent survey of the distribution of this plant in West Sussex was
carried out by members of the Sussex Botanical Recording Society and reported by Ron Clough
(1998). The author concluded that additional records on the Downs in this survey were almost
certainly due to diligent searching rather than an increase in frequency of the plant.

Phyteuma orbiculare is totally confined to the chalk downs, and the distribution map in the
Sussex Plant Atlas exactly overlies the line of the South Downs. Locally it is known as “The pride
of Sussex” — rather inappropriately, as travelling in Europe we find that it is widespread, being
recorded from 15 countries in Flora Europaea (Webb et al. 1964-1980). More distinctive to
Sussex is the wood of Taxus baccata at Kingley Vale, recognised as the finest yew wood in
Europe. Steep chalk downland slopes provide a stronghold for these woods, and at Kingley Vale
the slopes are south- and west-facing which, being dryer, are less favourable to the beech and ash
trees which are also typical of downland slopes. For many centuries the yew supplied the wood for
Britain’s strategic weapon, the longbow; Homer and Virgil had recorded that yew was one of the
best woods for bows. On the Reserve there are old gnarled trees and in the main valley at Kingley
Bottom there is the dark “sinister and fantastic” forest of Victorian times. Ring counts have shown
the older trees now to have been growing for 500 years (Williamson 1978).

PRESIDENTIAL ADDRESS 233

In contrast to the strict habitat requirements of these chalk species is Hypericum hirsutum.
Thinking of this as a downland plant we were surprised to find that the Sussex Plant Atlas
distribution map shows the species as widespread in West Sussex with many fewer downland
records in East Sussex. Norman Robson interpreted this for us by pointing out that the species
needs not only neutral soil and good drainage, but also some shelter, such as hedgerows and wood
edges rather than open fields. The bare downland of East Sussex is too windswept for Hypericum
hirsutum. Another Hypericum species, H. perforatum has been in the news as a treatment for
depression. Used externally in massage and as an effective wound healer for centuries, modern
research has revealed volatile oils, hypericins and prenylated phloroglucinol derivatives, and the
plant has been tested for internal use as an anti-depressant with encouraging success.

Ashdown Forest was the Sussex stronghold for Genista pilosa, but this species has dwindled in
the second half of this century and is now presumed extinct. David Coombe has suggested (pers.
comm.) that the heathland ecotype, the form which was found in Sussex and on the Suffolk heaths
(where it is also now extinct), was less vigorous than the more robust ecotype of the coastal heaths
in Cornwall; the eastern heathland form thus had less potential to recover after fire - a frequent
hazard in the Forest. Tim Rich reported on this in the Flora of Ashdown Forest (Rich et al. 1996).

Cardamine impatiens, another local plant restricted to woodland on neutral soil and to river
edges, germinates in sections of ash wood in the year following coppicing. We found it in cleared
soil near my home at Slinfold, when collecting springy sucker shoots of the ash (very suitable for
bow and arrow play). The Cardamine flowered in the second year, but by the third was crowded
out by the colonising vegetation. This local species is widespread in the northern hemisphere; I
have seen it growing in grykes on limestone pavement at Fell End Clouds in Cumbria (Halliday
1997), also in the Swiss and Italian Alps, and from a pony in a Himalayan wood in Kashmir.

Sussex is still a stronghold for Carex vulpina, now a Red Data Book species. Only separated in
Britain from C. otrubae in 1939, the two species can be difficult to distinguish; fruiting plants
apparently have a distinctive “jizz”, but comparison of the epidermal cells of the utricle under the
microscope, as described in the B.S.B.I. Handbook (Jermy, Chater & David, ed. 2, 1982) is
required by the Carex referees. The sedge is endangered in Sussex through loss of habitat,
especially now that so many river edges have been artificially built up with steep flood banks.

It is always exciting to find a completely unknown flower. Downingia elegans at Ardingly
Reservoir was a complete mystery, until the clues of Hordeum jubatum and a tall robust form of
Mentha pulegium (now presumed extinct as a Sussex native) suggested an American species.
Botanists at Wakehurst Place, Kew were able to find out for us that the edges of the newly
constructed reservoir had been sown with a grass mixture from the U.S.A., designed for
fluctuating water levels (Briggs 1986). Mary Elizabeth Parsons (1897) described these plants as
growing in wet places, “where they often make the ground blue. The showy white-centred flowers
are familiar along the roadsides upon the borders of puddles”. Another introduction is Elodea
callitrichoides, a waterweed from South America. The Sussex site in the Chichester Canal may
now be the only English record. There is also one in South Wales (Simpson 1986).

Historical records have a special fascination. In East Sussex Teucrium chamaedrys was recorded
on Camber Castle in 1690. J. Petiver and J. Sherard (1714) wrote “Upon that side of the castle wall
that looks to Rye we saw full plenty of chamaedrys”’. It could be seen there still until 1991, but in
this century the Castle has been excavated, first by the Ministry of Works, then by the Department
of the Environment. Despite the efforts of local plant recorders for 40 years, the last plants were
“cleaned off’ the castle walls with herbicide spray in 1991. In 1945 A. W. Graveson found T.
chamaedrys in short cliff-top turf at Cuckmere Haven. Francis Rose (pers. comm) accepts this as
native by comparison with chalk grassland sites in France.

A herbarium sheet in the Department of Botany, The Natural History Museum was first labelled
Polypodium vulgare var serratum, “Hbm. of J. Forbes Young MD 1796-1860”, and “from ye old
tower of Arundel Castle Sussex”. At the Castle, Sara Rodger researched her archives to identify
the “old tower” where the specimen was collected, as the present keep. On a more recent label
added to the herbarium sheet, a “redet.” reads: “P. australe Fee R. H. Roberts” - the species now
known as Polypodium cambricum L. It was renamed by our current Polypodium referee, R. H.
Roberts, as recent work had established the key character of paraphyses mixed with the sporangia,
enabling specific distinction. This well demonstrates the value of keeping herbarium specimens in
good storage, as reference from the past and for future work.

234 M. BRIGGS

As a professional pharmacist, my personal interest in medicinal plants has often involved those
which are poisonous. A local primary school phoned for advice when a small boy had eaten
Mercurialis perennis, fortunately with no lasting harm as he had been very sick. However, a tale is
told in Philosophical Transactions (Anon 1693) of the wife who went into the fields and gathered
M. perennis as a pot-herb to serve “Fryed with bacon (having first Boyled them)’. After eating this
all the family were ill, the children vomited and were purged and “slept from Thursday to
Monday” when sadly the youngest gave “2 sprunts” (convulsions) and died. The father, W.
Matthews, overslept too but went to work saying that “he thought his Chin had bin all the Day in a
Fire, and (he) was forced to keep his Hat full of Water by him all the Day long, and frequently dipt
his Chin in it as he was at Work.” Mother was reported to be “also very sick, but forced to rise to
look after her little family concerns”.

In 1983, in a modern case of poisoning by M. perennis the doctors described the reddening of
cheeks and jaw as a “curious malar erythema”. That time the M. perennis had been eaten in
mistake for Veronica beccabunga (an edible herb) (Rugman et al. 1983), highlighting the
importance of correct identification before eating.

In August 1977 three Dutch (botany) students youth hostelling and self-catering in Sussex added
Oenanthe crocata leaves to their stew, assuming it to be Apium graveolens which was familiar to
them from their local countryside at home. Although O. crocata is widespread in Sussex, and
throughout southern and western Britain and Ireland, it is a very rare plant in Holland. There it was
first recorded only in 1975. O. crocata contains highly dangerous oenanthetoxin and all the young
men collapsed within ten minutes and were taken to hospital; one seriously ill (after two helpings)
and in intensive care, but happily they all recovered.

Similarly an Eastern European family working in West Sussex went berry-picking and baked a
pie of mostly Atropa belladonna. All were seriously ill, but recovered after hospital treatment.

Looking to the future, the most advanced work on molecular botany, using characters which are
indistinguishable in the field, still relies in many cases on records from old collections, and living
material provided by local botanists. An example is the paper by Johannes Vogel, Fred Rumsey et
al. (1999), which describes Asplenium ruta-muraria subsp. ruta-muraria as widespread across
Europe and autotetraploid, but the diploid A ruta-muraria subsp. dolomiticum as coinciding in area
with the glacial refugia in northern Italy. The diploid subspecies needs at least two gametophytes
produced by different spores, giving a slower rate of colonisation which leads to a restriction of
diploid populations to ancient glacial refugia, and the colonisation of the rest of Europe by
polyploids. Collections of material by travelling and local botanists are critical for this ongoing
work on Asplenium.

Also looking to the future, B.S.B.I. records will be included in the World Database mapping
scheme at The Natural History Museum to provide information on biogeography. This, when the
number of sites, sizes of populations, climate and other factors are taken into consideration, will
lead to an improved evaluation of threats to rare species for conservation.

Petrorhagia nanteuilii is a Red Data Book species with its only British sites in West Sussex.
Morphological, cytological and geographical data strongly suggest that P. nanteuilii is an
allopolyploid derived from P. prolifer and the non-native P. velutina. The distribution of P.
nanteuilili is shown on the Atlas Florae Europaeae map as scattered along Mediterranean
coastlines, and on the Iberian Peninsula as widespread both on the coast and inland. In the British
Isles there are only two localities, both in West Sussex and coastal, at Pagham Harbour and
Shoreham; the latter is an old locality refound since the publication of the Red Data Book ed. 3
(Wigginton, 1999).

A most valuable section of B.S.B°I. activities is the publication of journals and identification
handbooks; these are a continual source of information on our plants, and most local floras are
written by B.S.B.I. members. Also informative, a series of B.S.B.I. surveys has been organised
through the years, some on a single genus or species e.g. Mistletoe, Atriplex, Sorbus torminalis
and Populus nigra; others on arable weeds, churchyards etc. All members can participate in these
network research surveys, which often lead to unexpected plant finds locally. The programme of
field meetings and conference meetings has enabled me to visit botanically most areas of the
British Isles during 39 years as a B.S.B.I. member, so that although this Address refers mainly to
my local county of Sussex I have had good opportunity to appreciate the wide variation in the wild
plants of Britain and Ireland.

PRESIDENTIAL ADDRESS 235

My family learned when young to identify ‘pin and thrum’ primroses; but only later at B.S.B.L.
lectures did I learn the intricate details of their morphology. Jack and Yolande Heslop-Harrison
brought scanning electron micrographs of pin and thrum stigma surfaces, showing the long
papillae of pin and the short papillae of thrum stigmas, which together with differences in size and
shape of the pollen grains effectively ensures that cross-fertilisation is the norm. Further papers
with diagrams on this by John Richards (1986, 1989) explained this complex mechanism simply.
What wonderful years to have been able to combine family life with the opportunity to learn such
detailed information on our plants! But I hope I have shown that, as well as providing such
personal enjoyment, careful study of a local flora can contribute to much wider knowledge, and
can put local botanists in touch with work of considerable scientific importance. This in turn
brings feed-back, leading to a deeper appreciation of the plants recorded locally. Through the
B.S.B.I. we all have the means to become active members of the wider botanical community.

ACKNOWLEDGMENTS

For the use of slides on 8th May 1999 [ would like to thank Miss Alison Paul, Department of
Botany; Prof. Chris Humphries, Biogeography and Conservation Laboratory, and the Photographic
Studio at The Natural History Museum, London, The British Library, Mrs Sara Rodger, Archivist
at Arundel Castle and Dr Alan Knapp. I would also like to thank all who helped with
encouragement and support.

REFERENCES

ANON., (1693). VIII. Part of a letter from Mr. T.M. in Salop, to Mr. William Baxter, concerning the strange
effects from the eating Dog-mercury. Philosophical Transactions 17(203): 875-876.

Briaccs, M. (1986). Downingia elegans refound in East Sussex. BSBI News 44: 20.

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Watsonia 23: 237—256 (2000) 237)

Mistletoe Viscum album L. (Loranthaceae) on oaks in Britain
J. D. BOX

17 Jockey Bank, Ironbridge, Telford, Shropshire TF& 7PD

ABSTRACT

Information on oaks with parasitical mistletoe (Viscum album L.) was sought from herbaria, county botanical
recorders and biological record centres in conjunction with site visits and an extensive search of published
records and reports. Currently there are eleven confirmed mistletoe-oaks in Britain. Evidence from past
records, reports and general accounts of mistletoe on oak indicates that the rarity of mistletoe-oaks in Britain
has not changed since the 17th century. The current results, together with the various reports of mistletoe-oaks
in the 19th century and early 20th century, suggest a population of mistletoe-oaks which is apparently fairly
stable in number but with some turnover as losses are compensated by the discovery of new sites. The existing
mistletoe-oaks are concentrated in Herefordshire and the current range is very much more restricted than the
historical range. Pedunculate oak (Quercus robur L.) is the most frequent host amongst the existing mistletoe-
oaks. The presence of mistletoe on red oak (Q. borealis Michx. f.) at three separate sites, as well as on a
scarlet oak (Q. coccinea Muenchh.), is notable. The estimated ages of the existing oak hosts range from 90 to
400 years. The existing mistletoe-oaks are not found in woodland but prefer hedges, woodland edges, parks
and a churchyard.

KEYWORDS: pedunculate oak, common oak, parasitism, plant distribution, red oak, Quercus.

INTRODUCTION

Mistletoe (Viscum album L.) has a wide range of hosts in Britain (Bull 1864a, 1864b, 1907;
Somerville 1914; Nicholson 1932; Perring 1973). The commonest hosts are apple (Malus
sylvestris (L.) Mill.), lime (Tilia x europaea L.), hawthorn (Crataegus monogyna Jacq.) and
hybrid black poplars (Populus x canadensis Moench) (Perring 1973).

Despite the traditional association between mistletoe, oaks and Druids in the minds of many
people (Frazer 1922; Kanner 1939; Box 1995), oaks have always been considered to be a rare host
for mistletoe in Britain (Evelyn 1664; Ray 1670, 1677; Withering 1796; Loudon 1838; Bull 1864a,
1864b, 1907; Nicholson 1932; Tubuef 1923; Perring 1973).

The Botanical Society of the British Isles (B.S.B.I.) carried out a survey of mistletoe and its
hosts from 1969 to 1972 using both B.S.B.I. members and the general public (Perring 1973).
Mistletoe was not recorded on sessile oak (Q. petraea (Mattuschka) Lieblin) and was recorded on
pedunculate oak (Q. robur L.) in only twelve 10 km squares in Britain (2-2% of the 539 10 km
squares recorded).

In contrast, the most recent survey of mistletoe carried out from 1994 to 1995 by Plantlife and
the B.S.B.I. produced some 140 sightings of mistletoe on oak, 1-8% of the 8,000+ sightings of
mistletoe sent in by the public and by botanists (Briggs 1995), although there are likely to be errors
due to the misidentification of oak in winter (Perring 1973).

An examination of historical records and the confirmation of existing sites of mistletoe-oaks
were undertaken in 1996 and 1997 in order to provide firm evidence of their current and historical
distribution in Britain.

METHODS

B.S.B.I. RECORDERS AND BIOLOGICAL RECORD CENTRES

B.S.B.I. vice-county recorders, the national Biological Records Centre at Monks Wood, and local
Biological Record Centres (usually county based) in England and Wales were asked for any past
and present records.

238 J.D. BOX

PUBLISHED SOURCES

Mistletoe-oaks referred to in the published literature were followed up, if there was sufficient
information to identify the site, through correspondence with the landowner and, if necessary, by a
site visit.

County Floras for England and Wales, from Victorian times to the present, were examined for
references to mistletoe on oak as were local Floras for Bristol (Swete 1854: White 1912) and
Chepstow (Shoolbred 1920) which are in mistletoe-rich areas.

Given the density of mistletoe records in Herefordshire and surrounding counties (Perring 1973:
Briggs 1995), sources of local natural history records were checked: botanical records published
by the Caradoc & Severn Valley Field Club (Shropshire) from 1892 to 1970 when the botanical
records petered out; the Transactions of the Woolhope Naturalists’ Field Society (Herefordshire)
from 1851 to the present; the Transactions of the Worcestershire Naturalists’ Club (1847 to the
present); and the Proceedings of the Somerset Archaeological & Natural History Society (1850 to
the present).

HERBARIA

Enquiries were made with those herbaria which were considered to have large collections, to be in
mistletoe-rich areas, or to be rich in historical material: Bristol Museum & Art Gallery
(BRISTM); Dept of Plant Sciences, Cambridge University (CGE); Royal Albert Memorial
Museum, Exeter (RAMM): Royal Botanic Gardens, Kew (K); Leicester Museum (LSR):
Liverpool Museum (LIV); Luton Museum (LTN); Maidstone Museum (MNE); Manchester
Museum, University of Manchester (MANCH): National Museum of Wales (NMW): Natural
History Museum (BM): Norwich Museum (NWH); Daubeny and Field-Druce Herbaria and
Morrisonian Collection, Oxford University (OXF); University of Reading (RNG); South London
Botanical Institute (SLBD; Warwickshire Museum (WARMS).

EXISTING MISTLETOE-OAKS

The oaks listed in Table | were visited with the permission of the landowner or occupier. The girth
of each tree was measured at 1-3 m above ground level (Hamilton 1975) and used to derive the
diameter at breast height (dbh). The age of each tree was estimated using the dbh taking into
account the location in which each tree was growing (White 1994); additional estimates of age
were derived from published accounts. Mistletoe is dioecious and the mistletoe on each oak was
examined for berries in December 1996/January 1997 through 12 x 25 binoculars: mistletoes
without berries were checked again in November 1998. Records of the existing mistletoe-oaks in
this paper include the 10 km square of the national grid and the nearest village. Further details are
deposited with the Biological Records Centre as confidential records.

RESULTS

BIOLOGICAL RECORDS CENTRE

There were 21 records of mistletoe growing on oak. One was the published report of mistletoe on
several oaks along the Towy and Cotti rivers in Carmarthenshire (Webster 1885) [see below]. The
other records date from 1963 to 1976 with the majority of records dating from 1969 to 1971 as a
result of the B.S.B.I. mistletoe survey (Perring 1973). These records were followed up wherever
possible in order to confirm the identification of the host tree and its location (Table 2).

PUBLISHED LISTS OF LOCATIONS ;

Lees (1842, 1851a, 1851b) gives brief details about various sites. Edward Jesse, Surveyor of the
Royal Parks & Palaces, lists four sites (Jesse 1844). He made enquiries about mistletoe on oaks in
the Royal forests and parks but without success; timber merchants assured him that they had never
seen it on oak. Bull (1864a, 1864b, 1907) gives details of various locations; Bull (1869, 1870)
gives accounts of newly discovered mistletoe-oaks together with a list of the other known sites
(with errors in the spellings). Anon. (1873a) gives a list of sites, and Webster (1885) gives a list
without sources (with errors in the spellings of sites) together with further details of one site.
Elwes & Henry (1907) give a list as well as some further details for some of the locations; Tubuef
(1923) gives a list based on Elwes & Henry (1907). Nicholson (1932) repeats these lists and adds
two more reports. Durham (1935) refers to some of these sites, notes that many of the recorded
specimens now no longer exist, and adds the report by Anon. (1933).

MISTLETOE ON OAKS IN BRITAIN

239

TABLE 1. CONFIRMED EXISTING MISTLETOE-OAKS IN BRITAIN

Dbh Estimated
Location Oak species (cm) age (yrs)! Brief Description
Brampton Bryan Q. coccinea 65 90° One “bunch” of mistletoe (probably male) c. 7
(Herefordshire) m up in the oak which is in a group of three
[SO/3.7] oaks in parkland.
Bredwardine Q. robur 185 300-400" Two “bunches” of female mistletoe c. 9.5—-10
(Herefordshire) m up in the hedgerow oak.
[SO/3.4]
Brinsop Q. robur 107 160-170 Three “bunches” of female mistletoe c. 9.5 m
(Herefordshire) and c.15 m up in the streamside oak in arable
[SO/4.4] land.
Deerfold Forest Q. petraeax 142 190-260° One “bunch” of female mistletoe c. 10.5 m up
(Herefordshire) Q. robur in the hedgerow oak.
[SO/3.6]
Eastnor Q. robur 85 210-240° Many female plants c. 8 m up in the woodland
(Herefordshire) oak which is adjacent to a drive.
[SO/7.3]
Frampton-on-Severn (QQ. robur 146 250-270° One “bunch” of female mistletoe c. 6 m up in
(Gloucestershire) the hedgerow oak.
[SO/7.0]
Gwehelog Common Q. robur 97 140-150 One “bunch” of mistletoe (probably male) c.
(Monmouthshire) 10 m up in the oak beside a lane in a small
[SO/3.0] copse.
Leintwardine Q. borealis 63 90 One “bunch” of female mistletoe c. 2m up in
(Herefordshire) the churchyard oak.
[SO/4.7]
Putley Q. borealis 74 110 One “bunch” of female mistletoe c. 8 m up in
(Herefordshire) the hedgerow oak.
[SO/6.3]
Stretton Sugwas Q. robur 180 380’ One “bunch” of female mistletoe c. 3 m up in
(Herefordshire) the hedgerow oak.
[SO/4.4]
Windsor Q. borealis 81 115 One “bunch” of mistletoe (probably male) c. 6
(Berkshire) m up in the roadside oak.
[SU/9.6]
Notes

' Estimated ages are calculated using the methods set out in White (1994) for common/sessile oaks based on
“average site’ conditions supplemented by additional estimates described below.

> Planted around 1910.

* Range includes the estimated ages calculated using both the present dbh and the girth given in Anon. (1870).

* Range includes the estimated ages calculated using the present dbh as well as the girths given by Bull (1869),
Anon. (1930) and Tonkin (1984).

> Range includes the estimated ages calculated on the basis of “inside normal woodland” conditions (White
1994) using both the present dbh and the girth given by Anon (1866a), as well as the estimated ages derived
from the reports by Lees (1842), Bull (1864a, 1864b, 1907) and Anon. (1866a).

° Range includes the estimated age calculated using the present dbh as well as the estimated age derived from
the report by Lees (185 1a).

” Age estimated without any allowance for the oak being a very old pollard which has been in a stag’s-horn
condition since at least the 1930s.

240

J.D. BOX

TABLE 2. RECORDS OF MISTLETOE-OAKS HELD BY BIOLOGICAL RECORDS CENTRE

Location (county)

Woodyates
Manor (Dorset)

Hatfield Forest
(Essex)

No location given
(Gloucestershire)

No location given
(Herefordshire)

Deerfold
(Herefordshire)

No location given
(Herefordshire)

No location given
(Herefordshire)

No location given
(Herefordshire)

No location given
(Herefordshire)

No location given
(Lincolnshire)

No location given
(Monmouthshire)

Neen Savage
(Shropshire)

Neen Savage
(Shropshire)

No location given
(Shropshire)

No location given
(Shropshire)

Rookery Farm,
Drinkstone
(Suffolk)

NGR

SU/01.19

TL/5.1

SO/7.0

SO/3.4

SO/3.6

SO/4.4

SO/6.3

SO/6.3

SO/7.3

TF/0.2

SO/5.1

SO/67.77

SO/67.77

SO/6.7

SO/6.9

TL/95.61

Details of record

June 1970. Recorder: M. P. Yule.
“Ancient oak tree growing on edge
of wood. Two plants obviously
dying fast though large’.

1970. Tetrad.J, Recorder: P.C.
Devlin (Bishop’s Stortford).

1974. Recorder: S. C. Holland. “In
lit 29/10/74”.

1971. Recorder: A. C. Powell.
“First recorded in 1870. Hfd. C. C.
Preservation Order 1954. Now has
1 large bunch - 7 small — some
berries”.

1963. “Extinct c. 1963. Miss M.
Bardel in lit. to D. McClintock
4/12/69”.

1971. Recorder: F. H. Perring.

1970+. Recorder: D. Moxley
(Abertillery).

1970. Recorder: N. Hadfield
(Putley). “Quercus borealis. One
bunch”.

1970+. Recorder: F. H. Perring.

1976. Recorder: P. B. Grimes
(Grimsthorpe, Bourne). “Oak (only
one and the tree has been felled by a
gale)”.

1971. Recorder: F H Perring.

1967. “In lit. to D. Mc Clintock”’.

1976. Recorder: Mrs Heywood-
Waddington. “Photograph given to
F. H. Perring”’.

1967. Tetrad T. Recorder: Thomson
in lit. to D. McClintock.

1970. Recorder: A. W. Smith
(Shrewsbury).

1971. Recorder: G. Thomson.

Current status

Followed up in 1997 by letter to the
owner of Woodyates Manor
(Woodyates, Verwood) without any
reply.

Followed up in 1997 with P. C.
Devlin who has revisited Hatfield
Forest but cannot find a mistletoe-
oak.

This is the Frampton-on-Severn tree.

This is the Bredwardine tree.

Presumably the Deerfold Forest tree.

This is the Stretton Sugwas tree (F. H.
Perring, pers. comm.).

Followed up in 1997 but Mrs Moxley
had moved away.

Followed up in 1997 by letter to Mrs
Hadfield but without any reply.
Presumably the Putley tree.

This is the Eastnor tree (F. H. Perring,
pers. comm.).

Followed up in 1997 but Mr Grimes’
successor at the Grimsthorpe Estate
has no knowledge of a mistletoe-oak.

Followed up in 1997 but Franklyn
Perring does not remember a
mistletoe-oak.

Presumably the same tree as below.

Branch with mistletoe fell off some
two years after record sent in (Mrs
Heywood-Waddington, June 1997).
This is probably the Neen Savage
tree:

Followed up in 1997 with Mr Smith
who cannot remember the oak.

Followed up in 1997 with present
owners of Rookery Farm who have
no knowledge of a mistletoe-oak.

MISTLETOE ON OAKS IN BRITAIN 241

TABLE 2. CONTINUED

Location (county) NGR Details of record Current status

Orvis House, East TM/07.34 1971. Recorder: K. Dodgson. Followed up in 1997 with the
Bergholt Mr F. R. Dodgson, the owner of Orvis
(Suffolk) House, who remembers mistletoe

growing on an oak from the 1930s
(when it was artificially planted on
the tree) until the late 1970s. There
are no other mistletoe-oaks in the

vicinity.
No location given SO/7.7 1974. Tetrad T. Recorder: F. Followed up in 1997 but Mr Fincher
(Worcestershire) Fincher (Bromsgrove). “Prob. Q. had moved away.

petraea in T approx 45 metres”’. “T This may be the Bewdley tree.
only visited this square to check up

on oak, the only Worcs Mistletoe

Oak known to me and seen some

years ago. The original branch fell

off about 3 years ago but Mistletoe

has been successfully re-established

and is now 3 years old, small but

app. healthy. This was the only one

to take of 30 attempts, all on oak”.

No location given SO/8.5 1972. Recorder: H. M. Caddick Followed up in 1997 with Ms

(Worcestershire) (Chipping Camden). Caddick who was not able to provide
precise information.

No location given SO/8.6 1970. Tetrad Q. Recorder: I. F. Followed up in 1997 but Mrs

(Worcestershire) Gravestock (Bristol). Gravestock has died.

RECORDS AND REPORTS

The sites of records and reports are given below in order of the relevant vice-county. For each site,

the first record or report is given. Thereafter, records and reports are only given if they add to the

knowledge about the mistletoe-oak; those accounts that just include the site in a general list are not

given. The following terminology has been used:

Recorded: first-hand observation of mistletoe growing on oak.

Reported: second-hand report of mistletoe on oak (these vary from reports of observations by
reliable observers to apparent mis-identifications of the host tree).

DEVON (V.CC. 3 & 4)

Plymouth. Gissing (1855) reports that Edwin Lees had seen a single bush in July 1854 in the
topmost branches of a lofty though scraggy oak in the first wood by the South Devon Railway
some 2-3 miles north of Plymouth. Dr R. A. Stevens (Nature Conservation Officer, Plymouth City
Council) has not found any mistletoe in the small woods along the old railway, which is now
abandoned (pers. comm., September 1996).

Tiverton. Nicholson (1932) notes a report of mistietoe on oak at Cruwys-Morchard by Mr H. M.
Fowler in 1910. The landowner at Cruwys Morchard House has no knowledge of this tree (pers.
comm., January 1997).

SOMERSET, NORTH (V.C. 6)

Bristol. Roper (1916) records a large bunch of mistletoe near the top of Q. intermedia Boenn. [Q.
robur x Q. petraea| in Leigh Woods; the oak was of moderate girth, about 50 feet (16 m) in
height, and probably more than a century old; it grew in a fringe of ancient woodland on the
Somerset side of the Avon gorge. In her Presidential Address to the Bristol Naturalists’ Society on
20 January 1916, Roper records that she had very recently found a magnificent bunch of mistletoe
growing near the top of a lofty Q. intermedia in Leigh Woods (Roper 19138).

242 J2D BOX

Roe (1981) notes that mistletoe has only ever been recorded on oak in Somerset by I. M. Roper
in 1916 in Leigh Woods. Currently, there is no local knowledge of this oak (Tony Robinson,
English Nature Site Manager for the Avon Gorge N.N.R., which contains Leigh Woods, pers.
comm., July 1996; Tony Titchen, local botanist, pers. comm., November 1996).

WILTSHIRE, SOUTH (V.C. 8)
Salisbury. A report in Webster (1885) of mistletoe on an oak at Clarendon Park. The estate
currently has no records of mistletoe on oak (pers. comm., December 1996).

DORSET (V.C. 9)

Good (1984) refers to one record of mistletoe on Quercus with no location given.

Edmonsham. The Dorset Environmental Records Centre has a record for Edmonsham from C.
Didham dated Nov. 1977—Mar. 1978 at SU/062.119 “on oak trees, about 12”. It seems likely that
this is the record referred to by Good (1984). A visit to Edmonsham House in February 1997
found three mature limes near the house and one mature lime on the edge of the wood to the north
of the house, all with many bunches of mistletoe, but no mistletoe on oak; the owner has
confirmed that mistletoe occurs on lime, maple, sycamore and apple but not on oak (Julia Smith,
pers. comm., March 1997).

Woodyates. Record in the Biological Records Centre dated June 1970 for Woodyates Manor
(SU/01.19) annotated “Ancient oak tree growing on edge of wood. Two plants obviously dying
fast though large.”

HAMPSHIRE (V.CC. 11 & 12)

Basingstoke. A herbarium sheet of mistletoe from BM is inscribed “On an Oak in Lord Bolton’s
Park at Hackwood, Hampshire. Rev. P. Roberts. 1808”; there is also a note which indicates that
this specimen originally came from Sir Joseph Banks’ herbarium.

Bull (1864a, 1864b, 1907) refers to a report of a mature oak with mistletoe at Hackwood Park
which was described by Sir Joseph Banks in the Philosophical Transactions. This reference could
not be found after searching both subject and author indexes to Philosophical Transactions at the
Royal Society using various keywords (Banks, Hooper, Lord Bolton, Hackwood Park, mistletoe).

Townsend (1883, 1904) refers to another herbarium record of mistletoe - “Hackwood park, on
‘the mistletoe oak’, 1864; Herb. Hill” and indicates that the source for this record was the
herbarium of R. S. Hill of Basingstoke which was in the possession of Winchester Natural History
Society. Hampshire County Council Museums Service has no knowledge of the Hill Herbarium
(pers. comm., Chris Palmer, September 1996) nor does Winchester College which has had a
natural history society since 1871 (J. M. Cooper, pers. comm., November 1996).

Morris (1917) reports receiving two recent photographs of the tree bearing a bunch of mistletoe.
Rayner (1929) states that “The famous plant on the Mistletoe Oak at Hackwood Park has now
(January 1928) disappeared, having apparently been carried off during the last few days, though
those on adjacent thorns were untouched.”

The present owner of Hackwood Park has no knowledge of mistletoe on oak (B. C. Spicer,
Agent to the Hackwood Estate, pers. comm., August 1996).

Winchester. Britten (1870) refers to mistletoe in K labelled “From the Oak near Winchester, T. O.
Duke.” Arnold (1887a) refers to a specimen in BM labelled “From the oak near Winchester,
1870.”

No record can be found of the Winchester specimen in K (Sandy Atkins, pers. comm.,
November 1996). The only herbarium records for mistletoe on oak in BM are from Maidstone
(Kent) and Hackwood Park (Hampshire) (Roy Vickery, Curator of Flowering Plants, pers. comm.,
September 1996).

SUSSEX, WEST (V.C. 13)

Chichester. Bull (1864a, 1864b, 1907) refers to a report of mistletoe on oak on an estate near
Shopwyke, Chichester.

Lurgashall. Wolley-Dod (1937) notes a report of mistletoe on an oak near Hempstead Farm,
Lurgashall, in 1913 but adds that none was seen in 1915 by E.W.S. on this tree.

Northchapel. Arnold (1887a, 1887b) records seeing mistletoe on a branch of oak in Petworth
Rectory which had been collected from Northchapel in the Sussex Weald and taken to the Rectory
in the 1840s. Northchapel is close to Lurgashall and may have involved the same tree.

MISTLETOE ON OAKS IN BRITAIN 243

KENT (V.CC. 15 & 16)

Ashford. Bull (1864a, 1864b, 1907) refers to a report of an oak at Mersham Hatch, Ashford, which
had been cut down.

Maidstone. Bull (1864a, 1864b, 1907) refers to a report from Mr Dickson in 1817 of mistletoe
growing on an oak some 4 miles (7 km) from Maidstone by the side of the Medway. A herbarium
sheet with mistletoe stems and leaves in BM is annotated “May 20 [28?] 1818. Found in company
with Mr Rishon the Mistletoe growing upon the oak about four mills [sic] from Maidstone Kent by
the side of the Medway. James Dickson.” [James Dickson was one of the founders of the
Horticultural Society of London and of the Linnean Society].

Sheldwich. Anon. (1873b) reports mistletoe on oak at Lord Sondes’ Park, Lees Court.

Lees Court is at Sheldwich, just south of Faversham. Sir George Sondes, Earl of Faversham, was
the owner in the 17th century and ownership continued in the family until late 19th century
(Canterbury Library and Faversham Library, pers. comms., December 1996). Lees Court Mansion
House and surrounding gardens were sold in 1971; there is no knowledge of any relevant records
neither for the formal gardens nor for the parkland (Lees Court Estate, pers. comm., April 1997).

SURREY (V.C. 17)

Dunsfold. Bull (1864a, 1864b, 1907) refers to a report of a 150 year old oak with one bunch of
mistletoe growing alone in a pasture at Burningfold Farm, Dunsfold. Salmon (1931) refers to the
report in Bull (1864b) and adds that “I saw it there in 1917 - oak is Q. pendunculata”’.

There is an herbarium sheet for mistletoe at SLBI which includes both a specimen of mistletoe
from an apple as well as a twig with oak leaves which are clearly Q. robur (= Q. penduculata) with
the annotation “Lower leaves from Oak at Burningfold, Surrey with a twig of the Mistletoe Oak.
28 August 1921. Coll. RWR.” This herbarium sheet was originally part of the London Natural
History Society herbarium and R. W. Robbins, of Limpsfield, Surrey, was Recorder of the
Botanical Section of the London N.H.S. (Peter Holland, Herbarium Curator, pers. comm., October
1996).

Lousley (1976) reports a failure to refind mistletoe on oak in Surrey, including the famous
mistletoe-oak at Dunsfold. The present owner of Burningfold Manor, which is the farmhouse, has
never seen mistletoe on the mature oaks (M. J. Simmonds, pers. comm., March 1996)

Farnham. Bull (1907) in a postscript refers to a report of mistletoe on an oak at the Bishop’s Park
at Farnham. Farnham Park was the private deer-park attached to Farnham Castle, home of Bishop
of Winchester, and now belongs to Waverley Borough Council. The Ranger has no knowledge of
mistletoe on oak (Ron Hills, pers. comm., April 1996).

Godalming. Jesse (1844) records part of an oak branch with mistletoe growing on it being sent to
him. Bull (1864a, 1864b) refers to this record but was not able to find out about this tree.
Norwood. Aubrey (1719; vol. II, p. 34) reports mistletoe on an oak which was felled about 1657.
Richmond. Maxwell (1900) notes a report of mistletoe on an oak in Richmond Park which he
could not confirm. There is no current knowledge of mistletoe on oak (Jayne Bragham, Assistant
Manager, Richmond Park, and Bill Cathcart, Crown Estate Office, Windsor Great Park, pers.
comms, June 1996).

ESSEX (V.CC. 18 & 19)

Britten (1870) reports Turner, without giving the source, as saying “I never sawe more plentye of
righte Oke miscel, then Hugh Morgan shewed me in London. It was sent to him oute of Essex:
where there is more plentye then in anye other place in Englande that I have ben in.”

Epping Forest. Jermyn (1974) notes that mistletoe still persists on oak in Epping Forest. The only
mistletoe currently present in Epping Forest is growing on apple, hawthorn and poplar at the north
end just outside the Forest boundary (Jeremy Dagley, ecologist with the City of London
Corporation which owns Epping Forest, pers. comm., April 1996).

Great Dunmow. Jermyn (1974) notes that current records include an oak at Marks Hill, Great
Dunmow [the records for this flora no longer exist]. A Phase I habitat survey of this wood in June
1993 found no mistletoe (Adrian Knowles, Essex Wildlife Trust, pers. comm., April 1996) and the
owner of this wood has no knowledge of a mistletoe-oak (R. Wallis, pers. comm., April 1996).
Hatfield Forest. Record dated 1970 in the Biological Records Centre. The recorder has revisited
Hatfield Forest but cannot locate the tree (P. C. Devlin, pers. comm., January 1998).

244 J.D. BOX

Woodford/Laughton. Warner (1771) reports that mistletoe is “Found on an oak, between
Woodford Row and The Bald Faced Stag, near The Ten Mile Stone: and on an apple-tree in an
Orchard in Loughton: and on several trees, many of them oaks, between that place and Mr
Conyers’s, Copped Hall.” Woodford, Loughton and Buckhurst Hill (Bald Faced Stag) are adjacent
to Epping Forest but are now part of the urban fringe around London and the mistletoe-oaks are
long gone (Dr K. J. Adams, B.S.B.I. Recorder, pers. comm., April 1996).

BERKSHIRE (V.C. 22)
Windsor. Maxwell (1900) notes a report of mistletoe on oak in Windsor Forest which he was not
able to confirm. There is a record of one bunch of mistletoe on red oak (Q. rubra, syn. Q. borealis)
growing as a roadside tree in Windsor Great Park; probably male as no berries present in January
1997 or November 1998 (Bill Cathcart, Crown Estate Office, Windsor Great Park, pers. comm.).
Visit on 12/6/97. Mature Q. borealis growing on light sandy soil with acid grassland and heather
nearby; tree is in good condition with a girth of 2:54 m at 1-3 m; one globose bunch of mistletoe
hangs down from a lateral branch at some distance from the main trunk and c. 5-6 m above the
ground.

SUFFOLK (V.CC. 25 & 26)

Drinkstone. Record dated 1971 for Rookery Farm, Drinkstone in the Biological Records Centre.
There is no local knowledge of a mistletoe-oak at Rookery Farm (D. & H. Donaghy, pers. comm.,
July 1997).

East Bergholt. Record dated 1971 in the Biological Records Centre. The recorder has stated that
mistletoe was artificially planted on an oak in the 1930s where it grew until the late 1970s (F. R.
Dodgson, pers. comm., June 1997).

NORFOLK, EAST (V.C. 27)
Aldeby, Woodbastwick & Shotesham. Trimmer (1866) reports mistletoe “On oak at Aldeby and
Woodbastwick,...” and ““..on the oak at Shotesham.”

CAMBRIDGESHIRE (V.C. 29)

Borley Wood. Evans (1939) reports that mistletoe is very scarce in the county, but seems to be
native on the oak at Borley Wood. The Ancient Woodland Inventory for Cambridgeshire (1987)
indicates that 48 ha of this 50 ha wood have been felled and replanted; the wood has been visited
by recorders for the Cambridgeshire Flora Project who have not recorded mistletoe (N. P. Millar,
Cambridgeshire Biological Information Service, pers. comm., December 1996).

NORTHAMPTONSHIRE (V.C. 32)

Rockingham Forest. Bull (1864a, 1864b, 1907) reports that mistletoe formerly grew on an oak
near Moorshay Lawn in Rockingham Forest. The current spelling is “Morehay Lawn”. It is a relic
from the grassy launds of the medieval compartmented Forest, and there is no mistletoe-oak (Jeff
Best, pers. comm., July 1996).

GLOUCESTERSHIRE (V.CC. 33 & 34)

Riddelsdell, Hedley & Price (1948) report mistletoe on various trees including oak around
Cheltenham, on oak at Frampton , on oak at Sedbury Park, and on oak at Drybrook and Aylburton.
Holland (1986) notes that the county was well surveyed during the B.S.B.I. Winter Mistletoe
Count of the 1970s; refers to the existence of the Frampton oak; but reports that the four other oak
trees mentioned by Riddelsdell et al. (1948) have not been located.

Cheltenham. Anon. (1873b) reports an account of mistletoe on oak some two miles (3 km) from
Cheltenham. Elwes & Henry (1907) were unable to get any confirmation of this report.

Chepstow. Bull (1864a, 1864b, 1907) refers to a report of an 85 year old oak on the northern
mound of Badams Court, Sedbury Park with one bunch of mistletoe with berries.

A visit to Badams Court in October 1996 found an oak (Q. robur, c. 200 years old) on the raised
ground to the north of the house, but no mistletoe; the father-in-law of the present owner moved
down to Badams Court seventy years ago and has never seen mistletoe on the oak; there is
abundant mistletoe (both sexes) on an apple tree next to the house and two bunches on a nearby
aspen. 7

MISTLETOE ON OAKS IN BRITAIN 245

Frampton-on-Severn. Lees (1851a, 1851b) reports a first-hand account by Professor Buckman of
Cirencester Agricultural College of a century old oak with mistletoe. Bull (1864a, 1864b) refers to
a report by Mr Clifford of Frampton Court [the landowner] that the mistletoe was dead and the
branch decayed. Subsequently, Bull (1907) refers to a report that the mistletoe had sprouted from
the trunk of the oak. Elwes & Henry (1907) report that the mistletoe-oak was still living in 1904 on
the authority of H. Clifford [the landowner]. Holland (1986) records that the ancient hedgerow oak
(Q. robur) on farmland near Frampton-on-Severn was still standing in 1974 with one large clump
of mistletoe near the trunk.

Visit in February 1997. Mature Q. robur growing in a hawthorn hedge with pasture on both
sides; appears to be an old pollard that has long grown out; in good condition with some dead
branches and a major split in the trunk (? lightning) which has healed; girth is 4-6 m at 1-3 m. One
straggly/globose bunch of mistletoe growing at base of a main branch on the north side of the tree
at c. 6 m above the ground. There were at least five insertions of mistletoe - two on underside of
branch, three on east side of branch; berries were definitely present on one of the insertions but it
was too late in the season to be certain about the others.

MONMOUTHSHIRE (V.C. 35)

Goitre. Jesse (1844) reports mistletoe on oak at Penporthleuny in the parish of Goitre. Bull (1864a,
1864b, 1870) refers to this report but notes that the mistletoe had been cut down some 25 years
previously.

Gwehelog Common. The only mistletoe-oak currently known in v.c. 35 is at Gwehelog Common
(Trevor Evans, B.S.B.I. Recorder, pers. comm., May 1996); this is the Q. robur reported without a
specific location in Evans (1995).

Visit on 15/2/97. Mature Q. robur growing with young hazel, holly, elder and bramble, near
other oaks and ash; the tree is on a slope in a small triangular area which appears to have once
been common land but is now fenced; the tree is some 50 m north of the fork in the track on the
eastern side of the eastern lane; girth is 3-05 m (at nght-angles to main axis of tree) at 1-3 m
(measured from ground level on upper side of tree) . One globose bunch of mistletoe with at least
five insertions on north side of main trunk at c. 10 m above ground level; probably male as no
berries present. Absence of berries confirmed in November 1998 (Derek Parks, pers. comm.).
Llangattock Lingoed. Bull (1870) includes a sketch and a description of the mistletoe-oak; the
sketch shows an oak with spreading branches growing in a wood. The accompanying text records
Q. sessiliflora [= Q. petraea] of around 80 years old with a girth of 4’6” at 5’ (1-37 m at 1-5 m)
growing in the hedgerow of a narrow coppice at the Hendre; it had one bunch growing from the
underside of a branch. Elwes & Henry (1907) report that this tree is not known to exist at the
Hendre.

There is an oak (c. 150-200 years old) growing on the east side of the lane some 200 m south of
entrance to The Hendre; it has been high pollarded but looks dead and is smothered with ivy; there
are two other oaks within 100 m, but both look less than 100 years old. The oak used to be part of
a small copse which was cleared for arable land; the family of the current landowner can still
remember this being called Gipsy Patch because gipsies would stop under the tree and collect
mistletoe up to 20 years ago (Sue Llewelyn, pers. comm., November 1996).

St Dials. Jesse (1844) includes St Dials in a list of mistletoe hosts and locations. Bull (1864a,
1864b) refers to this report but notes that the tree had been cut down about 12 years previously.
Usk. Mistletoe oak was reported near Usk by Jesse (1844). Bull (1864a, 1864b) refers to this
report but notes that the tree had been cut down some five years previously.

HEREFORDSHIRE (V.C. 36)

Brampton Bryan. Mistletoe was recorded on scarlet oak, Q. coccinea, in Brampton Bryan Park by
Geoffrey Messer, a local botanist, in an unpublished MS dated 1988 (Tom Wall, English Nature
Site Manager, pers. comm., June 1996).

Visit on 30/12/96. A group of three scarlet oaks (Q. coccinea) planted around 1910. The
mistletoe is on the northernmost oak; the girth is 2-04 m at 1-3 m. A well developed mistletoe plant
growing on the north side of the main trunk at the junction where a side branch has broken off (c. 7
m above ground level); straggly not globose; probably male as no berries present. Absence of
berries confirmed in November 1998 (Victoria Harley, pers. comm.).

246 JeDEBOX

Bredwardine. Anon. (1870) records the discovery of mistletoe on Q. pedunculata [= Q. robur|
about half a mile west of Moccas Park by Sir George Cornewall in conjunction with a photograph
and a description of the mistletoe-oak as one of the “The Remarkable Trees of Herefordshire.” The
photograph shows the oak as a mature tree with spreading branches which is on the edge of a field:
the oak is reported as having a girth of 11°6” at 5’ (3-51 m at 1-5 m), being in a hedgerow, and
having mistletoe growing in fifteen different places on the tree. Elwes & Henry (1907) record H. J.
Elwes being shown this mistletoe-oak by Sir George Cornewall in 1902. A record dated 1971 in
the Biological Records Centre states “Now has | large bunch - 7 small bunches - some berries”.
Two bunches of mistletoe were reported on the oak in the 1980s (Stephanie Thomson, B.S.B.I.
Recorder, pers. comm., April 1996). The mistletoe-oak is covered by the County of Hereford Tree
Preservation Order No. 7 made in 1953.

Visit on 31/12/96. Mature Q. robur with many staghorn and dead branches. It is one of a pair of
oaks on a fenceline at the top of a steep bank between two fields; the other oak shows signs of
pollarding and is not staghorned. Girth of mistletoe-oak is 5-82 m at 1-3 m above ground level on
the upper side of the bank. Two bunches of mistletoe were seen on the south-east side of the main
trunk at c. 9-5-10 m above the top of the bank; both bunches had berries and neither was globose.
Brinsop. Mistletoe on an oak was first seen in the 1990s (David Lovelace, pers. comm., June
1996); an oak with two bunches of mistletoe at the same place was seen in 1993 (Tom Wall, pers.
comm., March 1996); and one good bunch of mistletoe was seen on Q. robur by a small stream
(Stephanie Thomson, B.S.B.I. Recorder, pers. comm., April 1996).

Visit on 31/12/96. Three bunches of mistletoe on mature Q. robur growing on bank of a stream;

girth is 3-35 m at 1-3 m. One large globose bunch of mistletoe at c. 9-5 m above ground level
which may be more than one plant and is inserted either on the main trunk, or on a small branch
which it surrounds, or both; berries present. One of the upper bunches appears dead as the foliage
is very brown; there is a third bunch at c. 15 m above ground level which has berries.
Deerfold Forest. Bull (1869a, 1869b, 1907) includes a sketch and a description of the mistletoe-
oak; the sketch shows an oak with spreading branches growing in a hedge. The accompanying text
records one bunch of mistletoe growing on Q. sessiliflora [= Q. petraea] which was in a hedge and
was some 50-60 years old with a girth of 5’8” at 5’ (1-73 m at 1-5 m). Anon. (1930) refers to a
visit by the Woolhope Field Naturalists’ Club to the mistletoe-oak at Haven Farm which had a
circumference of 9’ 9 1/2” at 5’ [2-98 m at 1-5 m]. The oak was estimated to be 130 years old; the
mistletoe was observed to be growing in the same spot on the same branch as it was in 1870
although it was no longer near the top of the tree [the reference is to Bull (1869) which was issued
in 1870]. A record in the Biological Records Centre states “Extinct c. 1963”. Tonkin (1984)
describes a visit to the oak in March 1984, notes that the mistletoe was no longer there, and
speculates that the branch with mistletoe was possibly blown off in the gale on 2 January 1976; the
girth was measured as 13’ 9” at 5’ (4-19 m at 1-5 m).

Visit on 31/12/96. Mature oak (which appears to be a hybrid between Q. robur and Q. petraea)
with spreading branches in a hedgerow with hawthorn, blackthorn, honeysuckle, Rosa sp.; the oak
has no sign of staghorn branches or disease; girth is 4-47 m at 1-3 m. One bunch of mistletoe
comprising two major plants and some minor ones at c. 10-5 m above ground level: berries
present. The mistletoe is on the west side of the western of the two main trunks (which divide
around 6-5 m above ground level). This is different from the position shown in the 1869 sketch
(Bull 1869) but a major branch was lost prior to 1984 (Tonkin 1984).

Eastnor. Beaton (1837) records being given a shoot of oak with two mistletoe plants on it from an
oak on Earl Somers’ estate at Eastnor Castle. It is reported that there are several mistletoe plants,
one of which is of great age and nearly 5’ (1-5 m) in diameter. The editor of The Gardener’s
Magazine was sent the oak branch with the letter from D. Beaton and notes that the mistletoe
appears to be male and the oak appears to be Q. sessiliflora [= Q. petraea]. Loudon (1838) reports
an oak growing near Eastnor Castle with several plants of mistletoe, one of which was of great age
and vigour with its branches being nearly 5 feet (1-6 m) in diameter. A large and handsome
specimen of this mistletoe was sent to him by Mr Beaton in March 1837 and exhibited on 4 April
1837 at the meetings of the Horticultural Society and the Linnean Society. The former is
confirmed by Anon. (1837) who reports that mistletoe on oak from Mr Loudon, to whom it was
sent by Mr D. Beaton of Haffield, near Ledbury, was exhibited on 4 April 1837 at the London
Horticultural Society. The latter is recorded in the General Minute Book of the Linnean Society for

MISTLETOE ON OAKS IN BRITAIN 247

1830-1837, pp. 348-349. Lees (1842, 1851a) records that in June 1837 he saw a young oak, about
70-80 years old, with four fine bushes of mistletoe in Earl Somers’s park, at Eastnor, near
Ledbury, on the side of the Ridgeway; enquiries and a careful examination of 300 acres (120 ha)
of oak wood revealed no others.

Bull (1864a, 1864b, 1907) records a visit to the 80-90 year old oak on the side of the drive
leading from the Park up the Ridgeway hill towards Malvern; both male and female mistletoe was
present and it grew in three large bunches and in four other places on the tree. Anon. (1866a)
includes a photograph and description of the mistletoe-oak as one of a series of “The Remarkable
Trees of Herefordshire’; the photograph shows the oak growing in very open woodland (possibly
recently thinned) with a yew tree immediately adjacent to it. The accompanying text refers to
mistletoe growing in seven places on Q. pedunculata [= Q. robur| which is more than 100 years
old and has a circumference of 5’ 3” at 5’ [1-6 mat 1-5 m].

A herbarium specimen of mistletoe held at K is annotated “From the oak in Eastnor Park,
Ledbury. Two large plants grow on this oak, both are on one branch and run together. The oak was
in young leaf... J. D. Hooker, April 20, 1878”.

Hadfield (1974) refers to the mistletoe-oak reported by Loudon (1838) and confirms that it was
still present in 1973.

Visit on 31/12/96. Mature Q. robur growing in mixed deciduous woodland (ash, yew, oak, field
maple and hazel over dog’s mercury) on limestone; the woodland comprises c. 100 year old oak
standards over hazel/field maple regrowth; the girth of the oak is 2-67 m at 1-3 m. Luxuriant
growth of mistletoe, with many plants inserted into the top and the bottom of branches as well as
on the main trunk at c. 8 m above ground level on north/north-east side; berries present. The yew
tree next to the oak has a divided main stem and appears to be very similar to the yew in
photograph of the mistletoe-oak in Anon. (1866a).

Hereford. Anon. (1933, p. xv) reports mistletoe growing on oak in the garden of Vanham House,
Pengrove, within the city boundary. It has not been possible to locate Vanham House nor to trace
the name in contemporary reference sources and maps.

Ledbury. Beaton (1837) writing from Haffield, near Ledbury, notes a report of mistletoe on an oak
close to a willow loaded with mistletoe on a farm near Ledbury; the oak had been cut down six
years previously.

Leintwardine. Report of a mistletoe-oak in the churchyard of St Mary Magdalene, Leintwardine
(Paul Hand, pers. comm., December 1996).

Visit in December 1996. Red oak (Q. borealis) with a girth of 1-98 m at 1-3 m. One globular
bunch of mistletoe made up of a number of separate main stems fused together; inserted on the
top, the side and the bottom of an east pointing branch, c. 2 m from the main trunk and c. 2 m
above ground level; berries present.

Llangarron. Purchas & Ley (1889) report a large oak with mistletoe at Llangunnock Farm,
Llangarren (sic) discovered by Mr T. T. Mayos.

The owners of Llangunnock Farm for 30 years now live in a new house nearby and have not seen
mistletoe on the oaks in the area (Mr & Mrs Jenkin, pers. comm., September 1996).

Moccas. There is a report of a tree near Moccas House which was doing well in 1970 with a dozen
bunches of mistletoe (Stephanie Thomson, B.S.B.I. Recorder, pers. comm., April 1996).
Subsequent inspection of the tree found no mistletoe; but Ivor Saunders, the gardener at Moccas
Court, reports that there had been mistletoe on a branch which was blown off in a gale two or three
years ago (Stephanie Thomson, pers. comm., February 1997).

Morehampton. Bull (1864a, 1864b, 1907) refers to a report of a mistletoe on oak somewhere in
the parish of St Margaret’s, Moorhampton (sic). A farmer at Morehampton Park Farm had no
knowledge of such a tree (pers. comm., May 1996) neither had the vicar of St Margaret’s,
Morehampton (F. E. Rodgers, pers. comm., September 1996).

Putley. Record of one bunch on red oak (Q. borealis) by roadside (Stephanie Thomson, B.S.B.I.
Recorder, pers. comm., April 1996).

Visit on 31/12/96. Healthy, mature red oak (Q. borealis ) with a spreading crown growing by the
roadside in a row of six mature red oaks with hawthorn, hazel, ash and Prunus sp.; the girth is
2-31 m at 1-3 m. One globose bunch of mistletoe growing at the base of the main branch on east
side of tree at c. 8 m above ground level; inserted on top of the branch; berries present.

248 J.D. BOX

Stretton Sugwas. Anon. (1928, page lvi) gives a brief report without details. A herbarium
specimen of mistletoe at K is annotated “30.vu1.35. Stretton Sugwas. Viscum “from oak’. Coll.
30.7.35. Recd. August 1935. Dr Durham” [mistletoe on oak at Stretton Sugwas is mentioned
without any details by Durham (1935)]. There is a record dated 1971 in the Biological Records
Centre (see Table 2). In 1996 the tree is much as it appears in a photograph from the 1930s which
shows a stag’s-horn tree (Stephanie Thomson, B.S.B.I. Recorder, pers. comm., April 1996).

Visit on 31/12/96. Q. robur growing in a hedgerow; tree is a very old pollard covered in

brambles and ivy; upper branches fairly bare, lower ones showed good leaf growth; approx. girth
5-64 m at 1-3 m measured on the field side; one globose bunch of mistletoe on a lower branch at c.
3 m above ground level; many insertions on top and bottom of branch; berries present.
Tedstone Delamere. Lees (1851b) refers to a report from Rev. Canon Cradock of two young oaks
in the parish of Tedstone-de-la-Mere on banks of Sapey Brook near Knightsford’s Bridge. Bull
(1864a, 1864b, 1907) refers to the discovery in 1853 of a mistletoe-oak by Dr Cradock (Principal
of Brasenose College); he reports that the oak is 60 years old, has one bunch of female mistletoe,
and has been known about by the woodman for some 30 years. Anon. (1866b) includes a
photograph and description of the mistletoe-oak as one of the series “The Remarkable Trees of
Herefordshire’; the text refers to mistletoe growing in one bunch of five stems some 40 feet (c. 12
m) above the ground on Q. pedunculata (= Q. robur) in the corner of a wood on Primrose Hill; the
oak was recorded by Dr Cradock in 1851 and a woodman had known of it for more than 20 years
before; a Mr Evans (presumably the landowner) is recorded as cutting the ivy from the tree and
promising to remove the trees around the mistletoe-oak. Anon. (1867b) reports a visit to the
mistletoe-oak and correspondence with the landowner, Mr Bickerton Evans of Whitbourne Hall,
concerning the removal of ivy from the tree.

The parish is now incorporated into the parish of Greater Whitbourne. The oaks on the west of
the Sapey Brook were felled in the 1950s/1960s (Richard Belville, pers. comm., February 1996);
the owners of the woodlands on the other side of the Sapey Brook have looked at the oaks around
Primrose Cottage, but found no mistletoe (pers. comm., Julia & Bill Evans, April 1996).

Tretire. There is a herbarium specimen of mistletoe in BRIST held in BRISTM; annotated
“Viscum album on large oak tree in meadow on Tredunnock Farm, Tretire, Herefordshire. Sept.
29, 1927. E. Armitage” and “I saw the mistletoe on this tree many years ago, & there are a large
number of branches scattered about in the tree. E. A.” [Eleanora Armitage studied the flora of
Herefordshire and Gloucestershire].

The current owner of Tredunnock Farm has no knowledge of any oaks with mistletoe; there may
have been a fairly old oak in the hedgerows which were removed to make way for the airstrip at
the back of the farm (Julie Smith, pers. comm., March 1996).

Woodbury Hill. Cornewall (1903, pp. 104-105) records that he discovered a mistletoe-oak in
January 1903 on the west side of Woodbury Hill on the Moccas Estate. He estimated the age as
220 years old and noted that the tree was picturesque and gnarled; it was in company with a row of
large limes which had been felled.

There has been woodland thinning and conifer planting in the area, and the oaks on Windy
Ridge are only around 75 years old (Simon Quarn, farm manager at Wilmaston Farm, pers.
comm., June 1996).

WORCESTERSHIRE (V.C. 37)

Bewdley. Rea (1910, p. 272) records a mistletoe-oak growing on the county boundary at Dowles
parish. Salter (1928) records one from the grounds of Mr J. Steele Elliot, Dowles Manor, Bewdley.
A record by L. C. Lloyd is reported in Dallman (1939) as “A flourishing plant of some years’
establishment on an Oak tree (Q. robur L.) in Mr J. Steele Elliott’s grounds at Dowles Manor, just
inside the Shropshire border (Lloyd). The only instance of its growth on oak so far recorded in the
county” [the oak is in fact just in Worcestershire - see below].

John Robinson (English Nature Site Manager, Wyre Forest N.N.R.) was shown this oak by
Steele Elliot’s daughter, Mrs P. Tristram-Ede (pers. comm., December 1996). She can recall one
very large bunch of mistletoe on it in the 1940s & 1950s which was too high to pick. This was on a
branch which is now dead and has ropes for a swing hanging from it.

Visit in December 1996. It is a 180/200 year old oak which is a hybrid between Q. robur and Q.
petraea, although tending towards Q. petraea. The oak grows at the top of the north bank of the

MISTLETOE ON OAKS IN BRITAIN 249

Dowles Brook at a point which is just in Worcestershire (the county boundary deviates from the
brook at this point). The oak is in a stream corridor and near to a field.

Bromyard. Anon. (1878) records two oak trees with mistletoe in Buckenhill Coppice, near
Bromyard.

Droitwich. Report of Q. robur which used to have a single bunch of mistletoe, but the branch with
the mistletoe dropped off a couple of years ago; the oak was in the hedgerow which ran along
Green Lane, Hadzor, just to the east of the M5 at Droitwich (SO/910.622) (Andrew Fraser, pers.
comm., February 1996).

Site visits in March and September 1996 found two oaks in the hedge on the west of the lane,

but no sign of mistletoe, although there is an abundance of mistletoe on limes in the vicinity.
Knightsford. An herbarium specimen of mistletoe at K is annotated “Viscum album from oak.
Knightsford Worcester May 1875. ex Herbario Joannis Fraser MD. Presented by the Trustees of
the Shrewsbury Public Library, June 1954.” The bridge on the main road over the River Teme at
Knightwick is Knightsford Bridge. It is probable that this herbarium specimen came from the
Knightwick oak (see below).
Knightwick. Anon. (1871) records mistletoe on oak near Knightwick Church and reports that this
is a recent discovery. A local resident describes an oak between Knightwick and Suckley which
has always been known as the “mistletoe oak’; this tree is some 400 m from the old church which
was replaced by a mortuary chapel about 100 years ago; she does not remember mistletoe growing
on it and neither did her parents who were both born in the parish in 1885 (Mary Walker, pers.
comm., January 1997).

A visit in February 1997 found a mature Q. robur (c. 200 years old) in a hedgerow on east side
of the road from Knightwick to Suckley, approx. halfway between Knightwick Manor and the
main road. There was no sign of mistletoe on the oak and there were no other mature oaks around.
Lindridge. Anon. (1872) confirms a report of mistletoe on oak.

Stoulton. Anon. (1884) records mistletoe on oak; this was still extant in 1901/1902 (Rea 1903).

STAFFORDSHIRE (V.C. 39)

Stafford. An herbarium sheet with a stem and leaves of mistletoe in the Morisonian collection,
OXF, is annotated “Viscum quercinum ex agro Staffordiens, donum D. Thurstin, Orielens.
1690” [Translation: Mistletoe of oak from a field near Stafford, given by D. Thurstin of Oriel
College].

SHROPSHIRE (V.C. 40)

Sinker et al. (1985) indicate that 2% of the mistletoe records for the county are associated with
oaks without giving further details [in fact the data came from the 1969/70 B.S.B.I. mistletoe
survey (lan Trueman, pers. comm., January 1996)].

Neen Savage. Records dated 1967 and 1976 are in the Biological Records Centre. The branch with
mistletoe fell off some two years after 1976 record was sent in (Mrs Heywood-Waddington, pers.
comm., June 1997). A visit to the oak in June 1997 found no mistletoe.

WEST GLAMORGAN (V.C. 41)
Vale of Neath. Bull (1864a, 1864b, 1907) refers to a report of an oak which was 2 miles (3 km)
below Aberpergwm House but which was blown down in a storm.

CARMARTHENSHIRE (V.C. 44)

There are reports of mistletoe on several oaks in the remains of the old forest along the courses of
the Towy and Cothi rivers (Webster 1885). May (1967) cites this report in an appendix of species
presumed lost or extinct in Carmarthenshire. There are no recent records of mistletoe from the
Tywi or Cothi valleys nor any current records of mistletoe on oaks in Carmarthenshire (Pryce
1996).

CARDIGANSHIRE (V.C. 46)

Gogerddan. Salter records a mistletoe-oak on 26/12/27 in his unpublished diaries (National
Library of Wales MS 14445B) and records that the mistletoe-oak was adjacent to a tall poplar
which also had mistletoe (Salter 1928). Subsequently, he mentions the poplar as being felled and
refers to mistletoe growing on lime, but makes no reference to the oak (Salter 1935). Currently,
there is no mistletoe present at this site (A. O. Chater, B.S.B.I. Recorder, pers. comm., September
1996).

250 J.D. BOX

MONTGOMERYSHIRE (V.C. 47)

Trueman et al. (1995) report that a recent survey found mistletoe on oak, but no details are given.
The records came from David Orton who undertook a mistletoe survey in 1993/94 (Marjorie
Wainwright, B.S.B.I. Recorder, pers. comm., January 1996).

Abermule. Report from 1993/94 of a very small mistletoe plant on Q. robur at Penybryn; the
mistletoe is not growing there now (David Orton, pers. comm., June 1996).

CAERNARVONSHIRE (V.C. 49)

Anglesey. Lees (1842, 1851a) notes that Mr J. F. Dovaston reported in Loudon’s Magazine of
Natural History (vol. 5, p. 203) that he saw mistletoe growing on oak and “what is more singular,
hanging almost over a very grand druidical cromlech” in the Marquis of Anglesea’s park at Plas
Newydd, on the island of Anglesey.

It has not been possible to find the original report by Dovaston. Prior to 1842, there are two
complete series of Loudon’s Magazine of Natural History but there is no reference to Dovaston,
nor to mistletoe, in any of the volumes. There is no reference to Dovaston writing about mistletoe
in the Catalogue of Scientific Papers 1800-1863 compiled by the Royal Society of London (G. A.
Knowles, Manager, Current Information Services, The Royal Society, pers. comm., October
1996).

There are no longer any trees near the cromlech at Plas Newydd; most of the trees are believed
to have been felled around the turn of the century and the last by the 1950s (National Trust, pers.
comm., March 1996).

LEICESTERSHIRE (V.C. 55)

Leicester. Report by Pulteney (1795, p. clxxix) “In Quorn Wood. On the oak, and other trees. On
apple-trees in old orchards.” Pulteney (1795) contains a lot of records from Buddon Wood; the
entry for Rubus idaeus (p. clxxx1) states that it is found in Quorn or Buddon Wood and the names
were evidently interchangeable. Buddon Wood lies just to the south of the village of Quorn. It is a
large ancient wood which was clear-felled soon after the Second World War and mistletoe has not
been seen since Pulteney’s day (Michael Jeeves, B.S.B.I. Recorder, pers. comm., September
1996).

NOTTINGHAMSHIRE (V.C. 56)

Ordnoyo (1807) notes that mistletoe occurs on various hosts including oak but gives no details.
Sherwood Forest. Maxwell (1900) includes a report which he could not confirm. There is no
current knowledge of mistletoe on oak (Martin Freeman, Sherwood Forest Visitor Centre, pers.
comm., February 1997).

DERBYSHIRE (V.C. 57)

Staveley. Arnold (1887a) reports, without giving a source for the quotations, that “in one of
Colepeper’s MSS. at the British Museum, in a curious notice of Sir Peter Freschville’s house at
Staveley, Derbyshire, is this passage:- ‘Heare my Lord Freschville did live, and heare grows the
famous Mistletoe tree, the only oake in England that bears Mistletoe.’ And to this tree the
following letter, written between 1663 and 1682, from the Countess of Danby to Mrs Colepeper,
probably refers: - ‘Dear Cozen, -Pray if you have any of the mistleto of yo" father’s oke, oblige me
so far as to send sum of it to yo" most affectionat servant, Bridget Danby.’”’

The British Museum has a large collection of letters to and between Colepeper and his wife,
including the undated note of Lady Danby to Mrs Colepeper: “Deare Cosen / pray if you haue any
of the Miselto of yo' fathers oke oblidge me so fare as to send sume of it to y' most / affectionat
sarviant / Bridgett Danby” (folio 46, Harley MS 7005). Colepeper’s lengthy notes relating to
Staveley occupy folios 139-289 of Harley MS 7602, the volume covering the letter ‘S’ of his large
collection of miscellaneous notes and observations. A rapid search of the volumes covering both
‘M’ for mistletoe and ‘S’ for Staveley has not revealed any mention of mistletoe (W. H. Kelliher,
Curator of Manuscripts, The British Library, pers. comm., December 1996). [See record for
Sheffield under Yorkshire, south west (V.C. 63)].

YORKSHIRE, SOUTH WEST (V.C. 63) OR DERBYSHIRE (V.C. 57)
Sheffield. A report of an oak with mistletoe near Sheffield in the the 2nd edition of Ray’s
Catalogus Plantarum Angliae (Ray 1677) contains details which were omitted from the Ist edition

MISTLETOE ON OAKS IN BRITAIN 251

(Ray 1670). Ray (1677) has “Siquis viscum in Quercu crescentem videre desiderat, Sheffeldiam
adeat, a qua non procul inveniri eum a D. Fr. Jessop certior factus sum” [Translation: Anyone who
wants to see mistletoe growing on an oak should go to Sheffield, not far from where I have been
informed by D. Fr. Jessop, it can be found] whereas Ray (1670) only has “Siquis viscum in Quercu
crescentem videre desiderat’”’. Francis Jessop (1638-1691) of Broomhall, Sheffield, met John Ray
in 1668 and Ray published some of Jessop’s plant records (Desmond 1977). It seems probable that
this is the same tree as that referred to in the 17th century accounts concerning the mistletoe-oak at
Staveley in Derbyshire (Arnold 1887; Folio 46, Harley MS 7005, British Museum); Staveley is
only some 14 km from the centre of Sheffield.

PERTHSHIRE, EAST (V.C. 89)

Errol. Kanner (1939, p. 913) reports an old account of mistletoe growing in profusion on a vast
old oak in the neighbourhood of Errol, not far from the Falcon stone; the fate of the family of Hay
was reputed to be linked to the continued existence of this tree [see also Gurney (1848, pp. 576-
577) and Melville (1935, p. 156)]. Kanner (1939) also notes that the 13th century Scottish poet,
Thomas Rymour of Ercildowne (Thomas the Rhymer, 1220-1297), is credited with the authorship
of a poem which deals with the connection between this mistletoe-oak and the fate of the Hay
family:

While the mistletoe bats on Errol’s aik,

And that aik stands fast,

The Hays shall flourish, and their good grey hawk
Shall nocht flinch before the blast.

But when the root of the aik decays

And the mistletoe dwines on its withered breast

The grass shall grow on Errol’s hearthstane,

And the corbie roup [raven croak] in the falcon’s nest.

The estate was sold by the Hay family in the 1630s to cover the debts of the 10th Earl (Jeremy
Duncan, Local Studies Librarian, A. K. Bell Library, Perth, pers. comm., April 1997).

UNKNOWN LOCATION

Report of three plants of mistletoe on oak seen by a Mr Knowlton in August 1765 on the estate of
White Esq. at Watling Wells (Dillwyn 1843). Knowlton worked at Londesborough Hall in the East
Riding of Yorkshire (John Edmondson, pers. comm., October 1998), but it has not been possible to
trace Watling Wells despite the assistance of the Yorkshire Museum, the English Place-Name
Society and the Ordnance Survey Library.

DISCUSSION

The earliest known report of mistletoe on oak in Britain is the poem attributed to the 13th century
Scottish poet, Thomas the Rhymer, describing the mistletoe-oak at Errol in Perthshire (Kanner
1939; see also Gurney 1848 and Melville 1935). There are three references to mistletoe on oak in
the 17th century: the mistletoe-oak at Norwood (Surrey) felled in 1657 (Aubrey 1719); mistletoe
on oak near Sheffield (Yorkshire) reported by Ray (1670, 1677); and the herbarium specimen of
mistletoe from an oak from Stafford (Staffordshire) dated 1690.

There was considerable interest in mistletoe and its host species from the 19th century until the
early part of the 20th century and various accounts contain details of mistletoe-oaks in Britain
(Lees 1842, 1851a; Bull 1864a, 1864b; Anon. 1873a; Webster 1885; Arnold 1887; Purchas & Ley
1889; Bull 1907; Elwes & Henry 1907; Somerville 1914; Tubuef 1923; Nicholson 1932; Durham
1935). Bull (1864a, 1864b) built on the initial observations of Lees (1842, 1851a) and gives details
of six records/ authenticated reports of mistletoe-oaks. Footnotes are given in the later reprint of
this paper (Bull 1907) which extended the list of records/authenticated reports to eleven trees.
Later lists of mistletoe-oaks appear to be derived from earlier accounts, often combined with
additional second-hand reports without any first-hand evidence. Nevertheless, the various lists
compiled between the middle of the 19th century and the first part of the 20th century usually
describe between ten and twenty mistletoe-oaks.

252 J/DY BOX

Currently, eleven oaks with mistletoe growing on them have been located and verified in Britain
(Table 1 and Figure 1D). The majority of the mistletoe plants are female and the mistletoe-oaks are
concentrated in Herefordshire with single examples in Berkshire, Gloucestershire and Gwent. The
records cover ten separate 10-km squares with the Brinsop and Stretton Sugwas oaks being in the
same 10 km square. This figure is similar to the twelve 10 km squares for which mistletoe-oaks
were reported in the 1969-1972 B.S.B.I. survey (Perring 1973).

Written records go back at least a century for the existing mistletoe-oaks at Bredwardine,
Deerfold Forest, Eastnor and Frampton-on-Severn. Of these, the mistletoe-oak at Eastnor has the
longest recorded history being originally reported by Beaton (1837). Only four of the eleven
muistletoe-oaks reported by Bull (1907) are still in existence: Bredwardine, Deerfold Forest,
Eastnor (all in Herefordshire) and Frampton-on-Severn (Gloucestershire). Surprisingly, the very
old mistletoe-oak at Stretton Sugwas (Herefordshire) was not known to Bull and was not reported
in the literature until 1928. The other six existing mistletoe-oaks have not been recorded/reported
in the published literature.

These results, together with the lists of between ten and twenty mistletoe-oaks in the 19th and
early 20th centuries, suggest a population of mistletoe-oaks which is apparently fairly stable in
number, but in which there is a considerable turnover as losses are compensated by the discovery
of new sites and the parasitism of new trees.

Pedunculate oak, Q, robur, is the most frequent host amongst these existing mistletoe-oaks. The
presence of mistletoe on red oak, Q. borealis, at three separate sites, as well as on a scarlet oak, Q.
coccinea, is notable as the B.S.B.L survey recorded pedunculate oak as the only host species of
oak (Perring 1973). It is interesting to note that Somerville (1914) includes a report of mistletoe on
Q. rubra (syn. Q. borealis) in Worcestershire although no details are given. Mistletoe, however, is
found on “red oaks” (Q. rubra, Q. coccinea, Q. palustris) in other parts of Europe, particularly in
France (Timbal et al. 1994).

Pedunculate oak is also the most frequent species of oak identified in past records/reports of
mistletoe-oaks which now either no longer exist or where mistletoe is no longer present on the host
tree. These records/reports identify pedunculate oak, Q. robur, at four sites [Tedstone Delamere
(Herefordshire), Abermule (Montgomeryshire), Dunsfold (Surrey), and Bewdley (Worcestershire -
although direct observation of leaves from this oak indicates that the tree has characteristics which
are more those of a hybrid oak than of a pedunculate oak)]; sessile oak, Q. petraea, at one site
(Llangattock Lingoed, Gwent); and hybrid oak, Q. robur x Q. petraea, at one site (Bristol,
Somerset).

The estimated ages of the existing oak hosts range from 90-400 years. The hosts include mature
trees, but the hosts are not mostly very old, dying trees as the B.S.B.I. survey suggested (Perring
1973). Indeed, a photograph of the Stretton Sugwas oak (Herefordshire) taken in the 1930s shows
a stag-headed tree very similar to the present tree.

The existing mistletoe-oaks do not occur in woodland but prefer more open situations such as
woodland edges, hedges, parks and even a churchyard. Timbal et al. (1994) report a similar
finding with mistletoe on “red oaks” in France. Records and reports of other oaks with mistletoe
confirm that the trees occur in open situations; locations in woodland tend to be on the edge of the
woodland, for example “in the corner of a wood” (Tedstone Delamere, Herefordshire), or in “a
fringe of ancient woodland” beside the Avon Gorge (Bristol, Somerset), or near a stream in the
Wyre Forest (Bewdley, Worcestershire).

It seems likely that the distribution of early (pre-1800) records and reports of mistletoe-oaks
(Figure 1A) reflects the places visited by early botanists and the chance survival of records. There
has been no significant change in thé distribution of mistletoe-oaks recorded and reported in the
19th century (Figure 1B) compared to that for the 20th century (Figure 1C). The pattern of
distribution for all records and reports of mistletoe-oaks (Figure 1D) is similar to the distribution
for all mistletoe records given in Perring (1973) and Briggs (1995). However, the range of existing
confirmed mistletoe-oaks (Figure 1D, solid circles) is significantly more restricted than the
historical range [Figure 1D, all circles], but coincides with the concentration of mistletoe records
in Herefordshire and the surrounding counties (Perring 1973; Briggs 1995).

In conclusion, the verification of only eleven mistletoe-oaks confirms the rarity of mistletoe
growing on oak in Britain. Past records, reports and general accounts suggest that this rarity has
not changed significantly since the 17th century when it was commented on by Ray (1670, 1677)

MISTLETOE ON OAKS IN BRITAIN 253

FIGURE 1. A. Distribution of records and reports of mistletoe-oaks pre-1800. B. Distribution of records and
reports of mistletoe-oaks in the 19th century. C. Distribution of records and reports of mistletoe-oaks in the
20th century. D. Distribution of records and reports of mistletoe-oaks (O) and existing mistletoe-oaks (@).

254 J.D. BOX

and Evelyn (1664). Indeed, mistletoe from the oak at Eastnor was considered sufficiently
noteworthy to be exhibited in 1837 at both the Linnean Society and the London Horticultural
Society. Mistletoe (Viscum album) growing on native oaks (Q. robur and Q. petraea) is a rare
association throughout western Europe (various sources are cited by Tubuef 1923 and Grazi &
Urech 1983), although there are at least 150 sites for mistletoe on “red oaks” in France (Timbal et
al. 1994). The present range of verified mistletoe-oaks is, however, less than the historical range
based on past records and reports. This may be due to the clearance of woods and hedges resulting
from the intensification of agriculture and the extension of urban areas over the past 50 years.
Within the present range, the existing mistletoe-oaks are concentrated in Herefordshire which
coincides with the core of the present distribution of mistletoe in Britain (Perring 1973; Briggs
1995).

ACKNOWLEDGMENTS

The assistance of B.S.B.I. recorders throughout Britain is very gratefully acknowledged as is the
help given by the staff of local Biological Record Centres in England and Wales and the staff of
various herbaria and libraries. Particular assistance was given by Gill Gent (B.S.B.I. recorder for
Northamptonshire), Stephanie Thomson (B.S.B.I. recorder for Herefordshire), Paul Hand
(Shropshire), Sue Liptrot and her colleagues (Shropshire Libraries), Gina Douglas (librarian and
archivist at the Linnean Society of London) and Glenys Knowles and Mary Sampson (library of
the Royal Society). The owners and occupiers of the sites of mistletoe-oaks, both past and present,
have been particularly helpful both in correspondence and through arranging site visits in winter.
Jane Croft and Henry Arnold at the Biological Records Centre of the Institute of Terrestrial
Ecology at Monks Wood have given much help by computerising the records and reports and
generating the distribution maps using Alan Morton’s DMAP programme. Lastly, I must
acknowledge the assistance given by Franklyn Perring and Jonathan Briggs whose involvement in
surveys of mistletoe has added significantly to our knowledge of this species.

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(Accepted January 2000)

Watsonia 23: 257-267 (2000) 257.

A morphological comparison between some British Orobanche
species (Orobanchaceae) and their closely-related non-British
counterparts from continental Europe:

Orobanche reticulata Wallr. s. L.

Ma YorFOEEY

Department of Biological Sciences, Institute of Environmental and Natural Sciences, University of
Lancaster, Lancaster, LA] 4YQ

ABSTRACT

A morphological comparison has been made between lowland British plants and those of the European
mountains, both referable to O. reticulata s.l.. A brief summary is given of the previous taxonomic treatments
which have been applied to this group. It is concluded that two separate but closely-related taxa exist, and that
these are best treated at subspecific level, the British plant being referred to subsp. procera (Koch) Dostal. A
possible third taxon, perhaps no more than a pale colour form of one or both of the above, has been described
at various ranks under the epithet ‘pallidiflora’. No direct support for its continued recognition has been
established.

KEYWORDS: Taxonomy, root parasites, broomrape.
INTRODUCTION

BACKGROUND

Orobanche is a genus of obligate root parasites, which completely lack chlorophyll and are
therefore unable to photosynthesise and so independently sustain themselves. To do so they
parasitise a wide range of (usually) dicotyledonous, and very often herbaceous, hosts. The genus is
distributed throughout much of the world’s northern temperate latitudes and has a northern limit in
southern Scandinavia. Only relatively few species occur in the southern hemisphere and some of
these are probably introductions.

In comparison to continental Europe, the British Isles have a very limited Orobanche flora, only
ten species being recognised by Stace (1991). This is in contrast to mainland Europe where even
the conservative treatment in Flora Europaea (Chater & Webb 1972) describes 45 at species level.
Apart from the relatively widespread O. minor Sm., which in fact may have been largely
introduced into this country, all the other British species are apparently native and several are very
rare. Four of the latter have been accorded British Red Data Book status (Wigginton 1999):
O. artemisiae-campestris Gaudin (= O. loricata Rchb.), O. caryophyllacea Sm., O. purpurea
Jacq., and O. reticulata Wallr. In Europe, however, these and many others are much more
frequent, and the abundance and species diversity increases appreciably as one moves south, with
the Mediterranean area having an especially rich Orobanche flora. Many species show close
morphological similarities one to another and can therefore be difficult to separate taxonomically.

In this paper, the relationship is examined between British plants referable to O. reticulata
Wallr. and those from continental Europe. In doing so, much reliance has been placed on the
morphometric assessment of living populations of plants, with additional information obtained
from the study of preserved herbarium material.

PROBLEMS ENCOUNTERED IN THE STUDY OF THE GENUS

This is acknowledged to be a taxonomically difficult genus (e.g. Chater & Webb 1972, Pignatti
1982, Rumsey & Jury 1991) and meaningful work, whether in the field or in the herbarium, is
difficult to achieve. For instance, these morphologically very similar taxa possess few readily

258 Nia YOROLEY,

quantifiable characters. The above-ground plant comprises little more than an inflorescence, the
stem of which bears simple, sessile, scale-like leaves which rapidly become brittle soon after
emergence. This lack of normal leaves is therefore a major loss of potential taxonomic information
and the inflorescence is the only part of the living plant which it is feasible to study.

Being fleshy and lacking chlorophyll, Orobanche plants make poor herbarium specimens,
sometimes forming a rather amorphous mass so that important characters are lost. This is
especially true of corolla and stigma colours and subtleties of corolla shape. However, from well-
pressed specimens with accompanying field colour notes, useful information can be gained.
Unfortunately, a large proportion of specimens are not preserved to such a high standard, and so it
is better to study the genus as living plants.

There are also other difficulties. Many of the original authors’ descriptions are very brief and
inadequate or even ambiguous. In such cases, recognition of the taxon to which the description is
applied may be almost impossible to achieve. Very often descriptions refer to non-diagnostic
characters, such as leaf and stem, whilst information on much more important characters is
generalised or even omitted. Type specimens are rarely indicated by these authors and when they
are known, they may be very old and by now seriously fragmented, and so taxonomically almost
worthless.

OROBANCHE RETICULATA s. |.

O. reticulata s. I. is widespread but local throughout much of the montane/subalpine region of
Europe and is also recorded from lowland habitats elsewhere on the continent. However, it was not
until shortly after the start of the twentieth century that plants apparently corresponding with the
description of continental O. reticulata were recognised in Britain. These occurred (and still do so)
in a very limited area of Yorkshire, as lowland plants, usually on the magnesian limestone where
they mainly parasitise Cirsium arvense, growing on rough calcareous grassland and scrub, and
especially on river margins and flood plains. Since there must be many similarly suitable habitats
throughout the rest of the country, this very restricted British range is difficult to explain. The
discovery in Britain in 1907 is of some interest and has been summarised elsewhere (Foley, 1993).
I have examined both the British plant and those of the European mountains on several occasions
in the field and certain morphological differences between the two are apparent which suggest the
existence of separate taxa.

BASIC TAXA RELATED TO O. RETICULATA AND SUBSEQUENT TREATMENTS

O. reticulata was first described by Wallroth (1825) from plants collected near Toulouse, France.
His description is quite vague and does not especially differentiate it from other similar taxa. He
described the calyx as divided into simple, linear-lanceolate teeth, shorter than the tubular,
inflated, 20 mm long corolla. The lobes of the lower lip were rounded and equal in size and the
filaments and style were more or less glabrous. His most significant statement referred to the
purple coloration of the corolla caused by the reticulate veining of the upper lip, and this appears
to be a consistent character of O. reticulata s. l. as seen in the field. The type specimen (Toulouse,
1807, Fliigge) is held in LE(!) and, although comprising only the upper part of the inflorescence,
equates satisfactorily with living plants seen in mountain areas of southern France, but much less
so with those from Britain.

Shortly after Wallroth’s description of O. reticulata, Wimmer & Grabowski (1829) described
O. pallidiflora from a specimen collected in Poland. This plant was pubescent with flowers in a lax
spike. The calyx segments were described as ovate-acuminate and the corolla as curved, tubular-
ventricose, with glands and purple veins distally. The corolla lobes were crenulate margined and
the filaments pilose at the base and + glabrous above. There is very little information here to
separate it from Wallroth’s O. reticulata. Type material of O. pallidiflora has not been located but
specimens in several herbaria are referred to this species.

Another worker who described similar plants from Germany was Koch. His description of
O. procera (Koch 1833) stated that the calyx segments were uniformly divided or entire, the
corolla yellow-brown, campanulate, inflated towards the base and curved, and the lobes of the lips
denticulate and ciliate. The filaments, inserted at the base of the corolla, were sparsely hairy below
with a few glandular hairs above. In Koch’s opinion the plant lay taxonomically close to

OROBANCHE RETICULATA 259

O. rapum-genistae but the corolla and filament characters given clearly differentiate the two. The
plants upon which he provided his description apparently grew close to the Rhine and, from his
comments, it is clear that Koch knew the living plant from this same area.

At the same time as he published this description, he also described O. scabiosae (Koch 1833)
based on plants from a mountain area near Berchtesgaden, Germany, growing at an altitude of
5500 feet and apparently parasitising Scabiosa columbaria. His diagnosis described the calyx
segments as entire or divided into teeth of equal length; the corolla campanulate, slightly curved,
glandular-hairy with the margins of the lobes wrinkled, the filaments inserted low in the corolla
with scattered hairs in their lower part but almost glabrous above. He made interesting comments
on the corollas being basically bright ochre-yellow but tinged reddish-orange on the lip margins,
violet in the dorsal area and also violet-veined on the lips.

Most subsequent workers have treated O. scabiosae as a synonym of O. reticulata Wallr.. This
seems justified in view of Koch’s description, especially of the corolla colour and its montane
habitat. It is clear, however, that Koch considered it to be quite separate from his O. procera which
is described almost immediately before it in the same paper (Koch 1833). From this it might be
assumed that O. reticulata and O. procera are themselves morphologically quite separate. No type
specimens of either of the two species described by Koch have been located and none of either is
indicated as having been seen by Beck (1930).

Beck (1890) recognised three varieties within O. reticulata: the type (var. typica), and vars.
pallidiflora and procera. Later, he largely adhered to this treatment (Beck 1930), retaining the
three taxa at varietal level and providing a key to their identification where var. procera was
separated from the others chiefly by the shape of its broad, tubular-campanulate, completely
curved corollas, 15—18 mm in length. In the other varieties he considered the corollas to be tubular
or campanulate, longer than wide, with the dorsal line straight or only slightly curved, 20-23 mm
long or occasionally shorter. He separated var. pallidiflora from the type mainly by its paler
corolla, especially the degree of venation and presence of violet glandular hairs, and also in its
slightly more hairy filaments and choice of host (Cirsium and Carduus spp., rather than members
of the Dipsacaceae or other members of the Asteraceae).

In their French Flora, Rouy & Foucaud (1909) placed these taxa under O. platystigma Rchb.
based on a specimen of Reichenbach f. The first, var. scabiosae (based on Koch’s O. scabiosae)
had a tubular corolla 20-22 mm long, yellow at the base, then amethyst or purple distally and
covered with coloured glandular hairs. Another variety, var. procera, had a much broader,
subcampanulate, shorter corolla (16-18 mm long), weakly veined violet or purple but only on the
upper lip and generally with fewer, less strongly coloured glandular hairs.

Gilli (1965), working on central European taxa, recognised two subspecies of O. reticulata: the
type and subsp. pallidiflora. He described subsp. pallidiflora as having corollas which were
whitish or yellowish but pale lilac distally with few dark-coloured glandular hairs. In their upper
part, the filaments were sparsely glandular hairy to glabrous. He considered this taxon to occur
only in low-lying areas in Germany and Austria north of the Alps, where it parasitised species of
Cirsium and Carduus. Under subsp. reticulata he described the corolla as usually longer than
broad (20-23 mm long) and the dorsal line straight or only slightly curved. Within this he included
var. procera (Koch) Beck, characterised by a quite different corolla shape: broader, tubular-
campanulate and 15—18 mm long with the dorsal line completely curved. Gilli described the type
(subsp. reticulata) as differing from subsp. pallidiflora mainly in corolla colour, which was yellow
only at the base and otherwise intensely coloured violet to purple with dark veins and glandular
hairs.

In Flora Europaea, Chater & Webb (1972) described only one species, O. reticulata Wall.,
placing O. pallidiflora under synonymy; there was no mention of O. procera. However Pignatti
(1982) recognised two taxa in Italy: subsp. reticulata from the alpine area and subsp. pallidiflora
from lower altitudes. He separated the latter from the typical form by its paler corolla colour and
less glandular-pilose filaments. This was largely supported by Uhlich et al. (1995) when
considering European taxa as a whole but Hartvig (1991), who distinguished the two taxa in a
somewhat similar way, considered that for the montane areas of Greece most plants belong to var.
pallidiflora rather than var. reticulata. More recently, Kreutz (1995) recognised two taxa at species
level in Europe: O. reticulata from the alpine areas and O. pallidiflora from lower altitudes. He
placed the British plant under the latter whilst including var. procera within O. reticulata.

260 Me J. Y. FOLEY

From the varied treatments outlined above it is apparent that much confusion exists regarding
the taxonomy of O. reticulata s.l., but that it includes at least two taxa which differ mainly in their
corolla shapes. These are the montane/subalpine O. reticulata and the lower-altitude O. procera.
Plants with a paler corolla often appear to be referred to O. pallidiflora.

METHODS

A fundamental approach in this work has been to carry out detailed field examinations of the
morphology of living plants in their natural habitats. From personal field experience on the genus
and also from that of other workers (e.g. Beck 1890, 1930; Guimaraes 1904; Wilkomm & Lange
1870; Wilkomm 1893; Coutinho 1913; Schiman-Czeika 1964; Gilli 1965, 1982; Chater & Webb
1972; Feinbrun-Dothan 1978; Pignatti 1982; Meikle 1985; Valdés et al. 1987; Jones 1989;
Rumsey & Jury 1991), the characters described below and listed in Table 1 appear to be the most
useful in circumscribing closely related Orobanche taxa. Many taxa are best described by using
several characters in combination.

The methods used were as follows. A random selection was made of ten plants within a
population, all of which were in full flower. By “full flower” was meant plants in which the lowest
third of the inflorescence had fully open corollas, one of which was utilised for the morphological
evaluation of floral characters. For this purpose, only freshly-opened corollas were used. This
approach is similar to that previously employed for the assessment of taxa within other genera (e.g.
Roberts & Gilbert 1963; Roberts 1966, 1988; Bateman & Denholm 1983; Foley 1990) and has
also been recently applied to populations of British Orobanche taxa by Jones (1989). With
practice, selection bias can be avoided and a representative sample of plants assessed. The raw
data obtained from each of the populations studied are given in Foley (1998).

Measurements were achieved in two ways. Dividers used in conjunction with a ruler calibrated
to 0-5 mm were found convenient and very satisfactory for dimensions upwards of 20 mm. For
smaller values, a Leitz Wetzlar measuring lens possessing a built-in scale calibrated to 0-1 mm was
used. For the very largest dimensions such as plant height and inflorescence length it was often not
possible to measure reliably to less than 5 mm due to slight vagaries in stem contortions etc.. All
values were taken in a direct line and therefore lengths of curvature (for instance of corollas) were
ignored in such measurements.

Quantifiable numerical data obtained from the morphometric measurements of many of the
characters shown in Table | were also subjected to Principal Components Analysis (PCA) using
the Clustan 4 computer programme (Wishart 1987). By this technique, each specimen measured is
located in a multidimensional array, in which the number of dimensions is equal to the number of
characters measured, and where the most similar specimens are placed closest together. This
technique extracts the axes of greatest variation from the multidimensional space, thereby
simplifying the space to a few (usually 2 or 3) dimensions, so allowing the location of each
specimen to be visualised. Qualitative characters (34-45) and those of a binary nature could not be
used in PCA. Several of these are known to be important diagnostic characters for Orobanche taxa
and, of course, were fully considered alongside the PCA data and in the subsequent description
and delimitation of the individual taxa. Of especial significance are corolla colour and shape, calyx
shape and stigma lobe colour. Binary characters which clearly separate sections or groups within
the genus (e.g. presence or absence of stem branching (character 7) or presence or absence of
bracteoles (character 22), were not analysed further. The following characters (enumerated in
Table 1) were employed in the PCA: 1-6, 8-15, 18-20, 23-29b, 33. In the diagrams, the
component accounting for the greatest variance (PC1) is represented by the x-axis, that of next
significance (PC2) by the y-axis.

Photographic records were also made of plants within each population studied; many of these
are illustrated in Foley (1998). Whenever conservation considerations allowed, a typical specimen
was pressed for the herbarium (E, LANC) and inflorescences preserved in 70% alcohol (now in
LANC).

Most major British herbaria likely to contain material of the genus were visited during this work:
and a great number of loans were also obtained from European institutions. The internal flower
structures of preserved specimens were examined by detaching individual corollas and gently
boiling them for c. 10 minutes in water to which a small amount (<0-01%) of surfactant had been

——

OROBANCHE RETICULATA 261

TABLE 1. CHARACTERS AND SCORING SYSTEM UTILISED IN THE MORPHOMETRIC
ANALYSIS (ALL MEASUREMENTS IN MM)

1 PLANT HEIGHT: measured from ground level to the uppermost tip of the inflorescence.

2 STEM WIDTH: measured immediately below the lowest corolla in the inflorescence.

3 STEM PUBESCENCE: estimated on the scale shown below *.

4 INFLORESCENCE LENGTH: measured from the point of attachment of the lowest corolla to the
uppermost tip of the inflorescence.

5 INFLORESCENCE DENSITY: estimated on the scale:- 0 = lax; 1 = fairly lax; 2 = fairly dense;
3 = very dense.

6 COROLLA ATTITUDE: estimated on the scale:- 0 = almost erect to 3 = almost level, patent.

7 STEM HABIT: 0 = simple, unbranched; | = pedicellate at least below; 2 = branched.

8 © MAXIMUM LEAF WIDTH: as measured.

9 LEAF PUBESCENCE: estimated on the scale shown below *.

10 COROLLA LENGTH: measured in a direct line from base of the corolla to the extremity of the upper lip.

11 COROLLA VERTICAL HEIGHT: measured distally at a point just prior to reflexing of the lips.

12 DORSAL LINE CURVATURE: estimated on the scale:- 1 = almost straight; 2 = slightly curved;
3 = strongly curved; 4 = genuflexed.

13. DIVISION OF UPPER LIP: estimated on the scale:- 0 = entire; 1 = notched; 2 = slightly bilobed;
3 = strongly bilobed.

14. LOWER LIP CILIATE OR OTHERWISE: 0 = not; 1 = yes.

i LOWER LIP DENTICULATE OR OTHERWISE: 0 = not; 1 = yes.

16 & ATTITUDE OF UPPER AND LOWER LIPS OF COROLLA: estimated on the scale:- -1 = reflexed;

| 0 = upright; 1 = intermediately forwards; 2 = forward.

18 RELATIVE SIZE OF THE CENTRAL LOBE OF THE LOWER LIP OF COROLLA IN RELATION
TO THE LATERAL LOBES: ratio estimated.

19 SHAPE OF THE LOBES OF THE LOWER LIP OF COROLLA: estimated on the scale:- 0 = rounded,
distinctly obtuse; 1 = more or less acute.

20 COROLLA PUBESCENCE: estimated on the scale shown below *.

21 COROLLA CONSTRICTED DISTALLY: estimated as:- 0 = not; 1 = yes.

22 BRACTEOLE: 0 = absent; | = present.

23. BRACT LENGTH: measured from the point of attachment to the tip.

24 BRACT PUBESCENCE: estimated on the scale shown below *.

25 CALYX - LENGTH OF SEGMENTS; if divided, includes the longest tooth.

26 CALYX - SEGMENTS DIVIDED INTO SEPARATE TEETH OR OTHERWISE: Degree of division
estimated on the scale:- 1 = equally divided; 2 = unequally divided; 3 = very unequally divided;
4 = entire.

27 CALYX PUBESCENCE: estimated on the scale shown below *.

28 TYPICAL HEIGHT OF INSERTION OF THE FILAMENTS ABOVE THE COROLLA BASE:
measured from the point of attachment of the lowest

29a FILAMENT PILOSITY (UPPER PORTION): estimated on the scale shown below *.

29b FILAMENT PILOSITY (LOWER PORTION): estimated on the scale shown below *.

30 ANTHERS OR STYLES EXSERTED: estimated on the scale shown below *.

31 ANTHERS PILOSE: estimated on the scale shown below *.

32 STYLE PILOSITY: estimated on the scale shown below. *

33. STIGMA LOBE SEPARATION: estimated on the scale:- 0 = fused to 3 = well separated.

GENERAL SCALE: 0 = not, absent; 1 = slightly; 2 = appreciably; 3 = very, densely, excessively. (Values
intermediate between those given above may, in some cases, be arrived at).

NOTE: Additional characters (e.g. leaf length, bract width, maximum width of corolla tube, maximum
distance between the extremities of the lateral lobes of the lower lip of the corolla) were also measured during
the early stages of this work. However, some of these were later abandoned when the character states were
found to yield information of little value.

The following additional characters were recorded descriptively or photographically since they are not readily
quantifiable. They were recorded from fresh material in full flower.

34. STYLE COLOUR, 35. STIGMA LOBE COLOUR, 36. COROLLA SHAPE, 37. COROLLA COLOUR,
38. LEAF SHAPE, 39. LEAF COLOUR 40. BRACT SHAPE, 41. BRACT COLOUR, 42. CALYX SHAPE,
43. CALYX COLOUR, 44. FILAMENT BASE SHAPE, 45. STEM COLOUR.

262 M. J. Y. FOLEY

added. Formerly dry, brittle material could then be softened and more easily handled and
dissected. It was therefore possible to measure the height of filament insertion and the degree,
type, and distribution of pilosity along their length. Also, if the stigma and style were first softened
by a similar process and carefully pressed with clean filter paper, an indication of stigma lobe
colour could sometimes be obtained.

RESULTS AND DISCUSSION

A COMPARISON BETWEEN BRITISH POPULATIONS AND PLANTS FROM MONTANE/SUBALPINE HABITATS IN
EUROPE

A morphological evaluation was carried out on plants referred to O. reticulata from three British
populations within the plant’s very restricted British range in Yorkshire (see Table 2 for locality
details). These populations were compared one to another as well as to plants from montane
localities in eastern France. In the latter area, however, although several populations were located
and examined, only one population was found (Petit Mont Rond, Jura) which had plants in
sufficiently large numbers and in a suitable condition for a morphological evaluation to be made.
This was despite searches on several field visits over three seasons (1995-1997) and was mainly
due to their unpredictable flowering behaviour at higher altitudes.

Mean values and standard errors of the relevant characters are summarised in Table 2 and the
quantifiable numerical elements of these data have been subjected to Principal Components
Analysis (PCA). Examination of PCA data (Figure |) for the three British populations shows good
overlap on the PC1 axis but with some separation for the Aberford population on the PC2. This
appears to be mainly due to their less robust habit (reflected in reduced height, inflorescence
length, stem width, and leaf, bract and calyx dimensions (Table 2)). On the other hand there is very
clear separation on the PC] axis between the British and the French plants and an examination of

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FIGURE |. First two axes of Principal Components Analysis for individual plants from populations referred to
Orobanche reticulata from Yorkshire, England and France. PC] & PC2 jointly account for 43-6% of the total
variance.

OROBANCHE RETICULATA

263

TABLE 2. COMPARISON OF MORPHOLOGICAL CHARACTERS OF POPULATIONS OF
BRITISH AND ALPINE O. RETICULATA S.L.

Locality

Number
Date

Grid Reference/Coordinates

Plant height
Stem

Inflorescence

Leaf

Bract

Corolla

Upper lip
Lower lip

width
pubescence
colour
length
density
width
pubescence
shape
colour
length
shape
pubescence
colour

length
height
approximate
shape
attitude
dorsal line
pubescence
colour

divided

ciliate
denticulate
shape of lobes
relative size

(central to lateral)

Bracteole
Calyx

Filaments

Anthers
Stigma lobes

Apparent host

length
segment
division
pubescence
shape of teeth
colour

insertion height

pilosity above
pilosity below

pilose

separation
colour

BRITAIN

Aberford
Yorkshire

10
2.vii.1994
SE/43.35
407 + 16-5
8-0 + 0-4
2:5+0
fawn-violet
130-3 + 10-1
2:1+0-1
4:5+0-1
1-540

linear-lanceolate

cream
17:9+0-8

linear-lanceolate

2:1+0-1
cream-purple
20-4 + 0-4
6-8 + 0-3
+ campanulate

1:5+0
3:0 + 0
1:6+0-1
cream-purple

1:7+0-1

0

0

0
1-:1+0

0
11-4+0-4
2:5 + 0-3

2-0 +0
tapering
mauve-purple
3:-2+0-1
0-1+0
0-2 + 0-1
0
1:7+0-1
mauve-purple

Cirsium arvense

Mean values and standard errors (in mm)

BRITAIN

Oxclose
Yorkshire

10
9.vii. 1994
SE/36.46
529 + 33-2
9-2+0-5

2:0+0
cream-mauve
181 + 17-7
1-9+0-1
5-2+0-3
1-6+0

linear-lanceolate

mauve-cream
19-6 + 0:8

linear-lanceolate

2:0 + 0
cream
21-7+0:-5
8-1+03
+ campanulate

1-5 +0
cream-purple
1-8+0-1
0

0
0
10+0
0
12-6+0-5
2:9+0-3

1-6+0-1

narrow acuminate

cream-violet
3-2+0-1
0
0-4+0-1
0
1:7+0-1
brown-purple

Cirsium arvense

BRITAIN
Ripon
Yorkshire
5
13.vii.1994
SE/31.73
481 +51-0
10-4 + 0-7
2:0+0
deep violet
156 + 19-6
2-0+0-1
4-7 + 0-2
b5'+ 0

linear-lanceolate

violet-fawn
21-9+0-8

linear-lanceolate

2:0+0-1
mauve-violet

21-1+0-4

6:9 + (0-2

+ campanulate

1-5+0
2:6+0-1
1-520
cream-purple
1-6 + 0-2

0
0
0)
1-:0+0
0
15-1 +0-6
3-3 +0-3

1-8+0-1

broadly tapering

mauve-violet
3-4+0-2
0)
0)

0
1-9+0
brown-purple

Cirsium arvense

FRANCE

Petit Mont Rond
Jura
10
6.vill. 1995
46 20’N; 6 02’E
324 + 19-7
6:9 + 0-4
2:0+0
cream-mauve
155 + 14-5
2:5+0
4-0+0-1
15+0
linear-lanceolate
yellow-cream
16-6 + 0-9
linear-lanceolate
2:0+0
cream
18-5 + 0-4
6:7 + 0-3
tubular-tubular-
campanulate
10+0
2:2+0-4
1:-7+0-1

cream-yellow-purple

13+0-1

2:0+0

narrow, tapering

cream-yellow
2:-4+0-1
0-2 +0-1
0-3 +0-1
0
2:4+0-1
brown-purple

Carduus defloratus

264 MJY. gEOLEY:

Table 2 shows where the main differences lie. Other than in the different degree of robustness
mentioned above, the three British populations show close agreement in almost all important
characters (Table 2 and Figure 1). This is especially notable for those related to the corolla (shape,
size, height of filament insertion, filament pilosity and stigma lobe separation). Corolla shape,
being relatively broad in relation to length and strongly curved, is especially characteristic and
constant for the British plants and differs appreciably from the more tubular, much less curved
corollas of those from mountain areas of France. Being qualitative, however, this character is not
included in the PCA. The French plants also differ from the British in their lower level of filament
insertion, their wider stigma lobe separation and in the presence of ciliate, denticulate lobes to the
lower lip of the corolla. Corolla length in this population is also less than for British plants, but this
may not be significant as it was found to be somewhat longer in other French populations seen but
which were not morphologically assessed in detail. Broad similarities in corolla height of all four
populations did not reflect the characteristic corolla shape apparent in the British plants, i.e.
relatively broad in relation to length.

During the course of this work and earlier, 44 populations of the British plant have been
examined (Foley 1992a). All appeared to be morphologically similar but showed appreciable
variation in colour, especially that of the stem, calyx and corolla. A few populations had
individuals with very atypical coloration (Foley 1992b), some being uniformly rich yellow or
occasionally pure white. Apart from these exceptional plants, corolla colour differences were
usually due to the varying presence of mauve-purple glandular hairs and purple corolla venation.
This could vary widely and sometimes partially obscured the basic cream to greyish base colour of
the corollas. Plants seen in French montane/subalpine populations were similarly, but less variably,
veined and embellished with mauve-purple glandular hairs, usually over a deeper, almost yellow,
background.

The above observations suggest that the French montane/subalpine and British plants belong to
O. reticulata s. s. and O. procera respectively. Whether O. reticulata s. s. is a plant wholly
restricted to higher altitudes may be open to question since Wallroth’s type specimen was
collected from near Toulouse (France), some distance north of the Pyrenees, but no locality details
are indicated. Although not in an ideal condition, the specimen conforms satisfactorily to plants
seen by myself in the French mountains and, in corolla shape especially, it differs from the British
plant in ways similar to those mentioned above.

The British plants on the other hand largely agree with the description given by Koch (1833) for
O. procera, which occurs in lowland habitats in Germany and elsewhere. This applies especially to
corolla shape which is campanulate distally, inflated towards the base and curved; his description
is much closer to the British plant than to that from the Jura. A similar comment regarding the
distinctive corolla shape of O. procera is given by Rouy & Foucaud (1909) under O. platystigma
var. procera: “corolle plus large, subcampanulée, et plus petite’, i.e. broader and shorter; this in
comparison to other similar taxa within the group. Koch’s O. procera grew in fields close to the
River Rhine; the British plant is also most frequently found near river margins and flood plains
(Foley 1993) where it usually parasitises Cirsium arvense.

Despite similarities existing between the lowland British and the European montane/subalpine
plant, they can be readily distinguished by several clear-cut character differences. Due to this and
the fact that they appear to occupy quite distinct habitats, often parasitise a different host genus
and have a somewhat different geographical range, their appropriate taxonomic rank appears to be
at the subspecific level. The lowland plant of Britain and Europe therefore becomes O. reticulata
subsp. procera (Koch) Dostal and the montane/subalpine plant subsp. reticulata.

The status of O. pallidiflora Wiramer & Grabowski, however, remains uncertain. From its
original description it is unclear whether its affinities lie closer to subsp. reticulata or to subsp.
procera. By its specific epithet (= pale-coloured flowers) and the authors’ brief reference to corolla
colour, it would appear that this is its most distinctive character. If so, it may well be that light-
coloured variants of subsp. reticulata (and even subsp. procera) are routinely referred to it.
Although I have seen an appreciable number of herbarium specimens referred to O. pallidiflora
(placed at that or lower ranks), most of these are closer to subsp. reticulata than to subsp. procera.
In this genus, however, subtlety of colour is one of the first characters lost on preservation and it is
one reason why investigations into this taxon have not been pursued here. It is possible that taxa
referred to O. pallidiflora are mere colour forms of one or both subspecies of O. reticulata. No
living populations were discovered during this work which could be reliably referred to this plant.

OROBANCHE RETICULATA 265
CONCLUSIONS

There are two distinctive, yet closely allied taxa: (a) O. reticulata subsp. reticulata, a plant of
montane-subalpine occurrence, usually found at altitudes between 1000 and 2500 metres and
apparently widespread but local in suitable areas of continental Europe, where its main host is
Carduus defloratus, and (b) O. reticulata subsp. procera, a plant of lowland Europe (including
Britain), often found in agricultural areas and in the vicinity of major rivers or flood plains where
seed may dispersed by water-borne means. There are also other superficially similar lowland
European plants which have been referred to O. pallidiflora but which are somewhat problematic.
It is possible that these are simply colour variants and that such morphological differences as exist
are at least partly determined by the different hosts parasitised. J. Zazvorka (litt. comm.), based on
his experience in (the then) Czechoslovakia, goes even further and considers that all taxa embraced
by O. reticulata s. |. are really a single taxon whose differing morphology is host-determined.
There is no clear evidence for this, but the varying degree of robustness is one character which
may well be host-determined.

TAXONOMIC DESCRIPTION

OROBANCHE RETICULATA WALL. SUBSP. RETICULATA

(syn.: O. scabiosae Koch; O. pallidiflora auct., O. platystigma Rchb.; O. carlinoides Miégeville;
O. cardui Sauter; O. scabiosifixa St Lager).

Described from southern France “Jn agro Tolotono”. Type collected at Toulouse in 1807 by D.
Fliigge (LE!).

Typically 300-350 mm tall, with a fairly dense, many-flowered inflorescence. Stem: 6-8 mm
wide immediately below the inflorescence, distinctly glandular-pubescent, cream-mauve. Leaves:
maximum width c. 4 mm, linear-lanceolate, yellow-cream, glandular-pubescent. Bracts: 12-18
mm long, linear-lanceolate, cream, rapidly darkening with age. Calyx: c. 10-12 mm long,
segments usually unequally divided, sometimes entire, teeth narrow, tapering, cream-yellow,
glandular-pubescent. Corolla: 18-22 mm long, suberect, tubular to tubular-campanulate, yellow-
deep cream with mauve-purple veins and varying amounts of purple glandular hairs mainly
distally on upper lip, dorsal line not strongly curved. Upper lip slightly bilobed; lobes of lower lip
somewhat ciliate and denticulate, rounded, + similar in size. Filaments: inserted at 2-0-2-5 mm
above the corolla base, very slightly glandular-pilose to glabrous above, + glabrous below but
sometimes with a few hairs at the actual base. Stigma lobes: dark purple-brown with yellow
undertones, distinctly separated. Parasitic mainly upon Carduus defloratus, but probably also on
other taxa including Knautia spp.

A plant mainly of montane to subalpine habitats over a range of substrates, perhaps descending
to lower altitudes close to its main area of distribution. Flowers June-August (—September).
Widespread but local in suitable mountain habitats of mainland Europe including the Alps, the
Pyrenees and the Balkans. Its distribution outside this area is unclear at present.

OROBANCHE RETICULATA SUBSP. PROCERA (KOCH) DOSTAL
(syn.: O. pallidiflora auct.; O. cirsii Fries; O. mathesii Schultz: O. serotina Kirschleger).

Described from Germany “auf lehmigen Aeckern in Gegend von Mannheim in der Rheinpfalz...
[etc.]”. Specimens collected by Schimper but type not located.

Similar to subsp. reticulata but more robust and taller (400-550 mm); corolla: broader and
almost campanulate, quite strongly curved, lobes of lower lip not noticeably ciliate or denticulate;
filaments: inserted higher (3-0-3-5 mm) above the corolla base; stigma lobes: less widely
separated. Parasitic on Cirsium spp., especially Cirsium arvense and related genera.

A plant mainly of lowland habitats, river banks and flood plains, field borders, ruderal habitats,
roadsides, unimproved pastures, scrub, woodland clearings and perhaps occasionally reaching
submontane grasslands. Flowers June-August. Widespread throughout much of central Europe
north of the Alps, including Germany and Poland and with disjunct occurrences in eastern England
(Yorkshire).

266 VAY FOLEY

ACKNOWLEDGMENTS

I am most grateful to the trustees of the Welch Bequest Fund of the Botanical Society of the
British Isles and to the Botanical Research Fund (Kew) for financial assistance during this project.

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OROBANCHE RETICULATA 267

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FL Peo r,s de Rt Ie SE SORIA yo Soh MDE TCR em an REI a IRI TE OC RA LEE I LOE LOI I NE ORS AAI LOPLI IED AE AD DAYAR BO ELL TIS Ba {Se OPE EA mt Ie EUR As

*,

Watsonia 23: 269-278 (2000) 269

Viola rupestris and its hybrids in Britain
B. JONSELL*

Bergius Foundation, Royal Swedish Academy of Sciences, Box 50017, S-104 O05 Stockholm,
Sweden

I. NORDAL
Department of Biology, University of Oslo, P.O. Box 1045, N-0318 Oslo, Norway
and
F. J. ROBERTS

Eden Croft, 2 Wetheral Pasture, Carlisle CA4 SHU

ABSTRACT

Hybrids between Viola riviniana and V. rupestris were identified by means of isoenzymes from three of the
four localities for V. rupestris known from Britain. Differentiating characters between V. riviniana and
V. rupestris are primarily found in leaf shape and indumentum, flower colour, spur shape and capsule shape.
The ecology of V. rupestris is presented and compared with that of the hybrid and V. riviniana.

KEYWORDS: Viola riviniana x V. rupestris, isoenzymes, morphology, distribution, ecology.

INTRODUCTION

In Britain Viola rupestris F. W. Schmidt has a very restricted distribution with four locality groups
in northern England. Hybrids with V. riviniana (V. x burnatii Gremli) have been recognized or
suspected in those localities but the identification on morphological grounds has been regarded as
provisional owing to lack of knowledge about variation in the parental species. Valentine (1975)
still recognized the hybrid from only one of the V. rupestris localities, Widdybank Fell (Teesdale).
The aim of this study is to identify British hybrids by means of isoenzymes and to discuss their
morphology, ecology and distribution against that background.

V. rupestris has a wide palaearctic distribution, seemingly more-or-less continuous from eastern
Siberia to northern and central Europe. To the west it reaches the Pyrenees, south-eastern France,
western Germany and south-eastern Norway and has isolated locality groups on the Dutch dune
coast and in northern England. Montane populations in northernmost Scandinavia are
distinguished as subsp. relicta Jalas (Jalas 1950; Nordal & Jonsell 1998); otherwise no subspecies
have been described.

V. rupestris has the chromosome number 2n = 20, which is the lowest number known in the
section Viola, but evidence from isoenzyme patterns has revealed that the basic number is n = 5
(Marcussen & Nordal 1998). V. rupestris belongs to the subsect. Rostratae, which in Britain is
represented also by V. persicifolia Schreber (2n = 20), V. reichenbachiana Jord. (2n = 20),
V. canina L. (2n = 40), V. riviniana Reichenb. (2n = 37-46, according to Valentine 1949) and
V. lactea Smith (2n = 58). Crossing experiments between a number of these species were
performed by Moore & Harvey (1961), who made conclusions about the distribution and sharing
of genomes among them. V. rupestris was, however, not included in their study. For Britain
Valentine (1975) reported only V. riviniana x rupestris, while previous records of the hybrid with

*Address for correspondence: e-mail bengtj @bergianska.se

270 B. JONSELL, I. NORDAL AND F. J. ROBERTS

V. canina were discarded as being based on V. riviniana. Outside Britain V. rupestris hybrids have
been reported with V. canina and V. riviniana (and even with V. hirta L., which seems highly
doubtful). For combinations with the two first-mentioned evidence at least from Scandinavia
seems substantiated (Nordal & Jonsell 1998). The progeny of crossings undertaken between V.
canina and V. rupestris and between V. riviniana and V. rupestris have shown pollen fertility
between 0 and 5% and 0 and 14%, respectively (Bethke 1882; Erdner 1907; Becker 1910; Schofer
1954; Schmidt 1961; Harvey 1966).

In a number of species of Viola sect. Viola it has been shown that isoenzymes vary little if at all
within species and therefore can be used effectively as markers for the distinguishing of hybrids
(Marcussen & Nordal 1997; Nordal & Jonsell 1998).

MATERIALS AND METHODS

All known Viola rupestris localities in England (Fig. 1) were visited by the authors from 8-12
May 1998 (Table 1, sites 1-4, 8, 12-13). Permission was granted by English Nature and
landowners for very restrained collection of material. The intrapopulational homogeneity known
from Nordic representatives of the sect. Viola (subsect. Rostratae, cf. Nordal & Jonsell 1998)
justified restricted collection, an important consideration due to the relative rarity of the species.
Analyses were undertaken on an average of about four (in practice 1-7) individuals per site. In
most cases the sampling was non-destructive with removal of only a single leaf from the plant. A
few whole specimens were transferred to a greenhouse at the University of Oslo for comparative
cultivation. V. riviniana was sampled from the same or close sites (Table 1, sites 1-5, 8-13). In
addition a particular search was made for putative hybrids.

The removed leaves were placed in plastic bags and kept cold with freezing elements for one
week until they arrived at the laboratory. The transferred plants were immediately planted in pots
and grown under greenhouse conditions with an 18 hour day at 16°C and a 6 hour night at 12°C.

TABLE 1. LOCALITIES AND ACCESSION NUMBERS OF SAMPLED POPULATIONS

Site Grid reference V. rupestris hybrid V. riviniana
Ingleborough
1. Near Crummack Farm SDITT.T1 7650-2, 3 1652-15225
105 1=2, 3,453 7653-1
2. Long Scar SD/76.72 7654-1, 2, 3, 4,5 7655-1
3. Sulber Gate SD/77.72 7656-3, 5 7656-2, 4, 6
4. Near Crummack Farm SD/76.71 7659-1 7660-1, 2, 3
5. Near Crummack Farm SDT Tai 7664-1, 2, 3
6. Near Crummack Farm SD/77.71 37a 2.35
7. Long Scar SD/76.71 3773-1, 2
Teesdale
8. Widdybank Fell NY/81.29 7665-1, 2,4,7,8 7665-3 7666-2
7666-1, 3, 4 7667-15253
9. Widdybank Pasture NY/84.30 7668-1, 2, 3
10. High Force NY/88.238°". 767 141,2 7669-1, 2, 3
11. Cronkley Fell NY/84.28 7674-1
Long Fell (Brough)
12. Long Fell NY/76.19 7678-6, 7, 8 7676-1, 3 7676-2
7677-1, 2
7678-3, 4
Arnside Knott

13. Arnside Knott SD/45.77 1679-1, 2,354 7680-1, 2

VIOLA RUPESTRIS AND ITS HYBRIDS 2K

& Durham

é
A

Widdybank Fell

Ee
Long Fell

at

North
é ;

Yorkshire

Cumbria

8 Arnside Knott
@

Ingleborough

Morecambe

P 0 1 2 3 4 5 6 R4 8 9 0
FIGURE 1. 1-km distribution of Viola rupestris (solid circles) and V. rupestris + V. x burnatii (half-circles) in
Britain.

The field-sampled leaves and one from each of the transplanted specimens were crushed in a
grinding buffer (Morden et al. 1987). The crude extracts were absorbed into paper wicks for
subsequent horizontal starch gel electrophoresis (Wendel & Weeden 1990). Altogether, seven
enzyme systems were analysed: IDH (isocitrate dehydrogenase), MDH (malate dehydrogenase),
and PGM (phosphoglucomutase) were run on the so-called “D- system”; AAT (aspartate
aminotransferase), AMP (aminopeptidase), GPI (gluco-6-phosphate isomerase), TPI (triose-
phosphate isomerase) in the “A/B- system” (Selander & Yang 1969; Soltis et al. 1983, see also
Nordal & Jonsell 1998).

After the plants had been in cultivation under similar conditions from May to October 1998
leaves from the (surviving) plants were pressed for comparison.

ISOENZYME RESULTS

Three of the isoenzymes revealed patterns that gave different band phenotypes in Viola rupestris
and V. riviniana and an additive pattern in the hybrids: GPI, PGM and TPI (Fig. 2). The pattern
was totally consistent so that hybrid constitution in one isoenzyme system always inferred hybrid
constitution in the other two. The enzymes AAT, IDH and MDH did not differentiate between the
two species, and accordingly cannot be used in hybrid analyses. The results for AMP indicate
intraspecific variation in V. rupestris, which needs to be further elucidated.

The dimeric enzyme GPI showed activity in two regions (Fig. 2). In the anodal region two
bands, always blurred, were displayed in both species and the putative hybrids. In the cathodal
region a three-banded pattern was always found in V. rupestris, best interpreted as two allelic
bands with an intermediate heterodimer, in a pattern of fixed heterozygosity, which may be
expected in an allotetraploid. The octoploid V. riviniana always displayed a five-banded pattern, in

272 B. JONSELL, I. NORDAL AND F. J. ROBERTS

GPI PGM TPI

R; values

FIGURE 2. Isoenzyme results for gluco-6-phosphate isomerase (GPI), phosphoglucomutase (PGM) and triose-
phosphate isomerase (TPI) for Viola riviniana (RIV), V. x burnatii (x) and V. rupestris (RUP).

which the second and the fourth bands probably constitute heterodimers. With this interpretation
V. rupestris has the genotype BD and V. riviniana ACD in this complex “locus”. The hybrids
displayed an ABCD pattern with most of the constituent heterodimers, except the CD heterodimer,
present in the individuals.

In PGM (a monomeric enzyme) activity was also revealed in two regions (Fig. 2). In the anodal
area a fixed, identical two-banded pattern was shared by the two species. In the cathodal area
V. riviniana most often showed two bands (A and B), whereas V. rupestris only revealed the
B-band. In this case the hybrid pattern was identical with that of V. riviniana. In very few cases
V. riviniana displayed a band in a more cathodal (C) position, and one hybrid (7665-3) displayed a
triple-banded ABC pattern (not illustrated).

TPI (a dimeric enzyme) revealed a fairly complicated multibanded pattern, difficult to break
down into allelic interpretation. The bands may be referred to three regions. The most anodal
displayed one band shared by both species. Clear variation between the two species was found in
the next region, where V. riviniana had activity in front of V. rupestris (Fig. 2) and where the
hybrid revealed additivity. V. riviniana displayed two distinct bands in this region and V. rupestris
three. The latter is best interpreted as a fixed heterozygote (BD) with an intermediate heterodimer.
V.riviniana might have a third indistinct band and if so an AC-genotype, or the two distinct bands
may represent two monomorphic loci which do not interact. These alternative interpretations will
not influence the testing of the hybrid hypothesis. In the hybrids, with a putative AB(C)D
constitution, the bands tend to fuse, as visualised in Fig. 2. In the third region both species showed
a fixed three-banded pattern, - much weaker, however, in V. riviniana than in V. rupestris and the
hybrid.

MORPHOLOGICAL OBSERVATIONS

SEPARATION OF VIOLA RUPESTRIS FROM V. RIVINIANA

Although a distinct species, Viola rupestris is sometimes difficult to distinguish from V. riviniana,
especially when this species is represented by small-flowered forms (var. minor (Murbeck ex
Gregory) Valent.) as is the case in most British localities where it grows with V. rupestris. The
confusion is reinforced by the fact that V. rupestris is not consistently pubescent along petioles and
peduncles and on capsules as is generally stated in current Floras (e.g. Stace 1997).

VIOLA RUPESTRIS AND ITS HYBRIDS 273

The most reliable separating characters for British plants of V. rupestris from V. riviniana are as
follows:

i.

Leaf shape: the laminas of V. rupestris are ‘shovel-’ or “scoop’-shaped, with a clearly blunt
apex, and with a less strongly cordate base than in V. riviniana. In V. rupestris the sides of
mature leaves are often curled upwards (hence the ‘scoop’), whereas in V. riviniana it is the
basal lobes which are often upcurled. The areas of the lamina between the veins are somewhat
raised in most forms of V. riviniana, giving a + dimpled effect to the blade; in V. rupestris the
surface is flatter. The crenations at the leaf-edges are flat (i.e. within the plane of the leaf) in
V. rupestris, but in V. riviniana are slightly ‘crimped’, and thus more obvious. In Fig. 3 leaf-
silhouettes demonstrate, on limited material, clear differences particularly in the cordate shape
of the blade, the rounded blade apex in V. rupestris, and the intermediate nature of the hybrid.

Indumentum: the highly distinctive fine, very short indumentum of V. rupestris usually extends
up the peduncles, and up the petioles onto the margins of the basal lobes, leaving most of the
upper lamina essentially glabrous in the great majority of plants. (In some variants, the fine
indumentum extends all over the upper and lower surfaces of the lamina.) Where dense, the
indumentum produces the impression of a grey ‘fuzz’ to the naked eye.

It should be noted that some V. rupestris individuals are glabrous even along petioles and
peduncles. Such individuals were found, and confirmed by isoenzyme analysis, in all the
localities except for Arnside Knott. Furthermore, plants may often be found with indumentum
on some petioles but not on most.

The forms of Viola riviniana which accompany V. rupestris, while lacking this fine
indumentum of dense short hairs, almost invariably carry on the upper lamina sparse long
bristly hairs which are about 250-330 um long, and thus typically about six to eight times
longer than the indumentum ‘hairs’ of V. rupestris, which measure around 35-50 um. These
sparse, long, colourless hairs of V. riviniana can be inconspicuous, and are best observed by
looking with a lens across the upper surface of a leaf bent over the finger to create a convex
surface.

Flower colour: the corolla colour in V. rupestris is often paler and less blue than in
V. riviniana, being a ‘delicate pinkish-mauve’ in Teesdale plants (M. E. Bradshaw, pers.
comm. ), although the colour may vary in both species, and also alters with exposure, bleaching
and ageing processes. Looking into the flower from the front, in Viola riviniana there 1s
usually a narrow band of darker violet separating the colour on the lower lip from the white in
the throat; this band is apparently always lacking in Viola rupestris.

The flowers of the Arnside Knott colony, which have been sparsely produced in recent
seasons, are always pure white, as first noted by Valentine & Harvey (1961). The plant is
illustrated in Halliday (1997).

Spur: the spur of Viola rupestris is short, conical to a round tip without obvious groove or
projection. The spur in Viola riviniana is long, stout, and often vertically grooved at the end,
and often with a small projection from its upperside near the tip. (The colour of the spur varies
in both species.)

Ripe capsule: the ripe capsule of V. rupestris is almost globular with truncate apex in side
aspect, whereas that of V. riviniana is more elongated, and most often has an obtuse or even
acute apex.

V. rupestris produces rosettes on lateral branches, which are often hidden within the substrate.
Such branches are non-rooting, so that the rosettes do not become independent plants, although
often giving this appearance (Bradshaw & Doody 1978b). V. riviniana, however, has genuine
vegetative propagation in that adventitious shoots (soboles) may develop on the root system
(Valentine 1949).

It is worth noting that the forms of V. riviniana which occur in exposed sites close to V. rupestris
can approach that species. In comparison with V. riviniana forms in nearby, more sheltered, sites
such as block scree and fissures in limestone pavement they have a deeper green colour overall,
are more prone to developing purple tints, have a much more condensed form with short petioles
producing a tight rosette, have more leathery leaf texture, and smaller flowers.

274 B. JONSELL, I. NORDAL AND F. J. ROBERTS

eae

Viola rupestris

Viola riviniana x rupestris (V. x burnatit)

Viola riviniana

FIGURE 3. Leaf silhouettes of Viola rupestris, V. x burnatii and V. riviniana. Scale c. x 0-6.

HYBRID MORPHOLOGY

Detection of the generally rare hybrids in the often abundant and dense populations of the species
is not straightforward, due to variation of the parent species in all of the above characters, and the
low stature of the plants.

In plants examined to date, hybrids may be suspected by an intermediate leaf-shape (Fig. 3) and
by the presence both of the dense short indumentum of V. rupestris - albeit patchy - on petioles
and stems, and of the sparse longer hairs of V. riviniana on the upper leaf lamina. Some hybrid
plants can appear glabrous, as with a proportion of the V. rupestris parent; however, a few hairs,
sometimes of intermediate length and form, will generally be found on upper laminas, particularly
on the midrib or patchily towards the basal lobes. Where the indumentum is well-developed, it is
most abundant on the upper part of peduncles, but is less reliably found on petioles.

Chasmogamous (i.e. opening) flowers appear to be produced less frequently in most seasons
than by the parents. In colour and shape they tend to bridge the rather narrow differences between
the parents. Flower shape tends to approach V. rupestris rather than V. riviniana. The dark bar in
the throat usually characteristic of V. riviniana is in the hybrid faint or more-or-less absent. The
dark purple colour of the veins (honey-guides) varies in density, and where they run out onto the
mauve of the lower petal the pigmentation becomes more diffuse, contributing to the effect of a
bar. ;

Aborted flowers and lack of fruits late in the season (when cleistogamous flowers have been
produced) strongly indicate hybrid plants. The hybrid is thought never to ripen capsules, so that the
presence of ripe capsules in late summer, often abundant on mature plants, reliably indicates the
species. Hybrids can most easily be detected at this time when the plants attain their greatest
stature and leafiness.

The soboles of V. riviniana are apparently inherited by at least some hybrid plants, allowing
indefinite survival of individual plants. In the Long Fell (Brough) and Widdybank Fell (Teesdale)
localities, hybrid patches of considerable size (and perhaps age) are known, up to 1—2 metres in
diameter or more, and giving the appearance of clonal derivation from rare hybridisation events.

VIOLA RUPESTRIS AND ITS HYBRIDS 275

The production of adventitious shoots is particularly vigorous in the hybrid; on Widdybank Fell
densities of rosettes were recorded on trial plots of 656/m? for the hybrid, 149/m? for V. rupestris,
and 175/m? for V. riviniana (Bradshaw & Doody1978a).

ECOLOGICAL OBSERVATIONS

HABITATS OF VIOLA RUPESTRIS

In all its four English locations, Viola rupestris grows on exposures of Carboniferous limestone. V.
rupestris tends to occur in the more open and more exposed habitats available, on the thinnest
soils. It has two typical favoured habitats: one is on the eroding tops and flanks of low limestone
hummocks growing either in fine cracks in the bedrock, or in a ‘clitter’ of small stones lying over
the bedrock or as gently sloping screes; the other is in fine screes (more-or-less consolidated)
below low escarpments. It also occurs much more sparsely on the ‘clints’ (or flat tops) of
limestone pavements, but only where the rock surfaces are sufficiently fissured or the rock is
covered in ‘clitter’ - the loose stones resulting from degradation and fracture of the bedrock.

On the sites south-east of Ingleborough, some of the largest colonies are also on gently- to
steeply-sloping faces of drift deposits over limestone. Bare clayey soil, evidently calcareous, is a
feature of this habitat, the open nature of the sites apparently being maintained over long periods
by soil-creep and by stock trampling. The colonies lie between 320-425 m a.s.1.

On Widdybank Fell the plant occupies exposures of granular partially-metamorphosed limestone
(‘sugar limestone’), the rapid erosion of which creates zones of loose calcareous rubble and coarse
calcite ‘sand’. Viola rupestris appears to be an early colonist of such bare soils, with little else
besides a thin turf or occasional tufts of Festuca ovina L. and plants of Minuartia verna (L.) Hiern.
The species persists in closed Festuca turf, although flowering and fruiting less vigorously, and
only where some disturbance (such as by moles) maintains areas of open soil. The development of
denser closed swards appears eventually to oust the violet. Plants occur from 490-510 m a.s.1.

The Long Fell colonies lie at 575-610 m a.s.l. on a shelving S.S.W.-S.E.- facing escarpment, the
plants growing on slopes above cliffs, with a few patches on narrower ledges below the uppermost
low cliffs of the scarp. Eighteen colonies were located in 1998. The plants are consistently dwarf
in comparison with those at the other three sites; it is unclear whether this dwarfing is a response
to the greater exposure and altitude (Heaton 1999). The dwarf form appears to be retained in
cultivation.

Its most typical closest associates in the upland sites, whether on un-metamorphosed limestones
or calcareous boulder clay, are Carex flacca Schreber, C. panicea L., Carlina vulgaris L.,
Euphrasia confusa Pugsley, Sesleria caerulea (L.) Ard., and Thymus polytrichus Borbas. It can be
found in open turf of Festuca ovina and Sesleria caerulea, but rarely in dense turf of either
species.

At Arnside Knott there exists the only colony of Viola rupestris known off the ‘high’ limestone.
This site is on an exposed escarpment, in a very limited area free of scrub at 150 m a.s.]. (Lists of
associates in the Arnside, Long Fell, and Widdybank localities are given in Valentine & Harvey
1961).

HABITATS OF VIOLA RIVINIANA

Viola riviniana almost invariably occurs in the near vicinity at all four sites, but is seldom
intermingled. It obviously survives in closed turf of Festuca ovina, where V. rupestris is rare or
absent, but conversely it avoids open ‘clitter’ areas of small stones or fine scree, colonised by
V. rupestris. In a very dwarf and condensed form it is widely present also in the closed Agrostis/
Festuca grasslands between the limestone outcroppings. As a much larger and laxer plant with
large flowers it occurs, often abundantly, in the hollows between larger stones and in the
‘grykes’ (vertical fissures) of limestone pavement.

HABITATS OF HYBRIDS

The hybrid prefers deeper soils, and can tolerate denser turf, than V. rupestris, but has not yet been
seen far from colonies of the rarer parent. On Widdybank Fell the habitat consists of semi-open or
closed swards generally intermediate between those preferred by the parents. It also occurs with

276 B. JONSELL, I. NORDAL AND F. J. ROBERTS

Calluna on calcareous brown-earth soils, where moles are active (Bradshaw & Doody 1978b). On
Long Fell one patch of hybrid plants was on a gently-sloping south-facing patch of loose stones
immersed in black humus-rich soil, over bedrock. The few plants on Ingleborough have been
found in open patches on drift covered in loose stones, between areas of Festuca ovina turf, within
a few centimetres of the parent species.

POPULATION SIZES OF VIOLA RUPESTRIS AND THE HYBRID
Viola rupestris occurs most typically in local, dense colonies of a few to very many rosettes within
limited tracts of its preferred habitat, whereas V. riviniana occurs more dispersed, but over much
more extensive turf habitats between outcroppings of limestone. Work on establishing the
population sizes and spatial limits of V. rupestris is continuing. As previously remarked, due to the
difficulty of establishing the origin of rosettes, it has been usual to regard each rosette as an
individual for recording purposes. Surveys in 1998 on parts of the known distribution on the south-
east spurs of Ingleborough suggest that the population of the whole will amount to tens or
hundreds of thousands of individuals. The Widdybank sites are estimated to have thousands of
individuals (M. E. Bradshaw, pers. comm.), whilst the population on Long Fell was estimated in
1998 at between 1000 and 2000 plants. Arnside Knott had 56 plants in 1998 (Heaton 1999),
Hybrid numbers in ‘the high hundreds’ are known on Widdybank Fell, with ten dense colonies
and scattered groups and singles (M. E. Bradshaw, pers. comm.). It appears to be more localised
on Long Fell, although this area has been worked much less thoroughly than has Widdybank Fell;
the patches known in 1998 had a few dozen rosettes. On the Moughton/Crummack/Norber sites
south-east of Ingleborough, however, only a few individual hybrid plants have yet been located in
over 50 colonies of V. rupestris examined.

DISCUSSION

In the presented isoenzyme patterns of GPI and TPI, Viola rupestris in England displays exactly
the same band patterns as were found in Nordic populations of the same species (Nordal & Jonsell
1998). The A-band in PGM in the English plants is shared with plants belonging to the North
Scandinavian V. rupestris subsp. relicta and is different from the widespread ssp. rupestris. This
indicates an interesting connection which will be analysed in a wider geographical context (Jonsell
& Nordal in prep.). The V. riviniana plants analysed from the English localities have different
band patterns compared to Nordic material in all the isoenzymes presented. Our English
V. riviniana material consistently showing this isoenzyme pattern includes both small-flowered
forms which often grow together with V. rupestris and larger-flowered ones from sites in the
vicinity.

Based on enzymatic evidence hybridisation was revealed in ali but one of the four British areas
for V. rupestris. The exception is Arnside Knott though V. riviniana is frequent both on and close
to the spot. In two of them (Widdybank Fell and Long Fell) the hybrid was previously known,
while the record from Crummockdale (Ingleborough) is new (for details see Table 1). Of five sites
for V. rupestris sampled in the Ingleborough area, where V. riviniana is common throughout, the
hybrid was revealed only in one. This suggests that hybridization is comparatively rare there,
although one has to allow for the difficulties of distinguishing hybrids in the field. The hybrids
analysed give the impression of being F1 individuals and we have seen no traces of back-crossing
to parents. This is in contrast to the situation in parts of Scandinavia where, according to
preliminary observations (Nordal & Jonsell 1998), there are trends towards genetic swamping of
V. rupestris, where this species is rare. There, however, V. canina seems usually to be the other
parental species, though hybridisation with V. riviniana has also been documented (Nordal &
Jonsell 1998).

In many seasons V. rupestris is sparsely-flowered - at least as regards chasmogamous flowers,
which appear mainly during May. Thus, given the very exposed habitats in which it typically
grows, with little insect activity in the early season, the chances of cross-pollination of its
chasmogamous flowers with those of V. riviniana are clearly low. These factors may account for
the observed rarity of hybrids in most sites. Although both species appear to regenerate efficiently
from seed the vast majority of seeds are likely to result from selfing within cleistogamous flowers.

VIOLA RUPESTRIS AND ITS HYBRIDS 277

Whilst the fell-field habitats are undoubtedly over-grazed by sheep at present, the resultant erosion
helps to retain open unstable areas suitable for colonisation of V. rupestris, but less favourable for
propagation of hybrid clones, which need more stable substrates. The sterility of any hybrids, and
consequent apparent lack of back-crossing, coupled with the abundant seed-production of the pure
species, dictate against any swamping of Viola rupestris populations in the foreseeable future.

The close spatial juxtaposition of the habitats of the two species means that they may be found
in close proximity. The apparent tolerance of V. rupestris for eroding habitats with their mineral
soils, where it evades competition not only from V. riviniana but also the turf-forming grasses,
appears to have allowed its local survival. An observation by F.J.R. requiring further investigation
is that in other areas of the north of England where Carboniferous limestone is exposed in
apparently similar situations and climate, V. riviniana appears to occupy the types of microhabitat
preferred by V. rupestris in its few stations.

ACKNOWLEDGMENTS

We would like to thank the following for help in various ways: Dr M. E. Bradshaw, Erica
Donnison (English Nature, Cumbria Team), Peter Corkhill (Ingleborough National Nature
Reserve), Kristian Heaton (Edge Hill University College), Stephen Hewitt (Tullie House Museum,
Carlisle), John Hooson (National Trust), Chris McCarty (Teesdale National Nature Reserve),
Major Ken Miles (Range Officer, M.O.D. Warcop Training Range), and Jacqui Ogden (English
Nature, Cumbria Team). The distribution map in Fig. 1 was prepared using DMAP software
produced by Dr Alan Morton.

REFERENCES

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BRADSHAW, M. E. & Doody, J. P. (1978a). Chapter 2: Population-dynamics and Biology, in CLAPHAM, A. R.,
ed. Upper Teesdale, the area and its Natural History. Collins, London.

BRADSHAW, M. E. & Doopy, J. P. (1978b). Plant population studies and their relevance to nature
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HEATON, K. (1999). Morphological variation in the British populations of the Teesdale violet (Viola rupestris
F.W.Schmidt), pl. M.Sc. dissertation, Edge Hill University College

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botanici societatis zoologicae botanicae Fennicae ‘Vanamo’ 24: 1-362.

MARCUSSEN, T. & NORDAL, I. (1997). Viola suavis M. Bieb., a new species in the Nordic flora, with the
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European arosulate violets. New phytologist 60: 85-95.

MORDEN, C. W., DOEBLEY, J. & SCHERTZ, K. F. (1987). A manual of techniques for starch gel
electrophoresis of Sorghum isozymes. The Texas Agricultural Experimental Station. The Texas A and M
Univ. System, College Station, Texas.

NORDAL, I. & JONSELL, B. (1998). A phylogeographic analysis of Viola rupestris: three post-glacial
immigration routes into the Nordic area? Botanical journal of the Linnean Society 128: 105-122.

ROBERTS, F. J. (1977). Viola rupestris Schmidt and Juncus alpinus Vill. in Mid-W. Yorkshire. Watsonia 11:
385-386.

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Osterr. Bot. Zeitschr. 108: 20-88.

SCHOFER, G. (1954). Untersuchungen tiber die Polymorphie einheimischer Veilchen. Planta 43: 537-565.

SELANDER, R. K. & YANG, S. Y. (1969). Protein polymorphism and genetic heterozygosity in wild
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278 B. JONSELL, I. NORDAL AND F. J. ROBERTS

SOLTIS, D. E., HAUFLER, C. H., DARROW, D. C. & GASTONY, G. J. (1983). Starch gel electrophoresis of ferns:
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STACE, C. A. (1997). New Flora of the British Isles, Ed. 2. Cambridge Univ. Press.

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ed. British flowering plants and modern systematic methods (conference report). B.S.B.I. Pp. 48-53.
VALENTINE, D. H. (1975). Viola L., in STACE, C. A., ed. Hybridization and the flora of the British Isles, pp.

154-163. Academic Press, London.

VALENTINE, D. H. & HARVEY, M. J. (1961). Viola rupestris Schmidt in Britain. Proceedings of the Botanical
Society of the British Isles 4: 429-435.

WENDEL, J. F. & WEEDEN, N. F. (1990). Visualization and interpretation of plant isozymes, in SOLTIS, D. E.
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(Accepted March 2000)

Watsonia 23: 279-286 (2000) 79

Carex muricata L. subsp. muricata (Cyperaceae) - a review of its
present status in Britain

Me %oFOLEY*

Department of Biological Sciences, Institute of Environmental and Natural Sciences, University of
Lancaster, Lancaster LAI 4YQ

and
M. S. PORTER*

5, West Avenue, Wigton Cumbria CA7 9LG

ABSTRACT

The present status of the sedge, Carex muricata subsp. muricata, critically endangered in Britain, is reviewed
and, where relevant, the causes of its decline identified. A brief history of the plant at its few British colonies
is also given.

KEYWORDS: Prickly sedge, distribution, rarity, threats, conservation.

INTRODUCTION

It was only following the work of E. Nelmes (Nelmes 1947) that Carex muricata subsp. muricata
was recognised as a British plant, having been included within C. muricata s.|. by earlier authors.
Nelmes’ findings were based on herbarium studies in which he identified the plant in earlier
collections made at four widely scattered British sites in Gloucestershire, Denbighshire, Yorkshire
and Berwickshire. All other British material which he examined within C. muricata s.]. was
assigned by him to subsp. lamprocarpa (as C. pairaei). This plant, whilst locally frequent
throughout much of Britain, decreases rapidly moving eastwards in Europe where it is largely
replaced by subsp. muricata.

A taxonomic description of the two subspecies in Britain has been given by David & Kelcey
(1985). Important character differences between them include the more robust habit, rigid, erect
flowering stems, the much earlier flowering of subsp. muricata and, when immature, dark glumes
which are much shorter than the matt green utricles giving the inflorescence a chequered
appearance (Fig. 1). This contrast disappears as the fruit ripens, both the glumes and utricles
becoming dark brown or even blackish. On the other hand, in subsp. lamprocarpa the glumes are
initially pale yellow-green and almost the same length as the similarly coloured utricles, so that the
chequered contrasting pattern of subsp. muricata is initially lacking and not apparent until
maturity, when the glumes fade to white and the utricles darken. There are also ecological
differences. In Britain, subsp. muricata is a plant of calcareous habitats, occurring especially over
warm, sloping limestone scree, whereas subsp. /amprocarpa prefers an acidic substrate and is
more a plant of the lowlands (David & Kelcey 1985).

The present authors, familiar with subsp. muricata at its few known localities in northern
England, and having recently become concerned about the apparent decline of the populations and
the degeneration of the habitat at some localities, decided to investigate further the current status
and ecology of the plant. This work was later expanded and incorporated into a country-wide
project subsequently sponsored by English Nature. The original investigations were augmented by
an assessment of the local distribution and management. Vegetative material was also collected for

“Addresses for correspondence: e-mail m.foley @lancaster.ac.uk; m.porter@ wigton.cablenet.co.uk

280 M. J. Y. FOLEY AND M. S. PORTER

subsequent genetic studies involving DNA sequencing from individual plants at the extant
populations, and seed for storage in the Millennium Seed Bank at Kew. Further details relating to
this survey and to the location of all known populations are given in a confidential report to
English Nature (Foley & Porter 1999).

KNOWN BRITISH LOCALITIES

Nelmes’ critical work at K led to the initial recognition of subsp. muricata in Britain. In 1942 he
first identified as such, a specimen in Druce’s herbarium which had been collected in 1900 on
oolitic limestone at Woodchester, v.c. 34. Subsequently (Nelmes 1947), he identified three other
collections as the same taxon from the top of a limestone hill near Wrexham, Denbs., v.c. 50
(1840, Bowman), from the castle grounds at Lauder, v.c. 81 (1878, Brotherston), and from
limestone slopes and scree at Gordale, v.c. 64 (1934, Milne-Redhead & Sandwith; 1937, Lousley).

However, his work appeared to generate little interest, until R. W. David and J. G. Kelcey made
attempts to refind the plant at the original localities. Initially this was unsuccessful due to the
difficulty in locating such a rare, and somewhat insignificant plant in large areas of apparently
suitable habitat. However, in 1974 B. M. Mack found an unidentified Carex at Nympsfield,
Gloucestershire which she sent to David (David & Kelcey 1975) and which was subsequently
identified as subsp. muricata. Nympsfield is only a short distance from Woodchester, Druce’s
original locality, and may be the same.

In 1978, at Minera, near Wrexham, the plant was discovered by A. Newton. Despite claims that
this was a rediscovery at the original 1840 locality (Anon 1981), there can be no certainty that this
was So, as there is much suitable habitat elsewhere in this area.

Just prior to this in 1977, F. J. Roberts had found a small population growing on wooded
limestone pavement in upper Ribblesdale, v.c. 64. In the following year plants were seen at the
same locality and their identity confirmed by R. W. David but there was some doubt as to whether
this was Roberts’ original population (David 1979). At the previously known Yorkshire locality at
Gordale, searches were initially unsuccessful, but subsp. muricata was eventually refound there in
1984 by P. Jepson. His locality is probably slightly different to that of the earlier collections of
both Milne-Redhead & Sandwith and of Lousley. Up to the present, the original 1930s Gordale
localities have still not been refound and may now be extinct as this is a very popular area with
walkers and climbers who may have inadvertently eliminated the plant. Nevertheless, Gordale has
a large area of suitable, often inaccessible habitat where the plant may perhaps still remain
undetected.

When D. Millward was surveying for the Flora of Wensleydale in 1987, she discovered a
population of an unusual Carex in lower Swaledale. The plants were subsequently confirmed as
subsp. muricata by R. W. David and occurred in a fairly open habitat at the base of limestone
scree, the locality being some appreciable distance from other known sites. In 1999, S. J. Whild
and A. Lockton discovered another locality near Nantmawr, Shropshire, v.c. 40, in a rather similar
habitat to that at Swaledale and almost 30 km to the south of the Minera population.

Of Nelmes’ original four localities, it is only at Lauder where the plant has not been refound at
least within the general vicinity. However, here the habitat is not calcareous and appears to be
quite unsuitable. It is possible that Brotherston’s specimen may have been mis-labelled or, perhaps
more likely, may represent an accidental or even deliberate introduction. This will be discussed
later.

HISTORY AND PRESENT STATUS IN BRITAIN

In addition to Nelmes’ original localities, at least three others have been found relatively recently
in separate geographical areas, whilst the populations at Nympsfield and Minera may also be new
finds. The recent history of all of these and a description of the present habitat and status of the
populations are discussed below, largely in the sequence in which the populations were refound.
The information is also summarised in Table 1. It should be borne in mind that the plants often
occur in “clumps” so that it is difficult to decide whether one or several plants are present. Such
“clumps” are considered here to be a single plant.

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CAREX MURICATA SUBSP. MURICATA

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282 M. 3.-Y¥; FOLEY AND M. S-PORTER.

Full grid references for all known sites have been lodged with English Nature.

1. NYMPSFIELD (V.C. 34)

It is not clear whether the population discovered here in June 1973 by B. M. Mack is the same as
that known to Druce in 1900 (described as Woodchester) but is quite possibly so. It was recorded
at Dingle Wood, Nympsfield, on a warm slope (hanger) over oolitic limestone, when 21 plants
appeared after woodland clearance and re-planting. However, in 1976, it was described as hard to
find but was still there in 1979. The population declined further as the site became overgrown so
that there were only four plants in 1982. Fruitless searches were made thereafter and the site was
cleared in 1986 in the hope that plants would regenerate. None emerged during the following
spring and so subsp. muricata was considered extinct. Later that year (1987), three plants were
introduced to the site (grown at Cambridge Botanic Gardens from seed originating from Dingle
Wood) and whilst these persisted for a short while they rapidly declined in vigour and eventually
died. A single, caged plant survived until 1989 but the unattended cage became choked with
vegetation and the Carex succumbed. Material thought to originate from this site was grown on in
S. C. Holland’s garden and two plants were transferred to that of M. Kitchen after her death.
Further reintroductions within the area have been considered but have not yet been carried out.
The authors feel that the provenance of such material should be checked to ensure that it is of
Nympsfield origin.

2. MINERA, NEAR WREXHAM (V.C. 50)

When discovered in 1978, the habitat comprised light scrub with c.20 plants of subsp. muricata.
Shortly afterwards more were found close by and observations made soon afterwards referred to
good numbers occurring on loose limestone scree with associates which included Teucrium
scorodonia, a plant often indicative of stony habitats. In 1987 over 40 plants were reported but this
was reduced to 27 in 1992. Since then, the site has declined rapidly, the original Corylus scrub
developing into closed-canopy woodland. In consequence, litter and humus have accumulated and
the original loose scree is now completely obscured and a ground flora is present in which
Geranium robertianum and Sanicula europaea dominate. However, Hedera helix, which at one
time was a problem, is no longer present. The Carex population is much reduced in size and
vigour, the few plants surviving being in heavy shade and mainly vegetative. They also have to
compete with the gradually increasing ground flora. Flowering spikes are few and utricles rarely
seem to reach maturity. Typical associates of the Carex, such as Teucrium scorodonia, which were
present in the original, more open habitat, have also been eliminated.

When surveyed (1999) only three flowering plants were found, these possessing a total of eleven
inflorescences between them, most of which were weak and depauperate. A few probable (but
unconfirmed) seedlings were also present but seem unlikely to reach maturity unless conditions are
improved by a drastic reduction of the tree cover. At the time of the survey it was also noted that
cattle had broken through the protective fence and trampling had occurred. It is remarkable how
rapidly this site has deteriorated and this illustrates the need for regular monitoring and
management.

3. UPPER RIBBLESDALE (V.C. 64)

This locality on partly shaded, vegetated limestone pavement appears to be unique in Britain, all
other localities being, or having been, on sloping limestone scree. David (1979) questioned
whether the population from which he confirmed the plant in 1978 was the same as that first
discovered by Roberts in the previous year. However, his subsequent searches revealed only two
additional isolated plants and it is ‘more-or-less certain that the two records were for the same
population.

Initially there were only four plants, two in each of two populations in close proximity. An
additional plant was found by David some few hundred metres to the south but has not been seen
since. Meanwhile, the original colony was decreasing. The northernmost population lost one plant
in 1979 and the second by 1984 although an additional seedling was found situated between the
two populations; this could not be detected in 1987. The other two plants (probably those first
found in 1977) still survive. One of these is partially caged but still suffers from grazing by rabbits
which often remove the flowering spikes. An additional seedling was found very close to these in
1999. The plants occur in an open area within this relatively heavily wooded site, closely

CAREX MURICATA SUBSP. MURICATA 283

associated with such species as: Anthoxanthum odoratum, Dactylis glomerata, Geranium
robertianum, G. lucidum, Veronica chamaedrys, Filipendula ulmaria, Saxifraga hypnoides, Silene
dioica, Mercuralis perennis, Asplenium scolopendrium and A. trichomanes. Such associates
correspond closely to those seen by David in 1978, suggesting that conditions here have changed
little in the intervening period, in contrast to what appears to have happened at some other sites,
and especially at Minera. The upper Ribblesdale locality is situated within the Ingleborough
N.N.R.

4. GORDALE (V.C. 64)

The history of the plant at this site is a little confusing as the complex topography makes it
difficult to establish the precise whereabouts of early records. In 1985, the year after it had been
refound by P. Jepson, only four plants were recorded in an area of 4 x 4 m, two of which were
flowering poorly. These were on a steep, rocky apron at the base of a limestone cliff but were not
thought to be of the same population as those recorded in the 1930s which were apparently above
the waterfall. Subsequently, rumours circulated that the plants had gone from Gordale, but they
were refound by Jepson in 1997. In 1999 the present authors confirmed a thriving population on
the “apron”. Here there were at least 25 plants bearing a total of 128 flowering spikes, whilst a
short distance to the south, on the sides of a steep buttress, a further eleven plants (57 flowering
spikes) were found. The 45-60° sloping apron, whilst of an open, sunny aspect, was heavily
grassed, mainly with Sesleria caerulea, Dactylis glomerata, Brachypodium sylvaticum and
Festuca ovina, although the sedge still appeared to be flourishing there. Other close associates
included Glechoma hederacea, Torilis japonica, Hypericum hirsutum and Centaurea nigra with a
little Thymus, Teucrium scorodonia, Geranium robertianum and Campanula rotundifolia. On the
nearby steep buttress there was less competing vegetation and the sedge grew in close proximity to
plants typical of a more open habitat such as the last four mentioned above. On the “apron”, the
grasses are apparently becoming more dominant since there is little evidence of grazing with only
the occasional sheep roaming the area. There is, however, a great deal of other apparently suitable
habitat at Gordale where subsp. muricata may occur, but most of it is difficult and even dangerous
to explore. Past and present localities benefit from S.S.S.I. protection.

5. SWALEDALE (V.C. 65)

Carex muricata subsp. muricata was first found here in 1987 near the foot of a S.W. facing, 30-
35° vegetated scree slope below steep limestone crags. Initially there were about 15 plants, seven
of which had fruiting spikes, all occurring in an area of only a few square metres. The general
habitat was, and still is, open and sunny with scattered Sambucus shrubs, but latterly invasive
grasses strongly compete with the Carex with the result that, in 1999, the authors were able to find
only seven plants. A little distance above these plants, the vegetation of the slope quickly changes
and it becomes clothed in a dwarf, grazed, herb-rich community with Thymus, Pilosella
officinarum, Leontodon hispidus and Linum catharticum the dominant species. Here a few small
flowering plants of subsp. muricata are regenerating, apparently as a result of a reintroduction,
despite the fact that at the time (1994) the existing population was not under threat. Unfortunately,
it also appears that local plants were not used for this purpose and as a result the genetic make-up
of this isolated population will ultimately be affected. The dwarf, herb-rich community described
above would appear to be a more ideal habitat for subsp. muricata and was probably similar to that
present when the original, currently overgrown, population was first discovered. An associated
plant list given for 1990 appears to confirm this, as it includes such species of well grazed habitats
as Thymus and Galium sterneri, both of which are now absent from the immediate vicinity of the
native population. The increased competition here, probably resulting from a sharp reduction in
grazing in an area now enclosed for the sedge’s protection, gives some cause for concern. This
locality, which is situated on M.O.D. property, has S.S.S.I. protection.

6. NANTMAWR (V.C. 40)

This population was identified during the Spring of 1999 during a routine survey of an existing
local nature reserve. It had been noted as early as 1982 but had been wrongly referred to subsp.
lamprocarpa. The plant is locally very abundant here on open, sunny, lightly vegetated, limestone
scree with scattered Taxus and covers an area of approximately 20 x 30 m. The angle of the slope
is c.30-40°. Plants occur both singly and in relatively large clumps in minimal competition. They

284 M. J. Y. FOLEY AND M. S. PORTER

appear to be thriving and free flowering with abundant fruit set. It is very difficult to assess the
number of plants present since many grow close together, but there may be up to 100 which
produce several hundred inflorescences so making this the largest known British population.
Typical close associates at this lightly vegetated habitat include Teucrium scorodonia, Campanula
rotundifolia, Helianthemum, Thymus, Plantago lanceolata, Galium verum and Festuca ovina.
Very close by, in a similar open habitat, is a small population of the locally rare Galeopsis
angustifolia, a plant of loose stony substrates.

7. LAUDER (V.C. 81)

An examination of the parkland habitat at, and in the vicinity of, Lauder Castle confirmed that
there is no suitable, calcareous habitat present and that the 1878 record by Brotherston is either a
result of an error in specimen labelling or, if genuine, represents a plant or plants deliberately
introduced into the castle grounds. The latter would seem to be quite likely as other species,
including sedges, which are outside their normal area of distribution in Britain, have been recorded
at the castle (pers. comm. M. Braithwaite). The planting of aliens and exotics in estate woodlands
was fashionable in the past and it is likely that from such a source has come the Lauder record. It
should be considered only as a deliberate introduction here unless unequivocal confirmation can be
established otherwise.

CONCLUSIONS

Examination of the plant at its few known British sites suggests that it thrives best on sunny, dry,
limestone scree slopes in the presence of only minimum competition and shading. At the centre of
its distribution (Scandinavia/continental Europe) it is able to tolerate a wider variety of soils as
well as some shading (B. Jonsell, pers. comm.; B. G. Johannson, pers. comm.) but here in Britain,
on the edge of its range it appears to be more exacting in its demands. Where ideal conditions
occur such as at Nantmawr, robust clumps of plants exist which produce a large number of
inflorescences, whose fruits ripen readily. Smaller, apparently younger, fruiting plants also occur
on the open scree and on small ledges. Somewhat similar conditions occur at both Gordale and
Swaledale, although there the habitat appears to be changing through invasion by coarser grasses.
These populations should be examined annually in order to monitor for adverse change. In
contrast, the Minera site has deteriorated rapidly in less than ten years and is now severely
overgrown with Corylus woodland. In the area where the plants grow, sunlight is severely reduced
and humus has accumulated over the scree so that a dense ground flora now competes with the
sedge. In consequence, the few plants have become weak and appear unwilling to set much fruit.
This site is in need of immediate attention.

The other extant British locality is on limestone pavement at Upper Ribblesdale. This population
has never been known to contain more than a few plants and is now reduced to just two flowering
clumps together with a few seedlings. Rabbit grazing has been considered a problem elsewhere but
is probably not the cause of loss here. Investigations into the plant’s autecology in this unique
habit would be worthwhile.

Reintroductions are currently being considered at Nympsfield and Upper Ribblesdale. At the
former site, the authors feel that if the trees could be cleared from the original area it would then
be suitable for the plant which might even regenerate naturally from dormant seed. At both sites,
however, in order that genetic diversity is maintained, it is important that only seed or cultivated
plants originating from the site in question is used when this is attempted. During this survey, seed
from most of the extant sites was placed in the Kew Seed Bank, partly for this purpose. Leaf tissue
for DNA analysis has also been collected.

Unlike the commoner subsp. lamprocarpa, subsp. muricata is considered a very rare Red Data
Book plant in Britain and is listed as critically endangered (Roberts 1999). However, its widely
scattered, but very local, distribution (Gloucestershire to Yorkshire) suggests there may be
undiscovered populations. Much apparently suitable ground occurs on the Craven limestones of
Yorkshire, as well as in south Cumbria, Derbyshire, the Welsh borders and the Cotswolds and the
plant may be overlooked. It may also wrongly recorded, perhaps as subsp. lamprocarpa or C.
spicata, as this latter plant can also favour a similar habitat and grows with the Swaledale
population.

CAREX MURICATA SUBSP. MURICATA 285

Any plant of the Carex muricata aggregate found on warm, open limestone scree should be
carefully examined in case it is this taxon. C. muricata subsp. muricata is best recognised by its
relatively rigid, upright flowering spikes which appreciably exceed the leaves, and by the
(initially) contrasting dark glumes and lighter utricles (see Introduction). On maturity the
inflorescences may become decumbent and the fruiting head almost black. The utricles of subsp.
muricata are both more flanged and smaller than in subsp. Jamprocarpa and C. spicata whilst the
female spikelets are more rounded. To help in identification, typical plants and fruiting heads of
both subspecies are shown in the accompanying drawing (Fig. 1).

4

Figure 1. Comparative sketches of Carex muricata subsp. muricata (above) and subsp. lamprocarpa (below):
habit (x 1/5), inflorescences (x 2). A: Immature utricles and glumes (x 6) in June. B: Mature utricles and
glumes (x 6) in August.

286 M. J. Y. FOLEY AND M. S. PORTER

ACKNOWLEDGMENTS

We wish to thank the relevant vice-county recorders for their help with information on the various
sites and English Nature for allowing us to publish this paper.

REFERENCES

ANON (1981). Plant records. Watsonia 13: 342.

DAVID, R. W. (1979). Another British locality for Carex muricata L. sensu stricto. Watsonia 12: 335.

DAVID, R.W. & KELCEY, J. G. (1975). Carex muricata L. sensu Nelmes and Carex bullockiana Nelmes.
Watsonia 10: 412-414.

DAVID, R.W. & KELCEY, J. G. (1985). Biological Flora of the British Isles, 159. Carex muricata L. aggregate.
Journal of ecology 73: 1021-1039.

FOLEY, M. J. Y. & PORTER, M. S. (1999). A survey for Carex muricata subsp. muricata in Britain.
Unpublished report to English Nature, Cumbria.

NELMES, E. (1947). Two critical groups of British sedges. The Botanical Society and Exchange Club of the
British Isles, Report for 1947, 13: 95-105.

ROBERTS, F. J. (1999). Carex muricata L. subsp. muricata, in WIGGINTON, M. J. ed. British Red Data Book. 1.
Vascular Plants, 3rd ed. J. N. C. C. Peterborough.

(Accepted February 2000)

Watsonia 23: 287—291 (2000) 287

Choice of surveyor is vital to the reliability of
floristic change studies

A. OREDSSON*

Department of Systematic Botany, Lund University, O. Vallgatan 14-20, S-223 61 Lund, Sweden

ABSTRACT

The flora of an area of 46 km? of farmland and wooded hills surrounding the lake of Striern, Sweden was
surveyed. Fifty-six different habitats were selected from topographical and geological maps, within which 214
plots were chosen by random sampling. The plots, mostly 10 m x 10 m, were investigated in 1972 and 1998.
Each plot was recorded by walking through on parallel lines 1 m apart. Six of the best botanists from the
region carried out the field work and their recording capacity was estimated; apart from one individual,
variation between surveyors was negligible. Over the 26 years, the mean number of species recorded per plot
fell from 32-5 to 30-2; the downward trend probably reflects a true decline in the frequency of species.

KEYWORDS: Change in flora, different surveyors, Sweden.

INTRODUCTION

Ever since the times of Linnaeus, Swedish botanists have maintained a high level of knowledge of
the native vascular plants in Sweden. The tradition of writing Floras of provinces is flourishing,
often with many amateurs taking part in the field work.

Floras change. Since World War I, a marked increase in the standard of living in Sweden has
taken place which has had a massive impact on the natural environment, through air pollution and
indirectly by changes in life style, including migration from the countryside (Oredsson 1990).

Unfortunately there are few historical studies which have quantified the abundance of vascular
plants sufficiently accurately to allow a statistically reliable assessment of change. Consequently,
Red Lists of threatened species are rather more subjective than objective comparisons with the
past. One Swedish opportunity to assess change against quantitative historic data was by repeating
studies carried out by Hans Goransson as part of his quantitative survey of the flora related to
pollen rain in Ostergotland in 1972 in which I took part. As I had already demonstrated
considerable changes in northern Scania (the southern-most province of Sweden) after 25 years
(Oredsson 1990), it was assumed that any changes which had taken place in Ostergétland would
also be detectable. In this connection, a problem hitherto little discussed in Sweden has arisen,
namely the influence of individual botanists on the results of the field work.

This paper deals with a case in which six botanists were involved in recording the same habitats
26 years apart. The field work area covers 46 km’ of farmland and wooded hills surrounding the
lake of Striern, in southern Ostergétland, about 200 km S.W. of Stockholm.

METHODS

The 1972 survey was carried out by quaternary biologist Hans Géransson (HG) and the author
(AO), while the 1998 survey was carried out by me and four assiduous, proficient, amateur
botanists from Ostergétland (proof that Sweden is still teeming with Linnean pupils!) - Janne
Andersson (JA), Johan Bergstedt (JB), Dan Lindmark (DL) and Rolf Wedding (RW).

Before the 1972 survey, the study area was stratified using 1:50,000 topographical and
geological maps. 56 habitats were established, 44 of which combined information from both maps

“Address for correspondence: e-mail: Alf.Oredsson @sysbot.lu.se

288 A. OREDSSON

(e.g. forest on V4xjO granite, or open terrain on clay), and the remaining twelve from the
topographical map (e.g. church, or public roads in open terrain). Within each habitat, 1-10 plots
were chosen by random sampling depending on the size of the habitat. In 144 cases the general
position of the plot was located using 100 m co-ordinates, in 70 cases using 50 m co-ordinates. In
the field, the final position was then determined using further random sampling by assistants
without specialist botanical knowledge, and thus the final positions differed slightly between the
years, although the map co-ordinates were identical.

157 of the plots were 10 m x 10 m. On shorelines (22 plots) a 10 m x 20 m rectangle was
surveyed, half in water, half on land. Along roads and water courses (22 plots) the length was 10
m, whilst the breadth varied. These 201 plots were marked out with a cord, and at least the
terrestrial part of the plot was walked through along parallel lines 1 m apart. In the remaining 13
plots which included urban areas, the boundary of the garden, grounds or churchyard provided the
outer edge of the plot.

In each plot, all identifiable vascular plant species were recorded, no matter what stage in the
life cycle (e.g. seedling or withered) or number of individuals. Combining both years, 543 species
were found excluding cultivated species.

RESULTS

If all six botanists had a similar capacity to find almost every species in the plots, it would be
possible to make a reliable comparison in the flora between 1972 and 1998. The results for the
botanists were therefore investigated in detail.

1972 SURVEY

Hans Goransson surveyed 131 plots once between the end of June and early August and 22 plots
twice, the first time in May and the second in July or August (Table 1). He found an overall
average of 33-1 species per plot, but plots surveyed once had an average of 32-6 species and plots
surveyed twice an average of 36-0 species, though this was not statistically significant (The Mann-
Whitney U-test; p=0-14).

I surveyed 61 plots in early August, and found an average of 31-0 species per plot. To make
allowances for the different habitats surveyed, which affect the number of species, only habitats
investigated by both HG and AO were compared (viz. 107 plots in 21 habitats). For each habitat,
the median number of species was calculated, and then the number of plots which fell above,
equalling or below these medians summed (Table 2). Using these medians, the differences in
number of species recorded between HG and AO are not statistically different (Table 3). It is
concluded that HG and AO would have found approximately the same number of species if they
had investigated the same plot independently in 1972.

TABLE 1. TOTAL NUMBER OF RECORDS, NUMBER OF PLOTS AND
AVERAGE NUMBER OF SPECIES PER PLOT IN THE STRIERN AREA
SURVEYED IN 1972 AND 1998.

1972 1998
Surveyors Ga HGb HGS AOI total | AO2 DL JA JIB ~—s RW _ total

No of records 792 4265 5057 1892 6949 | 1991 453, 2188 1290 537 6459
No of plots 22 131 153 61 214 63 18 68 50 15 214

Average no
species/plot 36-0 32:6 ei 31-0 32:5 31-6 25:2 32:2 25:8 35-8 30-2

Surveyors in 1972: Hans Goransson (HGa), twice (12-28 May, 6 July to 5 August) or (HGb), once
(29 June to 11 August); and Alf Oredsson (AO1) (5-11 August).

Surveyors in 1998, 6-17 July: Alf Oredsson (AO2); Dan Lindmark (DL); Janne Andersson (JA);
Johan Bergstedt (JB); and Rolf Wedding (RW).

SURVEYORS AND FLORISTIC CHANGE STUDIES 289

TABLE 2. NUMBER OF PLOTS RECORDED BY DIFFERENT BOTANISTS IN 1972 AND
1998 WITH ABOVE, EQUALLING OR BELOW MEDIAN NUMBER OF SPECIES PER PLOT
FOR THE SAME HABITATS. SURVEYORS AS IN TABLE 1.

1972 1998
HGa HGb HG AOI AO2 DL JA JB RW
Above median 1 23 24 22 24 2 Di ily 8
Median 0 12 12 4 1 If 6 7 1
Below median 1 28 29 16 15 15 29 16 5

1998 SURVEY

In 1998, the 214 plots were surveyed between 6-17 July by myself (AO2), Dan Lindmark (DL),
Janne Andersson (JA), Johan Bergstedt (JB) and Rolf Wedding (RW). The average number of
species recorded per plot varied considerably; DL and JB recorded the least, AO and JA the
middle numbers and RW the largest numbers per plot (Table 1).

Again, comparisons between surveyors need to take into account the different habitats which
each surveyed (Table 2). Median species counts in 174 plots in 35 habitats were again
investigated. JA and JB recorded almost as many plots above as below the median, AO2 and RW
recorded slightly more plots above the median, and DL more below the median.

The Fisher Exact Probability Test indicated no significant differences in efficiency between the
surveyors, except for DL (Table 3). AO and RW had previous experience of field work of this
kind, and in a given plot might have observed one or two species more than the other botanists,
whereas DL came in late and so was less prepared for the task.

COMPARISON OF 1972 SURVEY WITH 1998 SURVEY
If HG recorded in 1972 with the same degree of accuracy as did jointly DL, JA, JB and RW in
1998, and if my capacity for recording species had not changed, my results from the habitat
comparisons in both surveys should have been alike, and they were. The number of plots which
fell above, equalling or below the medians in 1972 were 22—4—16; and in 1998 were 24—-1-15
(Table 2). Equalling values excluded, The Fisher Exact Probability Test gives p=0-17 (Table 3)
which is not statistically significant. Overall I came closer to the ultimate goal of finding all
species present in a plot than did the average colleague. As I recorded almost the same number of
plots in each year, this should not have affected the comparison between years species by species.
Between 1972 and 1998, the average number of species recorded fell from 32-5 to 30-2 species
per plot, a reduction of 7%. Despite the fact that six different botanists were involved in the field
work, the downward trend probably represents a true decline in diversity. Details of which species
changed will be published in another paper (Oredsson, in prep.).

TABLE 3. ESTIMATES OF DIFFERENT SURVEYORS’ RECORDING CAPACITY IN 1972,
1998 AND ONE SURVEYOR BOTH YEARS. DATA FROM TABLE 2 (EXCLUSIVE OF
MEDIAN) TESTED BY THE FISHER EXACT PROBABILITY TEST (SIEGEL 1956).
SURVEYORS AS IN TABLE 1.

LOFZISENO9S
AOQ2

0-005
JB 0-13 0-005 0-17
0-006 0-17

All figures are probability values

290 A. OREDSSON

DISCUSSION

DIFFERENCES BETWEEN SURVEYORS

In 1987-1988, the B.S.B.I. Monitoring Scheme engaged over 1600 botanists in a sample survey of
10 km squares in Britain and Ireland, during which time 2660 taxa were recorded (Rich &
Woodruff 1990). The problems encountered during a comparison with data from the Atlas of the
British flora prompted the harsh conclusion that ‘the widespread occurrence of recording bias
suggests that information about recorders and their behaviour should be collected and analysed as
carefully as information about the organisms themselves’ (Rich & Woodruff 1992)

Four tetrads (2 km x 2 km each) in West Sussex were investigated by 29 volunteers with a range
of botanical abilities representative of many national and county Flora projects in July 1992 (Rich
& Smith 1996). They worked in pairs recording vascular plant species for 2:5 hours in each square.
Individuals were then rotated to different squares and paired with a different botanist. In all, 634
species were recorded. From this experiment with different surveyors, Rich & Smith learnt that
different botanists were the key factor causing variation between the surveys.

In light of these two examples, it was obviously important to recruit the very best botanists
available for the 1998 survey of the Striern area, or the differences between surveyors might have
masked any floristic change. The results demonstrate it is possible to even out the differences
between surveyors provided they are of sufficient quality. This proved to be the case for
everybody involved in this project.

SIGNIFICANCE OF INTENSITY

From April to October 1982, six competent field botanists independently recorded the vascular
plants in three British woods, each about 30 ha (Kirby et al. 1986). Two different, methods were
used; a walk along 3—3-5 km long routes and quadrat survey in squares of 200 m’. It was found
that differences between surveyors or seasons were much smaller than differences in methods or
intensities of survey.

In the Striern surveys, the standard plot was walked through along ten parallel lines only 1 m
apart so few, if any, vascular plant species should have eluded a skilled botanist’s observation.
However, Goransson found on an average 3-4 more species in plots investigated twice than in plots
investigated once, and although this is not statistically significant it may indicate there is still some
variation due to the season.

ADVANTAGE OF STRATIFIED RANDOM SAMPLING

In August 1964, two professional botanists investigated the flora of the large peninsula of Lake
Toisvesi in central Finland, which was divided into 50 1 km squares. Each botanist tried to find as
many species as possible in every square. Altogether 362 species were recognised, eleven of which
were only found by one surveyor, and 14 only by the other. Moreover, the two surveyors were
shown to preferentially select different habitats to record which accounted for some differences
between lists for the same squares (Kyt6vuori & Suominen 1967). In the Striern area, the stratified
random sampling technique prevented the surveyors being drawn towards any favourite habitats.

Stratified random sampling is also relatively efficient. In a survey of mountain forest types in the
Swiss Alps, it was found that stratified random sampling provided an accurate picture of the small
scale vegetation pattern at low sampling effort in areas sized 10-50 km? (Godickemeier er al.
1997). It is also used successfully for the British Institute of Terrestrial Ecology Countryside
Survey (Barr et al. 1993).

A quarter of a century ago, I pointed out the advantages of stratified random sampling for
regional mapping of the vegetation (Oredsson 1974). Unfortunately, this method is still far from
being generally accepted. Sticking to vague ‘walk-about’ strategies is a horrible waste of human
capital, humiliating a large number of amateur botanists willing to spend so much of their spare
time on botanical research.

ACKNOWLEDGMENTS

Thomas Karlsson and Jimmy Persson, at that time colleagues at the Institute of Systematic Botany,

SURVEYORS AND FLORISTIC CHANGE STUDIES 291

Lund University, took part in the development of the method applied in the survey of the flora in
the Striern area. They both raised valuable objections to premature ideas of mine. Bj6rn Marklén
and Joachim Zethraeus (+) marked out the plots in the fields on 5—11 August 1972. In 1998, they
were followed by Romia Ahmed, Gustav Jonsson, Stefan Jonsson, and David Rovan. It was a great
pleasure to work together with all of them. Grants were gratefully received from the Lund
Botanical Society and the King Carl XVI Gustaf’s 50th birthday foundation for Science,
Technology and the Environment.

REFERENCES

BARR, C. J. et al. (1993). Countryside survey 1990. Department of the Environment, London.

GODICKEMEIER, [., WILDI, O. & KIENAST, F. (1997). Sampling for vegetation survey: Some properties of a
GIS-based stratification compared to other statistical sampling methods. Coenoses 12 (1): 43-50.

KIRBY, K. J. et al. (1986). Seasonal and observer differences in vascular plant records from British woodlands.
Journal of ecology 74: 123-131.

KYTOVUORI, I. & SUOMINEN, J. (1967). The flora of Ikkalanniemi (commune of Virrat, Central Finland),
studied independently by two persons. Acta Botanica Fennica 74.

OREDSSON, A. (1974). A new method of mapping vegetation (In Swedish). Forskning och Framsteg 1974 (3):
5-7.

OREDSSON, A. (1990). Changes in the flora of Matteréd, a parish in southern Sweden, between 1964 and 1989
(In Swedish, summary in English). Svensk Botanisk Tidskrift 84: 293-311.

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

RICH, T. C. G. & SMITH, P. A. (1996). Botanical recording, distribution maps and species frequency. Watsonia
21: 155-167.

RICH, T. C. G. & WOODRUFF, E. R. (1990). The B.S.B.I. Monitoring Scheme 1987-88. Chief Scientist’s
Directorate report no. 1265. Nature Conservancy Council, Peterborough.

RICH, T. C. G. & WOODRUFF, E. R. (1992). Recording bias in botanical surveys. Watsonia 19: 73-95.

SIEGEL, S. (1956). Nonparametric statistics for the behavioural sciences. McGraw-Hill Kogakusha, Ltd,
Tokyo.

(Accepted January 2000)

at.

“

Watsonia 23: 293-297 (2000) 293

Recording and the declines of the Nationally Scarce plants
Ajuga pyramidalis L. and Melampyrum sylvaticum L.

T. C. G. RICH
67 Heol Uchaf, Rhiwbina, Cardiff CF 14 6SR
and
C. SYDES

Scottish Natural Heritage, 2 Anderson Place, Edinburgh EH6 5NP

ABSTRACT

An assessment of the reasons for decline of the Nationally Scarce plants Ajuga pyramidalis and Melampyrum
sylvaticum was carried out by revisiting pre-1970 sites listed in the Scarce Plants Database. 17% of the records
were erroneous. Inadequate original information probably accounted for failure to refind another 25% of the
records. Excluding these records, 25% of Ajuga and 33% of Melampyrum pre-1970 sites were refound. Loss
of habitat has probably accounted for declines in 25% of Ajuga sites and 66% of Melampyrum sites. In 50% of
the Ajuga sites suitable habitat was still present but plants were not refound. Real decline has occurred in both
species, but the degree of change was masked by the problems of recording.

Keywords: conservation, botanical records, habitat loss, Lamiaceae, Scrophulariaceae.

INTRODUCTION

Nationally Scarce species are those which occur in 16-100 hectads (10-km x 10-km squares) in
Britain, and are of concern to conservationists as the second rarest group of species in the British
flora after the Red List species. Despite protection measures, local populations of rare and scarce
species are still liable to extinction, but the greatest loss of biodiversity has occurred, and appears
still to be occurring, amongst common but evidently vulnerable taxa (Sydes 1997). Between 1990
and 1993 a review of the Nationally Scarce vascular plant species was carried out (the Scarce
Plants Project), with a main aim of creating a computer database (Stewart et al. 1994). The
resulting Scarce Plants Database records were used to revise the list of Nationally Scarce species,
update distribution maps and assess changes in the frequency of species, and the results were
presented in Scarce Plants in Britain (Stewart et al. 1994).

The Scarce Plants review indicated that a number of species are declining rapidly, as judged
either from the high proportion of pre-1970 records shown on some maps, or the high percentage
of the post-1970 records compared to the records on the maps in the Atlas of the British flora
(Perring & Walters 1962). For instance, species such as Ajuga pyramidalis L., Betula nana L.,
Carex maritima Gunnerus, Melampyrum sylvaticum L., Mertensia maritima (L.) Gray, Meum
athamanticum Jacq. and Saxifraga nivalis L. are species which are fairly widespread in Scotland
but have significant numbers of pre-1970 records in Stewart et al. (1994). The causes of these
declines are not always clear, especially where they occur in the remoter parts of Scotland, and
might be attributable to natural metapopulation dynamics, under-recording, habitat change, or
combinations of the three. It is valuable to have some estimate of the relative importance of these
factors when assessing priorities for conservation, so as to concentrate on species whose habitats
are declining and not waste resources on species simply under-recorded.

To investigate the reasons for the large apparent declines in these seven species, in 1995 Scottish
Natural Heritage (S.N.H.) commissioned field work to revisit pre-1970 sites and assess possible
reasons for loss of sites. Of these species, the changes in Ajuga pyramidalis and Melampyrum

294 T. C.G. RICH AND C. SYDES

sylvaticum were quantified in the most detail, including searches for historical herbarium and
literature records to provide additional background information. They provide good examples of
the balance between decline and under-recording of species which occur in remote areas (e.g.
Ajuga) or are difficult to identify (e.g. Melampyrum). The aim of this paper is to summarise the
results of the work; full details are given in Rich et al. (1996).

METHODS

For both species, ten pre-1970 sites with known 1-km square or 6-figure grid references were
selected from the Scarce Plants Database to give a wide geographical spread in Scotland. Sites
with known 1-km squares or 6-figure grid references were selected to maximise the chances of
refinding the original site as many old records are vague and may not even be localised to a hectad.
Ajuga pyramidalis is widespread on the Inner and Outer Hebrides from which recent data are
particularly lacking, but these islands had to be excluded on the grounds of cost. Remote upland
sites for M. sylvaticum were also excluded for safety reasons. As there are few detailed pre-1970
records for Melampyrum, a number of sites had to be taken as ‘best guess’ 1-km squares (e.g.
‘Falls of Clyde’ was taken to be NS/88.41). In addition, recorders visited three known post-1970
sites to familiarise themselves with the species and their habitats, selected from personal
knowledge or where there were 6-figure references in the Scarce Plants Database. A few additional
sites were also investigated for both species.

The sites were visited during late June and July 1995, and the full 1-km square searched as far as
was practicable, concentrating on the most appropriate habitats. A wider area was searched where
the previous record was not clearly defined. Where the plants could not be refound, the field
workers assessed whether the plant was likely to have gone or not from the presence or absence of
suitable habitat. Most Melampyrum sites were surveyed by R. FitzGerald, often with the help of
the B.S.B.I. v.c. Recorders; the Ben Hiel site was surveyed by P. A. Smith. G. M. Kay surveyed
Ajuga pyramidalis sites. The areas searched and locations of extant populations were indicated on
1:25,000 maps. Where populations were not refound, notes were made on the likely causes for loss
or lack of re-discovery, and further enquiries were made about the locality from the original
recorder, if they could be traced.

Historical data from specimens in BM, BRISTM, E, GL, GLAM, K, LIV, MANCH and OXF
(abbreviations follow Kent & Allen 1984) were abstracted. A literature search of records in the
floras and journals held in the libraries at Kew and the Natural History Museum was carried out,
with additional references traced in BSBI Abstracts and Simpson (1960).

RESULTS

The results of the field survey are given in Tables 1 and 2, and are summarised in Table 3.
Erroneous records in the Scarce Plants Database, where either the species or grid reference was
wrong, are believed to be responsible for failure to find one Ajuga (9%) and three Melampyrum
(23%) pre-1970 sites (17% overall). In addition, insufficiently detailed original information was
thought to account for lack of discovery in another two of the Ajuga (18%) and four Melampyrum
(30%) pre-1970 sites (25% overall).

Excluding these sites, plants were refound in two of the pre-1970 Ajuga sites (25%), and two of
the pre-1970 Melampyrum (33%) sites (29% overall). Habitat change probably accounted for
losses at two Ajuga (22%) and four Melampyrum (50%) pre-1970 sites (35% overall). One of the
Ajuga sites appeared still suitable but plants could not be found after an extensive search indicating
they may have disappeared perhaps by chance demographic events, but some were found in a new
site nearby. Plants were not refound in the other pre-1970 sites either because they have
disappeared or because they were over-looked. One post-1970 site with 6-figure grid reference
was not refound for one of these same reasons.

166 (41%) historical records additional to the Scarce Plants Database were found for both
species (Table 4), which provided valuable additional information about distribution and ecology,
especially for M. sylvaticum where a number of specimens had been wrongly named (data not
presented; see Rich 1997; Rich et al. 1998; 1999).

DECLINES OF AJUGA PYRAMIDALIS AND MELAMPYRUM SYLVATICUM 295

TABLE 1. RESULTS OF 1995 FIELD SURVEY FOR AJUGA PYRAMIDALIS

Site

Loch Call an Uidhean
Torr Achilty
SW of Clais nan
Cruineachd
Traigh an T-Straithain
Creag na Faollinn
Aird a’ Chuilinn
Rhigolter/An Fhireach
Sanna Bay

Creag Clais nan
Cruineachd
Clais nan Cruineachd

Cnoc na Stri
Deadh Choimhead

Clashnessie
Loch Buine Moire
Thurso

Sanna Bay

Grid reference Last record Refound in 1995? Comments

6-figure
6-figure
6-figure

1 km
1 km
1 km
1km
6-figure
6-figure

1 km
6-figure

1 km

1992
1992
199]

1988
1967
1964
1964
1964
1961

1961
1961

1961

1954
1952
1916

New site

Wrong grid reference in database
Not refound, possibly still present
Wrong grid reference in database
Not refound, possibly still present
Not refound, possibly gone

Not refound

Possibly a different site

Inadequate information, later found to be
inaccessible site on cliff

Inadequate information, later found to be
inaccessible site on cliff

Two sites present

Possible gorse invasion caused loss

Not refound, possible gorse invasion or
agricultural improvement

Near old record

TABLE 2. RESULTS OF 1995 FIELD SURVEY FOR MELAMPYRUM SYLVATICUM

Site

Birks of Aberfeldy
Allt Doe
Glen Buck

Loch Ossian

Linn of Corriemulzie
Ballater Bridge
Reekie Linn

Struan, River Garry
Roy Bridge
Blackhall Woods

Den of Airlee

Glen of Reichip

Ben Hiel - Ben Loyal
Falls of Frenich

Falls of Clyde
Rowardennan

6-figure
6-figure
6-figure

1 km

1 km
6-figure
1 km?

6-figure
1 km?
1 km

1 km

1 km?
6-figure
1 km?

1 km?
1 km

1992
1984
LOT

1958
1957
1955
1950

1949
1896
1871

1846

1821

Undated
Undated

Grid reference Last record Refound in 1995? Comments

Confused information, actual site not
visited

Biggest population found

Grazing damage

Now densely shaded

Dense woodland and agricultural runoff
may have caused loss

Forestry caused habitat change

Possibly inadequate information

Forestry caused habitat change, but
imprecise original record

Imprecise original site, now densely
shaded

Possibly inadequate information, site
now densely shaded

Probable error for M. pratense

Possibly inadequate information

Error for M. pratense

Error for M. pratense

296 T. C. G. RICH AND C. SYDES

TABLE 3. CLASSIFICATION OF RESULTS INTO CATEGORIES RELATED TO
RECORDING

Melampyrum sylvaticum Ajuga pyramidalis

Pre-1970 sites
Sites refound in 1995 2
Sites not refound but possibly still present 2
Probable loss due to habitat change s)
Incorrect/inadequate information 3
Erroneous record 3
Post-1970 sites
Sites refound in 1995 2
Sites not refound 0
1
0

mere NO NbN

Incorrect information

Re ee LL

New sites
Total sites investigated 16 16

TABLE 4. NUMBERS OF RECORDS IN SCARCE PLANTS DATABASE AND ADDITIONAL
RECORDS TRACED AFTER CORRECTION FOR DUPLICATES AND ERRORS

Species Scarce Plants Database Additional records traced Total records

Ajuga pyramidalis Total no. records 330 Bil 361
Pre-1970 hectads 37 2 39
Post-1970 hectads 53 0 53

Melampyrum sylvaticum Total no. records 70 135 205
Pre-1970 hectads 22 26 48
Post-1970 hectads pH | i 22

DISCUSSION

Assessing the causes of the declines in Ajuga and Melampyrum was complicated by the problems
of recording and lack of detailed information. 13% of the records in the Scarce Plants Database
were found to be erroneous. Inadequate original information probably accounted for failure to
refind another 17% of the records. Where we were able to trace the original recorders, they kindly
provided valuable information about the records which would have been useful in the field, such
as occurrence on cliffs or the accuracy of the grid references. Similarly, the work on historical
records in herbaria and the literature provided valuable information. It is recommended for similar
projects in the future that this original information be obtained prior to commencing field work.

Ajuga pyramidalis and M. sylvaticum were refound in 29% of the pre-1970 1-km squares overall
(Table 3). Suitable habitat was still present in a further 29% of the squares overall, and we may
have over-looked the species in some of these sites. Searching the pre-1970 sites thoroughly is
very difficult, especially when they are as large an area as a 1-km square, and both species can be
difficult to find. A. pyramidalis tends to occur in very small, scattered populations, and localities
on cliffs are difficult to search adequately or safely. M. sylvaticum is also usually found in small,
discrete populations, often in steep, wooded ravines where access is difficult. It is always difficult
to be certain that a species has gone from a site. Nevertheless, ecological theory suggests that
populations can be expected to become extinct on some sites for a variety of reasons including
natural metapopulation dynamics.

Habitat change (e.g. scrub invasion, forestry) has probably accounted for loss of 22% of the
Ajuga pre-1970 1-km squares (Table 1), and 50% of the M. sylvaticum squares in eastern Scotland
(Table 2). Further discussion of the causes of the declines in these species are given in Rich ef al.
(1999) and Rich et al. (1998) respectively. It is concluded that both species have declined, but the
true extent of decline was masked by the problems of recording.

DECLINES OF AJUGA PYRAMIDALIS AND MELAMPYRUM SYLVATICUM 297

The data recorded for rare species have become more detailed in recent years. The range of
species recorded in this way should be broadened and the quality of information enhanced so that
conservation resources do not have to be diverted into checking data when the Scarce Plants in
turn become rare. The resource of historic records in Britain’s herbaria is under-utilised and could
greatly extend our knowledge of past change.

ACKNOWLEDGMENTS

We would like to thank those who gave access to land or helped with the field work or
information: Penny Angold, Elaine Bullard, Kathy Fallowfield, Ro FitzGerald, Sam Hallet,
Barbara Hogarth, Graeme Kay, Peter Macpherson, Douglas McKean, Henry Noltie, Fred Rumsey,
Alex Scott, Ros Smith, Edna Stewart, Neale Taylor, Agnes Walker, Keith Watson, Phyllis White
and Peter Wortham. We would like to thank the Keepers of BM, BRISTM, E, GL, GLAM, K,
LIV, MANCH and OXF for access to the herbaria and libraries, and an anonymous referee for
suggesting improvements to the manuscript.
The work was funded by Scottish Natural Heritage, contract SNH/001G/95 IBB.

REFERENCES

KENT, D. H. & ALLEN, D. E. (1984). British and Irish herbaria. Botanical Society of the British Isles,
London.

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

RICH, T. C. G. (1997). Using botanical records to interpret changes in frequency of British plants. The Biology
Curator 10: 8-12.

RICH, T. C. G., FITZGERALD, R. & KAY, G. M. (1996). Review and survey of scarce vascular plants. 3
volumes. Unpublished contract report to Scottish Natural Heritage.

RICH, T. C. G., FITZGERALD, R. & SYDES, C. (1998). Distribution and ecology of Small Cow-wheat
(Melampyrum sylvaticum L.; Scrophulariaceae) in the British Isles. Botanical journal of Scotland 50: 29—
46.

RICH, T. C. G., KAy, G. M., & SYDEs, C. (1999). Distribution and ecology of Pyramidal Bugle (Ajuga
pyramidalis L.; Lamiaceae) in the British Isles. Botanical journal of Scotland 51: 181-193.

SIMPSON, N. D. (1960). A bibliographical index of the British flora. Privately published.

STEWART, A., PRESTON, C. D. & PEARMAN, D. A., eds. (1994). Scarce plants in Britain. Joint Nature
Conservation Committee, Peterborough.

SYDES, C. (1997). Vascular plant biodiversity in Scotland, in FLEMING, U. V. & USHER, M. B., eds.
Biodiversity in Scotland: status, trends and initiatives, pp. 89-104. The Stationery Office, Edinburgh.

(Accepted January 2000)

y

+e

Watsonia 23: 299-303 (2000) 299

Dactylorhiza incarnata (L.) So6 subsp. ochroleuca (Wiistnei ex
Boll) P. F. Hunt and Summerh. (Orchidaceae): A comparison of
British and European plants

Mid: Y. FOLEY.

Department of Biological Sciences, Institute of Environmental and Natural Science, University of
Lancaster, Lancaster LAI 4YQ

ABSTRACT

A brief summary is given of the occurrence of Dactylohiza incarnata subsp. ochroleuca in Britain and
continental Europe and examples of the very few remaining native British plants have been compared to those
at three European localities. Although the sample size at the British locality was inevitably extremely small, it
is tentatively concluded that plants at all the localities are conspecific.

KEYWORDS: morphometrics, diagnostic characters.

INTRODUCTION

Known in Britain under the name Dactylorhiza incarnata subsp. ochroleuca, this plant has been
placed at differing taxonomic levels by many authors. Relatively rare and widely scattered from
central Europe to Scandinavia, the only genuine British records are from eastern England (East
Anglia) where it now appears to be reduced to one small population. As the subspecific epithet
implies, the flower colour is pale yellow and both in Britain and on the continent, it has been over-
recorded through confusion with yellow flowered, anthocyanin-lacking forms of other subspecies
of D. incarnata, especially subsp. pulchella. Flower colour alone, therefore, is not sufficient to
distinguish it and previous authors (e.g. Pugsley 1939; Nannfeldt 1944; Heslop-Harrison 1956;
Rajchel 1964; Lundqvist 1967; Bateman & Denholm 1983, 1985) have variously used a
combination of characters in its taxonomic delimitation. These include its relatively robust habit,
tall, broad stem, large leaves and bracts, and relatively large, pale yellow, unmarked and trilobed
labella, frequently having notched lateral lobes. It is a plant with a precise habitat requirement,
growing in the drier parts of rich fens where it occupies a distinct ecological niche.

As Orchis incarnata var. ochroleuca, it was first described by Boll (1860) from plants seen by
Wiistnei at Sternberg (in Mecklenburg), Germany in 1854. From his record (Wiistnei 1854), it is
not clear in exactly what type of habitat they grew and, because of this, reservations have been
expressed (Bateman & Denholm 1985) regarding the true identity of the plants upon which the
name ochroleuca is based. However, an examination of preserved specimens collected by Wiistnei
(Mecklenburg, Wiistnei, s.d. (W!)), similar to those on which Boll based his original description,
confirms that at least one of them is referable to this subspecies. This is the central specimen of the
three mature specimens on the sheet and bears the label “Orchis incarnata L. var. floribus
ochroleucis......In Mecklenburg. K Wiistnei”’.

The first definite British record was made in 1935 from the upper Ouse/Waveney valley area
(Blo Norton Fen, 23.vi.1935, Lousley (BM!)) and shortly afterwards it was also found at various
nearby fens within the same general area of East Anglia. Previous to this, however, the
Stephensons had tentatively recorded it from both Kidwelly, South Wales as well as from an
unlocalised site in East Anglia (Stephenson & Stephenson 1923). Although no preserved
specimens collected by the Stephensons from the latter area have been traced and the Kidwelly
locality has not been re-found, a Stephensons’ specimen, labelled ‘“(b)? v. ochroleuca’, does exist
(Kidwelly, 27.vi.1919 (TOR!)). In general habit this does show some affinity to less robust plants
of subsp. ochroleuca, but its labella are line-marked and almost entire and these are characters not
found in that plant. So, other than for the few known East Anglian localities, the plant has not been

300 M. J. Y FOLEY

confirmed elsewhere in Britain although erroneous records proliferate due to confusion with
similar coloured forms of other subspecies of D. incarnata.

Mainly due to habitat drainage, subsp. ochroleuca is now a highly endangered British plant.
There appear to have been no recent records at Blo Norton/Thelnetham and it may now be extinct
there. Further east at Redgrave there were reports of a few plants until the late 1980s but none have
been seen recently, although a single specimen was recorded at Market Weston Fen in June 1995
(M. Sanford, pers. comm., 2000). At Chippenham, perhaps now the only extant British site, there
were as many as 50 flowering plants in the early 1980s but now, even in a good year, there are
rarely more than five.

Subsp. ochroleuca is also known from several parts of continental Europe, especially Germany
(Lubs 1968; Fiiller 1983), Poland (Rajchel 1964) and the central Alps (Hegi 1939) and reaches as
far north as southern Sweden (Nannfeldt 1944; Wiefelspiitz 1976) and Estonia (Kuusk 1991) and
eastwards into Russia (L. Averyanov, pers. comm., 1990). Pale-yellow flowered plants from
species-rich fens in the Picardy area of north-eastern France have sometimes been recorded as
subsp. ochroleuca but these appear referable to pigment-lacking variants of the type or of subsp.
pulchella. Further evidence for this was gained during a visit to the area in 1991.

A MORPHOMETRIC COMPARISON OF BRITISH AND EUROPEAN PLANTS

Whether the British plant is the same as that of continental Europe has sometimes been questioned
(e.g. Stephenson & Stephenson 1923; Landwehr 1977). In order to examine this, the opportunity
has been taken to compare the morphology of the few remaining examples of the only apparently
extant British population with others occurring at localities in Scandinavia and mainland Europe.
Due to its great scarcity, however, the number sampled in Britain was extremely small and,
because of this, any conclusions drawn can only be tentative.

To carry out this comparison, plants from three European populations - Storsund, Gotland
(Sweden), Viidumae, Saaremaa (Estonia) and Murnau, Bayern (Germany), all apparently referable
to subsp. ochroleuca, were morphologically compared to those at Chippenham (England) for those
characters considered most useful in separating D. incarnata subsp. ochroleuca from closely allied
taxa. Under precise conditions and in a similar manner to the method of Bateman & Denholm
(1985), data were obtained in a non-destructive manner for the following characters: 1. plant

TABLE 1. MORPHOMETRIC COMPARISON OF BRITISH AND EUROPEAN PLANTS OF
DACTYLORHIZA INCARNATA SUBSP. OCHROLEUCA - POPULATION MEANS AND

STANDARD ERRORS
Chippenham Storsund, Gotland Viidumée, Saaremaa Murnau, Bayern
(England) (Sweden) Island (Estonia) (Germany)
No. of plants measured (n) =) 10 10 13

1. Plant height 338-3 + 23-8 235-1 + 20-3 321-8 + 19-1 291:5+7-9
2. Inflorescence length 42-3 +49 47-0 + 2-9 62:3 + 3-9 59-1 + 2:8
3. Stem width near base 6-1+0-6 6-8 + 0:5 7:54 0-4 5:7 40:3
4. Stem width below inflorescence 3-8+0-4 4:7+0-4 4:9 + 0-6 4-4+()-2
5. Longest leaf (length) LS 955 IZ7-1 2 7-7 117-6 + 6:3 108-5 + 3-7
6. Longest leaf (max. width) 17-8 + 0-4 220) & be] 22-4+ 1-0 22:9 + 1-1
7. Bract length 20:7:4'0:5* 33-9 + 2-8 29-4 + 1-3 26-1 + 1:0
8. Bract width 4-8 + 0-3* 6-8 + 0-6 6:5 + 0-3 4:5+0-2
9. Labellum width 6:5 +-0-1* 8-1 + 0-4 8:2 + 0:3 6-3 + 0-3
10. Labellum length 5-9 + 0-2* 6-4+0-1 6-4 + 0-2 6:3+0-1
11. Trilobed character 1-0 + 0-0* 0-9+0-1 0-64 0-1 0-7 + 0-1
12. Lateral lobe notching 1-0 + 0-0* 1:0 + 0-0 0:7 + 0-1 0-8 +0-1

(all measurements in mm)
P=

DACTYLORHIZA INCARNATA SUBSP. OCHROLEUCA 301

height, 2. inflorescence length, 3. stem width near base, 4. stem width below the inflorescence,
5. length of longest leaf, 6. maximum width of longest leaf, 7. length of lowest floral bract,
8. maximum width of lowest bract, 9. maximum width of labellum, 10. maximum length of
labellum, 11. presence of distinct labellum sinuses (i.e. a measure of the degree of the trilobed
character), 12. presence of notching on lateral lobes of labellum. The last two characters listed (11,
12) were scored as: 0 = character absent; 0-5 = present, but not highly developed; 1 = very
noticeably developed whilst, in all cases, the flowers measured were those towards the base of the
inflorescence. In addition all plants were checked for the presence/absence of line or dot markings
on the labella and for leaf spotting.

Wherever possible at least ten plants were assessed, chosen in as random a manner as possible
so as to avoid selection bias. Unfortunately in Britain, where the population is now very small
(five plants in that year), random selection could not be applied; furthermore, flowers of two of
these could not be fully assessed due to partial damage, possibly by vermin. At all the localities the
plants formed homogenous populations and were not interspersed with other subspecies of D.
incarnata or different colour forms.

At the British locality and at Storsund, populations were measured by the author. For the
Estonian and German plants, vegetative characters were, under rigorous guide-lines, respectively
measured by V. Kuusk (Tartu) and H. R. Reinhard (Zurich), whilst the floral characters were
assessed by the author on material supplied on the same plants. For reference purposes, the same
data (characters 1-12) were also obtained for a population of typical D. incarnata subsp. incarnata
of normal flower colour (Hodbarrow, Cumbria, England).

All these data were subjected to Principal Components Analysis (PCA) using the Clustan 4
computer programme (Wishart 1987) with the results shown in Figure 1. By this technique, each
specimen measured is located in a multidimensional array, in which the number of dimensions 1s
equal to the number of characters measured, and where the most similar specimens are placed
closest together. The axes of greatest variation are extracted from the multidimensional space,
thereby simplifying it to a few (usually two or three) dimensions and allowing the location of each
specimen to be visualised. The respective mean and standard errors were calculated for each
population (Table 1).

+5

+3

+1

5 -3 -1 +1 +3 +5

FIGURE 1. PC1 : PC2 for individual plants of four populations of Dactylorhiza incarnata subsp. ochroleuca
© Chippenham, Cambs. (England), LJ Vidumiae (Estonia), O Murnau (Germany), ® Storsund, Gotland
(Sweden), compared to one of typical D. incarnata subsp. incarnata @ Hodbarrow, Cumbria (England).

302 Md. YeROLEY

DISCUSSION

In Fig. 1 the variance accounted for by the PC1 axis is 47-3%, that for the PC2 axis is 15-1%. In
each case, characters making a significant contribution towards the variance include plant height
and stem, inflorescence and labellum dimensions. By examining the plotted values in Fig. 1, it can
be seen that the British plants lie especially close to those from Germany and plants of both these
populations appreciably overlap with those from Sweden and Estonia. There is, therefore, no
overall separation between the four populations on the PC1 axis but all are well separated from the
control population of normal D. incarnata subsp. incarnata on this same axis. On the secondary
axis (PC2) there is again broad overlap between the four subsp. ochroleuca populations with a
slight shift and a greater spread for those from Sweden and Estonia. For the characters measured,
therefore, the British plants lie morphologically very close to those from the three European
populations.

The character means of all four populations (Table 1) are also closely comparable. Perhaps
surprisingly, the mean plant height was not as great as is often quoted. Higher values for labellum
width have also been recorded (Bateman & Denholm 1985) but the British and German plants
were similar in this respect. A fully trilobed labellum is also apparently not mandatory for every
plant within a population but, despite this, it was most frequently present. Distinctly notched
lateral lobes were also not evident in every case and it is possible that the presence of both of these
characters may vary between the flowers on a single plant. However, most of the characters used
to separate subsp. ochroleuca from closely-related taxa were found to occur in all of the plants
sampled at the four populations. All of them were yellow-flowered and none had line-marked
labella or spotted leaves.

On the island of Oland (Sweden), plants of similar habit and with flowers of a deep violet
coloration with unmarked, dark-centred labella, and of similar shape to subsp. ochroleuca occur,
very occasionally, as isolated individuals amongst normal subsp. ochroleuca (Lundqvist 1967).
The existence of a saturated reddish purple-based colour form in subsp. ochroleuca had been
anticipated by Nannfeldt (1944) who predicted a coloration comparable to D. sambucina (L.) So6
rather than that typical of subsp. incarnata. So far, these plants appear to be known from only two
localities to the east of Farjestaden and have been observed only very intermittently. According to
Lundqvist (pers. comm.) and the colour illustration in Mossberg & Lundqvist (1994), they appear
to be otherwise indistinguishable from normal subsp. ochroleuca. This suggests the presence of
colour dimorphism in subsp. ochroleuca in a similar manner to that within other Dactylorhiza taxa
(e.g. D. sambucina and D. sulphurea (Link) Franco) but with a very much smaller incidence of the
violet form.

SUMMARY

Most populations of this quite rare plant are inevitably rather small. The three sampled in
continental Europe contained only 20-50 plants whilst at Chippenham there were only five
available. Admittedly, and especially for the latter, this is a very small number on which to base
any conclusions and those given for the British plant can only be tentative. Nevertheless, for the
diagnostic characters measured, the Chippenham plants were found to be morphologically very
similar to those from the three European populations, and all of them could be defined by the
characters attributable to D. incarnata subsp. ochroleuca. However, additional confirmation via
molecular data would be useful in supporting this conclusion.

ACKOWLEDGMENTS

I am very grateful to H. R. Reinhard and V. Kuusk for their cooperation and help with the
morphological measurements of their local populations; to M. S. Porter for assistance in the field
and with translations; to L. Farrell and M. Wright for permission to examine the Chippenham
plants; to the curators of the Natural History Museum (BM), Torquay (TOR) and Vienna (W)
herbaria for allowing me to examine their specimens, and to D. M. Turner Ettlinger, R. Lidberg,
A. Lundqvist, W. Wiefelspiitz and H. Wittzell for relevant information.

DACTYLORHIZA INCARNATA SUBSP. OCHROLEUCA 303
REFERENCES

BATEMAN, R. M. & DENHOLM, I. (1983). Dactylorhiza incarnata (L.) So6 subsp. ochroleuca (Boll) P. F. Hunt
& Summerhayes. Watsonia 14: 410-411.

BATEMAN, R. M. & DENHOLM, I. (1985). A reappraisal of the British & Irish dactylorchids, 2. The diploid
marsh-orchids. Watsonia 15: 311-355.

BOLL, E. (1860). Flora von Mecklenburg. Archiv des Verein der Freunde der Naturgeschichte in Mecklenburg
14: 307.

FULLER, F. (1983). Die Gattungen Orchis und Dactylorhiza. Orchideen Mitteleuropas 3: 92-93. Die Neue
Brehm-Biicherei, Wittenberg-Lutherstadt.

HEGI, G. (1939). Illustrierte Flora von Mittel-Europa 2. Carl Hanser, Miinchen.

HESLOP-HARRISON, J. (1956). Some observations on Dactylorchis incarnata (L.) Vermin. in the British Isles.
Proceedings of the Linnean Society of London 166: 51-82.

Kuusk, V. (1991). Dactylorhiza incarnata subsp. ochroleuca Eestissa. Orkidealehti 4: 98-99.

LANDWEHR, J. (1977). Wilde orchideeén von Europa 1. Vereniging tot Behoud van Natuurmonumentem in
Nederland, Amsterdam

Luss, E. (1968). Vorkommen von Dactylorchis incarnata, var. ochroleuca im Kreisgebiet Neustrelitz - 1967.
Naturschutzarbeit in Mecklenburg 11(1): 33-34.

LUNDQVIST, A. (1967). Orchideen Olands. Svensk Botanisk Tidskrift 61: 463-493.

MOSSBERG, B. & LUNDQVIST, A. (1994). Olandska angsnycklar. Svensk Botanisk Tidskrift 88: 85.

NANNFELDT, J. A. (1944). Orchis strictifolia Opiz var. ochroleuca (Boll) Hyl., en fodrbissed svensk
kalkkarrvaxt. Svedburg 1944: 601-622.

PUGSLEY, H. W. (1939). Recent work on dactylorchids. Journal of botany, London 77: 50-56.

RAJCHEL, R. (1964). Orchis incarnata L. subsp. ochroleuca (Wiistnei) O. Schwarz in Poland. Fragmenta
Floristica et Geobotanica 10: 193-197.

STEPHENSON, T. & STEPHENSON, T. A. (1923). The British forms of Orchis incarnata. Journal of botany,
London 61: 273-278.

WIEFELSPUTZ, W. (1976). Dactylorhiza incarnata subsp. ochroleuca (Boll) Hunt & Summerh. auf Gotland?
Jahresbericht Naturwissenschaft Verein Wuppertal 29: 103-105.

WISHART, D. (1987). Clustan user manual, 4th ed. Computing Laboratory, University of St. Andrews.

WUSTNEI, K. (1854) in Koch, F., Bericht tiber die am 9 Juni unternommene Excursion nach der Umgegend
von Sternberg. Archiv des Verein der Freunde der Naturgeschichte in Mecklenburg 8: 92-97.

(Accepted January 2000)

at

Watsonia 23: 305-310 (2000) 305

Conservation of Britain’s biodiversity: Hieracitum cambricum
(Asteraceae), Welsh Hawkweed

ec. Ga RICH=

Dept. of Biodiversity and Systematic Biology, National Museum & Gallery, Cardiff CF 10 3NP

ABSTRACT

Hieracium cambricum is a rare endemic hawkweed known from only three sites in Wales. It is a distinct,
polycarpic perennial which flowers mainly May—June, and regenerates readily from wind-dispersed seed. It is
probably a calcicole. Surveys of its sites were carried out in 1998. No plants were found at Graig-fawr (v.c.
41) where it may be extinct due to a rock fall. On the Great Orme (v.c. 49), where it is well known, 75 plants
were found in a c. 25% sample of sites suggesting a population of c. 300 plants. Thirty-eight plants were
found at Creigiau Eglwyseg (v.c. 50), the first time it has been recorded since 1907. Although rare, it does not
seem to be significantly threatened, and both extant sites are Sites of Special Scientific Interest.

KEYWORDS: Endemic, rare species, Wales.

INTRODUCTION

Hieracium cambricum (Baker) F. J. Hanbury, Welsh Hawkweed, is a very rare endemic plant,
known from only three sites in Wales (Fig. 1.). Sell & West (1968) reported that it was frequent on
the Great Orme, Caernarvon (v.c. 49) but had not been seen in its other two sites at Treorchy,
Glamorgan (v.c. 41) and Llangollen, Denbighshire (v.c. 50) for over 60 years. Along with a
number of other rare hawkweeds, it has recently been included in the 3rd edition of the Vascular
Plant Red Data Book (Wigginton 1999), but nothing was known about its current status. Therefore
in 1998 surveys were carried out to find out how many plants there were and determine its needs
for conservation (Rich 1999); the purpose of this paper is to summarise the work.

H. cambricum was first collected on the Great Orme by J. Ward in 1867, who sent material to
F. J. Hanbury who thought it was a new species. In 1876, J. E. Griffith gave some specimens to
W. H. Painter, who forwarded them to J. G. Baker for identification (Griffith 1895). This prompted
Baker, who had known of the plant for many years but had not seen it, to describe it as H. caesium
Fr. var. cambricum Baker (1879). Hanbury (1894) later upgraded it to a species, and placed it with
H. vagense Ley in his group Vulgata Caulescentia. Linton (1905) transferred it to sect. Oreadea,
its current position today. Pugsley (1948) included it in sect. Oreadea series Argentea.

H. cambricum is a distinct hawkweed not closely allied to any other British species, and is
morphologically constant in cultivation over decades (e.g. Hanbury 1894; Marshall 1901). Its
outstanding characters are that it is virtually glabrous (even from the first growth), has a pale
green-glaucous hue, narrow, sharply toothed or laciniate leaves with dull green or purplish spots
(well-developed on the larger leaves, less so on the smaller ones), and strongly imbricated, obtuse
and relatively hairless phyllaries (Pugsley 1948). The main source of confusion is likely to be with
H. vagense, from which it can be distinguished by the more divided, subglabrous leaves, grey-
green phyllaries, and lighter yellow ligules (leaves deeply toothed or sub-pinnatifid, pilose
beneath, phyllaries dark green and ligules yellow in H. vagense).

“Address for correspondence: e-mail: Timothy.Rich@nmgw.ac.uk

306

Date

v.c. 41
29/6/1903
2/7/1906

v.c. 49
6/1867
1869

(pre 1889)
30/8/1876
1876
7/1879
7/1879
7/1880
1885

7/1886
7/1888
1888
29/5/1889
7/1890
7/1891
7/1691

25/6/1891
8/1891
6/1892

3/8/1892
a/1892
6/1892
24/6/1896

1898
1898

18/6/1901
21/6/1901

21/6/1901

9/7/1904
ZHI 90S

6/1905
26/6/1905
17/6/1907
18/6/1907
8/1908
Zi A92
1918
13/6/1919
20/5/1920
27/5/1920

t. Cy G, RICH

TABLE 1. LIST OF RECORDS OF HIERACIUM CAMBRICUM

Collector

H. J. Riddelsdell

Site

Craig Fawr, Treorchy

Sources

BM, CGE, Riddelsdell (1907a)

H. J. Riddelsdell Craig Fawr, Treorchy Riddelsdell (1907b)
S. H. Bickham Llandudno CGE
J. Ward Great Orme’s Head; cultivated, root collected K
1867
J. Ward Great Orme’s Head K
J. Cryer Great Orme NMW
J. Griffith Great Orme Griffith (1895)
W. H. Painter Great Orme’s Head K; possible syntype?
J. Gniffith Great Orme CGE; possible syntype?
J. Griffith Great Orme CGE, K
E. Starker Orme’s Head, cultivated Croydon A. Bennett OXF, K, NMW, OXF
J. Griffith Great Orme BM, NMW, OXF
J. Griffith Great Orme BM
E. Starker Orme’s Head, cultivated Croydon A. Bennett OXF
R. W. Scully Great Orme BM
H. T. Mennell Great Orme BM
J. Griffith Great Orme BM
S.H.Bickham — Inravines amongst the rocks, Great Orme’s CGE
Head
J. L. Williams Great Orme NMW
S. H. Bickham Great Orme BM
E. F. Linton Great Orme’s Head, cultivated LIV; Set of British Hieracia (1896,
etc.) no. 92.
A. H. Wolley-Dod Great and Little Orme BM
J. Griffith Great Orme BM, K
J. L. Williams Great Orme BM
E.S. Marshall Great Orme’s Head, cultivated NMW, OXF; Set of British
Hieracia (1896, etc.) no. 92.
E. F. Linton Great Orme’s Head, cultivated CGE, K, OXF

E.S. Marshall

Az Ley
A. Ley

E. F. & W. R.
Linton

A. Ley

J. Griffith

C. P. Hurst

a
; =
Soy
{cox}
aq
a
iq?)
La}

coy Pee
=
S
S
2.

ANAM D> >
C
S
iq)
en

Q0
eS
“<

Great Orme’s Head, cultivated

Great Orme’s Head
South of Great Orme

South of Great Orme

North flank of Great Orme

Highest corner of south west cliffs of Great
Orme .

Orme’s Head

Great Orme

North flank of Great Orme

North flank of Great Orme

Tyn-y-Coed, Great Orme

Great Orme

Great Orme, roots cultivated at Sedbury

Great Orme, roots cultivated at Sedbury

Great Orme

Great Orme

NMW Set of British Hieracia
(1896, etc.) no. 92.

LIV, K; BEC reports

K, OXF, NMW;; Ley (1901); Set
of British Hieracia (1896, etc.)
no 165. Baker (1879).

BM: Baker (1879).

NMW
BM, K, OXF; BEC reports

HIERACIUM CAMBRICUM 307
TABLE 1. CONTINUED

Date Collector Site Sources
31/7/1922 H.W. Pugsley Great Orme BM
8/6/1923 S. H. Bickham Great Orme, roots cultivated at Sedbury K, OXF
23/6/1928 T. J. Foggitt Great Orme BM
19/5/1946 J. A. Whellan Great Orme BM
12/6/1947 J. E. Lousley Cliffs near light house, Great Orme K
3/7/1949 __—*~P. D.. Sell SW side of Great Orme CGE
26/6/1952‘ P. D. Sell NE side of Great Orme CGE
11/7/1954 V. Gordan Great Orme, limestone cliffs NMW
pHoM955 PF. Yeo Near Gogarth, Great Orme, limestone rocks CGE

c. SH/763.831, c. 350’
8/6/1964 U.K. Duncan Great Orme CGE

25/7/1969 W. Ramsden Great Orme’s Head on the plateau above the LIV
marine drive near the light house

14/5/1988 J. Bevan Great Orme’s Head, west facing cliffs fide D. McCosh Hieracium
database

29/5/1996 -V. Jones Great Orme’s Head, south facing cliff fide D. McCosh Hieracium
database

11/9/1998 T.C.G.Rich& Great Orme, cliffs above marine drive from NMW

W. McCarthy SH/767.842 to SH/756.835; 75 plants

wc. 50

25/6/1901 A. Ley Cefn Fedw, Denbigh BM, CGE, K, OXF, NMW: Set of
British Hieracia (1896, etc.)
nowk65:

16/7/1903 A. Ley Limestone cliffs opposite Eglwyseg Church, BM

Cefn Fedw
25/6/1907 A. Ley Cefn Fedw, opposite the new church, ridges CGE

10/9/1998 T.C.G. Rich Creigiau Eglwyseg SJ/22.45; 38 plants NMW

DISTRIBUTION

HISTORICAL INFORMATION

Locality and habitat information was taken from the literature and herbarium sheets at BM, CGE,
K, LIV, OXF and NMW. Information from the Countryside Council for Wales database of Red
Data Book species was provided by R. A. Jones, and from the Hieracium database by D. McCosh.
The records in Table 1 were used to direct field surveys in 1998. It is quite possible that it occurs
in other currently unknown localities.

Graig-fawr, Treorchy (SS/92.95 to SS/92.96), v.c. 41

H.. cambricum has only been collected once from Graig-fawr by H. J. Riddelsdell in 1903. Graig-
fawr is a large cirque above Treorchy composed of Pennant Sandstone, most of which is very
acidic but with occasional calcareous outcrops associated with coal and other seams. This site is so
different to and so disjunct from the two known sites on Carboniferous Limestone in North Wales
(see below) that the record looks erroneous, were it not for the consistency of Riddelsdell’s
specimens and literature references.

Graig-fawr is a huge site which is difficult to search thoroughly. It has been investigated for
hawkweeds on at least four occasions since the 1950s (e.g. Wade et. al. 1994) but H. cambricum
has not been refound. Riddelsdell (1907b) reported it growing with H. argenteum Fr., which is
now confined to one gully at SS/927.956. The rocks in this gully are unstable, and potentially a
small colony of H. cambricum could have been lost to a rock fall (one significant fall was noted).
H. cambricum is probably extinct at this site.

Great Orme (SH/7.8), v.c. 49
H. cambricum was first found by S. H. Bickham in 1867, and is still well known to local botanists.

308 T. C. G. RICH

On 10 and 11 September 1998, a search was made by T.C.G.R. and W. McCarthy stopping at
regular intervals along Marine Drive, and surveying sections of c. 50-100 m of cliffs, depending
on ease of access. In ac. 25% sample of sites along 2:5 km of cliff, 75 plants were found scattered
between SH/767.842 and SH/756.835. This suggests that the total population on the Orme is
approximately 300 plants.

Creigiau Eglwyseg (Cefn Fedw), Llangollen (SJ/22.45), v.c. 50

H. cambricum was found at this site in 1901 by A. Ley, and again in 1903 and 1907. There are no
more recent records (one 1952 record held at the Biological Records Centre is a result of confusion
with the Great Orme).

On 10 September 1998, Creigiau Eglwyseg was visited specifically following up the localities
on the herbarium sheets ‘opposite the new church’ and ‘Limestone cliffs opposite Eglwyseg
Church’. Thirty-four plants were rediscovered scattered over c. 200 m of the major cliff between
SJ/221.457 and SJ/222.459, at an altitude of c. 400 m. On the ridge above, another four plants
were found at c. SJ/222.458, 430 m altitude. The plants were generally tucked into tiny crevices in
the limestone on + vertical rock faces, with no competition from other species. There could be
more plants elsewhere on this massive site.

Unconfirmed sites

One 1892 specimen by A. H. Wolley-Dod is labelled ‘Great and Little Orme’ (Table 1). There are
no other records from the Little Orme, and the one label appears to have been used for all material
collected from both sites by Wolley-Dod.

Specimens distributed as the Set of British Hieracia no. 165 bear the label ‘south of Great
Orme’. Whilst there are limestone outcrops to the south of the Orme (e.g. Bryn Maelgwyn
SH/796.805, The Vardre SH/782.795), H. cambricum has not been reported when investigated for
other hawkweeds. The locality could mean south of the Great Orme summit, which would relate to
localities known on the Orme itself.

A 1908 specimen collected by F. A. Lees from Tyn-y-Coed, Great Orme (c. SH/774.826; this
distinguishes it from Tyn-y-Coed, a farmstead in the valley south of the Orme at SH/792.797).
W. McCarthy lives in Tyn-y-Coed and has never seen the plant, but it is possible that it could have
occurred there in the past.

HABITATS

Hieracium cambricum is a plant of dry rock ledges and crevices, with little competing vegetation
in open, exposed, predominantly ungrazed situations. It appears to tolerate at least light shade.

The soil in the rock crevices are impossible to extract, but on the limestones at least are likely to
be immature rendzinas with high pH. Soil collected from the base of the cliff at Creigiau Eglwyseg
with seedlings immediately below mature plants was a black, fine, organic rendzina, pH 7:8
(measured with a pHep2 Hanna pocket-sized pH meter in a 50:50 mixture with distilled water). At
the possible historical site on Pennant Sandstone in Graig-fawr, the soils were rankers derived
from shale, pH 6:8.

The vegetation of the limestone rocks at the Great Orme and Creigiau Eglwyseg is very similar,
with typical limestone ledge communities with Asplenium ruta-muraria, Solidago virgaurea and
Festuca ovina (referable to the OV39 Asplenium trichomanes - Asplenium ruta-muraria
community of British Plant Communities; Rodwell et al. 1991 et seq.). On the Great Orme, a few
plants were also found with Helianthemum canum in ledge communities transitional to the N.V.C.
community CGI Festuca - Carlina grassland, and rarely in the lower MC4 Brassica oleracea
maritime cliff community. At both sites, vegetative plants were seen in grazed CG10 Festuca-
Agrostis-Thymus grassland. At Graig-fawr, the vegetation of the calcareous ledge where
H. cambricum may have occurred is different to the acidic Calluna/Vaccinium-dominated
vegetation of the remainder of the site but was not investigated.

The more accessible plants had their inflorescences eaten by feral goats or rabbits on the Great
Orme but the basal rosettes otherwise appeared tolerant of grazing. There was little sign of slug or
snail herbivory in the wild, though seedlings are susceptible in cultivation. Plants at Creigiau
Eglwyseg had severe leaf roll caused by aphids.

HIERACIUM CAMBRICUM 309

FIGURE 1. National distribution map of Hieracium cambricum. @ 1998, O pre-1950. Plotted using DMAPW
by Alan Morton.

LIFE CYCLE

Hieracium cambricum is a perennial which regenerates by seed. The main flowering period is June
and early July, but plants at both Creigiau Eglwyseg and Great Orme had a second flowering
period in the autumn (this often happens in Section Oreadea; Sell & West 1968). Plants at
Creigiau Eglwyseg flowering in 1998 had some dead flowering stems from 1997 indicating that
they are polycarpic. Hawkweeds are usually regarded as obligate apomicts but this has not been
tested specifically in H. cambricum. The seeds are wind-dispersed, and tests in windless conditions
show that seeds fall at a velocity of c. 150-200 cm per second, but can be kept afloat in the air by a
breeze. There was ample evidence of regeneration from seed at the Great Orme, and some possible
seedlings at Creigiau Eglwyseg. Seedlings have broader, nearly entire, sparsely hairy leaves.

CONSERVATION

Hieracium cambricum is not protected under Schedule 8 of the Wildlife and Countryside Act
1981, but is included in the Vascular Plant Red Data Book (Wigginton 1999), which should draw
attention to its rarity and be sufficient to ensure its survival. All three sites also have statutory

310 T.€,G.RICH

protection as S.S.S.I.s. At both Creigiau Eglwyseg and the Great Orme the populations grow on
cliffs, where there is effectively no requirement for management (grasslands around both sites are
grazed).

H. cambricum does not appear to be under any immediate threat, though rock climbing, scrub
invasion, spread of alien plants, rocks falls and rock safety work could potentially affect the plants
in the longer term at both extant sites. It is suggested that the populations should be monitored
every five years.

Seed has been collected from Creigiau Eglwyseg and the Great Orme and deposited in the
Millennium Seed Bank at the Royal Botanic Gardens, Wakehurst Place.

ACKNOWLEDGMENTS

This work was jointly funded by Countryside Council for Wales and National Museum & Gallery,
Cardiff as part of a joint project into conservation and ecology of critical species. The Royal
Botanic Gardens, Kew contributed towards the cost of collecting seeds for the Millennium Seed
Bank.

I would like to thank the Keepers of BM, CGE, K, LIV and OXF for access to specimens, Alan
Orange for identification of bryophytes, Peter Sell for discussion, David McCosh for records from
the B.S.B.I. Hieracium database and John Edmondson, George Hutchinson, Andy Jones, Serena
Marner, Wendy McCarthy, Alison Paul and Sarah Smith for their help.

REFERENCES

BAKER, J. G. (1879). On a variety of Hieracium caesium from the Great Ormes Head. Journal of botany 17:
360-362.

ELLIS, R. G. (1983). Flowering plants of Wales. National Museum of Wales, Cardiff.

GRIFFITH, J. E. (1895). The flora of Anglesey and Carnarvonshire. Nixon & Jarvis, Bangor.

HANBURY, F. J. (1894). Notes on British Hieracia. Journal of botany 32: 225-229.

HUTCHINSON, G. (1986). Graig Fawr and Graig Fach above the Rhondda 13 July [1985]. Cardiff Naturalist’s
Society Botanical Section Newsletter 5: 5.

Ley, A. (1901). H. cambricum F. J. Hanb. Report of the Botanical Exchange Club of the British Isles 2: 18.

LINTON, W. R. (1905). An account of the British Hieracia. West, Newman & Co., London.

MARSHALL, E. S. (1901). H. cambricum F. J. Hanb. Report of the Botanical Exchange Club of the British Isles
2:18.

PUGSLEY, H. W. (1948). A prodromus of the British Hieracia. Journal of the Linnean Society of London
(botany) 54: 1-356.

RIDDELSDELL, H. J. (1907a). A flora of Glamorganshire. Journal of botany, 1907 (Supplement): 1-88.

RIDDELSDELL, H. J. (1907b). H. argenteum Fr. Report of the Botanical Exchange Club of the British Isles 2:
23%

RODWELL, J. S. et al., eds. (1991 et seq.). British plant communities. Cambridge University Press, Cambridge.

SELL, P. D. & West, C. (1968). Hieracium L., in PERRING, F. H., ed. Critical supplement to the Atlas of the
British flora. B.S.B.1., London.

WADE, A. E., KAY, Q. O. N. & ELLIS, R. G. (1994). Flora of Glamorgan. H.M.S.O., London.

WIGGINTON, M. J, (1999). British Red Data Books. 1. Vascular plants. 3rd ed. Joint Nature Conservation
Committee, Peterborough.

(Accepted August 1999)

Watsonia 23: 311-316 (2000) Bit

Conservation of Britain’s biodiversity: Hieracium tavense
(Asteraceae), Black Mountain Hawkweed

T. C. G. RICH*
Dept. of Biodiversity and Systematic Biology, National Museum & Gallery, Cardiff CF 10 3NP

and
L. HOUSTON

2 West Grove, Montpelier, Bristol BS6 5LS

ABSTRACT

Hieracium tavense is a very rare endemic species, restricted to one waterfall in the Upper Tawe Valley,
Brecon (v.c. 42), Wales. It was first found in 1899 but appears to have only been seen a few times
subsequently. In 1998, a survey by climbing found 13 inaccessible plants on one cliff. It is a polycarpic
perennial which flowers in late July-August and reproduces by seed. Although rare and with a critically small
population, it is not threatened except potentially by a severe rock fall.

KEYWORDS: Endemic, rare species, Wales

INTRODUCTION

Hieracium tavense (Ley ex W. R. Linton) A. Ley, Black Mountain Hawkweed, is a very rare
endemic species, restricted to a ravine in the Upper Tawe Valley, Brecon (v.c. 42), Wales. Along
with a number of other rare hawkweeds, it has been included in the 3rd edition of the Vascular
Plant Red Data Book (Wigginton 1999), but very little was known about its detailed distribution or
ecology.

Hi. tavense was first found by Augustin Ley in the Upper Tawe Valley ‘above Callwen’ in 1899,
and was sent unnamed to the Botanical Exchange Club (Ley 1900). Ley took specimens into
cultivation, and later submitted them to the B.E.C. as H. protractum Fr. (Ley 1903), but F. J.
Hanbury suggested the general appearance of the plants was that of H. rigidum Hartmann rather
than H. protractum. It was then described as H. rigidum var. tavense Ley ex W. R. Linton (Linton
1905). Later at Linton’s suggestion, Ley raised it to species noting it was close to H. latobrigorum
(Zahn) Roffey (H. auratum Fr.) (Ley 1909). Zahn (1921-1923) later relegated it to a subspecies of
A. laevigatum Willd., but this view was not adopted by Pugsley (1948) who retained it as a
species. It is currently accepted as a species by P. D. Sell (pers. comm. 1998).

This species has only been seen a few times. Pugsley (1948) had not seen it himself and noted
that it had apparently not been collected since first found by Ley. B. A. Miles refound it in 1957
and visited it in 1959 and 1960, and M. Porter saw it in 1975 (Ellis 1983). In 1998, a survey was
carried out to establish its current status and determine its needs for conservation (Rich 1999). The
purpose of this paper is to summarise the work.

TAXONOMY

Hieracium tavense belongs to sect. Foliosa (Fr.) Arv.-Touv. This is a small section of ten
microspecies, mainly distributed in the north and west of Britain (Fig. 1). Pugsley (1948) gives a
good description, which appears to be largely based on cultivated material. Syntypes are located in
LIV (annotated by Sell & West in 1959) and at BM.

*Address for correspondence: e-mail: Timothy.Rich @nmgw.ac.uk

312 T. C. G. RICH AND L. HOUSTON

H. tavense 1s almost certainly a neoendemic which may have evolved from another member of
the sect., such as H. latobrigorum, since the last Ice Age. There appears to be nothing particularly
special about the waterfall on which it occurs (many similar waterfalls occur throughout the
Brecon Beacons), and it is not especially isolated from other species in the sect. (Fig. 1). Whether
it evolved in situ or 1s relict in this site from a wider distribution is unknown.

At least five Hieracium species grow in Nant-y-Llyn. H. tavense can be distinguished from H.
diaphanum Fr., H. scoticum F. Hanb. and H. sparsifolium Lindeb. by having numerous stem
leaves, the middle ones of which at least partly clasp the stem. It can be distinguished from the
other, as yet unnamed, species of sect. Foliosa with which it grows at the waterfall by the linear-
lanceolate (not ovate) upper stem leaves (this other species is currently under investigation).
Another 23 Hieracium species are listed from the 10-km square SN/8.2 in Sell & West (1968), and
more occur to the south in SN/8.1.

DISTRIBUTION

Locality and habitat information were extracted from the literature and from BM, CGE, LIV.
OXF and NMW. Information from the Countryside Council for Wales database of Red Data Book
species was provided by R. A. Jones, and from the Hieracium database by D. McCosh. The
historical records indicate it has been found in one specific locality only, i.e. ‘waterfall at Nant-y-
Llyn’ (Table 1), though Ley cited it in several different forms. He also noted that it was ‘not yet
known from any other station other than those in the Upper Tawe Glen’ (Ley 1909) suggesting it
could occur elsewhere in this area, but no other stations are known.

Nant-y-Llyn is a ravine on the east side of the Black Mountain with a series of rocks and
waterfalls. In 1998, 13 plants of H. tavense were found confined to an unstable cliff c. 20 m high
on the north-facing side of a small cirque developed around the upper of the two larger waterfalls
at c. 510 m altitude (grid reference SN/844.207). The plants could only be reached safely by
climbing with ropes. One stem of one plant was removed as a voucher (NMW) and its identity
confirmed by P. D. Sell.

The rocks of the waterfall are composed of the Brownstones of the Lower Old Red Sandstone of
Devonian age (Geological Survey of Great Britain 1979; Barclay et al. 1988). The Brownstones
are red-brown and purple sandstones interbedded with red mudstones and siltstones, with
occasional intraformational conglomerates and less frequent calcrete nodules. They form beds c.
400 m thick, and outcrop over much of the Brecon Beacons. At the waterfall, there are a series of
hard strata c. 20-50 cm thick, interbedded with softer silts, resulting in a series of unstable and
overhanging ledges on which H. tavense grows. Lower down Nant-y-Llyn, green sandstone of the
Senni beds also outcrops to form a waterfall.

TABLE 1. LIST OF HIERACIUM TAVENSE RECORDS

Date Collector Site Source
26/7/1899 A. Ley Tawe gorge above Cellwen BM, syntype (Ley 1900)
3/8/1899 A. Ley Waterfall at Nant y Llyn near Cellwen BM (ex. herb. Hanbury)
3/8/1899 A. Ley Upper Tawe Valley BM, LIV; syntype (Ley 1900)
3/8/1899 A. Ley Head of the Tawe BM, CGE, OXF (Ley 1900)
3/8/1899 A. Ley Baggards Glen, Nant-y-Llyn, Upper Tawe BM, CGE
30/7/1902 A. Ley Upper Tawe, cultivated BM, NMW, OXF (Ley 1903)
18/7/1906 A. Ley Upper Tawe Glen, cultivated BM, CGE, LIV, NMW, OXF
1910 A. Ley Upper Tawe Glen, cultivated BM, CGE, OXF (Ley 1911)
26/7/1957 B.A. Miles rocks by the waterfall, 1650’, Nant-y-Llyn, CGE

Upper Tawe
1959 B. A. Miles Nant-y-Llyn, Upper Tawe; | plant Note on specimen in CGE
1960 B. A. Miles Nant-y-Llyn, Upper Tawe; 3 plants Note on specimen in CGE
1975 M. Porter Upper Tawe Ellis (1983)
28/8/1998 T.Rich& Nant-y-Llyn; 13 plants NMW

L. Houston

HIERACIUM TAVENSE B43

te]

FIGURE 1. Distribution of Hieracium tavense (@) and Hieracium Section Foliosa (=).

ECOLOGY

LIFE CYCLE

Hieracium tavense is a polycarpic perennial which reproduces by seed. In 1998, the mean number
of stems per plant was 2-9 (range 1-7) (Table 2). Nine out of the 13 plants (69%) were flowering,
and most (76%) of the stems had flowers. The mean number of capitula per inflorescence was 3-2,
which is smaller than the numbers observed on two specimens cultivated by Ley in NMW (eleven
and 14 capitula respectively) suggesting sub-optimal growth conditions in the wild compared to
cultivation. Some plants rooted in shallow crevices were vegetative, and most plants had some
aborted capitula within each inflorescence. Many plants also had dead stems from 1997.

Plants were just beginning to flower on 14 August and had virtually all finished flowering by 28
August 1998. Herbarium specimens have flowers from mid-July to the end of August. It is
assumed that H. tavense is an obligate apomict like other Hieracium species but this has not been
tested. Flowers seen on 14 August had not fruited by 28 August in the wild, but did so rapidly
when taken into cultivation. Seed ripening is thus estimated to take 2-3 weeks. The mean number
of seeds produced by each flowering plant was 225 (range 35-1015), and the total seed production
of the population was estimated as c. 2030 seeds.

314 T. C. G. RICH AND L. HOUSTON

TABLE 2. CHARACTERISTICS OF HIERACIUM TAVENSE AT NANT-Y-LLYN
ON 28 AUGUST 1998

No. live No. stems Number of capitula No. dead stems

Plant stems flowering ineachinflorescence from 1997 Notes
1 3 0 - 4
2 2 0 - 5 + 3 possible seedlings
3 7 a 3,3,4,4,4,5,6 y Huge clump
4 1 0 - 0 Stems growing downwards
5 4 3 22 6
6 2 2 ee) 6 Four capitula picked + soil collected
7 4 1 1 0 Under an overhang
8 1 0 - 0 Very small, + no soil, hanging out of cliff
9 4 ) 1,3,aborted 3 Voucher specimen collected 14/8/1998
10 1 1 8 0 Two capitula collected
14 1 1 1 0 Growing sideways under overhang
12 q 1 2 9 Growing well
13 1 1 Z 0 Plant dangling down, two capitula
collected
Mean 2-9 1-53 3-2 3:1

The seeds are wind-dispersed. Tests in windless conditions show that seeds fall at a velocity of
c. 40-50 cm per second, but are easily kept afloat in the air by a gentle breeze (e.g. gentle
blowing). Although the seeds have the potential for dispersal, the paucity of available habitat in the
Upper Tawe Valley, restricted calcareous outcrops and low seed production suggest that the
chances of H. tavense spreading are limited, though some possible reproduction by seed within the
site was noted during the survey (Table 2). All four seedlings which germinated in cultivation had
one of the cotyledons split into two, resulting in an apparent appearance of three cotyledons.

ASSOCIATED SPECIES AND SOILS

The species associated with H. tavense on the waterfall are listed in Table 3. An attempt was made
to relate the vegetation type on the rock face to the National Vegetation Classification (Rodwell et
al.1991 et seq.), but the site is somewhat heterogeneous. Most of the waterfall face appears to be
H12 Calluna vulgaris - Vaccinium myrtillus heath vegetation. H. tavense occurred on the more
base-rich ledges, often without any other species, and it has not been possible to ascribe it to a
clear N.V.C. type.

Plants were rooted in immature rankers on the ledges or sometimes directly into rock crevices
with no apparent soil. The soil from the base of one plant was composed of c. 70% large fragments
of Brownstone, reddish-brown mineral material and fine organic matter, pH 7-1 (measured with a
pHep2 Hanna pocket-sized pH meter in a 50:50 mixture with distilled water).

CONSERVATION

Although the population size is critically small, the only significant threat to survival of the species
is a large rock fall which could wipe. out the entire population. It is under no threat from collecting
due to the very difficult climbing conditions and requirement for roped access. Although the area
around the waterfall is intensively grazed, the cliff is inaccessible to sheep (H. tavense, like most
hawkweeds, is likely to be sensitive to grazing). No site management is currently required, and the
only potential management in the future might be control of Sorbus aucuparia trees to minimise
shade.

Seeds from eight flowering heads were sent to the Millennium Seed Bank at the Royal Botanic
Gardens, Wakehurst Place for long-term storage. Attempts are also being made to grow the plant
in cultivation.

Hieracium tavense is not protected under Schedule 8 of the Wildlife and Countryside Act 1981,
but is included in the Vascular Plant Red Data Book (Wigginton 1999), which should draw

HIERACIUM TAVENSE 315

TABLE 3. SPECIES ASSOCIATED WITH HIERACIUM TAVENSE, NANT-Y-LLYN,
28 AUGUST 1998 WITH ‘DAFOR’ COVER RATINGS

Agrostis capillaris R Oreopteris limbosperma R
Anthoxanthum odoratum R Oxalis acetosella O
Asplenium viride R Polypodium vulgare s.l. R
Athyrium filix-femina O Solidago virgaurea O
Calluna vulgaris F Sorbus aucuparia R
Carex pilulifera R Succisa pratensis O
Deschampsia flexuosa A Thymus polytrichus O
Digitalis purpurea O Vaccinium myrtillus O
Dryopteris dilatata O Valeriana officinalis R
Dryopteris filix-mas R Veronica officinalis R
Epilobium brunnescens O Viola riviniana R
Epilobium montanum R Bryophytes

Euphrasia sp. R Brachythecium plumosum O
Festuca ovina R Amphidium mougeotii F
Festuca rubra R Blepharostoma tricophyllum R
Galium saxatile R Ctenidium molluscum F
Hieracium sp. R Diplophyllum albicans O
Hieracium tavense O Dicranum scoparium O
Hypericum pulchrum R Fissidens taxifolius R

attention to its rarity and be sufficient to ensure its survival. The upper part of Nant-y-Llyn is
included within the Mynydd Du S.S.S.I, and is also part of the Brecon Beacons National Park.
These designations give a significant degree of protection to the site.

Thus H. tavense has no specific conservation requirements despite being very rare. The
population should be surveyed every five years to monitor the status quo.

ACKNOWLEDGMENTS

This work was jointly funded by Countryside Council for Wales and National Museum and
Gallery, Cardiff as part of a joint collaborative project into conservation and ecology of critical
species. The Royal Botanic Gardens, Kew, contributed towards the cost of collecting seeds for the
Millennium Seed Bank.

We would like to thank the Keepers of BM, CGE, LIV and OXF for access to specimens, Alan
Orange for identification of bryophytes, Peter Sell for discussion, David McCosh for Hieracium
identifications and records from the B.S.B.I. Hieracium database, John Edmondson, George
Hutchinson, Serena Marner, Alison Paul and Sarah Smith for assistance, and Andy Jones, Jamie
Bevan and Graham Motley of C.C.W. for their help. The base map was plotted using DMAPW by
Alan Morton.

REFERENCES

BARCLAY, W. J., TAYLOR, K. & THOMAS, C. P. (1988). Geology of the South Wales Coalfield, Part V, the
country around Merthyr Tydfil. 3rd ed. H.M.S.O., London.

ELLIS, R. G. (1983). Flowering plants of Wales. National Museum of Wales, Cardiff.

GEOLOGICAL SURVEY OF GREAT BRITAIN (England and Wales) (1979). Sheet 231, Merthyr Tydfil. Solid and
drift edition. 1:50,000. Ordnance Survey, Southampton.

KENT, D. H. & ALLEN, D. E. (1984). British and Irish herbaria, 2nd ed. B.S.B.I., London.

Ley, A. (1900). H.-. Report of the Botanical Exchange Club of the British Isles 1: 639.

LEY, A. (1903). H. protractum Fr. Report of the Botanical Exchange Club of the British Isles 2: 55.

Ley, A. (1909). Brecon and West Yorkshire hawkweeds. Journal of botany 47: 8-16 and 47-55.

Ley, A. (1911). H. tavense Ley. Report of the Botanical Exchange Club of the British Isles 3: 573.

LINTON, W. R. (1905). An account of the British Hieracia. West, Newman & Co., London.

316 T. C. G. RICH AND L. HOUSTON

PUGSLEY, H. W. (1948). A prodromus of the British Hieracia. Journal of the Linnean Society of London
(botany) 54: 1-356.

RicH, T. C. G. (1999). Distribution and conservation of Hieracium tavense, Black Mountain Hawkweed.
National Museums & Galleries of Wales, Cardiff.

RODWELL, J. S. et al., eds. (1991 et seq.). British plant communities. Cambridge University Press, Cambridge.

SELL, P. D. & WEST, C. (1968). Hieracium L., in PERRING, F. H., ed. Critical supplement to the Atlas of the
British flora. B.S.B.1I. London.

WIGGINTON, M. J. (1999). British Red Data Books. 1. Vascular plants, 3rd ed. Joint Nature Conservation
Committee, Peterborough.

ZAHN, K. (1921-1923). Hieracium, in ENGLER, H. G. A., ed. Das Pflanzenreich IV, 280: 1-1705.

(Accepted July 1999)

Watsonia 23: 317—322 (2000) BF

Conservation of Britain’s biodiversity: Rubus trelleckensis
(Rosaceae), Trelleck Bramble

R. D. RANDALL and T. C. G. RICH

Dept. of Biodiversity and Systematic Biology, National Museum & Gallery, Cardiff CF 10 3NP

ABSTRACT

Rubus trelleckensis is a rare endemic bramble found only in one 10-km square near Trelleck, Gwent (v.c. 35),
Wales. It is a perennial which flowers from mid-July to mid-August, and sets abundant fruit but probably does
not spread vegetatively. A field survey was carried out in 1998. Five small populations were found, all in
locations which were probably at one time either open heath or open Birch-Oak woodland but are now either
conifer plantation, or conifers mixed with broad-leaved trees. Plants were most frequent on acidic podzols in
sunny but sheltered spots on level ground. The main threats to its survival are change of land-use from forestry
or changes in forestry operations. Seeds have been deposited in the Millennium Seed Bank at the Royal
Botanic Gardens, Wakehurst Place.

KEYWORDsS: Endemic, rare species, Wales.

INTRODUCTION

Rubus trelleckensis Edees & Newton, Trelleck Bramble, is a rare Welsh endemic species with a
very restricted distribution centred on Beacon Hill near Trelleck in Gwent (v.c. 35, Monmouth).
Along with a number of other rare brambles, it has been included in the 3rd edition of the Vascular
Plant Red Data Book (Wigginton 1999), but as yet little is known about its detailed distribution or
ecology. In 1998 the available information was collated, combined with a field survey, to establish
its current status and determine its needs for conservation. A summary of the data are given below;
full details are given in Randall & Rich (1999).

TAXONOMIC HISTORY

R. trelleckensis, as we now know it, has had a long and somewhat confusing history. It is a
distinctive species but would probably not have received a name if it had not been distributed in
the Set of British Rubi and later identified, erroneously, with a number of European species.

It was first collected by A. Ley in 1885 from plantations on Beacon Hill near Trelleck,
Monmouthshire. Specimens were sent to W. O. Focke for naming, who commented ‘R. virescens
G. Braun, var. glandulosa. The typical plant has no glandular bristles on the flower stalks’. In
1886, Ley again visited Beacon Hill and collected further specimens which he submitted to the
Botanical Exchange Club (B.E.C.) as ‘R. Borreri Bell-Salt.’ (Linton 1887). C. C. Babington
commented that it was ‘Sprengelii probably’ and Focke agreed.

In 1888 Ley again visited Beacon Hill and this time selected two plants for submission to the B.
E.C. (Ley 1889). He suggested ‘R. Borreri, Bell-Salt. variety ?’ for one of them but Babington and
Focke were unable to place it (this taxon has received no name). The other plant was submitted as
‘R. virescens G. Braun’ on the strength of the name supplied by Dr. Focke for Ley’s 1885
specimen. Ley could not guarantee that he had collected the same plant as previously and, indeed,
Dr. Focke’s comments this time were less encouraging: ‘Leaves exactly as in virescens, but the
other characters do not agree. I can give no name’. Ley reported that both plants grew with R.
sprengelii in the plantation and that the ‘R. sprengelii var.’ partly showed the suberect habit of R.
virescens. He was later of the opinion that the two species hybridised there.

In 1893, Ley again visited Beacon Hill, accompanied by W. A. Shoolbred on 4 July and by E. F.
Linton on 3 August. Material from these visits was distributed as no. 60 in the Set of British Rubi,
labelled R. myricae Focke var. virescens G. Braun, forma glanduligera.

318 R. D. RANDALL-AND TC: G. RICH

When Focke visited Trelleck and saw living material he withdrew the name he had originally
suggested (there is a specimen at BM (Barton & Riddelsdell no. 4107) collected on 31 July 1894
by Ley, Focke and W. H. Purchas; a typed note states that ‘Part of this gathering is in Focke’s
scaber packet (159), without name or comment’ ).

As the distinctive plant was not clearly identifiable with any known species it was duly
described as new as R. orthocladus Ley (Ley 1896). Rogers (1900) pointed out that the name was
already in use, viz. R. orthocladus Boulay (1869), and that the Set of British Rubi No. 60 contained
representative material of Ley’s plant. At that time it was thought that M. Gravet had discovered
Ley’s plant at Louette St. Pierre in Belgium, and it was listed as R. orthocladus Ley in Sudre
(1908). Later he revised his views (Sudre 1910) and reduced R. orthocladus Ley to a synonym of
R. bracteosus Weihe ap. Le}. & Court. (Lejeune & Courtois 1831), an earlier name that had been
applied to the Belgian plant.

Watson (1932), at the time referee for the genus, associated a number of forms from around the
country under R. egregius var. effeminatus Focke, a name that had originally been given to a plant
from Oxfordshire. Included by Watson under this name was material collected by Ley from
Beacon Hill Woods on 4 July 1893, labelled R. orthoclados and specimens from an open heath on
Beacon Hill on 23 July 1897, labelled ‘R. orthoclados var.’.

At the time of the publication of Rubi of Great Britain and Ireland (Watson 1958) and Flore
Générale de Belgique (Legrain 1958) it was listed as R. bracteosus Weihe ap. Lej. & Court., which
was assumed to be present in both Britain and Belgium. Some specimens in herbaria were later
labelled R. euchloos Focke under which name it occurs in Flora Europaea (Heslop-Harrison 1968)
on Watson’s authority.

E. S. Edees and A. Newton assessed the validity of all the European species which Watson had
claimed to be present in Britain, and decided that the name R. bracteosus could not be applied to
the Trelleck plant and gave it a new name, R. trelleckensis (Edees & Newton 1978). It fits rather
uncomfortably into its current position in Subsection Rubus but would be equally ill-fitting in
Series Silvatici, or Series Mucronati.

Summary of synonymy

R. myricae var. virescens sensu Focke pro parte, non R. virescens G. Braun in Focke
R. orthocladus Ley, non R. orthocladus Boulay

R. euchloos auct., non Focke in Ascherson & Graebner

R. bracteosus auct., non Weihe ex Lej. & Court.

R. egregius var. effeminatus sensu W. C. R. Watson pro parte, non Focke

IDENTIFICATION

There are a number of species which resemble R. trelleckensis, a guide to which 1s given in Rich &
Randall (1999) (see also Edees & Newton 1988). Its distinctive characters are the broad-based
declining prickles, narrow green long-pointed leaflets, sub-racemose panicle, frequent unequal
stalked glands and acicles on peduncles, white flowers, long-pointed green clasping sepals and
carpels at first hairy. Hybrids with R. pyramidalis, R. scissus and R. sprengelii have been
suggested but require more detailed study.

DISTRIBUTION

Locality and habitat information was abstracted from the literature, and from herbarium sheets at
the Natural History Museum (BM) and the National Museum of Wales (NMW), which between
them include much Botanical Exchange Club material and the herbaria of W. M. Rogers, H. J.
Riddelsdell and E. S. Edees. The records indicated that R. trelleckensis is very rare, being confined
to woods and heathy ground between Trelleck and Tintern in Gwent, and these areas were
concentrated on for the field survey.

The survey was carried out between mid-July and early September 1998, the best time being
mid-July to early August when plants were flowering. About 80+ plants (i.e. distinct clumps) were
found in five sites in one 10-km square, SO/5.0, all in v.c. 35 (Figure 1).

RUBUS TRELLECKENSIS 319

Trelleck @7

05

00
50 55

Figure 1. Location of sites in western part of SO/5.0 10-km square searched for Rubus trelleckensis in 1998.
@ plants found. O no plants found. The inset shows the location of the map in Wales.

Site 1. Bargain Wood (SO/520.033)
This site was first found by A. Orange in 1997. Seven plants were found scattered along about 50
metres of a footpath in a recently replanted conifer plantation.

Site 2. Ninewells Wood (SO/509.037)

Two plants were discovered in open heathy birch scrub. This site is immediately south of Trelleck
Bog, an old site (see also below).

Site 3. Beacon Hill (SO/512.054)

This is the type locality for R. trelleckensis, where Ley (1896) described it (R. orthoclados) as
‘occupying a large area of woodland (some three square miles) on Beacon Hill, Monmouthshire’.
Beacon Hill is now largely conifer plantation, some parts of which were being felled in rotation
resulting in vigorous regeneration of the native vegetation, including brambles. About 50 plants
were seen scattered in various forestry compartments across the hill.

320 R. D-RANDALE AND* EC; GF RICH

Site 4. Beacon Hill - North Slope (SO/513.057)

In light of Ley’s comments about the relative abundance of R. trelleckensis on Beacon Hill, a brief
visit was made to the northern slopes. Three plants were found by a forestry track in a small patch
where Calluna was frequent.

Site 5. Vicar’s Allotment (SO/512.060)
A thriving population of over 20 plants was found along a bridleway through a recently felled
conifer plantation.

Sites searched without success

A number of other sites in the area were also visited (Fig. 1). There were historical records for
Trelleck Bog (now known as Cleddon Bog) but a check of the developing birch scrub on the south
and west edges of the bog failed to locate any plants. Church Hill Common, Penallt (SO/52.10)
was also searched as the soil maps (Soil Survey of England and Wales 1983) indicated that a
podzol similar to that exhibited on Beacon Hill and the Vicar’s Allotment was present; R.
trelleckensis has not been reported from this site and none was found.

Erroneous or unconfirmed records

The record for ST/2.8 in Edees & Newton (1988) is based on a specimen from Pennsylvania near
Castleton (v.c. 35) collected by A. E. Wade in 1941 (NMW). The specimen is not R. trelleckensis
but belongs to Series Anisacanthi and is probably R. hibernicus (Rogers) Rogers.

A plant from Mitcheldean Meend (v.c. 34), distributed by Ley through the B.E.C. as ‘R.
orthocladus Ley var.’ was considered by Sudre to be R. orthocladus x R. gratus and subsequently
described as a new species or hybrid R. dobuniensis Sudre & Ley (Sudre 1908). It is probably a
hybrid of R. gratus with a species other than R. trelleckensis, perhaps R. melanocladus (Sudre)
Riddelsd. which is abundant at Mitcheldean.

No specimens have been traced to support the unlikely records from Bewdley (v.c. 37) and
Staffordshire (v.c. 39) (Watson 1958), or Dol-y-bont, Llanfihangel and Llyfnant Valley (v.c. 46)
(Rogers 1908).

LIFE CYCLE

R. trelleckensis is, like all brambles, a polycarpic perennial. It is more-or-less evergreen, with
leaves often persisting until the new year. Panicles arise from buds in the axils of the previous
year’s leaves (as in other species) but vegetative branches were not seen and it is likely that
individual stems and branches die back after fruiting and are therefore biennial. The flowering
time is mid-July to mid-August. Fruits were abundant at the end of August 1998, and most
appeared to ripen about the same time. Dispersal of seed by birds and mammals probably occurs,
as in other brambles. The fruits are quite sweet.

TABLE 1. PROBABLE NATIONAL VEGETATION CLASSIFICATION TYPES AT SITES OF

RUBUS TRELLECKENSIS
Site National Vegetation Classification types
Bargain Wood H8e Calluna - Ulex gallii heath, Vaccinium sub-community developing in 5-year

old cleared and replanted conifer plantation

Beacon Hill U2 Deschampsia flexuosa grassland in recently cleared conifer plantation, H8e
Calluna - Ulex gallii heath, Vaccinium sub-community in replanted older
conifer plantation

Beacon Hill - north slope W416 Quercus - Betula - Deschampsia woodland on disturbed edge of possibly
original W10 Quercus - Pteridium - Rubus community.

Ninewells Wood W16 Quercus - Betula - Deschampsia woodland, replanted with conifers

Vicar’s Allotment H8e Calluna - Ulex gallii heath, Vaccinium sub-community developing in 5-year
old cleared and replanted conifer plantation

RUBUS TRELLECKENSIS 32M

Although many Rubus species spread vegetatively by adventitious roots from the ends of the
arching stems this does not appear to occur very often with R. trelleckensis. Most bushes observed
appear to be groups of stems from a single rootstock.

Population sizes appear to depend on the amount of competition from bracken and other species,
on the amount of shade, and on soil type and drainage. Plants were most frequent on the podzols of
Beacon Hill and the Vicar’s Allotment, with the best-growing plants in sunny but sheltered spots
on level ground where there was good moisture retention.

At Beacon Hill, the state of the population was closely linked to the forestry management. In
recently cleared areas there were scattered patches, mostly stunted due to the dry exposed
conditions. In an area replanted with young trees, there were well-grown plants intermixed with
other brambles, bracken and gorse. Where young trees were well established and beginning to
crowd out the associated underscrub, R. trelleckensis and other brambles were confined to the
margins of the plantation. Very few plants were found in the dense, well-established 3-4 m tall
plantation except in gaps.

HABITATS, VEGETATION AND SOILS

All sites seen in 1998 are either conifer plantation, or conifers mixed with broad-leaved trees. An
attempt was made to relate the vegetation types to the National Vegetation Classification (Rodwell
1991 et seq.), but this is difficult for recently disturbed conifer plantations (Table 1). The original
vegetation types of these sites may have been the W16 Quercus - Betula - Deschampsia woodland
type (possibly extending into other related woodlands such as the W10 Quercus - Pteridium -
Rubus community), and the H8 Calluna - Ulex gallii heath. Ley’s specimens and his comments on
its distribution suggest that R. trelleckensis thrived in woodland.

R. trelleckensis is typically distributed on the tops and upper slopes of hills where a peaty soil
has developed in situ over quartz conglomerate base rock. It was not found on deeper, more
mineral-rich soils or the carboniferous limestone areas of the Wye Valley. Soils collected from the
rooting zones in August 1998 are described briefly in Table 2. They are all heathy acidic podzols
pH 3-4-4-0, probably derived from the local quartzite rock. At Beacon Hill and Vicar’s Allotment
they are classified as the 631le Goldstone Association, a humo-ferric podzol, developed over quartz
conglomerate of the Old Red Sandstone, and in the Ninewells Wood - Bargain Wood area as the
541c Eardiston 1 Association, a typical brown earth, developed over the Tintern Sandstone Group
of the Upper Old Red Sandstone (Soil Survey of England and Wales 1983; Geological Survey of
England and Wales 1974).

TABLE 2. SOIL TYPES AT SITES OF RUBUS TRELLECKENSIS

Site Soils

Bargain Wood A fine grey-brown heathy soil with lots of organic matter, pH 4-0
Beacon Hill A fine brown heathy soil with organic matter, pH 4-0

Ninewells Wood A gritty brown organic soil, pH 3-6

Vicar’s Allotment A blackish organic soil with large quartz sand grains, pH 3-4

CONSERVATION REQUIREMENTS

Rubus trelleckensis is a rare plant. It is not protected under Schedule 8 of the Wildlife and
Countryside Act 1981, but is listed in the Vascular Plant Red Data Book (Wigginton 1999). None
of the sites found in 1998 are scheduled as S.S.S.I.s. Given that it is only known in five sites, most
with tiny populations, it would be worth trying to protect the colonies at some sites. Consideration
should be given to designating Beacon Hill, the type locality with the largest population, as a Site
of Special Scientific Interest for this rare endemic species.

The main threats to its survival appear to be changes in the existing forestry regime (e.g. weed
killing, longer periods of deep shade, fertiliser application to soils, etc.). The most favourable
forestry management is currently thought to be a regime similar to that exhibited at Beacon Hill,
where trees have been cropped in rotation with adjacent areas being felled several years apart. This

322 R. D. RANDALL AND T. C. G. RICH

allows the possibility of seed from a regenerating section being bird-sown into the most recently
felled area. The amount of regeneration on Beacon Hill shows that the seed-bank there is probably
sufficient to ensure R. trelleckensis will survive as long as current forestry management continues.

Blackberries from plants at Bargain Wood and Beacon Hill have been deposited at the
Millennium Seed Bank at the Royal Botanic Gardens, Wakehurst Place. One plant is held in
cultivation by A. Orange.

ACKNOWLEDGMENTS

This work was jointly funded by Countryside Council for Wales and National Museum and
Gallery, Cardiff as part of a joint project into conservation and ecology of critical species. The
Royal Botanic Gardens, Kew, contributed towards collecting blackberries for the Millennium Seed
Bank. We would like to thank Andy Jones and Lindi Rich of C.C.W. for their help, and Alan
Orange and Jessica Carey for help with field work. We would like to thank the Keeper of BM for
access to the herbarium.

REFERENCES

EXSICCATAE

BOULAY, J. N. (1864-1869). Ronces vosgiennes. Fasc. 1-5, nos. 1-152. Rambervillers. (Fasc. 5, no. 142,
1869, R. orthocladus Boulay).

LINTON, E. F. & W. R., MURRAY, R. P., ROGERS, W. M. & LEY, A. (1892-1897). Set of British Rubi. Fasc.
1+4, nos. 1-106 and supplement, nos. 106A—134. (no. 60, 1894, R. myricae Focke var. virescens G.
Braun, forma glanduligera).

SUDRE, H. (1903-1917). Batotheca Europaea. Fasc. 1-15, nos. 1-750. (no. 218, R. orthocladoides Sudre).

SUDRE, H. (1908-1912). Rubi rari vel minus cogniti exsiccati. Fasc. 1-5, nos. 1-140. Albi. (Fasc. 1, no. 7,
1908).

LITERATURE

EDEES, E. S. & NEWTON, A. (1978). Amendments to the British Rubus List. Watsonia 12: 133-136.

EDEES, E. S. & NEWTON, A. (1988). Brambles of the British Isles. Ray Society, London.

FOCKE, W. O. (1902). Rubus L., in ASCHERSON, P. & GRAEBNER, P. Synopsis der mitteleuropdischen Flora 6
(1). Leipzig.

GEOLOGICAL SURVEY OF ENGLAND AND WALES (1974). Sheet 233, Monmouth. Solid and drift edition.
1:50,000. Ordnance Survey, Southampton.

HESLOP-HARRISON, Y. (1968). Rubus L., in TUTIN, T. G. ef al., eds. Flora Europaea 2: 12. Cambridge
University Press, Cambridge.

LEGRAIN, J. (1958). Rubus L., in ROBYNS, W., ed. Flore Générale de Belgique 3: 49. Bruxelles.

LEJEUNE, A. L. S. & COURTOIS, R. J. (1831). Compendium Florae Belgicae, etc., 2: 162. Bruxelles.

Ley, A. (1889). R. Borreri Bell-Salt variety? and R. virescens, G. Braun. Report of the Botanical Exchange
Club of the British Isles 1: 207.

Ley, A. (1896). Herefordshire Rubi. Journal of botany 34: 159-160.

LINTON, E. F. (1887). R. Borreri Bell-Salt. Report of the Botanical Exchange Club of the British Isles 1: 148.

RANDALL, R. D. & RICH, T. C. G. (1999). Distribution and conservation of Rubus trelleckensis, Trelleck
Bramble. National Museums & Galleries of Wales, Cardiff.

RODWELL, J. S., ed. (1991 et seg.). British plant communities. Cambridge University Press, Cambridge.

ROGERS, W. M. (1900). Handbook of British Rubi. London.

ROGERS, W. M. (1908). R. orthoclados, Ley. Report of the Botanical Society and Exchange Club of the British
Isles 2: 367, .

Soil Survey of England and Wales (1983). 1:250,000 Soil Map. Sheet 2. Wales. Ordnance Survey,
Southampton.

SUDRE, H. (1908). Rubi Europae, etc.. Paris.

SUDRE, H. (1910). Les Rubus de Belgique. Bulletin de la Société royale de Botanique de Belgique 47:
185-250.

WATSON, W. C. R. (1932). R. egregius Focke *var. effeminatus Focke. Report of the Botanical Society and
Exchange Club of the British Isles 9: 645-646.

WATSON, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland. Cambridge University Press,
Cambridge.

WIGGINTON, M. J. (1999). British red data books: vascular plants. 3rd ed. J.N.C.C., Peterborough.

(Accepted March 2000)

Watsonia 23: 323-326 (2000) 323

The nativeness and non-nativeness of species
M. B. USHER*

Scottish Natural Heritage, 2 Anderson Place, Edinburgh EH6 5NP

ABSTRACT

It has been usual to classify species as native or non-native. However, in relation to biodiversity conservation
and species recovery programmes, it has been recognised that there are “shades of nativeness” between these
two extremes. Scottish Natural Heritage has recently adopted a classification of six categories - native,
formerly native, locally non-native, long-established, recently arrived and non-native. These six categories are
described and examples given.

KEYWORDS: alien species, introduced species, native species.

INTRODUCTION

Webb (1985) made a systematic attempt to establish a series of criteria which could be used to
reach a decision on whether a plant species is native or not. His eight criteria were fossil evidence,
historical evidence, habitat, geographical distribution, frequency of known naturalisation, genetic
diversity, reproductive pattern and possible means of introduction. A ninth criterion,
entomological evidence, was added by Preston (1986). There are two particular features of the use
of this set of criteria. First, implicitly there is a time scale; this is taken as the ending of the last
glaciation 10,000 or so years ago and hence the criteria are not used for species that might have
been in the geographical area being studied in inter-glacial periods. Second, the criteria are
indicative rather than definitive; thus assessment of “native” or “non-native” status depends upon
the balance of probabilities, weighing evidence from all of the nine criteria.

Much of the work related to implementation of the United Kingdom’s Biodiversity Action Plan
(Anon. 1994) is implicitly, or at times explicitly, concerned with discriminating between native
and non-native species of plants and animals. Similarly, species recovery programmes are based
on enhancing populations of native species, or aim at re-introducing native species that have
become extinct locally or nationally. Examples are the successful re-introduction of Milvus milvus
(red kite) into England and Scotland and of Haliaeetus albicilla (white-tailed eagle) into western
Scotland, or the on-going studies into re-introducing Castor fiber (European beaver) into Scotland
(Anon. 1998b). In contrast, site management often targets non-native species for control; such
species include Rhododendron ponticum (Usher 1986) and Fallopia japonica (e.g. Pope 1997).

Whereas Webb’s (1985) criteria were designed to discriminate between native and non-native
species, Scottish Natural Heritage recognised that these were two of a number of possible
categories. The British Ornithologists’ Union, for example, recognised rather more categories in
compiling The British List of bird species (Anonymous 1998a). The aim of this study was
therefore to review species that occur in Scotland, and to propose a classification of nativeness that
would be useful as a factor in conservation practice.

CATEGORIES OF SPECIES

A geographical scale needed to be defined. For the purposes of this study the whole of Great
Britain was used because of its island character. The classification, in Table 1, is therefore related
to the whole island, but it could equally be applied to any defined geographical area and portions
of that area.

* Address for correspondence: e-mail MBU @snh.gov.uk

324 M. B. USHER

Native species are those that are presumed to occur in Great Britain by natural means. Examples
of such native species would include such common plants as Bellis perennis and endemic species
such as Primula scotica. Both of these species are native in Great Britain as a whole: however, if a
smaller geographical area is used, both species are native in Scotland, whereas only one, B.
perennis, is native in Wales.

Formerly native species are those that are known to have occurred in Great Britain since the last
ice age, but which have become extinct. Many of the species in this category apparently became
extinct because the environment changed (category 2a in Table 1). Birks (1997), for example,
listed a number of such plants, including Papaver sect Scapiflora and Cassiope spp. Other species
in this category might have become extinct through human activity rather than environment
change (category 2b in Table 1). Examples include Trichophorum alpinum, whose only British site
was destroyed by marl digging in the 13 years following the species’ discovery in 1791, and
Pinguicula alpina, whose only habitat was encroached by agriculture, subsequently invaded by
coniferous trees, and not assisted in the mid-19th century by the avariciousness of plant collectors
(Lusby 1998).

Locally non-native species are those that are native in part of Great Britain, but which have been
introduced to the local area being considered. An example is Luronium natans, which is native in
England and Wales but not native in Scotland where it has been introduced to at least three sites
(Stewart et al. 1994): whereas it is native in Great Britain it is non-native in Scotland. There are
few botanical instances where this category might be important from a biodiversity point of view,
but in some cases locally non-native animals present a serious threat to native species. An example
is the occurrence of Erinaceus europaeus (hedgehog) in the Outer Hebrides. It never naturally
reached those islands, but since its introduction it has been creating serious problems for ground-
nesting birds.

Long-established species have often been described as “naturalised”; they have become part of
food webs where the majority of species are native. An example of a long-established species in
Britain is Acer pseudoplatanus, which Salisbury (1961) presumed to have been introduced by the
Romans of the order of two thousand years ago. This species is long-established in Great Britain,
whereas a species such as Fagus sylvatica, which is native in the southern part of Britain, is locally
long-established in many northern areas.

Recently arrived species are those for which there is as yet insufficient evidence to decide
whether they arrived “naturally” (category 5b in Table 1) or were assisted by some human agency
(category 5a in Table 1). In a sense, it is a temporary category, a useful classification pending
further scientific evidence that would allow the species to be assigned to category | (native) or to
category 6 (non-native). An example would be the discovery of Serapias parviflora in Cornwall in
1989. Both Sell & Murrell (1996) and Stace (1997) inclined to the view that this orchid species
was deliberately introduced (category 5a), whereas French er al. (1999) considered it likely to be a
natural colonisation (category 5b) because it occurs in dry maritime grassland on islands off the
Brittany coast. Until there is more evidence, the species cannot be assigned with certainty to
categories | or 6.

Finally, non-native species are those where all of the evidence points to the fact that they have
been brought to Great Britain by people, either intentionally or unintentionally. Logically, if a
species has never been native in any part of Britain, then the application of the term can only be to
Britain as a whole rather than to any part of Britain.

These six categories are not mutually exclusive. Categories 1, 2 and 3 in Table | could be referred
to as British native species because, at some time during the last 10,000 years or so, they have
occurred naturally in at least some part of Great Britain. Similarly, categories 4 and 6 in Table 1
could be referred to as non-native since neither category has ever occurred naturally in the country.
Category 5 is more problematic, but in general 5a would fall into the non-native group whereas 5b
are more likely to be treated as native. Although there are therefore certain ambiguities in the
definition of the six categories and their subdivisions, they nevertheless prove useful in thinking
about the conservation and management of species.

NATIVE CATEGORIES 325

TABLE 1. SIX CATEGORIES, TWO SUBDIVIDED, FOR CLASSIFYING SPECIES
ACCORDING TO THEIR NATIVENESS. THE CATEGORIES CAN BE APPLIED TO THE
WHOLE OF GREAT BRITAIN (B) OR JUST TO A LOCAL PART OF BRITAIN (L).

Category Application Sub-divisions
1. Native Bly -
2. Formerly native BE a. Extinct because of environment change
b. Extinct due to human activity
3. Locally non-native le -
4. Long-established BL -
5. Recently arrived Bye a. Arrival suspected of being human assisted
b. “Natural” means of arrival suspected
6. Non-native B -
DISCUSSION

The categories of nativeness (Table 1) have been developed for nature conservation purposes.
Both habitat loss (and fragmentation) and the effects of non-native species are seen to be the two
primary causes of the loss of biodiversity on the Earth (e.g. Heywood 1995). It is therefore
important to consider attitudes to the “nativeness” of species and to consider the conservation
actions that might follow from such categorisation in the geographical area of concern.

The six categories listed in Table 1 were conceived in relation to Great Britain, a single island
that is in many respects a single political entity. Inevitably with national biodiversity action plans
and with national conservation agencies, political boundaries tend to predominate in
considerations of nativeness. However, it might be preferable to conceive of more naturally-
defined boundaries, such as the archipelago of islands off the coast of north west Europe (i.e. the
whole of the British Isles) or, preferably, biogeographically distinct areas (cf. Usher & Balharry
(1996) for a division of Scotland into twelve biogeographical zones). The efforts of the European
Commission to establish trans-national conservation, on the basis of a small number of
biogeographical regions encompassing all 15 Member States, is certainly an attempt to move from
a national to a more natural perspective for the conservation of species and habitats (Anon. 1992).
Logically, natural boundaries should be favoured over political ones, but, pragmatically, this is
unlikely to be realised!

Table 1 can also be compared with the British Ornithologists’ Union’s categorisation of birds for
inclusion on the British List (Anon. 1998a). That categorisation includes native species (A),
formerly native species (B, with a date of 31st December 1949 as the decision point), introduced
species that have established breeding populations (C), species that would be in another category
but about which there is some doubt (D), and species that have been introduced and which have
not established self-sustaining populations (E). In a comparison with Table 1, categories A and 1
are similar, as are B and 2. Category C relates to both 4 and 6, and possibly also to 5a, whereas
category D relates primarily to 5b. Category E relates to 6. Nothing in the BOU list equates to
category 3 in Table 1, but this could be predicted with a highly mobile (and migratory) taxonomic
group such as the birds.

Williamson’s (1996) consideration of biological invasions led him to prepare a conceptual
framework, of which two points are particularly pertinent. First, most invasions fail, with only
about 10% of introduced species establishing a population. Second, most invasions, probably
around 90%, have only minor consequences. This implies that only about 1% of introduced
species become “pests”. Does this matter? If one takes Clement & Foster’s (1994) book, which
lists 3,586 non-native species recorded in the British Isles, of which 885 are established, then
between 36 species (1% of 3,586) and 88 species (10% of 885 species) might be expected to pose
problems for native species and natural plant communities. There is therefore potentially a large
task to control and manage such a number of problematic non-native species.

Nature conservation, or biodiversity action plans, will focus on native species (category 1 in
Table 1). Because a proportion of the non-native species (category 6) can become pests, it is useful

326 M. B. USHER

to think of controlling them, or even eradicating them, as soon as they are noticed. Re-introduction
programmes can be focused on the formerly native species, but only if environmental conditions
are still suitable for them to establish self-sustaining populations (i.e. category 2b). Generally there
are risks in introducing species beyond their native range (category 3), but care needs to be
exercised in deciding to control long-established species (category 4) because they may, over the
centuries, have become part of cultural landscapes. Caution is needed with species that have
recently arrived; there may be benefits in treating category 5a as if they were category 6, but it has
to be remembered that species distributions are dynamic, and that distributions will change
naturally under the various scenarios of climate change.

There is an increasing scientific basis for adopting a precautionary approach to dealing with the
arrival and establishment of non-native species (and, by analogy, also with non-native genotypes).
Williamson’s (1996) study documented the problems that biological invasions can pose to a
territory’s biodiversity. There is also an increasing practical basis for the early control of non-
native species. Once a non-native species has become established, and starts to have an impact on
local biodiversity, it can be extremely expensive to mount a control campaign, even if control is
ultimately possible.

REFERENCES

ANONYMOUS (1992). Council Directive 92/43/EEC of 21 May 1992 on the conservation of natural habitats

and of wild fauna and flora. Official Journal of the European Communities L206: 7-50.

ANONYMOUS (1994). Biodiversity: the UK Action Plan. H.M.S.O., London.

ANONYMOUS (1998a). The British List: the official list of birds of Great Britain with lists for Northern Ireland

and the Isle of Man. British Ornithologists’ Union, Tring.

ANONYMOUS (1998b). Re-introduction of European beaver to Scotland: a public consultation. Scottish

Natural Heritage, Perth.

BirKS, H. J. B. (1997). Scottish biodiversity in a Scottish context, in FLEMING, L. V., NEWTON, A. C.,
VICKERY, J. A., & USHER, M. B., eds. Biodiversity in Scotland: status, trends and initiatives, pp. 21-35.
The Stationery Office, Edinburgh.

CLEMENT, E. J. & FOSTER, M. C. (1994). Alien plants of the British Isles. Botanical Society of the British
Isles, London.

FRENCH, C. N., MURPHY, R. J. & ATKINSON, M. G. C. (1999). Flora of Cornwall. Atlas of the flowering
plants and ferns of Cornwall with notes on some species recorded in the Isles of Scilly. Wheal Seton
Press, Camborne.

HEYWOOD, V. H., ed. (1995). Global biodiversity assessment. Cambridge University Press, Cambridge.

Lussy, P. (1998). On the extinct plants of Scotland, in LAMBERT, R. A., ed. Species history in Scotland, pp.
45-62. Scottish Cultural Press, Edinburgh

POPE, P. J. (1997). The spread of Japanese knotweed and Himalayan balsam. B.S.B.I. News 76: 62.

PRESTON, C. D. (1986). An additional criterion for assessing native status. Watsonia 16: 83.

SALISBURY, E. (1961). Weeds and aliens. Collins, London.

SELL, P. & MURRELL, G. (1996). Flora of Great Britain and Ireland. Volume 5: Butomaceae - Orchidaceae.
Cambridge University Press, Cambridge.

STACE, C. (1997). New flora of the British Isles, 2nd ed. Cambridge University Press, Cambridge.

STEWART, A., PEARMAN, D. A. & PRESTON, C. D., eds. (1994). Scarce plants in Britain. Joint Nature
Conservation Committee, Peterborough.

USHER, M. B. (1986). Invasibility and wildlife conservation: invasive species on nature reserves.
Philosophical transactions of the Royal Society, series B 314: 695-710.

USHER, M. B. & BALHARRY, D. (1996). Biogeographical zonation of Scotland. Scottish Natural Heritage,
Perth. .

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

WILLIAMSON, M. (1996). Biological invasions. Chapman & Hall, London.

(Accepted March 2000)

Watsonia 23: 327-330 (2000) i,

A new record for Salix x angusensis (Salicaceae) Rech. f. from
Ainsdale Sand Dunes National Nature Reserve, S. Lancs. v.c. 59

R. D. MEIKLE

Ranscombe Lodge, Wootton Courtenay, Minehead, Somerset TA24 SRA

and
N. A. ROBINSON

3 Abbey Drive, Natland, Kendal, Cumbria LAY 7QN

ABSTRACT

Salix x angusensis Rech. f. (S. cinerea subsp. oleifolia x S. repens var. argentea X S. viminalis), a very rare
hybrid, previously known only from the type locality, has been found on Ainsdale Sand Dunes National
Nature Reserve. Type material of the hybrid has been located, and an amended description drawn up from
additional foliage and catkin material. The paper includes notes on the population and ecology of the hybrid.

KEYWORDS: willow, hybrid.

INTRODUCTION

In July 1993 N.A.R. sent R.D.M. some freshly collected specimens of willows from Ainsdale Sand
Dunes National Nature Reserve, S. Lancs. v.c. 59. Amongst these were Salix repens L. var.
argentea (Sm.) Wimm. et Grab. and the rare $. x doniana G. Anderson ex Sm. (S. purpurea L. x
S. repens L.), both of which were known to occur in the area. But a third specimen (Robinson 65)
defied immediate identification, though it was evidently a hybrid, with S. repens as one of its
parents. This puzzling specimen was successfully rooted and grown in R.D.M.’s garden, and
developed into a slender, erect shrub about 1 m high, with small, greyish, acuminate leaves, and
numerous narrow-cylindrical female catkins, with hairy ovaries, and remarkably elongate styles
and stigmas, superficially not unlike the styles and stigmas of the arctic-alpine S. Japponum L., and
very distinct in this respect from other lowland willows. At first, to account for this unusual
feature, R.D.M. thought it just possible that the puzzling willow might be S. myrsinifolia Salisb. x
S. repens L. (S. x felina Buser ex Camus & A. Camus) but S. myrsinifolia is not known to occur
anywhere in the vicinity of Ainsdale, nor apart from the elongate styles and stigmas, was there any
evidence of S. myrsinifolia in the make-up of the hybrid.

Further searches by Mr Tony Duckels in 1996 and 1997 failed to furnish additional material of
the puzzling willow, though S. cinerea L. subsp. oleifolia Macreight (S. atrocinerea Brot.), S.
repens var. argentea and S. cinerea subsp. oleifolia x S. repens var. argentea was received from
the same area of the dunes. Happily, the puzzling plant was relocated by N.A.R. on 10 August,
1998, by which time R.D.M. concluded that it must be a triple hybrid, S. cinerea subsp. oleifolia x
S. repens var. argentea x S. viminalis L., the presence of the last-named being necessary to
account for the acuminate leaves of the puzzling plant. This triple hybrid has only once been
recorded, as S. X angusensis Rech. f. (Rechinger 1950) from Angus v.c. 90, “Barry Links near
Carnoustie, September 1947, K. H. Rechinger, no. 45”. Rechinger’s short description of habit,
twigs and foliage did not exactly tally with that of the Ainsdale plant, so it became necessary to
examine the type, stated by the author to have been deposited in BM. R.D.M. had earlier tried,
without success, to locate this type when preparing an account of Salix for Stace (1975), and a
further effort by Mr Roy Vickery (BM) was likewise unsuccessful. An enquiry to E was no more
fruitful, though Mr Douglas McKean drew attention to the fact that Rechinger (op. cit.) had also

328 R. D. MEIKLE AND N. A. ROBINSON

stated, “Numbers quoted refer to dried specimens, duplicate sets of which have been deposited at
the British Museum (Natural History) and at Angus Herbarium”. At Douglas McKean’s suggestion
R.D.M. wrote to Miss Rachel Benvie, Curator, Montrose Museum and Art Gallery, Montrose,
Angus, but was informed that no material of S$. X angusensis was to be found in the collections
there. Almost on the point of giving up hope of finding the missing types, a good photocopy of the
type was received from W. Examination of the photocopy satisfied us that the Ainsdale Salix and
S. X angusensis are indistinguishable.

Since Prof. Rechinger’s original description is brief, and based exclusively on foliage, I subjoin
a more detailed description of this rare and interesting hybrid.

DESCRIPTION

Salix X angusensis Rech. f. in Watsonia, 1(5): 275 (1950); Meikle in Stace, Hybridization and the
Flora of the British Isles, p319 (1975). Salix cinerea L. subsp. oleifolia Macreight (S. atrocinera
Brot.) x S. repens L. var. argentea (Sm.) Wimm. ex Grab. (S. arenaria L.) x S. viminalis L.
Holotype: Scotland; Angus (v.c. 90), “Barry Links near Carnoustie, September 1947, K. H.
Rechinger no. 45” (W).

Slender, erect or sprawling shrub, c. | m high, but probably taller when more mature; twigs at first
densely appressed-pubescent, but soon becoming glabrous or subglabrous and rather lustrous dark
reddish-brown. Leaves numerous and rather close together, lanceolate-acuminate, (2-5—)3-5—6(—7)
cm long, (O-5—)1—2(—2:5) cm wide, dull green and thinly pubescent above, densely appressed
sericeous-pubescent below, margins flat or undulate, entire or subentire, midrib prominent below,
lateral nerves numerous, close together ascending; petiole short, usually less than 5 mm long;
stipules lanceolate, 3-5 mm long, 1-5—2-5 mm wide, fairly conspicuous on young growths but soon
shed. Female catkins numerous and rather close together, suberect or spreading, narrowly
cylindrical, shortly stalked, 2—3-5 cm long, 0-4—0-6 cm wide; bracts 3—5, spreading acuminate, 4-7
mm long, 2-3 mm wide, green above, sericeous below; catkin-scales oblong-ovate, acute, about
1-8 mm long, 0-8 mm wide, fuscous except towards base, clothed with long, silvery-silky hairs;
ovaries crowded, narrow-ovoid, tapering to apex, 2-2-5 mm long about 1-5 mm wide, shortly
stalked, densely subappressed silvery-silky; nectary oblong, truncate, about 0-8 mm long, 0-4 mm
wide, style distinct, slender, about 1-5 mm long; stigmas 4, filiform, to about 0-8 mm long,
recurving at maturity. Male catkins and seeds not seen.

ECOLOGICAL AND DISTRIBUTIONAL NOTES

The first specimen was collected in July 1993 in Slack 65 in the north-western corner of Ainsdale
Sand Dunes National Nature Reserve, at SD/295.118 by Michael Gee, Site Manager, and N.A.R.
The slack is situated at the seaward edge of the fixed dunes, just behind the more mobile frontal
dune ridges. The plant community in the slack is known in the National Vegetation Classification
(Rodwell 1991-1996) as “Salix repens-Calliergon cuspidatum damp slack”, in which the most
conspicuous plant is the leggy, creeping Salix repens and the most consistent plant on the ground
surface is the moss Calliergon cuspidatum (Hedw.) Kindb. Other plants present, typical of damp
slacks, are Mentha aquatica L., Equisetum fluviatile L., Eleocharis palustris (L.) Roem. & Schult.,
Epilobium palustre L. and Lythrum salicaria L. The water table varies greatly and the slack is
usually flooded during the winter, sometimes into the spring, but can be quite dry in summer.
Collections of willows in the slack taken in 1996 and 1997, det. R.D.M., were found not to
contain S. X angusensis, but established that although the willow in the slack was mainly S. repens
var. argentea, S. cinerea subsp. oleifolia was also present as scattered, stunted bushes, and as a 4
m tall clump at the edge of the slack. The second gathering, which was determined as S. x
angusensis, was made on 10 August 1998 from a group of six bushes within the stand of S. repens
var. argentea, centred about 7 m south of the tall S. cinerea subsp. oleifolia. The bushes had the
same sprawling growth with much horizontal branching, as the surrounding S. repens var.
argentea, and, at 0-7 m, were about the same height. In consequence they did not stand out from
the rest and were not visible from the edge of the slack. They could be distinguished form the

SALIX Xx ANGUSENSIS 329

FIGURE 1. Salix x angusensis Rech. f. del. P. Halliday © 2000

Drawn from N. A. Robinson 65 (cult. Ranscombe Lodge, Wootton Courtenay, Som.) Leaves 4 August 1995;
catkins 15 April 1997. A: Foliage, B: Mature leaf, C: Stipules, D: 2 Catkins, E: 2 Flower.

330 R. D. MEIKLE AND N. A. ROBINSON

surrounding willows by the longer leaves on the new growth — leaves on the old growth appeared
to be much the same. The bushes looked old and broken down, but new growth was coming from
the base and also new shoots could be seen round about, presumably from suckers. The fact that
they were growing in a compact group suggested that they had arisen from a single initiate, within
the stand of S. repens var. argentea. The third parent involved in the S. x angusensis triple hybrid,
namely S. viminalis, is known to be present in the reserve, though not in the immediate vicinity,
and the rare hybrid S. x friesiana Anderss. (S. repens X S. viminalis) is widespread and common in
the v.c. 59 dunes. Some distance away, in the same slack, there is a clump of S. x doniana Sm. (S.
purpurea X S. repens). This hybrid was not known outside Scotland until it was found on this part
of the coast in 1947 (Savidge et al., 1963). It is present at several locations in the Reserve, and has
been found at some other places in the coastal dunes.

These willows are in parts of the Reserve which are not generally open to the public. Anyone
wishing to see them, or to take samples, should contact the Site Manager, Ainsdale Sand Dunes
National Nature Reserve, 2 West End Lodge, Pinfold Lane, Southport.

ACKNOWLEDGMENTS

We are grateful to Mr Michael Gee and to Mr Tony Duckels for their help in searching for S. x
angusensis; also to Messrs Roy Vickery (Natural History Museum) and Douglas McKean (Royal
Botanic Garden Edinburgh), and Miss Rachel Benvie (Montrose Museum and Art Gallery) for
assistance in our pursuit of the missing type material, and our thanks to Dr Emst Vitek
(Naturhistorisches Museum, Botanisches Abteilung, Vienna) for tracing the missing specimens
and supplying photocopies. Our thanks also to Miss P. Halliday for the illustration which
accompanies this paper.

REFERENCES

RECHINGER, K H. (1950). Observations on some Scottish Willows. Watsonia 1(5): 275.

RODWELL, J. S., ed. (1991-1996). British Plant Communities, 14, Cambridge University Press, Cambridge.

SAVIDGE, J. P.. HEYwoop, V. H. & GORDON, V. (1963). Travis’s Flora of South Lancashire. Liverpool
Botanical Society, Liverpool.

STACE, C. A. (1975). Hybridization and the Flora of the British Isles, pp. 312-319. Academic Press, London.

(Accepted July 1999)

Watsonia 23: 331—334 (2000) 331

Discovery of male plants of Najas marina L. (Hydrocharitaceae)
in Britain

R. J. HANDLEY* and A. J. DAVY

School of Biological Sciences, University of East Anglia, Norwich, NR4 7TJ

ABSTRACT

The dioecious, annual macrophyte Najas marina is represented in Britain only in the Norfolk Broads.
Although it was discovered in 1883 (at Hickling Broad), only female plants had previously been recorded and
all seed production was assumed to be apomictic. Following detailed study of the population biology of this
species, the occurrence of male plants at Hickling and Upton Broads in the summer of 1998 is reported.
Photographs of the inconspicuous male flowers and pollen are provided. Possible reasons for the plants
remaining unreported for so long are discussed. The potential for sexual reproduction has implications for the
conservation of this Red Data Book species in Britain.

Keywords: Annual, aquatic macrophyte, dioecy, Holly-leaved Naiad, rarity.

INTRODUCTION

Najas marina (Holly-leaved Naiad) is a dioecious annual which grows completely submerged in
shallow lakes (1-3 m deep), rooting extensively into the bottom sediments. In Britain, N. marina is
only found in the Norfolk Broads and there only in a handful of lakes, where it forms dense beds
and can predominate over large areas. N. marina has a long history in the British flora, with fossil
evidence for populations being far more widespread during the last climatic optimum than at
present (Godwin 1975). In 1883, Arthur Bennett found N. marina for the first time as a living plant
(Bennett 1884, 1909) and since then there has been a well reported increase in the number of sites
at which it has been found (Barry & Jermy 1953) although there have rarely been more than seven
sites in any one year.

Najas marina is listed as Vulnerable in the British Red Data Book and in the IUCN European
Red Data Categories and is therefore specially protected under the Wildlife and Countryside Act
1981 (Wigginton 1999). Despite considerable interest in N. marina, arising from its rarity value in
Britain and Europe, there has been little detailed knowledge of its autecology and life history. In
particular, there have been no reports of male individuals of N. marina in Britain and, in the
apparent absence of male flowers, there has been speculation that its prolific seed production is
entirely apomictic (Stace 1997; Preston & Croft 1997). This paper reports that male plants do in
fact occur in Britain and illustrates the features of male flowers.

THE IDENTIFICATION OF MALE PLANTS

The discovery of male plants of Najas marina was made while snorkel diving to collect plant
material in Hickling Broad in East Norfolk (Nat. Grid ref: TG/418.209). Subsequently, plants
bearing male flowers were found at Upton Broad, another well established site for this species.
Hickling Broad is the largest of the Norfolk Broads and the site at which Bennett first recorded N.
marina (Bennett 1884).

During 1998, Hickling and Upton Broads were visited at fortnightly intervals throughout the
year in order to survey the populations of N. marina. In June and July, only plants bearing female
flowers were observed; despite meticulous examination of a considerable quantity of plant
material, male flowers could not be found. Plants apparently had only female flowers or no floral

*Author for correspondence, e-mail: r.handley @uea.ac.uk

332 R. J. HANDLEY AND A. j. DAVY

structures at all. However, in August 1998 male flowers were identified for the first time while
diving. These flowers were evident for only a few weeks.

The dense beds, with near-continuous cover, and the branching growth form of Najas marina at
Hickling and Upton Broads combine to create considerable difficulty in distinguishing individual
plants. The bases of the stems often become covered in sediment and root at the covered nodes
making the identification or extraction of individual plants impossible. Nevertheless it was clear
that the ratio of female to male flower bearing stems in these beds was high, perhaps as high as
10:1, and that male stems occurred in distinct patches, suggesting that isolated, individual males
were surrounded by females. With some care it was possible to collect entire stems of N. marina.
Examination of these revealed that each stem bears either exclusively male flowers or exclusively
female flowers. As an individual plant may comprise several stems arising from the initial node it
is not possible to say categorically that all the stems of an individual plant were of the same sex.
However, there is no evidence from work on these populations to query the traditional view that N.
marina 1s dioecious.

FLORAL MORPHOLOGY

The flowers of Najas marina arise singly, or sometimes several together, on a fertile shoot with
suppressed internodes within a sheath axil (Rendle 1899). The male flower is ellipsoid, c. 2—4 mm
long and 1—2 mm wide at the widest point. It possesses a thin membranous spathe which forms a
tight envelope about the flower and protrudes above it to form a narrow crown with characteristic
spines (Fig. la). The four locules of the single anther are discernible within the perianth (Fig. 1b,
lc). The locules are densely packed with pollen grains, which are for the most part ellipsoid, but
some are spherical (Fig. 1d). The morphology of these structures conforms well with the classical
literature on the Najadaceae (Bailey 1885; Campbell 1897; Rendle 1899).

Initial field observations were of male flowers that appeared conspicuously pale in colour,
apparently because the white pollen grains were visible through the anther wall and _ thin,
membranous perianth. Subsequently, darker male flowers were found. The pale flowers were close
to dehiscence, with mature pollen. In these flowers the pollen-filled anther was particularly visible,
because elongation of the filament prior to dehiscence caused the anther to split the spathe from
the apex downwards (Rendle 1899). Flowers were also found in which the anthers had clearly
dehisced, leaving the anther filament and the vestiges of the four locules.

Male flowers are similar in size and shape to the maturing fruits of the female plant, and
therefore they can be difficult to distinguish in the field. The phase during which the male flowers
have a pale colour provides a reliable field diagnostic feature but this was evident only during
August.

DISCUSSION

Given the small size and superficially similar appearance of male flowers to the developing fruits
in Najas marina, it is perhaps not surprising that male plants of this dioecious species have gone
unrecorded for so long in the British flora. The annual “window” during which the sexes are
reasonably distinct appears to be narrow: outside this period, only very close scrutiny would reveal
male flowers still within the spathe or the remains of dehisced anthers. In addition, the use of
diving equipment and the unusually good water clarity in 1998 would have improved the chances
of discrimination. Recently the macrophytes of the Norfolk Broads have been the subject of
intensive study (Schutten et al. 1997; Kennison et al. 1998) without male plants of N. marina
being reported; it is possible that the high ratio of female to male plants requires a sampling
method more sensitive that the double-headed rake commonly used.

We cannot be sure, however, that male plants have always been present in the British population
and therefore have simply been overlooked previously. An examination of N. marina herbarium
material at K did not reveal male flowers that might have been overlooked by previous collectors.
However the preservation in herbarium specimens of the minute flowers of N. marina is poor.
There are alternative explanations. Sex expression in plants can be fickle and subject to strong
environmental influences (Richards 1997). There may be an influence on N. marina that could
make the expression of maleness a rare or uncommon event, or may have resulted in males in the
British populations for the first time recently. The origins of the current British populations are not

MALE PLANTS OF NAJAS MARINA 333

200 um

a os ™
. : rf : -
: i. ~ ie
f - A t

=

FIGURE 1: (a) Photograph showing the male flower of Najas marina with spathe protruding above the anther
to form characteristic spines, a second immature male flower can be seen to the left obscured by the leaf.
(b) Photograph showing anther of N. marina with perianth wall removed but still visible, two locules of the
anther can be seen through the anther wall. (c) Photograph showing the anther of N. marina with perianth and
anther wall removed and one locule of the anther punctured spilling pollen. (d) Photograph of pollen grains of

N. marina showing mostly elongate and some spherical grains.

334 R. J. HANDLEY AND A. J. DAVY

known: it may have gone unrecorded for many centuries before Bennett’s discovery in 1883 or it
may have been a relatively recent colonist from continental European populations, transported in
the guts of migrating wildfowl (Agami & Waisel 1986). It is possible that there have been several
or repeated colonization events, giving rise to plants of different gender by chance or under
environmental control.

Najas marina in Britain is an obligate annual; therefore, even a temporary deficiency of males
requires an alternative, apomictic mechanism for seed production. In 1998 fruit formation began
well before the first male flowers were observed in the field, which suggests the possibility of
facultative apomixis, with seed production early in the summer being apomictic and later seed
production at least partially sexual (Asker & Jerling 1992).

At present it is impossible to say why male plants of Najas marina have not been recorded
before. Perhaps the most parsimonious explanation is that male flowers simply went unnoticed
until now. More significant is the fresh light shed on the nature of its breeding system, which is of
fundamental importance in determining the genetic variability and survival potential of the British
populations. The demonstrable occurrence of males gives a new perspective to our strategy for
conserving this rare species.

ACKNOWLEDGMENTS

The collection of Najas marina was carried out by license granted under the Wildlife &
Countryside Act 1981. We thank the Biotechnology and Biological Sciences Research Council
(BBSRC), The Broads Authority, The Environment Agency and English Nature (Species
Recovery Program) for financial support, the Norfolk Wildlife Trust (Hickling) and Norwich
Union Fishing Club (Upton) for use of boats while surveying, and the Curator of K for access to
herbarium material.

REFERENCES

AGAMI, M. & WAISEL, Y. (1986) The role of mallard ducks (Anas platyrhynchos) in distribution and
germination of seeds of the submerged hydrophyte Najas marina. Oecologia 68: 473-475.

ASKER, S. E. & JERLING, L. (1992) Apomixis in plants. CRC Press, Florida, USA.

BAILEY, C. (1885) On the structure, the occurrence in Lancashire and the probable source, of Naias graminea,
Delile, var. Delilei,. Magnus. Memoirs of the Manchester Literary and Philosophical Society: Third
Series 10: 29-75.

BARRY, D. H. & JERMY, A. C. (1953) Observations on Najas marina. Transactions of the Norfolk and
Norwich Naturalists’ Society 17: 294-297.

BENNETT, A. (1884) Plants new to Norfolk with notes on other species. Transactions of the Norfolk and
Norwich Naturalists’ Society 3: 633-636.

BENNETT, A. (1909) Najas marina, L., and Chara stelligera, Bauer, as Norfolk plants. Transactions of the
Norfolk and Norwich Naturalists’ Society 9: 47-50.

CAMPBELL, D. H. (1897) A morphological study of Naias and Zannichellia. Proceedings of the California
Academy of Sciences: Third Series Botany 1: 1-61.

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

KENNISON, G. C. B., DUNSFORD, D. S. & SCHUTTEN, J. (1998) Stable or changing lakes? A classification of
aquatic macrophyte assemblages from a eutrophic shallow lake system in the United Kingdom. Aquatic
conservation: marine and freshwater ecosystems 8: 669-684.

PRESTON, C. D. & CROFT, J. M. (1997) Aquatic plants in Britain and Ireland. Harley Books, Colchester.

RENDLE, A. B. (1899) A systematic revision of the Genus Najas. Transactions of the Linnean Society of
London: Second Series Botany 5: 379-444.

RICHARDS, A. J. (1997) Plant breeding systems, 2nd ed. Chapman & Hall, London, England.

SCHUTTEN, J. et al., (1997). Factors affecting water plant recovery, Technical Report for EC LIFE Project
No. 92—93/UK/031, BARS 14c/P-89. Broads Authority/Environment Agency, Norwich.

STACE, C. A. (1997) New Flora of the British Isles, 2nd ed. Cambridge University Press, Cambridge.

WIGGINTON, M. J., ed. (1999) British Red Data Books. 1. Vascular plants, 3rd ed. Joint Nature Conservation
Committee, Peterborough.

(Accepted August 1999)

Watsonia 23: 335-348 (2000) 385

Notes

CORYNEPHORUS CANESCENS (L.) P. BEAUV. (POACEAE) AT KINVER EDGE,
STAFFORDSHIRE: A RE-ASSESSMENT

In his review of the distribution and status of Corynephorus canescens (L.) P. Beauv. in Britain,
Trist (1998) describes the site at Kinver Edge, Staffordshire, which holds one of the few British
inland colonies of this grass, and states that “the population of c. 550 plants has remained fairly
constant since 1977.” Though his site description is reasonably accurate, Trist’s estimate of the
population size of C. canescens fails to take account of its recent dramatic increase following
sympathetic site management. This note, therefore, aims to update Trist’s account and trace the
effects on the C. canescens population of conservation measures taken over the last twelve years.

BACKGROUND HISTORY

C. canescens was discovered on the National Trust’s property at Kinver Edge by I. C. Trueman in
the 1970s. Following the appointment of one of the authors (M. E. Blunt) as Warden in 1986, the
Trust has pursued active measures to conserve and increase this population.

In 1986 C. canescens was present on two areas some 300 m apart near the south-eastern
boundary of the property, where the dip-slope of the escarpment inclines gently to the south-east.
Site 1 consisted of a bowl of bare sand of approximately 200 m’, heavily trampled by the public;
C. canescens survived here as a narrow rim around the bowl. Site 2 was a flatter area of about 140
m*, where the grass was reasonably well established. Trist’s (Joc. cit.) estimate of c. 550 plants is
not unreasonable for the size of the population at this date. The two Corynephorus sites were
bounded on the south-east by a sandy track and fence, beyond which lay a grassy field in private
ownership; this field received periodic treatments of fertilizer and was cut for sileage up to 1986
but not thereafter. Professor Trueman (pers. comm.) has stated that a small quantity of C.
canescens was present in this field when the main colonies were discovered. On its other sides the
C. canescens locality was surrounded by birch/oak/pine scrub encroaching extensively on lowland
Calluna vulgaris (L.) Hull heath.

MANAGEMENT SINCE 1986

With C. canescens fairly well established on Site 2, priority in 1986 was given to halting the
erosion of Site | and encouraging the expansion of the species across the eroded area. Initially
barriers of pine brashings were constructed around the rim of this site to protect existing plants
from trampling; but after three seasons there was no noticeable increase in the size of the C.
canescens colony. Therefore, in winter 1989/90 pine brashings were spread thinly across the open
sand, their branches barely touching to allow good penetration of light. This technique largely
eliminated trampling and within a season C. canescens seedlings had colonised the entire area of
open sand. At this point the pine brashings were removed, the grass continued to flourish, and
public pressure on this site has remained light ever since.

The main problem with maintaining C. canescens populations at Kinver Edge is the presence of
mat-forming Polytrichum mosses which rapidly establish themselves on sand stabilised by the
grass. Polytrichum is followed by a succession of other species, notably Rumex acetosella L.,
Deschampsia flexuosa (L.) Trin., Festuca rubra L. and Hypochaeris radicata L. After some three
years seedlings of Calluna vulgaris and Ulex europaeus appear, and within six the C. canescens
population shows some deterioration. Therefore, in the mid-1990s an experiment to reverse this
succession by ploughing the whole of Site 1 with a Dutch harrow was tried, with successful
results. The moss thatch was broken up, fresh sand was turned over and C. canescens re-colonised
from plants which survived in the gaps between the harrow’s blades.

Site 2 has had no direct management since 1986 except for regular light trampling from visitors
using the adjacent path, and since 1997 from cattle introduced to graze the surrounding heath. This

336 NOTES Watsonia 23 (2000)

trampling sufficiently disturbs the sand to allow a few new plants of C. canescens to establish
themselves; but on the whole the population of Site 2 is deteriorating slowly and gorse scrub has
occupied much of the site.

In 1994 a strip of sandy track c. 30 x 4 m parallel to Site 2 and 3 m from it was fenced off and
covered with pine brash as described above. Here, too, Corynephorus seedlings covered the entire
area within a season and now form a closed sward. This is Site 3 in Table 1 below. In the early
1990s ownership of the field next to the Corynephorus sites passed to the National Trust, since
when light grazing by sheep and cattle has been introduced to encourage the establishment of
heathland communities. By 1997 it was evident that C. canescens was spreading in two areas of
this field adjacent to Sites 1 and 2: these are Sites 4 and 5 listed in Table 1. Other C. canescens
populations at Kinver Edge consist of a small area just to the west of Site 1 (numbered Site 6 in
Table 1) and individual plants scattered widely along track sides and among the Calluna heath
near the main populations.

ESTIMATES OF THE CURRENT POPULATION

In 1997 and 1998 counts were made of the whole and parts of the Corynephorus sites on Kinver
Edge in order to estimate the total population. In summer 1997 M. E. Blunt, assisted by S.
Anderson, ran four transect lines across Site 4, finding C. canescens up to 13 m into the field at its
furthest point of colonisation. Quadrat counts were made at 2 m intervals along the transects.

In July 1998 M. E. Blunt, assisted by S. Fereday, measured the dimensions of all the
Corynephorus sites and made population estimates from quadrat counts on all sites. In September
1998 the two authors made further estimates of the numbers of plants in the three main populations
(Sites 1, 2 and 3) from random quadrats. The results of these counts are shown in Table 1.

The September 1998 estimates were made the day after a severe thunderstorm washed away
parts of Site 1, doubtless accounting for the discrepancy between the two estimates for this Site;
being on more level ground Sites 2 and 3 were little affected by the storm. The July 1998 estimate
for Site 4 was taken over a larger area than that of 1997, since C. canescens was found up to 21 m
into the field in 1998: the area given in the Table for this site is that calculated in July 1998.

It may be deduced from the Table that the total population of C. canescens at Kinver Edge
currently lies between 40,000 and 50,000 plants.

CONCLUSIONS

In its Kinver Edge stations, C. canescens is the pioneer colonist of open sand; it appears to require
a fairly stable surface to establish itself successfully, and seems unable to do so where sand is
regularly disturbed, for example by trampling or the activities of rabbits (Oryctolagus cuniculus):;
though a few plants do establish themselves on the less disturbed edges of rabbit warrens. Once
C. canescens is established, succession of the habitat to gorse (Ulex europaeus) and heather
(Calluna vulgaris) scrub begins within three years and C. canescens populations begin to decline
within six years unless there are further exposures of bare sand. In short, at Kinver Edge

TABLE 1: ESTIMATED NUMBERS OF CORYNEPHORUS CANESCENS PLANTS ON

KINVER EDGE
Area (m°) 1997 July 1998 Sept. 1998
Site 1 202 ; 19,400 12,700
Site 2 145 . 8,100 8,700
Site 3 129 : 10,900 11,000
Site 4 90 1,400 3,900 -
Site 5 40 : 500 -
Site 6 =15 : 3,900 -

Scattered - - 1,600 -

NOTES Watsonia 23 (2000) 337]

2

LA;
5
ip
Co

¢.
oF,
7.
Lasy
UN blepy
=e
—
~
Le —) 7

an 3 Corynephorus =
\ ifs canescens =
:~\v? sites

36 heathland oo

NT Warden's
&? Lodge

= A

Figure |. Plan of Kinver Edge, Staffordshire. Scale approx. 1:2500

C. canescens exhibits a narrow niche requirement, viz. periodic disturbance of sand followed by a
cessation of disturbance over a cycle of 3-6 years. Current studies suggest that trampling, wind-
blow, rainstorms and the activities of rabbits may, depending on their frequency, both create
conditions favourable to C. canescens and also destroy them. Where conditions remain
unfavourable over a period, C. canescens seems able to survive as individual plants scattered along
the edges of paths, particularly in the lee of overhanging vegetation where they receive little
trampling.

Observations further suggest that seed production is overwhelmingly the means whereby C.
canescens colonises new ground at Kinver Edge; once established, it may continue to propagate
itself either from seed or vegetatively. This contrasts with Trist’s (loc. cit.) studies which
suggested vegetative means as the grass’s chief method of propagation in Britain. The role of pine
brashings in encouraging germination is of interest: besides reducing trampling they may also
protect tiny plants from frost and drought, considered by Marshall (1967) as critical factors in
seedling survival. Observations also indicate that C. canescens is not selectively grazed at Kinver
by sheep, cattle or rabbits, and seeds freely at the grazing intensities encountered there.

The authors’ studies of the ecology of C. canescens at Kinver Edge and its response to various
management techniques form part of an ongoing programme. It was, however, felt worthwhile
bringing current findings to the attention of those interested in this species and its conservation, as
well as updating the published record of what now appears to be a major population of this scarce
grass in Britain.

ACKNOWLEDGMENTS

The authors wish to acknowledge the help and advice of Professor I. C. Trueman of the University
of Wolverhampton throughout these studies, and for his comments on the draft of this paper.

338 NOTES Watsonia 23 (2000)
REFERENCES

MARSHALL, J. K. (1967). Biological Flora of the British Isles: Corynephorus canescens (L.) Beauv. Journal of
ecology 55: 207-220.

TRIST, P. J. O. (1998). The distribution and status of Corynephorus canescens (L.) P. Beauv. in Britain and the
Channel Islands with particular reference to its conservation. Watsonia 22: 41-47.

A. G. BLUNT
University of Wolverhampton, Molineux Street, Wolverhampton WV1 ISB

M. E. BLUNT
Warden’s Lodge, Comber Road, Kinver, Stourbridge, West Midlands DY7 6HU

A REANALYSIS OF THE MISTLETOE (VISCUM ALBUM L.; LORANTHACEAE)
SURVEY DATA FROM THE 1970S AND 1990S

In 1993, the late John Knight and I, exasperated by the Plantlife 1993 Christmas card depicting
unseasonal Bluebells, decided to find a more appropriate species for the following year, and had
the idea of repeating the 1970s B.S.B.I. Mistletoe (Viscum album L.) survey reported by Perring
(1973). Independently, Jonathan Briggs had also already been investigating the possibility of
repeating the survey. Discussions between the two societies resulted in a joint Plantlife/B.S.B.I.
survey in the mid 1990s whose main aim was to determine if Mistletoe was declining. The 1994
Plantlife Christmas card included the Mistletoe survey form.

For the 1969-1972 survey, data on the occurrence of Mistletoe and its hosts were requested
from the B.S.B.I., the Northern Horticultural Society and the general public throughout Britain
(Perring 1973). For the 1990s survey, records were again collected from throughout Britain by
B.S.B.I. and Plantlife members and the general public, and the survey received extensive press
coverage (Briggs 1999). When the two surveys were compared, there was large increase in records
since the 1970s. Briggs (1999) was unable to draw any conclusions about possible declines or
increases because of the uncertainty of whether there had simply been more records sent in for the
1990s survey (i.e. the increase in records was a result of more recording). I had previously had
exactly the same problem with the B.S.B.I. Monitoring Scheme data and have spelt it out in some
detail (Rich & Woodruff 1990, 1992, 1996).

Fortunately, some further analysis of the data presented by Briggs (1999) is possible. Crucially,
the tetrad map for England and Wales (Perring’s Fig. 7) had areas marked with hatched lines
where Mistletoe had been searched for “intensively but had not been found”. By assuming that the
areas had indeed been surveyed comprehensively during the 1970s it is possible to assess
statistically the probability of any significant increases or decreases in these areas.

METHODS

The number of tetrad records were counted for areas marked with hatched lines in Perring (1973)
and for the same areas from the maps in Briggs (1999). The poor reproduction of the map in
Perring (1973) meant that determining the limits or exact locations of some hatched areas was very
difficult and areas where there was any doubt were rejected. Data for 39 discrete hatched areas
which range in size from groups of four tetrads to whole counties representing a total of 2817
tetrads (c. 7% of the tetrads in England and Wales) were available, excluding the large area of
Gloucestershire and Herefordshire.

The differences in the number of tetrads were tested for significance using a two-tailed test of
equality of the two percentages (Sokal & Rohlf 1969) with 95% confidence limits.

RESULTS

There were 468 tetrad records in the 1970s hatched areas (16-6% of tetrads surveyed intensively),
and 562 tetrad records in the same areas in the 1990s (19-95% of tetrads surveyed intensively in

NOTES Watsonia 23 (2000) 339

the 1970s). This is a significant increase (p<0-001). Of the discrete hatched areas marked as
intensively recorded in the 1970s, Mistletoe has increased in frequency in 22 areas, showed no
change in six areas, and declined in eleven areas.

DISCUSSION

The re-analysis depends on the assumption that the hatched areas marked in Perring (1973) have
been surveyed comprehensively. Although Perring indicated areas which had been surveyed
sufficiently “intensively” to be reasonably certain that Mistletoe was indeed absent in the 1970s, I
have doubts whether hatched areas as large as Surrey were really surveyed thoroughly. Indeed the
map of the 1970s survey in Briggs (1999) includes more records showing this to be the case (some
from the extension of the survey into 1973). None-the-less, it is reasonable to accept Perring’s
hatched areas at face value as a reasonably thorough baseline survey. Briggs (1999) rightly makes
no claim that any areas had been surveyed intensively.

As it is likely that on average the hatched areas were less intensively recorded in the 1990s than
the 1970s, despite the greater number of people participating in the survey, if there was no change
or a decline in Mistletoe there should be fewer records. The fact that there are significantly more
records despite a less intense survey allows a firmer conclusion to be drawn that there is no
evidence for decline for tetrad frequency data in these areas. It is not possible to determine whether
Mistletoe has increased or stayed the same because the intensity of the survey in the 1990s
compared to that in the 1970s is not known. The inclusion of the Gloucestershire/Herefordshire
area, where there is a large apparent decline, may also affect this conclusion.

Assuming that the hatched areas are a representative random sample (they are scattered across
the country though with local concentrations), it is possible to extend the conclusion to no
evidence of decline at a national level for tetrad frequency data. Rich & Woodruff (1990, 1996)
found no statistically significant change in frequency of Mistletoe at a 10-km square level between
1930-1960 and 1987-1988.

A more appropriate form of survey to determine change than an ad hoc national survey would
have been to re-record the hatched areas intensively. Alternatively, a properly designed sample
baseline survey including estimates of recorder effort could have been established against which
future changes could have been assessed more accurately. The application of standardised survey
techniques, as we have for Ashdown Forest (Rich er al. 1996; Rich 1998), would minimise these
problems. Once again, the problems of recording bias may have masked real changes in wild
plants in the countryside.

REFERENCES

BRIGGS, J. (1999). Kissing goodbye to Mistletoe? The results of a national survey aimed at discovering
whether mistletoe in Britain is in decline. Plantlife, London.

PERRING, F. H. (1973). Mistletoe in GREEN, P. S., ed. Plants: wild and cultivated. B.S.B.I. Conference
Proceedings 1972. pp. 139-145. E. W. Classey, London.

RICH, T. C. G. (1998). Squaring the circles - bias in distribution maps. British wildlife 9: 213-219.

RICH, T. C. G. et al. (1996). Flora of Ashdown Forest. Sussex Botanical Recording Society, East Grinstead.

RICH, T. C. G. & WOODRUFF, E. R. (1990). B.S.B.I. Monitoring Scheme 1987-1988. Chief Scientist’s
Directorate Report No. 1265. Nature Conservancy Council, Peterborough.

RICH, T. C. G. & WOODRUFF, E. R. (1992). The influence of recording bias in botanical surveys: Examples
from the B.S.B.I. Monitoring Scheme 1987-1988. Watsonia 19: 73-95.

RICH, T. C. G. and WOODRUFF, E. R. (1996). Changes in the floras of England and Scotland between 1930-
1960 and 1987-1988: The B.S.B.I. Monitoring Scheme. Biological conservation 75: 217-229.

SOKAL, R. R. & ROHLF, F. J. (1969). Biometry. W. H. Freeman, San Francisco.

T. C. G. RICH
Dept. of Biodiversity and Systematic Biology, National Museum & Gallery, Cardiff CF 10 3NP

340 NOTES Watsonia 23 (2000)

THREE NEW BRITISH SITES FOR CAREX DEPAUPERATA WITH. (CYPERACEAE)
REPRESENTED IN THE IRISH NATIONAL HERBARIUM, GLASNEVIN

In the Irish National Herbarium at Glasnevin Botanic Garden, Dublin (DBN), there are four
herbarium specimens of Carex depauperata from previously unreported localities in Britain, one
of which is of doubtful origin.

V.C. 17 SURREY

The first specimen is labelled ‘near Farnham, Surrey, August 1846, ex herb. W. Mclvor’. William
Graham Mclvor (?-1876) collected a number of plants in Surrey, but does not seem to have
communicated the records to the Surrey botanists of the time (e.g. J. D. Salmon or J. A. Brewer).
There is plenty of suitable woodland on calcareous soils in this area (e.g. Hog’s Back), not far
from the well-known Godalming sites, and this record is accepted.

The second specimen is more problematic. It is labelled ‘Leith Hill, Surrey, 30.6.1890, per J.
Leitch’. The fact that the label is not in Leitch’s hand-writing, the potential for confusion of the
popularly-known ‘Leith Hill’ with the well-known C. depauperata locality at Frith Hill,
Godalming, and the predominantly acidic nature of soils on Leith Hill, suggests the specimen may
be mislabelled. John Leitch (1849-1896) appears to have collected widely in Kent and Cumbria,
but there are few Surrey specimens. His herbarium at Carlisle (CLE) contains two sheets
cultivated at Bournemouth on 21 June 1890 (presumably by E. F. Linton), and another from the
Royal Botanic Gardens, Edinburgh, June 1871. This record is not accepted.

V.C. 83 EDINBURGH

A third specimen is labelled ‘Wood above Bonelly (ex. herb. W. R. McNab)’. We interpret the
locality as Bonally (sometimes spelt Bonaly), near Edinburgh, grid reference c. NT/21.67, now
partly a country park. Dr William Ramsay McNab was born in Edinburgh in 1844. He worked at
the Royal Botanic Garden, Edinburgh, lectured in botany in Dumfries in 1869 and in Cirencester
from 1870, and became the Scientific Supervisor at the Glasnevin Botanic Gardens in 1880, where
there are many of his specimens (Nelson 1990; in press). He died in Dublin in 1889. The
handwriting on the herbarium sheet is different to other McNab specimens and it may not have
been collected by him, but his local Edinburgh connection reinforces our interpretation of the site.
This record is accepted, though it 1s conceivable there may be links between Bonally, McNab and
material grown in Edinburgh (cf. above).

V.C. 90 FORFAR

The final specimen is labelled ‘Forfarshire, T. McFarlane (ex herb. Rev. H. G. Carroll)’ but is
undated. We have been unable to trace anything about T. McFarlane to give us any more direct
clues about its origin. However, the Rev. Henry George Carroll was an Irish botanist born in c.
1810 and died in 1902, so the specimen was probably collected sometime before 1900. Carroll had
obtained other notable specimens from a range of collectors and his herbarium sheets, with their
characteristic taxonomic headings and handwriting, represent a significant component of DBN.
We would like to know more about his possible connections with Scottish botanists.

The specimen also casts new light on the record in Hooker (1821) ‘Woods near Forfar, G. Done
accepted by Gardiner (1848) but rejected as doubtful by Ingram & Noltie (1981) as it could have
originated from Don’s garden (the source of several other confusing records). We believe this
apparently independently-collected specimen establishes C. depauperata as a Forfarshire species.

There are no other specimens collected in Scotland in ABD, DEE, E, GLAM or PTH.

This now brings the total number of sites for C. depauperata in the British Isles to 14. Although
the British collections in DBN are relatively poorly known, it is none-the-less surprising that at
least three new records should turn up in the same herbarium folder. We hope that these specimens
lead to further information and, above all, to the rediscovery of C. depauperata in these sites.

ACKNOWLEDGMENTS

We are grateful to David Allen, Richard Brinklow, Stephen Hewitt, Douglas McKean, Henry
Noltie, Mark Simmons, Joyce Smith and Keith Watson for their help.

NOTES Watsonia 23 (2000) 341
REFERENCES

GARDINER, W. (1848). The flora of Forfarshire. Longman, Brown and Green, London.

HOOKER, W. J. (1821). Flora scotica. Archibald Constable & Co, Edinburgh.

INGRAM, R. & NOLTIE, H. J. (1981). The flora of Angus (Forfar, v.c. 90). Dundee Museums and Art Galleries,
Dundee.

NELSON, E. C. (1990). William Ramsay McNab’s herbarium in the National Botanic Gardens, Glasnevin
(DBN). I. Its early history and acquisition. Glasra 1: 1-7.

NELSON, E. C. (in press). William Ramsay McNab’s herbarium in the National Botanic Gardens, Glasnevin
(DBN). II. A catalogue of specimens from the Royal Gardens Kew (1805-1810) relating to Hortus
Kewensis (edition 2), the Botanic Garden, Edinburgh (c. 1815) and Scottish native species. Glasra.

Ts G.G. Rico
Dept. of Biodiversity and Systematic Biology, National Museum & Gallery, Cardiff CF 10 3NP

R. A. JONES
Countryside Council for Wales, Plas Gogerddan, Aberystwyth, Dyfed SY23 3EE

M. JEBB
Herbarium, National Botanic Gardens, Glasnevin, Dublin 9, Ireland

ELEOCHARIS PARVULA DISCOVERED IN SCOTLAND

On 29 July 1999 we visited the Cromarty Firth S.E. of Pitglassie, Dingwall, E. Ross (v.c. 106),
where the River Conon flows in a number of channels separated by low-lying islands with marshes
and pools. The marshes and pools contain some species characteristic of fresh water and others
which typically occur in more brackish conditions. When we arrived at high tide the channels were
full, but as we left we could see that the tide had gone out to reveal extensive mud banks beside
them. It was while looking at one of these banks, at grid reference NH/553572, that we noticed a
small Eleocharis which turned out on further examination to be E. parvula (Roem. & Schult.) Link
ex Bluff, Nees & Schauer. This species has hitherto been recorded in Britain on the coasts of
southern England (Devon, Dorset, S. Hampshire) and North Wales (Merioneth, Caernarvonshire).
The occurrence on the Cromarty Firth therefore represents the first record for Scotland and an
extension of its British range some 500 km northwards. The identification has been confirmed by
S.M. Walters and voucher specimens have been deposited in CGE and E.

The Eleocharis parvula population at Pitglassie grew on both sides of the channel in a substrate
of fine mud, or in predominantly stony areas with mud in the interstices of the stones. On the date
of our visit it would have been submerged under 50-75 cm of water at high tide. In places it grew
under a very sparse stand of Bolboschoenus maritimus, but the most frequent associate was
Callitriche stagnalis, which grew as small, prostrate, fruiting plants which were abundant on the
mud banks but did not form a complete cover. Other associates were very occasional plants of
Armeria maritima, Cochlearia officinalis and Triglochin maritimum. In late July we could find
only vegetative stems, and there was still no sign of flowering when Stewart Angus, Ro Scott and
Peter Wortham revisited the site on 31 August 1999, when Peter Wortham returned on 30
September or when Ro Scott went again on 22 October 1999. Even in July the characteristic white
turions of E. parvula were present, albeit sparingly, on the slender rhizomes. By the end of
September the plants were described as yellowing.

The habitat of Eleocharis parvula at its English and Welsh sites has recently been described by
Byfield (1999). As in the Cromarty Firth, it grows “in large tidal rivers ... on firm bare muddy
substrates ... close to the upper limits of tidal influence, avoiding the strongly saline conditions
associated with many saltmarshes.” However, the occurrence of FE. parvula with frequent
Callitriche stagnalis does not appear to have described elsewhere in Britain: no Callitriche species
are recorded with Eleocharis parvula by Rhind & Jones (1994) in their account of the E. parvula
saltmarsh community at Glaslyn Marsh, Merioneth, for example. Indeed, we have not traced any
description in the standard British sources of tidal mud bank communities dominated by
Callitriche species. Our very limited experience suggests that these are a feature of the large
estuaries in N.E. Scotland: on 2 August 1999 we visited the River Beauly E.S.E. of Tomich House,

342 NOTES Watsonia 23 (2000)

NH/54.47., where C. stagnalis formed an abundant green sward on the sloppy mud banks and flats
alongside the large single channel of the river. Rodwell (1995) does not describe these
communities, though he mentions the occurrence of Callitriche stands in temporarily or seasonally
flooded sites such as tracks or woodland rides, the edges of fluctuating ponds or the summer-dry
upper reaches of chalk streams. C.D.P. was similarly unaware of the tidal mud communities when
preparing the habitat descriptions for Callitriche species in Preston & Croft (1997). However, R.
M. Burton (in litt., 1999) has since told us that he has seen C. stagnalis on estuarine mud sheltered
by eyots by the River Thames in west London.

The absence of flowers in the Scottish population of E. parvula in 1999 is consistent with its
behaviour elsewhere in Britain and Ireland: Byfield (1999) notes that “in many localities the plant
flowers and fruits very poorly’, and this certainly appears to be true of the more northerly
populations: “the plant rarely flowers in Wales” (Rhind & Jones 1994) and the population by the
River Bann in Co. Londonderry “apparently does not flower” (Praeger & Megaw 1938).

Although the discovery of E. parvula in northern Scotland would not have been predicted from
its known British distribution, it is not surprising when viewed in a wider, European perspective. It
is perhaps most frequent in Europe around the coast of southern Sweden and the Baltic Sea,
including the Gulf of Finland and the southern part of the Gulf of Bothnia, where it grows at
higher latitudes than the recently discovered Scottish site (Hultén & Fries 1986; Mossberg 1997).
Indeed, its northernmost sites are further north than Shetland. It is, as Byfield (1999) writes, “an
inconspicuous plant that usually grows in an uninviting habitat” and it may well have been
overlooked elsewhere in Scotland.

ACKNOWLEDGMENTS

We are grateful to B. Goater, G. M. Kay & D. A. Pearman for their help on fieldwork in Scotland,
to R. Scott and P. Wortham for providing details of their later visits to the site, to R. M. Burton for
his observations on estuarine Callitriche stagnalis, to S. M. Walters for checking our material and
to D. A. Pearman for comments on a draft of this note. C.D.P. thanks Arto Kurtto, Pertti Uotila and
Leena Helynranta, who demonstrated swards of Eleocharis parvula at Uusimaa, Gulf of Finland,
on an Atlas Florae Europaeae excursion in 1997, thus increasing his awareness of this
inconspicuous species.

REFERENCES

BYFIELD, A. J. (1999). Eleocharis parvula (Roemer & Schultes) Link ex Bluff, Nees & Shauer (Cyperaceae),
in WIGGINTON, M. J., comp. & ed., British red data books I vascular plants, 3rd ed., p. 134. Joint Nature
Conservation Committee, Peterborough.

HULTEN, E. & FRIES, M. (1986). Atlas of north European vascular plants north of the Tropic of Cancer. 3
vols. Koeltz Scientific Books, K6nigstein.

MOSSBERG, B., STENBERG, L. & ERICSSON, S. (1992). Den Nordiska Floran. Wahlstr6m & Widstrand, Solna.

PRAEGER, R. L. & MEGAW, W. R. (1938). A Flora of the north-east of Ireland, 2nd ed. Quota Press, Belfast.

PRESTON, C. D. & CROFT, J. M. (1997). Aquatic plants in Britain and Ireland. Harley Books, Colchester. .

RHIND, P. M. & JONES, A. (1994). Brackish saltmarsh communities in the Glaslyn Marsh Trust Reserve. Field
Studies 8: 373-384.

RODWELL, J. S., ed. (1995). British plant communities, 4. Aquatic communities, swamps and tall-herb fens.
Cambridge University Press, Cambridge.

T. D. DINES
Rhyd y Fuwch, near Bethel, Caernarfon, Gwynedd, LL55 3PS

C. D. PRESTON
Institute of Terrestrial Ecology, Monks Wood, Abbots Ripton, Huntingdon, Cambs. PE28 2LS

NOTES Watsonia 23 (2000) 343

SUAEDA VERA FORSSK. EX J. F. GMEL. (CHENOPODIACEAE), SHRUBBY
SEA-BLITE, PRESENT IN ANGLESEY (V.C. 52), WALES

On the B.S.B.I. field meeting of 13 June 1999, T. C. G. Rich, M. and. J. Davidson and J. A. Green
discovered a new locality for Suaeda vera Forssk. ex J. F. Gmel., Shrubby Sea-blite, on Anglesey
(v.c. 52). Four large plants were found at the high water mark at the top of a stony beach on the
north side of the eastern end of Black Creek on the Bodior Estate, Rhoscolyn, Holy Island (grid
reference SH/293.757). The plants were to c. 1-3 m tall, were much branched at the base and
evidently old, and were just coming into flower and hermaphrodite. They were tucked into a very
sheltered rocky corner opposite a small stone fisherman’s hut and under low cliffs. It is no surprise
that they have not been seen before as they are hidden from view except in the immediate vicinity
and the site is on a private estate away from public footpaths. No further plants were found during
additional searches of the coastline in this area in June and August. A side branch was collected as
a voucher and has been deposited in the National Museum of Wales (NMW).

Suaeda vera is generally regarded as native around the coasts of South-east England from
Lincolnshire to Dorset, with occasional introductions with ships ballast in North-east England and
South Wales (for a recent map, see Stewart et al. 1994). The Anglesey locality is highly disjunct
from these other sites, but the information available to date does not suggest it was deliberately
introduced by man. Other typical saltmarsh species occurred nearby including Juncus maritimus
Lam. var. atlanticus J. W. White and Zostera angustifolia (Hornem.) Rchb., and no introduced
species other than Spartina anglica C. E. Hubb. were present in this pristine area. There are no
records of planting by the estate. No ripe seed was found even in August, and it is possible that the
four plants have originated by layering (cf. Chapman 1947). We thus regard this locality as the
first native site in Wales.

We presume that the plants have either arisen from long distance dispersal by sea water or
wildfowl, or are relict from a previously wider, more continuous distribution (it is certainly widely
distributed in Europe, Asia, Africa, and in Atlantic islands such as Madeira, Canaries and St
Helena, on the coast as well as inland). Ridley (1930) states that Suaeda seeds appear to sink very
speedily, but quotes no data for S. vera. To provide more information on seed floatation, 30 ripe
seeds were collected from bushes on Chesil Beach, Portland in September 1999 and placed in a
beaker of sea water, with or without calyces depending on how they had fallen off the plants. After
stirring to simulate wave action, the number of seeds floating were recorded at intervals for one
month (Table 1). Most seeds sank within 5 days, but one seed floated for at least one month. No
germination occurred in sea water. To test that the seeds were still viable after immersion in sea
water for a month, they were rinsed in fresh water and placed on damp tissue paper in a petri dish
and left to germinate. Five seeds (17%) germinated within two weeks but none thereafter. The
poor flotation time suggests that plants are unlikely to have arrived at this site directly by sea water
transport, and are more likely to have been brought by wildfowl which regularly winter on
Anglesey.

A number of other essentially maritime species with disjunct distributions also reach their
northern limit on Anglesey including Juncus capitatus Weigel, Asparagus officinalis L. subsp.
prostratus (Dumort.) Corb. and Rumex rupestris Le Gall. Frankenia laevis L., another south-

TABLE 1. NUMBER OF SEEDS OF SUAEDA VERA FLOATING IN SEA WATER
WITH TIME (N = 30).

Days Number floating (%)
0 30 (100%)
3 20 (67%)
4 8 (27%)
=) 5 (17%)
6 4 (13%)
10 1 (3%)
30 1 3%)

344 NOTES Watsonia 23 (2000)

eastern species of saltmarshes and rocks with a virtually identical distribution to S. vera, was found
nearby on Anglesey in 1965 (Roberts 1975); the discovery of the Suaeda suggests it could also be
native.

ACKNOWLEDGMENTS

We would like to thank Frank Roberts and the Bodior Estate for access to the private land.

REFERENCES

CHAPMAN, V. J. (1947). Biological flora of the British Isles. Suaeda fruticosa Forsk. Journal of ecology 35:
303-310.

RIDLEY, H. N. (1930). The dispersal of plants throughout the world. L. Reeve & Co., Ashford.

ROBERTS, R. H. (1975). Frankenia laevis L. in Anglesey. Watsonia 10: 291-292.

STEWART, A., PEARMAN, D. A. & PRESTON, C. D. eds. (1994). Scarce plants in Britain. Joint Nature
Conservation Committee, Peterborough.

i CsG: Rich
Dept. of Biodiversity and Systematic Biology, National Museum & Gallery, Cardiff CF10 3NP

N. BROWN
Treborth Botanic Garden, University College of North Wales, Treborth, Bangor, Gwynedd
LLS572RQ

A PROBABLY NATIVE AND REGENERATING POPULATION OF
TILIA PLATYPHYLLOS SCOP. IN BEDFORDSHIRE

Evidence that Tilia platyphyllos is native in scattered localities in England and Wales was
reviewed by Pigott (1981), where most of the sites discussed are on cliffs or steep slopes of hard
limestone or volcanic rocks in the Welsh borders, Derbyshire and Yorkshire and where annual
rainfall exceeds 800 mm. Since 1981 as many as 23 localities have been discovered on the chalk of
the South Downs in west Sussex and Hampshire (Abraham & Rose; unpublished report 1998).
These populations are usually of large old trees, or a mixture with younger trees, but there is
almost no regeneration. Quite characteristically in this drier part of England the sites are at the foot
of the north-facing escarpment of the South Downs where the trees grow close to the spring-line,
often at the head of coombes and along their flanks.

In 1993 Graham Bellamy, manager of the Barton Hills National Nature Reserve in Bedfordshire,
told me that there were three large lime-trees in Leat Wood on the steep east-facing slope of the
chalk coombe south of Barton-le-Clay (51°57'N 0°25'W; GR 52/08.29). Shoots were obtained
from the emergent crowns by climbing the trees, and proved to be of typical Tilia platyphyllos
Scop. subsp. cordifolia (Besser) Schneider.

The upper part of Leat Wood is dominated by beech (Fagus silvatica), which was probably
planted about 1830, but the steep slope is occupied by a mixture of ash (Fraxinus excelsior),
sycamore (Acer pseudoplatanus) and horse chestnut (Aesculus hippocastanum). There is patchy
regeneration of ash, beech and sycamore with scattered saplings of lime. The largest lime had a
basal diameter of almost 3-0 m. As an indication of its age it may be noted that this is as large as
the tree of T. platyphyllos which was planted in 1565 in the open and in a more fertile site at
Burghley House, Northamptonshire, and was therefore 450 years old in 1995. Leat Wood was
shown on the OS. map (1:10560 sheet 30NW) of 1891 (surveyed in 1880-81) and also on the pre-
enclosure map of 1778, where it is called “The Lead’.

Below Leat Wood, a stream flows northwards down the valley. Its source is two vigorous
springs at the head of the lower coombe at 52/089.295, and beside them grows a tree of
T platyphyllos in a typical natural habitat for the species. The tree originally had two trunks but in
1995 one was a stump and the other 18 m tall with a diameter at 1-3 m above the ground (dbh) of
1-09 m. A section cut from the stump gave a ring count of 173 but the outer 70 mm was decayed.

NOTES Watsonia 23 (2000) 3.45

From the average ring width in the adjacent wood this represents another 30 years, to give an
approximate age of 203 years. Having two trunks, the tree had probably been cut and regrown and
was therefore older.

Above the spring is the most remarkable feature of the site which is perhaps unique in Britain.
On the steep north-east facing slope at 52/088.294 is an area of woodland containing 32 trees of
T platyphyllos, mixed with ash and beech of similar size. Most of the trees are maidens (with a
single stem) and variation in diameter (dbh) is approximately normal with a mean of 0-31 m, SD +
0-09 m (March 1995). The immediate impression was that the trees were of one age and might
therefore have been planted.

On the pre-enclosure map of 1778 the area was south of the boundary of Leat Wood and shown
as “common pasture’. On the O.S. map it was in 1880-81 still treeless and shown as rough pasture.
In Dony (1953) a photograph (plate 20), which was probably taken about 1945, shows large trees
around the spring and all the slopes above are downland, while the particular slope has sparse
scrub above and below the prominent footpath.

The hills were part of the estate of Hexton Manor and the present owner Mr J. A. Cooper told
me that before World War 2 the hillsides were grazed by sheep, which were later confined to the
east of the sharp boundary of the scrub in Dony’s photograph. Parts of the scrub were cut but at no
time this century were trees planted on the slope.

Cores were taken in March 1995 with a Pressler corer from close to the base of the trunk of a
sample of lime trees. Ring counts showed that the trees were, in fact, of several ages. The oldest
trees (dbh 0-40—0-49 m) had 51-55 rings and therefore date from 1939-43, several trees (dbh
0-22—0-35 m) had 41-45 rings and date from 1949-1953, which is before the demise of rabbits
following myxomatosis, and the youngest tree sampled (dbh 0-14 m) dated from 1966. One of
several large hawthorns (Crataegus monogyna) dying beneath the limes dated from 1942. The
field-layer under the limes was sparse and included species from the original downland (Primula
veris) but consisted predominantly of woodland species dispersed by birds and mammals (Arum
maculatum, Geum urbanum, Hedera helix, Sanicula europaea). Mercurialis perennis, which was
abundant in Leat Wood and near the spring was conspicuously absent from most of the new
woodland.

The evidence that 7. platyphyllos is native at Barton is circumstantial and is essentially the age
of the oldest trees and their presence at a spring-head site. It must, however, be recognised that T.
platyphyllos sets plentiful fertile seeds and in most years seedlings are common in England, so that
it has the potential to regenerate and to be dispersed. Indeed, at Barton the colonisation of
ungrazed downland and the ability to spread and become established in suitable habitats is
dramatically demonstrated.

ACKOWLEDGMENTS

I thank Mr Graham Bellamy for his enthusiastic support and for cutting the slice of the stump, Dr
Jonathan Spencer and Mr Ian Evans for sampling the crowns of the old trees, Mr J. Ashley Cooper
for giving me information about the history of the site, Dr Eric Till for confirming the date of
planting of the great lime at Burghley. and Dr Francis Rose and Miss Frances Abraham for
allowing me to quote from their unpublished data.

REFERENCES

Dony, J. G. (1953). Flora of Bedfordshire. Corporation of Luton, Museum and Art Gallery.
PIGOTT, C. D. (1981). The status, ecology and conservation of Tilia platyphyllos in Britain, in SYNGE, H, ed.
The biological aspects of rare plant conservation, pp. 305-317. Wiley, Chichester.

C,DsPiGOrE
Greenbank, Cartmel, Grange-over-Sands, Cumbria LAI1 7SQ

346 NOTES Watsonia 23 (2000)
FURTHER BRITISH SPECIES OF RUBUS L. (ROSACEAE) IN NORTH-WEST FRANCE

Following the discovery in the Cotentin Peninsula of west Normandy of eight Rubus species
previously unknown outside the British Isles (Allen 1966), wider exploration of north-west France
as Well as further study of material already collected have resulted in the detection of several more.
A number of British species already known there have also been found to occur more widely.

In this second list first published records for mainland Europe are highlighted by an asterisk, as
before. Determinations assented to by A. Newton are indicated by an obelisk (7+) against the
locality in question. For départements renamed in recent years the previous name is adhered to, for
the sake of consistency with earlier records — by analogy with the British Isles vice-county system.
My specimens have all been lodged in BM.

R. nobilissimus (W. C. R. Watson) Pearsall Calvados: Bois de St.-André, Falaiset. New to
France, but collected in Belgium in 1970 by H. Vannerom, det. Newton (Vannerom 1998).

R. oxyanchus Sudre_ Calvados: a colony in wood at west end of Rochers des Parcs, Clécy. This
trans-Channel species seems much rarer in France than in England.

*R. plymensis (Focke) Edees & Newton Manche: margin of heath, La Pernelle, near Quettehour.
This Cornubian species has also recently been found in Guernsey.

R. pyramidalis Kalt. Extends westwards at least to central Brittany (Ille-et-Vilaine: Forét de
Paimpont).

*R. riparius W. C. Barton ex Newton Calvados: abundant both in heathy scrub at west end of
Rochers des Parcst and in Bois de St.-Clair, Clécyy. Only recently recognised in south Wessex,
before which it was believed endemic to north-west Wales.

*R. viridescens (Rogers) T. A. W. Davis Loire-Inférieure: patch in a boggy ditch, wood on west
side of Herbignac, det. Newton.

R. rubritinctus W. C. R. Watson Locally common in Cotés-du-Nord, where Rogers turns out to
have collected it in 1897 (BM). Very local in central Normandy.

R. hylophilus Rip. ex Genev. Rare. Calvados: foot of Rochers de Brisevieille, Cossesseville; Bois
de St.-André, Falaise. Cotés-du-Nord: Lande de Fréhel. Single bushes or patches in each case.

R. neomalacus Sudre One of the commonest species of the region at the mouth of the Loire, its
French headquarters, yet, puzzlingly, in Britain virtually confined to Surrey.

R. stenopetalus Lef. & P. J. Mueller Calvados: Bois de St.-André, Falaise.

R. adscitus Genev. Mostly thinly distributed. Seen in plenty only in certain woods and plantations
in Calvados, Sarthe, Maine-et-Loire and Cétes-du-Nord.

*R. thurstonii Rilstone Manche: Mont du Roule, Cherbourg, locally frequent on lower slopes‘.
Collected by Newton in 1984 near Quettehou, in the north-east of the Cotentin Peninsula.

R. vestitus Weihe Orne: Chateau d’Or plantation, Montrée, on clay. The nearest this basicole
seems to come to the granite massif covering most of north-west France.

*R. coombensis Rilstone Manche: Mont du Roule, Cherbourg, locally common in heathy scrubr*;
wood margin, La Glacerie Eglise, near Cherbourg’.

R. echinatus Lindl. Rare or absent north of the Loire valley, where it becomes plentiful from
Angers eastwards.

R. leightonii Lees ex Leighton Though seemingly absent from the Cotentin Peninsula proper.
locally common in Calvados, Cétes-du-Nord, Maine-et-Loire and Indre-et-Loire.

R. bloxamii (Bab.).Lees Increases southwards to become locally comon in Orne and Mayenne.
but apparently absent from Brittany.

NOTES Watsonia 23 (2000) 347

R. botryeros (Focke ex Rogers) Rogers The supposed first record from the European mainland
(Allen 1996), near Cherbourg in Manche, turns out to have been anticipated by Rogers in 1897 at
Dinan in Cétes-du-Nord+(BM).

*R. cantianus (W. C. R. Watson) Edees & Newton Manche: Bois du Mont du Roc, near
Cherbourgy. A surprising occurrence in this strongly Atlantic florula of this otherwise south-east
England species, unknown west of Wiltshire.

R. flexuosus P. J. Mueller & Lef. Not seen outside the Cotentin Peninsula.

R. insectifolius Lef. & P. J. Mueller Calvados: locally common on gorse-covered hillside,
Rochers des Parcs, Clécy. Apparently rare in north-west France,

R. asperidens Sudre ex Bouvet Calvados: one clump at east end of Rochers des Parcs, Clécy.
Apparently very rare in Normandy and absent from Brittany, coming in quantity only in Anjou.

*R. rilstonei W. C. Barton & Riddelsd. Morbihan: Lanmeur, 1969, H. Vannerom, det. Newton.
Cétes-du-Nord: common on heath margins, Lande de Fréhel*.

*R. tamarensis Newton Calvados: in three places near Clécy — locally common on north bank of
R. Orne at foot of Rochers de Brisevielle, Cossessevillet, Bois de St.-Clair, Meslay+; hedge
bottom by La Vigne, Culey-le-Patry.

R. tumulorum Rilstone Manche: Bois de Pépinvast, Val de Saire+. Second record for the Cotentin
and mainland Europe.

R. scaber Weihe Ille-et-Vilaine: Forét de Paimpont, one patch (new to Brittany’).

R. nemorosus Hayne ex Willd. Penetrates west to the Loire mouth, where it is common in the
fens of the Parc des Briéres.

R. transmarinus D. E. Allen The wider distribution claimed by Rogers, earlier doubted (Allen
1994), has proved to be the case after all: thinly distributed in Calvados, this is also the prevailing
member of Sect. Corylifolii in the coastal parts of C6étes-du-Nord (where it may be the plant
collected by Coilliot at Le Val-André which Sudre described as R. x namneticus).

Of the 208 species included in the monograph by Edees & Newton (1988) that were believed at
that time to be endemic to the British Isles, no fewer than 36 have now been detected on the
European mainland. Of these, 17 have been found as yet only in France and 14 in the Low
Countries (cf. Vannerom 1988). A further seven are also now known to extend to the Channel
Islands, which belong to the mainland geographically rather than to Britain.

REFERENCES

ALLEN, D. E. (1994). Four new Anglo-Sarnian species of Rubus L. (Rosaceae). Watsonia 20: 143-151.

ALLEN, D. E. (1996). British species of Rubus L. (Rosaceae) in the Cotentin Peninsula of Normandy.
Watsonia 21: 129-133.

EDEES, E. S. & NEWTON, A. (1988). Brambles of the British Isles. Ray Society, London.

VANNEROM, H. (1998). Wijzigingen aan de Rubus-inventoris (Sect. Corylifolii Lindl. en Sect. Rubus) in de
derde editie van de ‘Flora van Belgie’. Dumortiera 70-71: 45—49.

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

RUBUS ANGUSTICUSPIS SUDRE (ROSACEAE) IN SCILLY
In a recent revision (Allen 1997) of Rubus fruticosus agg. in the Isles of Scilly, v.c. la, the granite

archipelago 45 km south-west of Land’s End, 18 morphotypes were listed as occurring, of which
five appeared to be undescribed. Two of the latter, both members of ser. Hystrix Focke, are locally

348 NOTES Watsonia 23 (2000)

abundant. One of these, already known to be widespread in Brittany, has subsequently been
described as R. venetorum (Allen 1998); the other has now belatedly been recognised as an
unusually robust version of R. angusticuspis Sudre, a British endemic hitherto recorded only from
the southern end of the Welsh Marches, North Somerset, v.c. 6, and one wood in the Isle of Wight,
vce 0:

The discovery of this species in Scilly is of interest for three reasons apart from the major
extension to its known range that this represents. For a start, its local profusion on the two largest
islands, St. Mary’s and Tresco, and the robustness of its growth, in full sun no less than light shade
and despite the thinness of the soil and exposure to strong westerly winds, would seem to indicate
that it finds here conditions conducive to its optimal development. The other English and Welsh
populations thus have to be regarded as for the most part depauperate by comparison. The Scilly
plants, however, differ from the generality of those not only in dimensions - the terminal leaflet of
the stem leaves ranges up to 11 x 6 cm as against 8 x 5 cm given as the limit in Edees & Newton
(1988) — but also in two qualitative characters: in the specimens examined the styles were noted as
yellowish, not “red-based”’, and the petals instead of being “bright pink” are uniformly more or
less white. In this last respect R. angusticuspis in Scilly parallels R. mollissimus Rogers, another of
the islands’ Rubus species with a much more easterly range in Britain. It may be that the high
actinic content of the sunlight for which the islands are noted places pink flowers at a selective
disadvantage there.

A third point worthy of note is the coexistence in quantity in this comparatively remote island
group of members of three quite different Rubus florulas: R. iricus Rogers, otherwise all but
confined to the western half of Ireland, R. venetorum D. E. Allen, a species predominantly of
Brittany and R. angusticuspis, a member of the Severn Bay Florula (Newton 1980). This is one
further illustration of the value of Rubus for the finer discrimination of phytogeographical
affinities than most other biota are able to provide.

REFERENCES

ALLEN, D. E. (1977). Rubus L. (Rosaceae) in the Isles of Scilly: a revised list. Watsonia 21: 355-358.
ALLEN, D. E. (1998). Five new species of Rubus L. (Rosaceae) with transmarine ranges. Watsonia 22: 83-96.
EDEES, E. S. & NEWTON, A. (1988). Brambles of the British Isles. Ray Society, London.

NEWTON, A. (1980). Progress in British Rubus studies. Watsonia 13: 35-40

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

Watsonia 23: 349-358 (2000) 349

Book Reviews

A Rum Affair. Karl Sabbagh. Pp. ix + 223. Allen Lane, The Penguin Press, 1999. £16.99,
ISBN 0-713-99277-8.

Karl Sabbagh is a Cambridge graduate and a television producer but not a botanist. This is a book
about a botanist, about the unmasking by fellow botanists of fraudulent botanical records made by
Professor J. W. Heslop Harrison in the Hebrides during his pioneering researches there with
students from Newcastle University in the period 1935-51. In particular, it focuses on the part
played by J. E. Raven and his visit to Rum in 1948, after he had been let down by the distinguished
botanists that it was originally planned would accompany him. Raven found persuasive
circumstantial evidence of plants being fraudulently introduced which he wrote up in a riveting
manuscript that he brought back to his fellow dons at Kings College, Cambridge, where it was
hushed up for 50 years. John Raven, the kindest and most sensitive of people and an expert and
meticulous botanist found himself in the most invidious position. He alone of all those who had
doubted Heslop Harrison’s records had found evidence of fraud in the field, others just failed to
refind the plants and insects claimed by Heslop Harrison. Raven had been a conscientious objector
in the war, he would never have launched a personal vendetta.

Sabbagh has taken the trouble to research the nature of botanists, and field botanists in
particular, he perceives that “botany is very different to a love of cultivating plants. In fact, there
seems sometimes to be very little overlap” and, quoting from John Raven’s friend, Tim Clutton-
Brock, “my impression is that botanists are really fascinated with the details and the facts, whereas
zoologists like working in bigger, broader-brush, theoretical frameworks”. Heslop Harrison came
to zoology before botany. Sabbagh’s perceptions of us botanists, and his misconceptions, are
refreshing, though some of the misconceptions might have been avoided if his research had
included a day on a B.S.B.I. field meeting where he would have learnt an appreciation of the
endless scope for errors in fieldwork.

The book is a vignette on the social structure of British botany in the nineteen thirties and
forties. There are class prejudices, there is the Oxbridge-Redbrick divide, there are amateurs and
professionals, there is Rum itself as a personal fiefdom of the Bullough family. There are cliques
and rivalries but, transcending all, there is a culture in which there was a great reluctance to expose
a fellow scientist especially when he had a family of successful scientists following him. There is
much excellent biographical detail not only on the Heslop Harrisons and Ravens but also of other
prominent naturalists of the day, including Frank Balfour-Browne, “Willie” Clark, J. E. Dandy,
Dick David, E. B. Ford, George Taylor, Max Walters and A. J. Wilmot.

It is not clear if Sabbagh grasps why publication of an exposé was in a sense unnecessary. The
botanical grapevine was sufficient. The relatively few who had a need to know because it might
affect their research were tipped off. Inevitably, the insinuations sometimes suggested too black a
picture. Be that as it may, many of Heslop Harrison’s “dubious” records have since been refound.

Many a botanist has left a trail of dubious records behind him, and much trouble they have
caused, but fraud, as such, is usually thought to be rare. Sabbagh studies the motives for scientific
fraud, which he illustrates with other examples and suggests it to be much more common than
generally thought. Quoting Felix Franks “If ever the delicate balance between making a living,
searching for the truth, and obtaining the approbation of one’s peers is upset, then there is the
danger of deviant science”. Nowhere, however, does he suggest that advancing years could have
had something to do with it, or that some of the arrogance and dogma with which Heslop Harrison
ruled his university department could have affected him under the maxim “power corrupts,
absolute power corrupts absolutely”.

This book is a first rate read, you will not put it down until the end. And, if you are a botanist,
you will be left, fraud aside, with personal heart searching as you seek a humble approach to the
intractable intricacies of nature.

M. E. BRAITHWAITE

350 BOOK REVIEWS

A Naturalist’s Shetland. J. Laughton Johnston. Pp xii + 506. T. & A. D. Poyser. 1999. £27.95,
ISBN 0-85661-—105-0.

It is some 20 years since the first publication of Sam Berry and Laughton Johnston’s monograph
on “The Natural History of Shetland” in the Collins New Naturalist series. Since then, much has
changed in Shetland and much of this change has had a major effect on both the islands’ natural
heritage and our understanding of it. Of course this change had been largely anticipated at the time
of writing, as the oil industry was already having a tremendous impact on the social,
environmental and economic fabric of the islands. It is therefore timely that the state of the
environment and our knowledge of it should be revisited, and who better than one of the original
authors of the New Naturalist book.

The book is not simply a re-working of the previous book: indeed it has been thoroughly revised
and an enormous amount of new material added, especially information and understanding gained
from the many ecological and environmental studies that have taken place in the intervening years.
Although the extent and scope of the book remains as broad as it was 20 years ago, the structure of
the book remains largely unchanged (despite different chapter headings), moving through the
physical influences on the landscape; the history of colonisation; and then on to the nature of the
biological communities that make up Shetland’s natural environment. The new information is
crucial though; for the botanist, the publication of Walter Scott & Richard Palmer’s The Flowering
Plants and Ferns of the Shetland Islands, published in 1987 after the original New Naturalist
volume is a case in point, and Laughton Johnston has incorporated a substantial amount of recent
botanical work including the improved understanding of the Alpestria hawkweeds (now
‘protected’ under the U.K. Biodiversity Action Plan) and the many studies on the enigmatic, but
endlessly fascinating, serpentinite soils of the Keen of Hamar N.N.R. The final chapter of the
book, reflecting perhaps the time we live in, deals with issues of sustainability: issues previously
dealt with under the chapter devoted to conservation! The influence of the oil industry on the
natural history has been all-pervasive. In some cases the effect has been direct, in terms of built
development e.g. the giant oil terminal at Sullom Voe) and the effect of spilt oil on the coastal and
marine environments, especially from high profile incidents such as the Esso Bernicia and the
Braer. But the oil industry has brought wealth to the islands, which has been used to fund much
other change. The influence of grants to the agricultural industry is one of the most noticeable in
terms of its impact on Shetland’s semi-natural vegetation. Laughton Johnston deals with these
subjects effectively, and brings us right up to the present and the challenges that the islands (and
their environment) face as we start another century.

The book is a tour de force. Its format and presentation have been well thought out, and the
insertion of many new figures, sketches, and colour photographs give it a sense of quality that
justifies its price. Some of the photographs have been strangely cropped and appear too small, but
this rarely detracts from the overall feel of a real gem of a book, that will last well for at least
another 20 years. Buy it if you are at all interested in Shetland, or plan to go at some time in the
future. There’s everything in it that you’d ever want to know about Shetland and a lot more
besides!

A. DOUSE

British Red Data Books I Vascular Plants 3rd edition. Compiled and edited by M. J. Wigginton,
Pp. 465. J.N.C.C. Peterborough 1999. £30, ISBN 1-86107—-45 1-4.

It is 16 years since the publication of the 2nd edition of the Red Data Book for Vascular Plants,
and in this time there have been many field surveys, county Floras, and local red data books. The
previous slim volume was very much a basic factual account which stimulated more detailed work
and helped set the scene for better things to come. This book, with 465 textual pages, expands
significantly on the autecology, distribution and changes in populations of the species that have
taken place especially in the last 20 years. Comments are also provided on the European or world-
wide status.

BOOK REVIEWS efoul|

The introductory pages set out the background and context of the book and summarise the all-
important data sources. Without accurate record-keeping and meticulous checking, it would not
have been possible to produce a meaningful and rigorous account for each species, or the summary
tables. Accurate records also provide the essential platform for the analysis of numbers of
populations, spatial changes, and colonisation of new areas in a truly scientific, rather than
conjectural, manner.

There are 289 native or probably native species and subspecies that are considered to be
threatened or near-threatened in Britain addressed in this volume, compared with 317 in RDB 2nd
edition. This is due to the fact that 44 taxa have been excluded because they are non-native,
introduced, casual, escaped, varieties, hybrids or of dubious status. There are now 20 extinct
species as opposed to 19 in the 2nd edition; the most recent extinctions being Agrostemma
githago, Crepis foetida and Neotinea maculata. On the other hand species identified by Scarce
Plants in Britain as being rare have been added.

A significant change is in the revised I.U.C.N. threat categories, which have now been agreed
after much debate, and which aim to provide clearer guidelines for evaluating rarity. Before, there
were four categories and now there are eight. They are intended to help assess priorities for
conservation on an international scale.

For each species the text follows a similar format, beginning with habitat, associates and
reproductive biology. Historical and present geographical ranges are then described with
population details. Information on threats and conservation management is provided and, finally,
commentary on the distribution in a wider context i.e. European and/or world-wide. These
accounts were contributed by many botanists, both amateur and professional, but the majority have
been distilled by the editor, Martin Wigginton. His dedication and meticulousness have ensured
that much of the existing and new information has been incorporated into the book. Margaret
Palmer’s patience and guidance ensured that these efforts came to fruition. The valuable
contributions from other authors, following the excellent example set by the Nationally Scarce
Plants volume, ensure that the individual accounts are unique, and provide variety for the reader.
Some accounts are more erudite and detailed than others, reflecting the author’s depth of
knowledge and experience of their species. But this does not unduly detract from the work, as we
know there is still much to be discovered about our rarer plants, and it leaves room for additional
notes in the 4th edition!

Some of the most useful information is contained in tabular form e.g. the list of species
protected by the various international and national regulations, and a list of endemics, 71% of
which are threatened or near-threatened. Occurrence of species in different habitats shows the
importance of our coastal areas, calcareous grasslands and mountains. Planners will be delighted to
see the rare species listed for counties and unitary authorities.

In the conservation section, the many practical and positive steps that have been taken during the
past 20 years through the activities of English Nature’s Species Recovery Programme and
Plantlife’s Back from the Brink campaign, plus the Wildlife Trusts, the National Trust and the
Wildflower Society, are discussed. Threats from habitat destruction, pollution, collection and
global warming continue to exist but, on the whole, the decline of the rarest species has been
halted and the status of many has become more secure.

Recently, thoughts have been focused on genetic aspects, and this is a subject which has been
neglected botanically. Now there is an increased awareness of the conservation of genetic diversity
and a section summarises current aspects and highlights this as an area of work that needs
increased effort.

There are a few drawbacks to note. The volume is definitely too heavy to carry far and the maps
for some species could be considered a waste of space (those with one or two dots or numbers).
There is also the exclusion of Ireland and the Channel Isles (but there is the Irish Red Data Book
1988 to compensate).

I am well aware of all the hard work and collaboration that is needed to produce such a volume,
and I have no hesitation in saying well done to all concerned, on producing a work which is
informative, interesting to read and, on the whole, well laid out, and a substantial improvement on
the previous editions.

L. FARRELL

352 BOOK REVIEWS

Sir Joseph Dalton Hooker, traveller and plant collector. R. Desmond. Pp. 286. Antique
Collector’s Club, Woodbridge, with the Royal Botanic Gardens, Kew. 1999. Hardback £29.50.
ISBN 1—85149-305-0.

No British botanist can have started life with so favourably auspicious a pedigree as the younger
Hooker: a professor of botany who would shortly become Director of Kew, for a father, Dawson
Turner for a grandfather and James Dalton for a godfather. He would subsequently reinforce that
by marrying the daughter of Henslow and, when she died, the widow of Scotland’s leading
naturalist, Sir William Jardine. Various other scientific luminaries can be found slotted into that
dynasty on all its sides. Though very different in temperament from his father, it must have
seemed foreordained that he would have a no less illustrious career in the same line of work.

That Ray Desmond should write this major biography seems almost to have been foreordained
too. After many years as librarian and archivist of Kew he has produced in a steadily more
impressive stream a history of the India Museum, a history of the botanical exploration of India
and the long-needed, definitive history of the Royal Botanic Gardens themselves. Now this
massive accumulation of expertise has eventually all come together in this scholarly yet highly
readable, lusciously illustrated account of the man who will always be associated first and
foremost with the exploration of the flora of the Himalayas and the enrichment of our gardens with
so many of their finest rhododendron species.

As early as the age of five or six Joseph Hooker gave signs of having inherited the family’s
taxonomic blick by identifying the moss Bryum argenteum on a Glasgow wall. In addition to that,
though, he turned out to have enormous physical stamina, acute powers of observation, that
painstaking industriousness typical of so many Victorian ‘workaholics’ and, not least, a shyness
which bred a complete self-sufficiency. This combination of qualities ideally equipped him for the
role of explorer, the role which he himself must surely have recognised as the one most central to
his being. This took him first to Antarctica, as a naval surgeon in his early twenties; then came the
years in India — the most momentous experience of his life, to which Desmond fittingly devotes no
fewer than six chapters, the main bulk of the book — and finally, as late palliatives for his enduring
restlessness and love of travel, shortish trips to the Middle East, North America and Morocco. As a
result he acquired a wider knowledge of the World’s flora at first hand than any of his
contemporaries and was able to make important pioneer contributions to plant geography. In
particular, he supplied valuable ammunition to Darwin, his long-time friend and confidante, who
chose him as the first to hear of his conversion to the view that species are not immutable.

Unlike the elder Hooker’s achievements, which were pre-eminently in building institutions, as a
teacher and administrator, the son’s were essentially in his individual capacity as a scientist. Kew
was not necessary to his career except in providing him with a permanent base as a taxonomist,
and it was not indeed till he was nearly forty that that base became a secure one, when at last the
post of Assistant Director was created for him by the government. That was only a half-solution,
however, for he was not at all cut out by temperament for life as a public administrator: short-
fused and often brusque, he was tactlessly impetuous and inflexible in dealings with civil servants
and politicians where his father had been the soul of diplomacy. An academic position would have
suited him far better, if only he had not been precluded from that by a lifelong aversion to speaking
before audiences.

If the author has one weakness as a historian and biographer, it is in closing his eyes overmuch
to the frailties of his subjects. It is only in the very last paragraphs that he allows us to learn of
Hooker’s petty and shameful treatment of W. H. Fitch, the artist to whom he owed so much (and
some of whose superlative hand-coloured lithographs are reproduced in this book). And we are not
told at all of his unpardonable subverting of Welwitsch in that heroic man’s legal battle with the
Portuguese government over his Angolan collections.

Of more moment to readers of this journal, though, is the absence of any reference to Hooker’s
publications on British botany. That there were indeed any of these at all is, on the face of things,
surprising, for he would appear to have scarcely looked at a wild plant in these islands after his
student days were over. Rather, as H. C. Watson was moved to complain to J. G. Baker in 1866, he
seemed to hold “British botany in a sort of contempt”. It is all the odder, therefore, that he
undertook The student’s flora of the British Islands for Macmillan’s around that very time, a
period, moreover, when he was seriously behind with the Flora of British India and having to
carry the entire load of Kew’s administration (for Thiselton-Dyer only arrived in 1875 to relieve
him of much of that). It is known that his then solitary scientific assistant, Baker, whose

BOOK REVIEWS 353

knowledge of British plants was profound, contributed at least the excellent section on Rosa and
there must be a suspicion that he ghosted a good deal more. For the work certainly met with a
favourable reception, went to three editions and several reprintings and was still the preferred text
of some field botanists as late as the 1950s, when Clapham, Tutin and Warburg finally edged it
into extinction.

Hooker’s name was to become very much more familiar, though, as one of the duo to whom that
all-too-hardy evergreen, the Handbook of the British flora, was accredited from the 5th edition
onwards. The creation originally of George Bentham, who had knocked it off back in 1858, as
only that incredible man could have done, as a before breakfast relaxation (“for the ladies”, as he
once admitted), this brought to bear on the taxonomy of this country’s vascular plants the broad
species concept advocated by those, at Kew and elsewhere, who were struggling to impose order
on the almost overwhelming mass of novelties pouring in from newly-explored regions of the
globe. The ruthless ‘lumping’ that this approach involved conveniently resulted in a much
simplified text which held a particular appeal to beginners. But by lending his authority to what
amounted to a dismissal of years of critical work by those who knew our flora far better, Bentham
did serious and lasting harm to British field botany. On the credit side, however, was the equally
lasting influence of the companion volume of illustrations by Fitch and Worthington Smith, first
introduced in 1880. Those black-and-white figures, which cried out to be painted in, were to send
hundreds over the years scouring the British Isles on a lifetime quest to register by that means the
finding of as many as possible of the species depicted — as a substitute for the herbarium
specimens which would previously have served as trophies.

Again, this was an incongruous publication for Hooker to have taken on, though it is not at all
clear whether its editing required much from him in the way of either work or knowledge.
Probably he did so in part out of loyalty to Bentham’s memory and in part because in his
retirement he felt in need of the money that this even steadier seller could be relied on to continue
to bring in. It is ironical, and sad, that so misguided a work, in which his involvement may well
have been tangential at most, outlasted by far everything else that he published and served to keep
his name before the public. For a man who ranks as one of Britain’s all-time botanical greats it was
very far from being an appropriate memorial.

D. E. ALLEN

Atlas Florae Europaeae. Distribution of Vascular Plants in Europe. J. Jalas, J. Suominen, R.
Lampinen & A. Kurtto, eds. Committee for Mapping the Flora of Europe & Societas
Biologica Fennica Vanamo, Helsinki, 1999, vol. 12 Resedaceae to Platanaceae, pp. 1-250.
ISBN 951—9108—12-2.

Volume 12 is the latest in the series from the Atlas Florae Europaeae Committee. As I shall
explain later, it is much more than a set of maps, although there is no shortage of these in the
present work (343 in all). Although eleven families are dealt with altogether (Resedaceae,
Sarraceniaceae, Droseraceae, Crassulaceae, Saxifragaceae, Parnassiaceae, Hydrangeaceae,
Escalloniaceae, Grossulariaceae, Pittosporaceae and Platanaceae) most of the volume is taken up
with treatments of Sedum and Saxifraga. Taxa are mapped down to the level of subspecies, with
spots and other symbols indicating presence/absence, extinction/probable extinction, native/alien
status, etc. A cut off date of 1930 is used to distinguish recent from older records. This does not
give an especially accurate picture of present-day distribution, such has been the pace of
extinctions in some taxa, although even with this criterion the vulnerability of some taxa is clearly
evident, e.g. the chronic decline of Saxifraga hirculus across much of its continental distribution.

The authors/editors of the respective accounts have taken the opportunity to revise the
treatments provided in Flora Europaea (editions 1 and 2) and, at least as far as Saxifraga is
concerned, they have included not only necessary nomenclatural changes, but also new taxa that
have been recently recognised or discovered in places as far apart as Spain and northern
Scandinavia. In many such cases a certain narrowing of the species concept can be detected from
the earlier Flora Europaea account. In my view, this can often be a good thing, because scrutiny of
the resulting distribution maps reveals interesting patterns of geographically correlated variation
that would otherwise have been obscured. Analysis of these patterns from a phylogeographical
perspective is now possible using the various molecular techniques that are available, and by this
means we learn more about how our flora has evolved and become distributed.

354 BOOK REVIEWS

Apart from the detailed maps, the notes on distribution, the revised nomenclature and updated
taxonomy, what else have the authors/editors provided for us? The answer is an extraordinary
amount of information by way of literature references not only to the nomenclature and taxonomy
but also to studies of biosystematics (which here includes almost any aspect that has a bearing on
variation or relationships), with chromosome numbers being singled out for detailed attention. The
treatment of chromosome numbers deserves special praise because it is both comprehensive (citing
a list of the countries from which the counts were made) and critical (indicating aberrant,
problematic or otherwise erroneous counts), and includes references to the original literature.
Speaking as someone who has attempted such a review of the genus Potamogeton, I know just
how much work is involved in this task alone.

In short, the authors and editors are to be congratulated on a most impressively detailed and
accurate piece of work which will join its predecessors as an essential reference for anyone with an
interest in plant geography, evolution, floristics and conservation.

R. J. GORNALL

Flora of Cornwall. Atlas of the Flowering Plants and Ferns of Cornwall, with notes on some
species recorded on the Isles of Scilly. C. N. French, R. J. Murphy & M. G. C. Atkinson. Pp. 400.
Wheal Seton Press, Camborne. 1999. £40, ISBN 0—953461 3-0-0.

The need for an entirely new Cornish Flora has been keenly felt for some while. The present work,
unlike its predecessors, is an Atlas. It is based on systematic tetrad recording by a small team of
volunteers begun in 1987, along with a wealth of old and not-so-old records gleaned from other
sources. The scope of the Flora is formidable, with entries for some 2,600 vascular plant taxa. Its
treatment of aliens and critical groups like brambles and dandelions is particularly impressive.
Many sub-taxa not mentioned in Stace (1997) are included, though the taxonomic value of some of
these must be questionable. (What, for example, is Festuca ovina var. hispidula?)

The volume is in A4 format, with the usual introductory sections covering geology, soils,
climate, vegetation history and so on, plus descriptions of the main plant habitats and essay-length
pieces on four regions (Bodmin Moor, the Lizard, West Penwith and the Isles of Scilly). The
habitat/vegetation accounts are detailed and well-rounded but, rather oddly, only “wetlands” and
heaths are described in terms of their NVC communities.

The book attempts to deal with the whole of Cornwall although, as the authors admit, coverage
of the Isles of Scilly is considerably less thorough than the rest. The islands are (inexplicably)
omitted from the distribution maps, which means that for species with no mention of Scilly in the
text one is left not knowing whether they occur there or not. For example, on p. 58 Ranunculus
bulbosus, R. parviflorus, R. sardous and R. sceleratus are all found on Scilly, but (unlike
R. repens) are not listed as such (and incidentally R. marginatus, given as “thought to be extinct”,
is still extant on St Martins). The introductory essay on the Isles of Scilly (p. 31) lists certain
“missing” plants that are common on the mainland, including at least two (Stachys officinalis and
Dipsacus fullonum) that do occur there. This reviewer, for one, ended up wondering whether the
Isles of Scilly would have been better left out of the book altogether.

The species accounts provide useful summaries of habitat preferences, conservation status,
distribution and abundance. Tetrad dot maps - showing old (pre-1980) and recent (post-1979)
records - are placed within the text, and for all the less common species there are lists of selected
localities. Production constraints have meant that maps are limited to four per page. This is not a
problem when the unmapped species are rarities with all their localities listed in the text, but it is
slightly annoying that some “less -rare” plants are without maps too (e.g. Cerastium
semidecandrum, Ranunculus sardous, Picris hieracioides and Persicaria amphibia).

Assessments of “conservation status” are based on Scarce Plants in Britain (1994) and,
unfortunately, the badly out of date 2nd edition of the Red Data Book (1983) rather than the 3rd
edition (1999). Readers should therefore take the categorisation of rare species (and the maps on
p. 6) with a hefty pinch of salt! The “locally scarce” list on p. 381 is an updated version of one
produced for the Red Data Book for Cornwall and the Isles of Scilly (1997), but the rationale
behind the selection of species is not stated and the omission of Brachypodium pinnatum, Carex
disticha, Catabrosa aquatica, Neottia nidus-avis and Trifolium fragiferum, for example, is hard to
understand.

BOOK REVIEWS 395

Floristic similarities and differences with neighbouring Devon, and beyond, would perhaps have
merited further commentary and analysis. To anyone living “up-country” it must seem strange that
in Cornwall Berula erecta, Briza media, Dactylorhiza fuchsti, Hordeum secalinum and Leontodon
hispidus are all extremely rare, while Valeriana dioica and Limonium vulgare (to name but two)
are missing altogether. Each county has its own special quirks, of course, but Cornwall seems to
have more than its fair share!

Even the most thoroughly researched Flora will fall short of being the last word on its subject.
As the authors rightly say, “this Flora ... should be considered as part of an ongoing study” (p. 1).
It is an important milestone that will, in its turn, stimulate a new wave of botanical recording in the
county. And, yes, I am pleased to report that there are plants out there still waiting to be
discovered: a brisk walk along the coast between Millandraeth, Looe and Portnadler Bay in
October produced a list of 200-odd species, including new tetrad records for Atriplex
portulacoides and Verbena officinalis, and post-1979 “firsts” for Asplenium marinum and
Geranium rotundifolium.

At £40 this Flora is not cheap, but as a turn-of-the-century guide to the botanical delights of this
strange and wonderful county it is probably indispensable. Despite its few shortcomings, it stands
as a fitting tribute to the enormous effort of all those involved in its production. Anyone with more
than a passing interest in Cornwall’s plants should make sure they get hold of a copy.

S. J. LEACH

Flora of Cornwall on CD-ROM. Cornwall Business Systems. 1999. £40.00 from CBS House,
Albany Road, Redruth, Cornwall, TRI15 2HY

By producing the Flora of Cornwall on CD-ROM format, it is possible to include much more than
appears in the book. All of the introductory chapters and appendices can be read on screen if
desired, but the bonus is that the database on which the Atlas section of the Flora is based is also
included, from which maps can be generated, including species not mapped in the Flora. The CD-
ROM database also contains some records extra to the Flora. Many more photos than could be
produced in the book are also included.

Species are searched for using Latin or common name lists arranged alphabetically. On
selection, the species caption from the flora is reproduced, one or more photos of the species
appear, if available, the map is generated and the records listed. The map can be generated on one
of a choice of six bases including a geological base, at one of four sampling scales, 10-km, 5-km,
tetrad and 1-km scales. The records for the species are listed giving grid reference at tetrad level,
locality, date and recorder(s) and can be ordered by grid reference or year. Some limited querying
of the records can be undertaken on a date basis, such as records for a particular year, before or
after a chosen year, or for a date range.

A gazetteer of Cornwall is included, searchable by place name or by grid reference, and
pinpointing places on the map. This links to the Flora database, making it possible to list the
records for a particular 1-km square, or to produce a printable species list for the square. Like the
Flora records and map, these lists can be limited to particular date choices.

Other add-ons of interest are a tourist map of the county and photographs and information on
local landmarks. The species photographs can be searched and viewed on their own.

Using a recent PC with a fast CD-ROM, the map and records were generated fairly quickly for
the rarer species, less than 15 seconds for species such as Polygonum maritimum or Rumex
rupestris, which have about 100 or less records. However for commoner species, there is some
waiting time involved; to generate Agrostis capillaris, listed as the 100th most commonly recorded
species, took 90 seconds if the records were listed by year or 160 seconds if listed by grid
reference. Other than a few error messages on installation, which were notified with this review
copy, the only problem some users may have is resetting their usual display resolution on use to be
able to view the CD-ROM full screen.

As the first county Flora to be produced on CD-ROM as well as traditional book format, this is
surely an indicator of the future. This CD-ROM will be invaluable for any botanist regularly
requiring more detail than can be gleaned from the printed media.

A. STEWART

356 BOOK REVIEWS

Britain’s Rare Flowers. Peter Marren. Pp. xv + 334. Poyser Natural History, London. 1999.
£24.95, ISBN 0-85661-114—X.

Over a decade ago a book published with the title Wild Flowers in Danger caused widespread
concern because it was mainly a guide to where to find our rarest plants and the book itself added
to their danger. So, you might expect Britain’s Rare Flowers would take you down the same path
to the exact spot — but you would be wrong. Peter Marren has written a celebration of our 300
rarest species bringing them to life along with all those botanists, past and present, who have found
them and enthused about them over the last four centuries — a book for the armchair rather than the
field.

What the author set out to do, and achieved so triumphantly, was, as the American writer John
Burroughs said of Gilbert White, ‘to seize the significant and interesting features and put the
reader into sympathetic communication with them’. He takes you on a journey, first to meet the
fathers and elders of British botany, from William Turner to George Claridge Druce, who
discovered our flora, and then to areas like the Lizard Peninsula, Avon Gorge, Breckland, Upper
Teesdale and the Scottish Highlands which hold the highest concentrations of rarities. During the
journey he manages to convey the excitement of discovering or rediscovering a rare plant in a wild
place even after a tiring day in ghastly weather.

Later he looks at individual species and considers the value of research in assisting their
conservation. He can be wise and challenging and points to a growing gap between theoretical
scientists and practical conservationists in the last 30-40 years (partly a direct result of the
disastrous break-up of the Nature Conservancy Council). However, his admission that he is not a
scientist perhaps explains his ambiguity about the effects of collecting and a failure to appreciate
that the picking of daffodils can be beneficial and that banning it has caused decline. On the other
hand his non-academic view that long-established introductions like the spring crocus and spring
snowflake deserve protection and were properly included in Eds 1 & 2 of British Red Data Book
1: Vascular Plants and should not have been excluded, as they were in Ed. 3, is surely to be
welcomed.

The non-scientific background also shows in his failure to appreciate variation within species
e.g. that there are native and introduced forms of sainfoin, the former in the ancient turf of the
Wiltshire Downs and only the latter brought in as a fodder crop in the 17th century, or that there
are native forms of weed species on beaches and river banks and others which have come in as
followers of man. Elsewhere he gets into a muddle with Batrachian Ranunculus hybrids placing
R. aquatilis x tripartitus in the New Forest where the taxon is R. omiophyllus x tripartitus (R. x
novae-forestae S. D. Webster).

It is the mixture of fact and anecdote which make the book so palatable and informative but, as
these examples show, it cannot be used as an entirely reliable source. There is a wicked story of
one of the 14 remaining wild cotoneasters on the Great Orme being cut down by ill-trained
conservationists but the truth is that there are only seven native bushes and none of these or any
translocated plants have been destroyed. Inaccuracies have crept into the captions (a non-botanical
picture-editor?) so that Welsh botanists will be surprised to learn that Chamaemelium (sic) nobile
is ‘now restricted to scattered localities in southern England’ when there are 8 10-km square
records on their coast north to Lleyn. The photo of Lundy cabbage, Coincya wrightii, given as on
p. 121 in the index turns up on p. 124 with a caption reading ‘two kinds of beetles seem to eat
nothing else’ whilst the adjacent text names three.

The index is the worst feature of the book — totally user-unfriendly. It is arranged in alphabetical
order of English names starting with -the first word so that to find references to our mountain
saxifrages you need to look under Alpine, Drooping, Marsh and Tufted, but there is no
consistency — both native Limes are together! Moreover, recommended English names are not
always used: to find Goldilocks Aster, Aster linosyris, you would need to know that the Marren
name is Hair-leaved goldilocks. There is no alphabetical list of Latin names.

So if you want reliable facts about Britain’s rare plants (and no index at all!) turn to Ed. 3 of the
British Red Data Book (Wigginton 1999 see Review p. 350), but it will not compare as a good
read with Peter Marren’s compelling story — but regard some of that as fiction — and the index is
friction. ;

F. H. PERRING

BOOK REVIEWS 357

Vegetation of the British countryside — the Countryside Vegetation System, Ecofact Volume 1.
R. G. H. Bunce, S. M. Smart, H. M. van de Poll, J. W. Watkins & W. A. Scott. Pp. 224. Dept. of
Environment, Transport and the Regions, London. 1999. Price £48.00. ISBN 1—85112—155-2

Measuring Change in British vegetation, Ecofact Volume 2. R. G. H. Bunce, S. M. Smart, H. M.
van de Poll, J. W. Watkins & W. A. Scott. Pp. 144. Dept. of the Environment, Transport and the
Regions, London. 1999. Price £10.00. ISBN 1—870393-47-3

Ellenberg’s indicator values for British plants, Ecofact Volume 2 Technical Annexe. M. O. Hill,
J. O. Mountford, D. B. Roy & R. G. H. Bunce. Pp. 45. Dept. of Environment, Transport and the
Regions, London. 1999. Price £5.00. ISBN1—870393-48-1

One of the biggest problems faced by distribution ecologists and conservation campaigners alike is
that the British vascular plant flora (impoverished though it is by continental standards) appears
too vast in terms of species to survey quantitatively. Where is the plant equivalent of the common
bird census, plant conservationists cry — well, this may be it.

These three volumes are the first major publication resulting from the Countryside Survey: the
results from the Countryside Surveys of 1978 and 1990 undertaken by the Institute for Terrestrial
Ecology have been analysed and compared, giving for the first time a quantitative, objective and
repeatable survey of the vegetation of the British countryside. The data are collected from 508
1-km squares containing over 13000 plots, selected by stratified random sampling so that the full
range of landscape types are included. The data collected include species lists with abundance
from quadrats and the results have been analysed in a multitude of ways. So, what do we get in
these three publications? To start with, 37 groupings of plants by their ecological preferences, and
100 broad vegetation classes called the Countryside Vegetation System or CVS. Does this replace
the National Vegetation Classification (NVC)? No — the two systems are very different — the NVC
covers semi-natural plant communities in detail and the sample plots were selected deliberately
rather than randomly; the CVS aims to cover the vegetation we encounter in the wider countryside,
with the random samples allowing statistical analysis for monitoring change in the British
vegetation.

In Volume 1, there is an abundance of coloured distribution maps of vegetation classes, pie
charts and histograms of landscape associations, soil and land cover, together with comparisons
with and relationships to the NVC and also Grime’s plant strategy characterisations (based on
three basic strategies — competitor, ruderal or stress-tolerant, or combinations of these three).
Volume 2 provides lists of the vegetation classes and indicator species groupings. There are even
Statistics on the decline of butterfly (larval) food plants and effects on lowland farm birds. The
second volume is also supported by a technical annexe containing re-calibrated Ellenberg values
for the British flora, which I have already found invaluable. If you haven’t encountered Ellenberg
values before, these constitute a set of ecological indicator values based on a plant’s tolerances of
temperature, light, pH, etc. and are particularly useful in attempting to quantify some of the vaguer
aspects of descriptive plant ecology.

Overall, the amounts of data contained within are staggering, perhaps even overwhelming, but
the project is well on its way to confirming what many of us have suspected for some time — that
nitrogen-loving competitors are on the increase at the expense of less competitive species. There is
no doubt that the data can be used as serious campaigning tools for changes in modern agricultural
methods, but my impression is that these messages are almost lost in the mass of information;
hopefully we will see a series of papers published as extracts from these reports. The programme is
being repeated as part of Countryside Survey 2000, so we can expect further updates on the state
of British vegetation, in addition to Volume 3, Causes of change in British vegetation (in prep.).
Following Countryside Survey 2000, it will be interesting to see if the periodicity of the surveys is
right. The point of monitoring is to detect change and ten years may tell us of change and of loss —
but will it give us enough time to do anything about it?

In the introduction to Volume 1, the NVC, the Phase I Handbook and the Flora Britannica are
listed as essential items for the ecologist’s tool box. While I might take issue with the inclusion of
Flora Britannica, I would suggest that these volumes are essential reading for any professional
plant ecologist, especially those involved in plant conservation. However, at over £60 for the three
softback volumes, together with the dense technical content, it will be the truly dedicated amateur
botanist who dips a toe into the Countryside Vegetation System.

S. J. WHILD

358 BOOK REVIEWS

The liverwort flora of the British Isles. Jean A. Paton. Pp. 626. Harley Books, Colchester. 1999.
£52.50, ISBN 0—946589-60-7.

Liverworts in the British Isles (BI) grow in an almost continuous range of habitats from the
lowlands to the highest mountain summits. Of about 6,000 species known in the world, the number
in these islands, 296 species, two subspecies and five varieties, seems quite insignificant, but they
comprise nearly 70 per cent of the European liverwort flora, and their richness and diversity in the
western parts of Britain and Ireland is of great international significance.

This is not the first liverwort book for the BI to be produced this century; but it is a definitive,
modern work. The pages are approximately A4, the text is in two columns and each of the 314 text
figures contains many individual drawings, all by the author. After the preliminaries an
Introduction of 21 pages contains sections on Collection and Preservation, and Examination of
Material. One on Habitats and Distribution lists species of diverse habitats and then species
characteristic of five more specialised habitats. There are sections on Conservation and on how to
use the text and figures. Ending the book is a useful illustrated Glossary, a list of Vice-Counties of
the British Isles with a map, a Bibliography and an Index.

The bulk of the book is the Flora wherein every taxon at present known in the BI is keyed out,
described and illustrated. Full descriptions, even including unpublished chromosome counts, are
followed by statements on habitat, ecology, altitude, associates and distribution both within the BI
and without. Then there are notes on distinguishing features, comparisons with other species and
observations on morphological variations. Each taxon has a fine Figure or Figures which
accompany the descriptive section and illustrate everything one would want. For example, that of
the common liverwort Cephalozia bicuspidata which is *... in almost any moist or wet, shaded or
insolated habitat ... from sand, loam, peat ...; also on rotting wood ...’, has over 60 individual
drawings which include several habit figures (female, male and sterile), leaves, cells and stem
sections.

Primarily an identification manual, its importance to those studying plant communities in the BI
and Europe will be profound and cannot be over-emphasised: the rich accounts of the ecological
requirements and associated species were gathered from the author’s intimate field studies of the
plants over many years. With the excellent, all-encompassing illustrations, keys and full
descriptions, this book is a highly significant event in the study of European liverworts.

Through the publication in the 1900s of several important books on mosses and liverworts in the
BI, and an active British Bryological Society, our bryological flora is probably the best known in
the world. It is because of the intense interest thus aroused and of the expertise of dedicated
bryologists like Jean Paton that species are still being added to the British and Irish lists, some of
them new to science. This hardback with covers bearing splendid colour photographs is not for the
pocket: it weighs over 2:5 kg. Harley Books has done well to add this attractively produced work
to their already distinguished list.

Jean Paton is to be congratulated on a splendid achievement. This is a veritable tour de force
that will compel the respect and admiration of bryologists world-wide and become the most used
liverwort book in the British Isles and Europe in the new century.

A. R. PERRY

Watsonia 23: 359-372 (2000) 359

Obituaries

JOAN WENDOLINE CLARK
(1908-1999)

Joan Clark (née Rust) passed away on 6 July 1999 at Fort William, Inverness-shire. She had lived
in the area for some 60 years and was known and loved by all sectors of the wider community.
During the last 25, she developed a deep interest in the botany of North-West Scotland and the
Islands.

Joan was born on 6 October 1908 at Benholm, Kincardineshire, the only child of Wendoline and
John Rust, City Architect of Aberdeen. Her father died when she was ten, and Joan and her mother
went to live in Worthing, Sussex, to be near her mother’s family. She continued her education in
Worthing before going to a ‘finishing’ school in Switzerland. As a result she was good at
languages, and on returning to England went to a college in Brighton to study French, German,
and the traditional ‘Shorthand and Typing’. After working in the Foreign Office in London, and in
the British Embassy in Paris for two years she came back to London, where her Scottish roots
surfaced and she joined the newly formed Scottish Nationalist Party. She was a staunch supporter
of the SNP until her death, attending annual conventions almost without fail until 1998. In 1932, at
the London meetings, she met Branch Secretary Alastair Clark, (a Glaswegian of Mull origins)
who was to become her future husband; they were married in London in April 1935.

They moved to Glasgow where Alastair set up a fur-trading business but the outset of World
War II in 1939 severely restricted his work. They decided to move north to Loch Leven where
they bought and successfully ran the Loch Leven Hotel, a hostelry well-known by many a traveller
botanist who had queued for the Ballachulish ferry! Here they were the ideal Highland hosts,
making many life-long friends, both locally and from those who came to stay at the hotel during
the next 19 years. After the untimely death of Alastair, in 1959, Joan sold the hotel and moved to a
bungalow. Tigh Solais, built on nearby land they had owned.

I first met Joan in 1972 when she came to Kindrogan Field Centre to the first Sedges Course I
had been asked to tutor. There were only four students: a very fit sixth-former preparing for
university (with whom I just about coped), two ‘B.S.B.I. Ladies’ (who knew much more about
sedges than I did!), and this quiet Scottish soul who had recently joined the Wild Flower Society
and wanted the challenge of a critical group. Her advancement to Valhalla was being monitored by
Mary MacCallum Webster (which Joan later admitted was a wonderful, albeit terrifying training)
and Joan was determined to be a model student. We were two days into the course before she
produced a walking stick and timorously admitted she was waiting for a hip replacement! I learnt
afterwards that she thought I might send her straight home if it had been apparent from the outset.

When we came to a day walking up Lawers Burn, she struggled to the weir where we had lunch.
I took her aside and said I felt she might find the scramble through deep heather and bog to the
Yellow Corrie (then in the descending cloud) to see the Carex microglochin, too much. I could see
her disappointment but she, seeing my real concern, said “I will sit here, and will not move”. The
rest of us went on, and having found the species we had come to see and dropped back over the col
to return to the Burn, through the mist Joan Clark appeared lifting one leg at a time over the deep
sedge litter. I could not disappoint her again and we went up to see the ‘new’ species and joined
the other three at the Burn for tea.

This determination to ‘complete the course’ and to do as good a job as she could was, I was to
find out, the hallmark of Joan’s contribution over the next 20 years to Scottish botany. She had
‘graduated’ from the W.F.S. with a sound understanding of Scottish wild flowers. As a W.F.S.
‘interne’ with ‘Mary Mac’, Joan had offered to type Mary’s MS for the Flora of Moray, Nairn &
Easterness (published in 1978). This took many weeks but Joan persevered, and learnt much from
the exercise.

Joan joined the B.S.B.I. in 1977. She took full advantage of the Society’s Specialists and soon
realised the value of voucher specimens and pressed material for checking. Her eldest daughter
Sine lived out in N. Uist, and Joan made regular visits to this land of low-altitude nutrient-rich

360 OBITUARIES

lochans and extensive coastline. With her own little Morris Minor and son-in-law Archie’s boat,
she covered the island. Water plants became a special interest and she sent much material to J. E.
Dandy at the Natural History Museum. He was once chiding me for not taking the trouble to ‘float
out’ a grass-leaved Potamogeton, saying there “were few botanists of the calibre of Joan Clark and
Dorothy Cadbury, when it came to collecting pondweeds”. As some will realise, this was indeed a
great compliment to Joan.

With the pondweeds sorted out Joan took to other specialist collecting, sometimes ferns when
we were looking for Dryopteris expansa, and then, in a big way, dandelions, where she again took
great trouble in arranging the plants in the press and arranged her visits to be there in May. By this
time she was in regular correspondence with the R.B.G. in Edinburgh and sent many herbarium
specimens to the Scottish National Collections. Rubus and Hieracium were other genera that
received her careful attention. Her fieldwork in Uist was further encouraged by Richard Pankhurst
and others at the “~BM’ when writing the Flora of the Outer Hebrides and her contribution was
considerable.

When John Cannon relinquished the v.c. Recordership for Mull in 1978, Joan Clark took over
v.c. 103. She took her stewardship seriously and made extensive annual visits. We encouraged her
to maintain our mini-vice-counties as a recording base on Mull and she added many new records.
She also took to recording plants of Coll and Tiree seriously, especially the former where she
made several friends through her regular annual visits.

Although she was an Aberdonian Scot from the East, she took to the West and its islands with
eagerness. Being naturally friendly she wanted to chat with people she met and in the Islands she
realised this meant speaking Gaelic. She studied diligently and passed her O-grade. She quickly
saw that vernacular Gaelic names given to plants varied from area to area and made a plan to
collect these as widely as possible. Between 1975 and 1985 she circulated lists of what names
were in the literature, asking for additional ones in use in her contacts’ area. She was asked by
Richard Pankhurst to provide names for the Hebridean Flora and she used her knowledge of
Gaelic to coin names for taxa not 1n everyday use. For this she enlisted the help of another Uist
man, Ian MacDonald (now Director of the Gaelic Books Council) and between them they came up
with a definitive master list for all vascular plants growing in Scotland. This 20-year labour-of-
love (Gaelic Names of Plants - Ainmean Gaidhlig Lusan, was finally published (at her own
expense but with some help from the Gaelic Book Council and the B.S.B.I.); although Joan saw it
all, including the coloured cover in final proof, in spite of all stops being pulled out by the printers
it reached Tigh Solais seven days after her death.

Joan will be greatly missed by many in Lochaber. She was a devoted mother, grandmother and
great-grandmother to Sine, Anna and Eilidh and their respective families. She was also a
wonderful friend and correspondent to many a botanist who will mourn her passing and remember
her with warm affection.

A. C. JERMY

DAVID EDWIN COOMBE M.A., Ph.D.
(1927-1999)

Dr David Coombe died in Cambridge on 28 June 1999. Although his name will be known to many
B.S.B.I. members for his studies of the Lizard peninsula, and of the British and European species
of Trifolium, few will be familiar with the wide range of his interests and the depth of the
scholarship with which he pursued them.

David Coombe was born in Bath on 9 March 1927, the only child of Charles William and
Phyllis Coombe (née Weaver). His father, a ‘motor cycle specialist’, ran a small workshop. He
won a scholarship from South Twerton primary school to King Edward’s Grammar School, Bath,
and then went on to Christ’s College, Cambridge, in 1945. After gaining a first-class degree and
Ph.D., he worked in the Botany School, University of Cambridge, for the rest of his career, being
appointed a University Demonstrator in 1952 and a University Lecturer in 1957. He was elected a
Fellow of Christ’s College in 1951 and subsequently served his college in many capacities,
including Vice-Master. As a bachelor he was able to live in college rooms until he retired
prematurely in 1989, after a prolonged period of ill-health.

Coombe’s interest in the British flora developed at an early age. He was given his first Flora,
Skene’s Flower Book for the Pocket, on his 10th birthday. He was able to botanise in Dorset as

OBITUARIES 361

well as in his native Wiltshire in the 1930s, as at least 20 of his mother’s relatives lived near
Portland. “Several of the couples were childless,” he recalled in 1991 in a letter to David Pearman,
‘and I was taken out in turn by these kind people on long summer holidays - with all sorts of
rituals - the train from Easton to Sandsfoot, and a picnic at “Vetch Corner’ (where Vicia bithynica,
V. lutea and Lathyrus nissolia have still been so abundant in recent years).” By 1939, aged 12, he
was already noting pure and albino Stork’s-bill, Sea Holly, Sea Samphire, Sea Purslane and the
Shrubby Sea-blite at Small Mouth, as well as the abundance of butterflies, especially Painted
Ladies, but this holiday was abruptly terminated on 27 August 1939 when all visitors were advised
to leave Portland because of the imminence of war. In 1944 he started a detailed card index of the
plants he had seen, but he was unable to find time to keep it up after his appointment to a teaching
post in 1952.

Coombe’s first botanical studies in Cambridge used analyses of plant growth to explain the
distribution of species in the field. His Ph.D. thesis, Plant Growth and Light in Woodlands (1952),
was supervised by Dr G. C. Evans. He used Impatiens parviflora as an experimental plant, solving
the problem of germinating this awkward species and growing it in different light regimes at
Madingley Wood near Cambridge. (The plants he grew spread and the species is now well
naturalised in the wood.) Coombe and Evans developed the technique of measuring the complex
light climate of woodland by hemispherical photography of the canopy. At first they used the
camera with the original ‘fish-eye’ lens designed by Robin Hill, who lent it to them until
commercial models became available. Coombe continued his work on plant growth after he
became a University Demonstrator. Visits to Nigeria and the Cameroons led to studies of the
tropical shrub Trema guineensis and the tree Musanga cecropioides, rapidly growing plants of
abandoned farmland and forest clearings.

Even when he was working for his Ph.D. thesis he developed the interest in the Lizard peninsula
which he retained throughout his life. His first trip to the Lizard was made with his parents in the
spring of 1950. They had struggled to scrape together enough petrol coupons for the journey, and
when they arrived at the rather squalid village round Lizard Green his mother burst into tears of
disappointment. However they soon began to appreciate the scenery and botanical interest of the
area. In June 1950 Coombe led the first of numerous Botany School excursions to the Lizard.
Lewis Frost, then an undergraduate at Emmanuel College, joined this excursion and subsequently
the two teamed up to study the flora and vegetation of the area. Later excursions included several
held at Easter to study bryophytes, which were jointly led by Dr Harold Whitehouse.

It was on excursions to the Lizard that many students were introduced to Coombe’s remarkable
ability to synthesise knowledge from many different fields into a ‘holistic’ appreciation of the
ecology of an area. His understanding of the Lizard was based on the critical interpretation of
evidence from such diverse sources as Cornish place-names, Anglo-Saxon boundary charters, tithe
redemption maps, the accounts of Victorian travellers, published tourist guides and the memories
of the people of the Lizard themselves, as well as the more usual ‘scientific’ sources such as
meteorological, geological and archaeological studies, aerial photography, botanical records and
his own knowledge of the soils of the region and its vegetation at all seasons of the year. He
developed an acute ability to ‘read’ a landscape. The detailed, long-term observations (including
the repeated recording of permanent quadrats) carried out with Lewis Frost provided great insight
into the ecological processes at work in the Lizard vegetation, and in particular the way in which
rare species at the edge of their range were able to exploit small areas where the vigour of the
dominant plants was reduced by different combinations of summer drought, winter flooding and
human disturbance. All this information was committed to Coombe’s prodigious memory, and
could be recalled at will.

Coombe and Frost’s first papers on the Lizard described the heathland vegetation in relation to
the underlying soils. A crucial discovery was the fact that, whereas some of the heaths were rooted
in soils derived directly from the underlying serpentine rock, the Agrostis curtisii heaths grew over
wind-blown loess which had originated elsewhere. Although these papers were perhaps their most
influential, a later paper by Coombe alone provides a greater insight into his approach to less
Strictly scientific subjects. In a bibliographic study of C. A. Johns’ classic guide A week at the
Lizard, he drew on a wide range of sources to clarify the complex publishing history of this work,
which was incorrectly described in all the standard bibliographies and major library catalogues.
These conclusions were not written up in a dry bibliographical paper but in a highly readable
account full of detailed observations and spiced with humour.

362 OBITUARIES

To British botanists, the Lizard is famous for its rare annual Trifolium species, and Coombe and
Frost were particularly interested in their population dynamics. It was while on fieldwork at the
Lizard on 27 September 1957 that Coombe noticed a perennial clover which he described in his
field notebook as “‘a thick leaved, +glaucous, hairy-petioled Trif. repens (collected). This turned
out to be a diploid species which he described as Trifolium occidentale in 1961. (His original
intention to call it 7. atlanticum was dropped when H. Gilbert Carter pointed out that strictly
speaking this epithet applied to the Atlas Mountains.) Few British species can have been described
with such a wealth of observational and experimental detail, and the paper is in effect a worked
example of the biological species concept. Coombe got considerable satisfaction from the fact that
botanists elsewhere in western Europe soon came to recognise T. occidentale, and those who knew
it in the field had no difficulty in accepting it as a distinct species. (Many of its characters are lost
in the herbarium.) He later prepared the account of the large genus Trifolium for Flora Europaea.
It was characteristic of his approach that, rather than base his descriptions on published accounts
or herbarium material, he grew as many species as possible at Cambridge from seed distributed by
botanic gardens all over Europe (much of which turned out to be misidentified). As a result he was
able to write an accurate account and a key that actually works. When I first visited the Lizard with
David in September 1976, the clovers were germinating in abundance after the heavy rain which
followed the droughts of 1975 and 1976. He was able to find all the rare and many commoner
species, and to show me how to identify seedlings with only a single trifoliate leaf.

Coombe’s field studies were usually backed up by the cultivation of plants in the Botanic
Garden at Cambridge, many of them grown in the experimental glasshouses which he had helped
to design. He developed methods of growing species such as Isoetes histrix and Ophioglossum
lusitanicum, and cultivated many dwarf or prostrate ecotypes from the exposed Cornish coasts. He
took into cultivation all the surviving plants of Juniperus communis at the Lizard, rooting them
from cuttings (an operation which required horticultural skill and great patience). This provided
the basis of a public exhibit in the Botanic Garden where these plants were grown with progeny
raised from seed from one of the Lizard plants and with other junipers from all over Britain. The
differences between the columnar plants from southern England and the dwarf plants from Fair
Isle (which Coombe likened to a Wilton carpet) were very striking, and the display had a great
impact on visitors to the garden. Erica x williamsii, the hybrid between E. tetralix and E. vagans,
was another special interest. Coombe is the only person to have found more than one plant of this
hybrid (only eleven have ever been discovered, all at the Lizard) and he grew all the surviving
clones at Cambridge. These were introduced into commercial cultivation, one as cultivar ‘David
Coombe’. He also carried out a long series of experiments at the Garden on soil samples collected
in the 1950s, which demonstrated the long dormancy of the seeds or spores of several rare or
scarce species, including /soetes histrix, Juncus capitatus, Juncus pygmaeus, Lythrum hyssopifolia,
Riccia beyrichiana and R. crozalsii. Unlike many botanists he was a good gardener, and he acted
for some years as Hon. Garden Steward at Christ’s, designing a garden which contained all the
plants mentioned in Milton’s poem Lycidas. His interest in garden plants led him to realise that
Limonium companyonis (now L. hyblaeum), discovered at Rottingdean and originally reported as
an “enigma of doubtful origins’, was widely cultivated and almost certainly a garden escape in
Britain.

Although Coombe did not find the East Anglian countryside as attractive as that of counties
further south or west, he nevertheless acquired a detailed knowledge of the ecology of Wicken
Fen, Breckland, the chalk grassland and woods of East Anglia and the Chilterns, to all of which he
led student excursions in the Long Vacation. His approach to botany proved inspirational to many
students. When he was in his prime his lectures to the Part II botany students were superb, and
Peter Grubb remembers two absolutely outstanding formal papers to British Ecological Society
meetings. After a day in the field, or a discussion in his college rooms, one left his company
exhilarated by his insight into plant ecology. He influenced many who passed through Cambridge
University in the period when the Botany School was one of the leading departments for whole-
plant biology. One might have thought that as a “congenital conservative” (his words), and
someone who could talk about his subject at great length, he might have seemed a remote figure to
students, or even a bore. In fact he developed a great rapport with many students, and some have
even commented that he was the one don they met during their time at Cambridge whom they
really admired. They doubtless responded to his great enthusiasm and knowledge, his transparent
integrity and his willingness to spend time with them. He also treated students as individuals and

OBITUARIES 363

was able to see good points in almost
all of them. He responded very
generously to requests for information
from other botanists, professionals and
amateurs alike. In many cases where
he failed to respond to letters it was
not because of any lack of desire to
help, but because he started on a very
detailed exposition of the subject in a
letter which he was unable to find time
to complete. He could, however, be
scathing in his criticisms of
professional botanists whose work did
not measure up to his own standards.

Coombe applied the same academic
rigour to all he undertook. He
appeared to be incapable of
considering anything superficially, and
he usually seemed to be either an
expert in a subject or completely
ignorant of it. As a young lecturer he
had been required to teach a wide-
ranging introductory course to
students, during which he had obtained
a detailed knowledge of several
cryptogamic groups, including algae
and bryophytes. (In March 1958
Coombe and Whitehouse discovered
the moss Hennediella_ stanfordensis,
new to Europe, on one of the Lizard
bryophyte excursions.) Coombe had a
very good command of German,
having spent 3 months in Heidelberg
aS a research student, and he
collaborated with Peter Bell on a translation of Strasburger’s textbook of botany. To his botanical
interests were added many others, including books (inevitably), English furniture and the visual
arts. His tastes were traditional, and he responded particularly to authors like Thomas Hardy and
painters like John Constable who shared his sense of place. As a relaxation he composed limericks
and clerihews.

It was a matter of great regret to many of Dr Coombe’s friends and colleagues that much of his
work remained unpublished. They knew how great a contribution he was capable of making, and
tried in vain to persuade him to write more. In his early career he published a series of excellent
papers on diverse topics. Unfortunately his output then slowed, and he published little in the late
1960s and 1970s. The detailed Biological Flora accounts of the Lizard rarities which he planned to
write, for example, were never completed, and his studies of the semi-natural vegetation and the
crop yields on the patterned soils of Breckland were not written up for publication. No doubt his
heavy teaching load, his service on innumerable college, university and other committees and his
work as an editor of Journal of Ecology all contributed to this. The editorial work, in particular,
made great demands on his time and energies as he was meticulous in his attention to detail and
haunted by the prospect of misprints. His very appreciation of the interaction of factors which
made him such an astute ecologist also appeared to make it difficult for him to pursue a single line
through to publication. Added to this was an innate perfectionism: when I once encouraged him to
prepare a Biological Flora account of Juncus pygmaeus he said he would not do so, as although it
would be better than many accounts in the series it would not be as good as the best. However,
none of these factors really seem to provide a sufficient explanation for his failure to publish more
at the peak of his career.

ate

David Coombe at Traunsee, Austria, | August 1956 (photo.
taken by Dipl. Ing. B. Weinmeister on the 11th International
Phytogeographical Excursion).

364 OBITUARIES

A more tangible impediment to Coombe’s botanical work arose in 1978, when he developed a
lung disease which was diagnosed as cancer. It was only after a major operation that it was
recognised as a fungus (Aspergillus) infection. The adverse effects of this unnecessary operation
remained with him for the rest of his life: in particular his right arm was often painful, especially in
cold weather. In these later years he turned from his Cornish work to a series of studies of East
Anglian topics. Although limited in breadth, these were all pursued with the same depth of
scholarship. Several were written up for Nature in Cambridgeshire, where Gigi Crompton
provided initial encouragement and Philip Oswald was a sympathetic ‘midwife’ and editor. Two of
these papers resulted from his discovery of new populations of Veronica spicata on Newmarket
Heath. This led him to a consideration of the lost chalk heaths of Cambridgeshire, which he
reconstructed from the fragmentary surviving evidence. He also published a detailed paper on
‘Milton’s Mulberry’ in his college magazine. This tree was said to have been planted at Christ’s
College by the poet when he was an undergraduate. Coombe not only demonstrated that it was
almost certainly planted many years after Milton’s death, but went on to develop a theory that
many of the connections between great men and ancient trees were invented during the Romantic
period. Sadly his health prevented him from writing up several of the topics he had been working
on in this period - William Turner’s Cambridgeshire plant records (which had been neglected by
authors of successive Cambridgeshire Floras), the changing vegetation of Ringmere and the
history of the Lombardy Poplar in Britain. The poplar paper would have been particularly
interesting, as much of the evidence was drawn from landscape paintings, prints and literature
references.

In the last years of his life Coombe’s health deteriorated still further, and he was only rarely able
to get into the field. He made his last two visits to the Lizard in 1994, to help survey populations of
Juncus capitatus and J. pygmaeus for the Red Data Book and to follow up a developing interest in
Pyrus cordata. In 1999 he learnt that he was suffering from Parkinson’s disease and began to
prepare for the inevitable disabilities it would bring. However in June 1999 he contracted a
bacterial infection which failed to respond to treatment, and he died peacefully in hospital only a
few days later.

David Coombe will be remembered as an ecologist with an unrivalled insight into the flora and
vegetation of southern England. He left a substantial corpus of published work, all completed to
his own high standards and some of it of outstanding merit. However, his major contribution
perhaps lay in his teaching. By great good fortune one of his pupils was Oliver Rackham, who was
thus introduced to historical ecology. Dr Rackham has developed and extended Coombe’s
approach and in numerous books and papers brought it to the attention of a wider audience. By this
and other routes the holistic studies which Coombe pioneered came to influence an entire
generation of ecologists, and to change the way in which we all look at the English countryside.

PUBLICATIONS BY DR D. E. COOMBE

1947
The Sixty-one Society. Christ’s College Magazine Easter Term 1947: 16-17.

1951
(with F. White) Notes on calcicolous communities and peat formation in Norwegian Lappland. Journal of
Ecology 39: 33-62.

dz

The Wordsworths and botany. Notes and queries 5 July 1952: 298-299.

Clapham, A. R., Tutin, T. G. & Warburg, E. F. Flora of the British Isles. [Review.] The Cambridge Review
74: 88-90.

Ellenberg, Heinz. Landwirtschaftliche Pflanzensoziologie. Vol. 1. Unkrautgemeinschaften als Zeiger fiir
Klima und Boden. [Review.] Journal of Ecology 40: 409-410.

1954

Carex humilis Leyss, in C. D. Pigott & S. M. Walters, On the interpretation of the discontinuous distributions
shown by certain British species of open habitats. Journal of Ecology 42: 111-113.

Ancient earthworks. Letter to The Times 26 April 1954: 7.

Early Devon botanists, in W. Keble Martin & T. J. Wallace (eds), Forty-sixth report on the botany of Devon.

Report and transactions of the Devonshire Association for the Advancement of Science, Literature and
Art 86: 284.

OBITUARIES 365

1956

(with L. C. Frost) The heaths of the Cornish serpentine. Journal of Ecology 44: 226-256.

(with L. C. Frost and mineralogical data by M. le Bas & W. Watters) The nature and origin of the soils over
the Cornish serpentine. Journal of Ecology 44: 605-615.

Biological Flora of the British Isles. Impatiens parviflora DC. Journal of Ecology 44: 701-713.

Notes on some British plants seen in Austria. Veroffentlichungen des Geobotanischen Instituts Riibel in Ziirich
35: 128-137.

Botany, in Britannica Book of the Year 1956, pp. 88-89. Encyclopaedia Britannica, London.

1957
The spectral composition of shade light in woodlands. Journal of Ecology 45: 823-830.
Botany, in Britannica Book of the Year 1957, pp. 82-83. Encyclopaedia Britannica, London.

1958
Botany, in Britannica Book of the Year 1958, pp. 84-85. Encyclopaedia Britannica, London.

1959

(with G. C. Evans) Hemispherical and woodland canopy photography and the light climate. Journal of
Ecology 47: 103-113.

(with F. H. Perring & S. M. Walters) Lythrum hyssopifolia L. Proceedings of the Botanical Society of the
British Isles 3: 286-288.

(with F. H. Perring & S. M. Walters) Lythrum hyssopifolia L. [Abstract.] Proceedings of the Botanical Society
of the British Isles 3: 327.

Botany, in Britannica Book of the Year 1959, pp. 94-95. Encyclopaedia Britannica, London.

1960

An analysis of the growth of Trema guineensis. Journal of Ecology 48: 219-231.

(with G. C. Evans) Hemispherical photography in studies of plants. Medical and Biological Illustration 10:
68-75.

1961

Trifolium occidentale, a new species related to T. repens L. Watsonia 5: 68-87.

1962
(with W. Hadfield) An analysis of the growth of Musanga cecropioides. Journal of Ecology 50: 221-234.

1965

(with P. Bell) Strasburger’s textbook of botany, rewritten by R. Harder, W. Schumacher, F. Firbas & D. von
Denffer, translated from the 28th German edition by P. Bell & D. Coombe. Longmans, Green & Co.,
London.

1966

The seasonal light climate and plant growth in a Cambridgeshire wood, in R. Bainbridge, G. C. Evans & O.
Rackham (eds), Light as an ecological factor, pp. 148-166. Blackwell Scientific Publications, Oxford &
Edinburgh.

1967
(with P. Morisset) Further observations on Trifolium occidentale. Watsonia 6: 271-275.

1968

Trifolium L., in V. H. Heywood (ed.), Flora Europaea, Notulae systematicae ad Floram Europaeam spectantes.
Feddes Repertorium 79: 54.

Trifolium L., in T. G. Tutin et al. (eds), Flora Europaea, vol. 2, pp. 157-172. Cambridge University Press,
Cambridge.

J. A. Paton. Wild flowers of Cornwall and the Isles of Scilly. [Review.] Cornwall Naturalists’ Trust Newsletter
October 1968.

1969
In free time. Letter to The Times 2 December 1969: 11.

1970

The editions of A week at the Lizard by the Revd Charles Alexander Johns. Journal of the Society for the
Bibliography of Natural History 5: 259-269.

Which “Week at the Lizard”? The Lizard 4(2): 11-17.

(with D. P. Dymond, M. D. Hooper & O. Rackham) The hedge’s worth and age. Letter to The Times 6 April
1970: 9. Reprinted in K. Gregory, The first cuckoo: a selection of the most witty amusing and memorable
letters to The Times 1900-1975, pp. 303-305. George Allen & Unwin & Times Books, London, 1976
(hardback) and Unwin Paperbacks, 1980 (paperback).

366 OBITUARIES

1973
The prostrate junipers at Gew Graze. The Lizard 5(1): 7-12.
Thomas Johnson, in Dictionary of Scientific Biography, vol. 7, pp. 146-148. C. Scribner’s Sons, New York.

1974
An anthology of very wild flowers round the coast. The Lizard 5(2): 12.

1975
An anthology of very wild flowers round the coast. Part 2: ‘I sent thee late a rosy wreath’. The Lizard 5(3):
15-17.

1976

(with P. Bell) Strasburger’s textbook of botany, rewritten by D. von Denffer, W. Schumacher, K. Magdefrau
& F. Ehrendoefer, translated from the 30th German edition by P. Bell & D. Coombe. Longman, London
& New York.

1977

Air photography and plant ecology, in J.K.S. St Joseph (ed.), The uses of air photography, 2nd ed., pp. 86-
102. John Baker, London.

The Lizard, Cornwall, in D. A. Ratcliffe (ed.), A Nature Conservation Review, vol. 2, pp. 146-148. Cambridge
University Press, Cambridge.

Miltonic plants in Library Court. Christ’s College Magazine May 1977: 3-6.

1979
A botanical hazard at the Lizard. The Lizard 6(3): 15.

1981
(with A. F. G. Douse & C. D. Preston) The vegetation of Ringmere in August 1974. Transactions of the
Norfolk and Norwich Naturalists’ Society 25: 206-217.

1982
The Rottingdean Limonium. |Abstract.] Watsonia 14: 228.

1983

(with L. C. Frost & M. H. Martin) The effects of the cessation of grazing and of drought on the grasslands in
the Caerthillian valley. University of Bristol Lizard Project Report no. 9. The University, Bristol.

Sir John Plumb’s mastership. Christ’s College Magazine 72: 8-10.

1984

Stony ground. The Lizard 7(1-2): 28.

Grid reference SW 752182. The Lizard 7(1-2): 31.

1987

Of Milton and mulberries. Christ’s College Magazine 76: 15-20.

Spiked speedwell, soil stripes and polygons, and the vanishing chalk heaths of Cambridgeshire. Nature in
Cambridgeshire 29: 26-37.

Longevity of spores of some terrestrial Jsoetes species. [Abstract.] B.S.B.I. News 45: 38.

1988

More about stone stripes, polygons and fairy rings. Nature in Cambridgeshire 30: 13-15.
Hayward, John. A new key to wild flowers. [Review.] Endeavour n.s. 12: 48-49.
Propogation of Polygonum maritimum L. [Abstract.] B.S.B.I. News 49: 43.

1989

Vegetative propogation of Lythrum hyssepifolia. [Abstract.] B.S.B.I. News 51: 46. [See also correction in
B.S.B.I. News 52: 3 (1989).]

1990

(with T. J. N. Mills & P. Upchurch) The history of Madingley brickpits: contrasts in succession. Nature in
Cambridgeshire 32: 3-14.

Lythrum hyssopifolia L.: two postscripts. [Abstract.] B.S.B.I. News 54: 47.

(with G. Crompton) Ferula communis L.: mythology and biology. [Abstract.] B.S.B.I. News 54: 47.

1992
Cymbalaria muralis forma toutonii. [Abstract.] B.S.B.I. News 60: 62.
Ophioglossum lusitanicum: 31 years in cultivation. [Abstract.] B.S.B.I. News 60: 62.

OBITUARIES 367

1994

‘Maritime’ plants of roads in Cambridgeshire (v.c. 29). Nature in Cambridgeshire 36: 37-60.

Trifolium occidentale Coombe, Trifolium ochroleucon Hudson, Trifolium ornithopodioides L., Trifolium
squamosum L. and Trifolium suffocatum L., in A. Stewart, D. A. Pearman & C. D. Preston (comps &
eds), Scarce plants in Britain, pp. 414-418. Joint Nature Conservation Committee, Peterborough.

(with C. Chatters) Scilla autumnalis L., in A. Stewart, D. A. Pearman & C. D. Preston (comps & eds), Scarce
plants in Britain, p. 374. Joint Nature Conservation Committee, Peterborough.

(with S. J. Leach) Trifolium glomeratum L., in A. Stewart, D. A. Pearman & C. D. Preston (comps & eds),
Scarce plants in Britain, p. 413. Joint Nature Conservation Committee, Peterborough.

1996
(with O. Rackham) Madingley Wood. Nature in Cambridgeshire 38: 27-54.

1999

Erica vagans L., in M. J. Wigginton (comp. & ed.), British Red Data Books 1 Vascular plants, 3rd ed., pp.
139-140. Joint Nature Conservation Committee, Peterborough.

(with L. E. Friday & P. J. Grubb) The Godwin Plots at Wicken Fen: a 55-year record of the effects of mowing
on fen vegetation. Nature in Cambridgeshire 41: 3346.

OBITUARY NOTICES

An obituary appeared in The Independent newspaper on 30 July 1999 and further obituaries will be
published in Christ’s College Magazine and Nature in Cambridgeshire.

ACKNOWLEDGMENTS

I am very grateful to Mrs G. Crompton, Dr G. C. Evans, Lady Rosemary FitzGerald, Dr P. J.
Grubb, P. H. Oswald, Mrs J. A. Paton, D. A. Pearman, Dr O. Rackham and the late Dr H. L. K.
Whitehouse for help in preparing this obituary or for comments on a draft of the text.

C. D. PRESTON

LEWIS CHARLES FROST
(1926-1998)

Lewis Frost was very proud to have been born and bred in Norfolk and remained deeply attached
to the county throughout his life even though in later life he rarely returned. He came from farming
stock with a Belgian mother to whom he was very close. He was to have been called Louis but the
Registrar disapproved of foreign names and wrote Lewis on the birth certificate. Hence, he was
known as Louis to his family, as Lewis to most in the academic world, to those of us on more
familiar terms as Lew and Uncle Lew to our children. At one time it was intended that he should
enter the priesthood and whilst he continued his connection with the church during his early
undergraduate days at Emmanuel College, Cambridge, it would seem that national service
followed by his academic successes in the natural sciences tripos persuaded him to follow a
scientific and academic career.

After attending school in Bungay he joined the Royal Army Educational Corps in 1944 and
when he was demobbed in 1948 went up to Cambridge. His career in Cambridge was
distinguished. He obtained First class in all three parts of the tripos and this led to starting a Ph.D
in genetics in the school of agriculture in 1952 under the supervision of D. G. Catcheside. After
one year he was appointed Demonstrator in Botany (a post he held until 1957); H. L. K.
Whitehouse became his supervisor when Catcheside left Cambridge. His Ph.D was awarded in
1956. In October 1957 he was appointed Lecturer in Genetics within the Department of Botany at
Bristol University.

In Bristol Lewis taught genetics to a wide variety of undergraduates. He was for many years the
only genetics lecturer in the university, so he taught medical and veterinary students as well as
biologists. He taught genetics until his retirement in 1983 with a further three years of part-time re-
engagement. However, he also developed courses on nature conservation as his research interests
in genetics waned and were overtaken by his deep commitment to conservation ecology. Indeed,
Lewis will probably be remembered by most biologists not for his genetics teaching, but for his
ecological and conservation activities primarily within two areas: the Avon Gorge and the Lizard
in Cornwall.

368 OBITUARIES

In Cambridge Lewis had teamed up with David Coombe and together they developed an interest
in the Lizard area of Cornwall; their joint work commenced whilst Lewis was still an
undergraduate, during a field course led by David Coombe in June 1950. Lewis’s passion for the
Lizard (and the associated Cornish cream teas) lasted throughout the rest of his life and he would
often visit the Lizard two or three times a year. In 1956 David Coombe and Lewis published their
classic paper on the heaths of the Cornish serpentine - even today this is still regarded as the
definitive paper and known to most ecologists as “Coombe & Frost”. This was followed by
another paper on the soils of the Lizard in which they made the startling discovery of wind blown
deposits overlying the serpentine of the Goonhilly Downs. Today it is accepted that such loess
deposits are wide-spread in the south west but at the time many treated the findings with
scepticism. Lewis took groups of Bristol students to the Lizard to imbue in them his enthusiasm
and idiosyncrasies. Those who attended these trips soon found themselves referred to as “Lizard
kiddies”. A favourite diversion was the grade one cream teas: these were taken at the farm on the
main road to the Lizard where a side road is signposted to “Grade 1’. A highlight of every visit to
the Lizard was the systematic visitation of all the serpentine turners in the village and the
bargaining with them for a few pence off the choice items of serpentine ornaments. Throughout his
career Lewis was a great battler for conservation causes. He raised considerable funds for the
Cornwall Wildlife Trust’s purchase of land and establishment of a nature reserve on the Lizard and
the Cornwall Wildlife Trust made him an honorary member. He inveigled, bullied, and persistently
pestered the Nature Conservancy Council to do the same, with the result that eventually substantial
areas of the Lizard became National Nature Reserves and Sites of Special Scientific Interest. In
Bristol he set up the Lizard Appeal for funds to carry out research. This received good support and
allowed numerous conservation and research projects to be undertaken, which are documented in a
long series of over 20 Lizard Reports, and the engagement of a series of Ph.D students including
John Hopkins, Marion Hughes and Andrew Byfield. His own research concentrated on the rarer
plant species such as Trifolium incarnatum ssp. molinerii and in particular Herniaria ciliolata ssp.
ciliolata.

During his early life in Bristol Lewis was a sub-warden in Wills Hall and he recognised the
Avon Gorge as a vast untapped study area on the doorstep of Wills Hall. He quickly formed an
informal field club and together with a trusty and dedicated group of students set about a wide
range of botanical studies, including detailed examination of the ecology of species such as Trinia
glauca, Arabis scabra, Hornungia petraea and Carex humilis. These interests culminated in the
establishment of the Avon Gorge Appeal Fund, this attracted considerable sums of money and
helped finance innumerable studies on the Avon Gorge and its rare plants, with Clive Lovatt and
Libby Houston as collaborators. After his retirement Lewis remained as an Honorary Research
Fellow in the School of Biological Sciences and continued producing research reports right up to
the time of his going into hospital.

Locally Lewis played an active role in the early days with the Gloucestershire Wildlife Trust and
took part in many of the early surveys of sites later to become nature reserves in that county.
Another highlight was his long involvement with the Badgeworth Nature Reserve and the Adder’s-
tongue Spearwort. The fact that this reserve was officially declared the smallest nature reserve in
the world and appeared as such in the Guinness Book of Records gave him great pleasure.

Most of us will remember Lewis for his dry sense of humour; even a few days before he died we
were able to have a joke together and that impish glint came to his eyes. We will also remember
the ways that Lewis always savoured a bargain. It gave him immense pleasure to be able to
bargain a few pence off virtually any purchase from relatively expensive hi-fi equipment, to a new
pair of shoes, even to a packet of biscuits. But this aspect of his character changed totally with
friends and colleagues who were in trouble when he would be enormously generous, helpful and
considerate. Those of us who were fortunate to experience this side of Lewis were indeed
privileged.

M. H. MARTIN & M. D. AMES

Postscript: At the time of his death David Coombe was in the process of writing an obituary for
Lewis Frost. Unfortunately due to his illness he did not manage to write this, but the authors are
aware of how much more David would have included. We are however, very grateful to M.H.M.
and M.D.A. for taking on the writing of the obituary in these circumstances. Ed.

OBITUARIES 369

HUGH ALLISON LANG
1920-1999

Dr Hugh Lang died on 19 November 1999 in Newton Stewart where he had been a much respected
single-handed family doctor for 27 years. He joined the B.S.B.I. in 1973 although he had botanical
interests going back many years. He was essentially a field botanist and very much an individualist
who became well known to members of the society in Scotland, where he had served on the
B.S.B.I. Committee. The late Olga Stewart described him in her “Flowering Plants of
Kirkcudbrightshire” as a “mountaineer and mountain plant botanist” which summed up his love of
wild and remote places and the plants which grew there. He was a large man who had great
strength and stamina and enjoyed scrambling up to and exploring particularly inaccessible habitats
and was an expert photographer who recorded most of his finds on film. He was also a keen
observer of many aspects of the natural world and these included astronomy, geology and
meteorology.

He was born in Sauchiehall Street, Glasgow and entered Glasgow University in 1937. After a
science degree he obtained his medical degree in 1943 and immediately joined the airborne
division of the Royal Army Medical Corps and was posted to India. He was one of the first doctors
parachuted into the Japanese prisoner of war camps in Burma and saw at first hand the appalling
condition of the Allied prisoners there. He was demobbed in 1947 and after entering General
Practice in Stranraer and Barrhill, he eventually set up his own practice in Newton Stewart in 1958
helped by his wife Anne, whom he married in 1953. He was ideally suited to attending
emergencies in the hills and put his organisational experience to good use, being instrumental in
founding the Galloway Mountain Rescue Team and in 1975 becoming its first chairman. He was
also a past president of the local Rotary Club. As a side line he made an impressive collection of
aircraft wreckage collected from military jet crash sites. He retired from his practice in 1985.

Hugh knew the rocky coastline and hills of Galloway and the location of its botanical rarities
better than anyone and was especially fond of the crags of the Merrick, the highest point in
Southern Scotland. He told me how early on he had met the late Dr Humphrey Milne-Redhead,
another doctor/botanist and vice-county recorder, and showed him a willow collected high in the
hills. Humphrey pronounced it to be Salix lapponum (Downy Willow) an extremely rare plant of
the district and was duly impressed. He exhorted Hugh to take his botany seriously and this paid
off for in 1973 he added Saxifraga oppositifolia (Purple Saxifrage) to the flora of the Galloway
hills; a third site for Southern Scotland. However, Hugh’s great discovery had come previously in
1971 when exploring the remote hills of Easter Ross, he found Artemisia norvegica (Norwegian
Mugwort) in a third British station. It is Hugh’s photograph of this species which appears as the
frontispiece in Ursula Duncan’s “Flora of East Ross-shire”. It is intriguing to note that it was
Humphrey Milne-Redhead who found the second British site for this species in West Ross 17
years before. His enthusiasm was such that only a few months before his death he had managed to
photograph Arabis alpina (Alpine Rock-cress) at its recently discovered second site in the Cuillins.
Hugh served on the Torrs Warren reserve committee and was undertaking a photographic survey
of this nationally important dune system. He also monitored the sole remaining colony of Woodsia
ilvensis (Oblong Woodsia) in the Moffat Hills.

The arctic and the history of its exploration had always fascinated him and on retirement he was
at last able to visit the high arctic regions of North America and particularly Greenland. He didn’t
miss a single summer visit in the 14 years between his retirement and death. His tiny blue tent was
a constant feature of these expeditions and there was always speculation as to how he managed to
fit his large frame into it. His ultra-long sturdy home-made trekking pole gave him extra stability
especially when crossing glacial torrents. He had a very good eye for the small plants of the arctic
flora and noted anything which looked unusual including the occasional lichen and fungus. His
collecting was unorthodox but effective, using a notebook in his hip pocket and his collections
were sent to the Botanical Museum of the University of Copenhagen and to Geoffrey Halliday at
Lancaster University. It was from Hugh’s high quality photograph of a Pedicularis sp. (a
Lousewort) from north-western Greenland that Geoffrey identified P. sudetica, new to the
Greenland flora. It had been collected previously, but was misidentified as the flower colour fades
in the herbarium and it resembled a closely related species. He was delighted on his last expedition
to see and photograph Potentilla stipularis, a very rare and elusive Siberian and Greenland species

370 OBITUARIES

of Cinquefoil. His archaeological interests were stimulated by ancient Eskimo remains and he was
in correspondence with the Danish authorities about possible new sites. On expeditions Hugh was
a steadfast companion who never let bad conditions get on top of him. He had an inexhaustible
fund of verse and quotation to suit any situation and this was delivered in his own inimitable way.
This was sure to ‘break the ice’ with strangers and boost flagging morale among friends. It was
this same side of his personality that made his slide shows at the Scottish Exhibition meetings so
memorable with descriptions which could reduce the audience to helpless laughter. He loved
Drambuie and his ceremonial decanting from bottle to plastic container in the airport departure
lounge to save weight, was a regular pre-expedition feature, causing many an eyebrow to be
raised. He was a member of the Arctic Club and served on the committee of the Scottish Arctic
Club.
He is survived by his son Angus. Sadly, his wife Anne predeceased him four years earlier.

R. W. M. CORNER

BERTRAM EVELYN SMYTHIES
(1912-1999)

Bertram Evelyn Smythies, usually known as Bill, died on 27 June 1999, aged 86. He had been a
member of the B.S.B.I. since 1974 and was elected a Fellow of the Linnean Society in 1969.

Bill was born in India on 11 July 1912, at Naini Tal, Uttar Pradesh. His father worked for the
Forest Service as the Assistant Conservator of Forests for the United Provinces and his mother was
a noted tiger huntress. Thus, he was introduced to natural history at a very early age. He was sent
to a preparatory school back in England at the age of eight, but during the holidays stayed with his
grandfather (who had himself worked in India teaching botany at the Dehra Dun forest school) in
North Devon.

Some years ago, Bill sent me a Christmas card by Elizabeth Twining, daughter of Richard
Twining (tea merchant) and Elizabeth Smythies, illustrating the Grossulariaceae. He added the
note that “My grandfather had the 2 vols of our ancestor’s work in his library, and I used to lie on
the floor and learn the names of the “Natural Orders’ by heart - no TV or even Radio in those days!
Sadly the artistic gene was not passed to me, but Elizabeth Twining’s work may have aroused my
interest in bot. Ill., along with Fitch’s line drawings in ‘Bentham & Hooker’”. It was no surprise
that he went up to Balliol College, Oxford to read Botany and Forestry. It was here that he met G.
K. Yates, a leading bird photographer, who taught Bill bird photography and how to prepare a bird
skin. Bill went off camping and photographing rare birds at the nest in Caithness and Sutherland,
skills he was to put to good use later.

After graduating, Bill went in 1934 to work for the Colonial Forest Service in Rangoon, Burma,
where he was quickly recruited by H. C. Smith to collect bird skins for the British Museum
(Natural History). This led to production of the Birds of Burma, published by the Mission Press,
Rangoon, in 1941. Before leaving Burma in February 1942, Bill managed to rescue the copper
printing blocks from the abandoned Mission Press and made possible the production of an
enhanced second edition in 1953. Some copies of the original edition were sold to expatriate
Europeans, but most were taken back to Japan where they were destroyed in an air raid: it is thus a
sought-after rare work.

After the war, Bill returned to Burma, but left in 1948 when the country left the Commonwealth.
He then took up a post in the forest service in Sarawak, publishing a 300-page Annotated Checklist
of the Birds of Borneo in 1957. This was the foundation for his glorious Birds of Borneo (1960), a
classic work that was later revised with a second edition in 1968 and a third in 1981. Although he
also worked for a while in Brunei in the 1950s, he returned to Sarawak to produce Sarawak Forest
Trees, published in 1965 by the Borneo Literature Bureau. It aimed to enable foresters to identify
tree species and is a fundamental work for the new Tree Flora of Sabah and Sarawak currently
being prepared.

In 1964 Bill married Jill Rogers and shortly afterwards retired from the Sarawak Forest Service
to live in Estepona, southern Spain. It was here that he pursued his love of botany, taking part in
some of the John Carr Expeditions to Andalucia. He travelled widely in the area and produced a
Flora of Spain: Checklist of Vascular Plants, published in three parts (1984, 1984, 1986) as

OBITUARIES 374

volume three of the periodical Englera of the Botanischen Garten und Botanischen Museum
Berlin-Dahiem. This phenomenal catalogue enumerates 7138 taxa compared to the 5948 in Flora
Europaea (and an estimated 8600 in Flora Iberica) and illustrates Bill’s industry. This increase
was not achieved by simply taking a more radical view of species, but by a careful review of the
extensive recent literature and consulting specialists.

However, it is not for this fundamental work that botanists most remember him, but for Flowers
of South-West Europe: a field guide, a very successful popular botanical guide produced in
collaboration with Oleg Polunin and published in 1973 by Oxford University Press. It is a credit to
them that even today it is still also a best seller in its Spanish edition.

Part of the success is due to the clear, double-page spread line drawings of plants by Jill
Smythies. Jill became a competent botanical artist as is also shown by her archive of drawings and
paintings of Spanish plants presented after her death in 1994 by Bill to the University of Reading
Herbarium. Following an accident to Jill’s right hand which ended her drawing, Bill endowed the
Linnean Society of London with the Jill Smythies award for botanical drawing which contributes
to plant identification.

After moving back from Estepona to Surrey, Bill became a frequent visitor to the Linnean
Society's rooms. Although always of few words, he was a good correspondent, and it was here that
his wit and humour was seen: “In my grandfather’s day, a century ago at Dehra Dun the mail was
carried by a dak runner trotting along from village to village; for long distances horse transport
was used. I suspect the Italian postal service is at the same state of development - your card
postmarked Rome 9 Dec. took three weeks to get here. In the days of Imperial Rome it would
surely have taken less!”

With Bill’s death, the story does not end, for botanists will continue to benefit through the
legacy of his library to the Linnean Society of London and the greater part of his estate to
botanical charities.

S: L. JURY

ERIC VERNON WATSON
1914-1999

Dr Eric Watson, the distinguished bryologist and greatly respected university lecturer died at his
home in Goring on 25 October 1999. He had been his usual self and working in the University
during the summer. Although he found lately that he had been getting very tired, he had kept up
his botany at home, almost to the last. He was diagnosed as having been suffering from leukaemia.

He joined the staff of the Department of Botany at the University of Reading in 1946 as
Lecturer, later becoming Senior Lecturer before his retirement in 1979. He was very well known in
bryophyte circles, especially for his books (The Structure and Life of Bryophytes, Hutchinson
Universal Library, London, 3 eds and British Mosses and Liverworts, Cambridge University Press,
3 eds). These were particularly successful in getting students of all ages interested in and able to
understand and know bryophytes. He had amassed a considerable bryophyte herbarium, putting
specimens in both the University collection and his own private one (this is left to the Royal
Botanic Garden, Edinburgh; the University of Edinburgh was his old alma mater). However, it
must be said that in fact he was an excellent all-round botanist and knew the British flora well,
having been a member of the B.S.B.I. for almost 50 years.

I remember vividly ‘demonstrating’ to students in his anatomy classes in the 1970s, when the
class was expected to trace the vascular tissues through a plant node by making a series of sections
and interpreting the results as a 3-D drawing. All the examples (Vicia, Lamium and strawberry
runners) came from his garden. His teaching abilities were such that most students were inspired to
work incredibly hard to do the most difficult example (Vicia, the stipules complicate the structure
enormously).

His tutees were regularly invited out to his house for supper and a slide show. He held regular
moss forays starting with morning coffee at his house and ending back for afternoon tea. These
invariably started with the species to be found in the garden and over the years a considerable
checklist of species was made. Generations of students will remember him with affection and
many had kept in touch with him — he was a great letter writer.

372 OBITUARIES

Dr Watson was a keen natural historian, accomplished ornithologist (founder member of
Reading Bird Club after the war), gardener, artist (watercolourist: he was a leading member of
Reading Guild of Artists), golfer and earlier a very good tennis player. Taking part in so many
local societies and activities made him well-known locally.

He will be greatly missed. We send his widow Joyce and family our condolences.

Sek. JURY

Watsonia 23: 373-375 (2000) 373

Report

ANNUAL GENERAL MEETING, 8 MAY 1999

The Annual General Meeting of the Society was held at the Royal Pharmaceutical Society, |
Lambeth High Street, London, at 11.50 a.m. The President, Mrs M. Briggs took the Chair in the
presence of 102 members.

Apologies for absence were read and Minutes of the 1998 Annual General Meeting, published in
Watsonia 22: 297-298 (1999), were accepted as correct, approved and signed by the President.

REPORT OF COUNCIL

The President took members through the Report of the Council, which had been previously
circulated to members, commenting on the main achievements of the Society during the year. She
mentioned especially the progress of Atlas 2000 and the setting up, with the Joint Nature
Conservation Committee (J.N.C.C.) and the Country Agencies, of the Threatened Plants Database
(T.P.D.B.) and the appointment of Mr A. Lockton and Mrs S. J. Whild to run this and act as B.S.B.
I. Co-ordinators. She finally thanked all those who worked so hard for the Society on a voluntary
basis and congratulated Mr R. G. Ellis on producing another excellent Report. The adoption of the
Annual Report was proposed by Mr C. R. Boon, seconded by Mr R. M. Walls and accepted
unanimously.

HON. TREASURER’S REPORT AND ACCOUNTS

The Treasurer reported that as usual, only summary accounts had been published in the Annual
Report, full sets of accounts were available at the meeting and on application to him. He
commented that only 30% of the resources used in the Society’s activities were derived from
subscriptions, much of the remainder coming from other bodies in the form of grants for Atlas
2000, T.P.D.B., etc. He was again pleased to report a small surplus for the year but, with the
disappointing returns from our recent Covenant campaign, in order to continue to cover
expenditure by income a modest increase in subscriptions was needed. He then proposed the
adoption of the report, which was seconded by Mr J. Ounsted and carried unanimously. The
President thanked the Treasurer for all the hard work he put into managing our financial affairs so
competently.

RULE CHANGES

In introducing the proposed rule changes, the President commented that the change to Rule 2 was
aimed at allowing the Society to involve itself with the study of British taxa in Europe and
elsewhere, while the change to Rule 4 would replace the now disbanded Conservation Committee
with the newly formed Science and Research Committee as a Permanent Working Committee.
Members were assured that the Society would continue to take an interest in plant conservation,
mainly through the S & R Committee, and the revised Terms of Reference for that Committee
were available at the meeting.

Proposed amendment to Rules 2 ( Aims of the Society)

That Rule 2 of the Society’s Rules be deleted and the following be substituted as recommended
by Council:

374 REPORT

AIMS AND OBJECTS

2. The aims of the Botanical Society of the British Isles shall be those of a learned society of
professional and amateur botanists dedicated to the study and conservation of the British and Irish
vascular plant and charophyte flora.

The objects of the Society shall be

a) To promote the study of and interest in the British and Irish vascular plant and charophyte
flora.

b) To support, encourage, carry out and participate in research into the taxonomy, ecology,
biogeography and conservation of the British and Irish flora and to co-operate with Continental
and other botanists in matters of mutual interest and concern.

c) To provide a forum for the dissemination of knowledge about the British and Irish flora, by
means of publications, Information Technology, meetings and advice.

d) To support and encourage the training of botanical skills.

e) To collaborate with statutory and other voluntary bodies engaged in the conservation of the
British and Irish vegetation and flora.

f) To maintain a botanical recording network in Britain and Ireland.

g) To provide, for the benefit of members, access to identification and other expert services for
the study of wild plants.

h) To support studies of the British and Irish flora or related taxa elsewhere, in furtherance of the
Society’s aims.

The change to Rule 2 as set out above was proposed by S. L. Jury, seconded by E. F. Greenwood
and carried unanimously

Proposed amendment to Rule 4 (Permanent Working Committees)

That Rule 4 of the Society’s Rules be amended. Council, in consultation with the Committees
concerned, decided that the furtherance of the Society’s aims for conservation would be better
achieved by the Committee reorganisation as follows:

(c) Conservation be deleted and replaced by
(c) Science and Research

The change to Rule 4 as set out above was proposed by Dr N. K. B. Robson, seconded by Dr F. H.
Perring and carried unanimously

INCREASE IN SUBSCRIPTIONS

The Treasurer proposed an increase from January Ist 2000 as follows (current rates in
parenthesis):

For members in Britain and Ireland

Ordinary £20.00 (£18.00)
Family £2.00 (£2.00)
Junior £10.00 (£9.00)
Senior £13.00 (£12.00)
Institutional Member £20.00 (£18.00)

For other overseas members £22.00 (£20.00)

This was seconded by Mr A.O. Chater and carried unanimously.

ELECTION OF PRESIDENT-ELECT

The President commented that it was a pleasant duty to propose Council’s nomination of Dr G.
Halliday, who, as Chairman of Records Committee, had been involved with many of the Society’s

REPORT a5

activities in recent years. The nomination was seconded by Mr M. Walpole and carried
unanimously with acclamation.

ELECTION OF VICE-PRESIDENT

The election of Mr D. J. McCosh, as nominated by Council, was proposed by Dr P. Macpherson,
seconded by Mr R. D. Pryce and carried unanimously.

RE-ELECTION OF HON GENERAL SECRETARY AND HON. TREASURER

Council had nominated Mr R. G. Ellis and Mr M. E. Braithwaite for re-election to their respective
posts. This was proposed by Mr P. H. Oswald, seconded by Miss A. P. Conolly and carried
unanimously with acclamation.

REAPPOINTMENT OF EDITORS, INDEXERS AND REPRESENTATIVES OF COMMITTEES

In proposing the reappointment of Editors, Indexers and Representatives of Committees, the
President praised them for the hard work they carried out, voluntarily, on behalf of the Society.
She gave special thanks to Mr P. J. Fry (Assistant Secretary) who gave so freely of his time in
answering postal and telephone queries at the B.S.B.I. desk in the Natural History Museum, and to
Mr P. Thomson (Council Minuting Secretary). These were received with acclamation.

ELECTION OF COUNCIL MEMBERS

In accordance with Rule 11 nominations had been received for Mr P. H. Oswald (proposed Dr C.
D. Preston, seconded Mr R. G. Ellis), Mrs J. M. Robertson (proposed Mr A. C. Jermy, seconded
Mr R. M. Walls) and Dr B. S. Rushton (proposed Mrs M. Briggs, seconded Mr R. G. Ellis).
Profiles had been published and election of these members was carried unanimously.

ELECTION OF HONORARY MEMBERS

The President mentioned that three members had been nominated for Honorary Membership of the
Society this year. All had, in different ways, made outstanding contributions to the Society.
Sponsors for all three gave short appreciations of their respective candidates (to be published in
BSBI News) and their election was carried unanimously with warm applause.

The three new Honorary Members (and sponsors) were: Mr E. J. Clement (Mr T. B. Ryves, read
by Mr R. G. Ellis), Dr B. S. Rushton (Mr M. Walpole) and Prof. G. A. Swan (Dr A. J. Richards,
read by Mrs M. Briggs).

RE-ELECTION OF HONORARY INDEPENDENT EXAMINER

The President warmly thanked Mr John Coats, of Greaves West & Ayre, Chartered Accountants,
Berwick upon Tweed for his exemplary examination of our accounts. His re-election was proposed
by Mr M. Walpole, seconded by Mrs M. Lindop and carried unanimously.

ANY OTHER BUSINESS

In reply to a written query by Dr T. C. G. Rich concerning the non-availability of a pre-publication
offer for the new Red Data Book published by J.N.C.C., the President reported that attempts to
persuade J.N.C.C. to give a discount to B.S.B.I. members had been unsuccessful.

There being no other business, the meeting closed at 12.47 p.m.

GWYNN ELLIS

—

CHWNIADARWN TS

72

West Cornwall
Scilly

East Cornwall
South Devon
North Devon
South Somerset
North Somerset
North Wiltshire
South Wiltshire
Dorset

Isle of Wight
South Hampshire
North Hampshire
West Sussex
East Sussex
East Kent

West Kent
Surrey

South Essex
North Essex
Hertfordshire

. Monmouthshire
. Glamorgan

. Breconshire

. Radnorshire

. Carmarthenshire

SCOTLAND

V3.
74.
1D:
76.
TT.
78.
79.
80.
81.
82.
83.
84.
S2;

IRELAND

a.
H2.

M3:
H4.

HS.
H6.
H7.
H8.
H9.
H10.
H11.
H12.
H13.
H14.

Dumfriesshire
Kircudbrightshire
Wigtownshire
Ayrshire
Renfrewshire
Lanarkshire
Peeblesshire
Selkirkshire
Roxburghshire
Berwickshire
East Lothian
Midlothian
West Lothian
Fife

South Kerry
North Kerry
West Cork

Mid Cork

East Cork

Co. Waterford
South Tipperary
Co. Limerick
Co. Clare

North Tipperary
Co. Kilkenny
Co. Wexford
Co. Carlow |
Laois

NAMES OF VICE-COUNTIES IN WATSONIA

pa
22,
29:
24.
2D:
26,
27
28.
Zo:
30.
eae
a2.
33:
34.
36.
ah
38.
39.
40.
40.
D2:
45.
46.
47.
48.
49.

86.
87.
88.
89.
90.
91.
v2,
93;
94.
93:
mG.
96b.
a
98.

H15.
H16.
BET,
H18.
lathe aa
H20.
H2 1.
H22.
HZ3:
H24.
H25:
H26;
R27.
H28.

ENGLAND
Middlesex

Berkshire
Oxfordshire
Buckinghamshire
East Suffolk

West Suffolk

East Norfolk

West Norfolk
Cambridgeshire
Bedfordshire
Huntingdonshire
Northamptonshire
East Gloucestershire
West Gloucestershire
Herefordshire
Worcestershire
Warwickshire
Staffordshire
Shropshire
Shropshire

South Lincolnshire

Pembrokeshire
Cardiganshire
Montgomeryshire
Merionethshire
Caernarvonshire

Stirlingshire

West Perthshire

Mid Perthshire

East Perthshire
Angus
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BOTANICAL SOCIETY
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BRITISH ISLES
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The B.S.B.I. traces its origin to the Botanical Society of London founded in 1836 and has a membership
of 2,850. It is the major source of information on the status and distribution of British and Irish
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BSBI handbooks

Each handbook deals in depth with one or more difficult groups of British and Irish plants.

No. | Sedges of the British Isles
A. C. Jermy, A. O. Chater & R. W. David. Revised edition, 1982. 272 pp., with descriptions,
drawings and distribution maps for all 73 species of Carex. Paperback. ISBN 0 901158 05 4.

No. 2 Umbellifers of the British Isles
T. G. Tutin. 1980. 200 pp., with descriptions and drawings of 73 species of Apiaceae (Umbel-
liferae). Paperback. ISBN 0 901158 02 X.

No. 3 Docks and knotweeds of the British Isles
J. E. Lousley & D. H. Kent. 1981. 208 pp., with descriptions and drawings of about 80 native
and alien taxa of Polygonaceae. Paperback. ISBN 0 901158 04 6. Out of print. New edition with
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No. 4 Willows and poplars of Great Britain and Ireland

R. D. Meikle. 1984. 200 pp., with descriptions and drawings of 65 species, subspecies, varieties
and hybrids of Salix and Populus. Paperback. ISBN 0 901158 07 0.

No. 5 Charophytes of Great Britain and Ireland
J. A. Moore. 1986. 144 pp., with descriptions and drawings of 39 species and varieties of
Characeae and 17 distribution maps. Paperback. ISBN 0 901158 16 X.

No. 6 Crucifers of Great Britain and Ireland
T. C. G. Rich. 1991. 344 pp., with descriptions of 148 taxa of Brassicaceae (Cruciferae), 129 of
them with drawings, and 60 distribution maps. Paperback. ISBN 0 901158 20 8.

No. 7 Roses of Great Britain and Ireland
G. G. Graham & A. L. Primavesi. 1993. 208 pp., with descriptions and drawings of 13 native
and nine introduced taxa of Rosa, descriptions of 76 hybrids, and 33 maps. Paperback. ISBN
0 901158 22 4.

No. 8 Pondweeds of Great Britain and Ireland
C. D. Preston. 1995. 352 pp., with descriptions and drawings of all 50 species and hybrids of
Potamogeton, Groenlandia and Ruppia, most of them with distribution maps; detailed
introductory material and bibliography. Paperback. ISBN 0 901158 24 0.

No. 9 Dandelions of Great Britain and Ireland
A. A. Dudman & A. J. Richards. 1997. 344 pp., with descriptions of 235 species of Taraxacum,
most of them illustrated by silhouettes of herbarium specimens; drawings of bud involucres of
139 species and 178 distribution maps. Paperback. ISBN 0 901158 25 9.

No. 10 Sea beans and nickar nuts
E. Charles Nelson. 2000. 156 pp., with descriptions of nearly 60 exotic seeds and fruits found
stranded on beaches in north-western Europe (many illustrated by Wendy Walsh) and of the
mature plants (some with drawings by Alma Hathaway), accounts of their history and folklore,
growing instructions etc. Paperback. ISBN 0 901158 29 1.

Other publications

English names of wild flowers
J. G. Dony, S. L. Jury & F. H. Perring. 1986 (2nd ed.). 126 pp. Recommended English names
for British and Irish vascular plants (Latin-English and English—Latin). Paperback. ISBN
0 901158 15 1.

List of vascular plants of the British Isles
D. H. Kent. 1992. 400 pp. Nomenclature and sequence as in Clive Stace’s New Flora of the
British Isles (1991, 1997), with selected synonyms. Paperback. ISBN 0 901158 21 6. Supplied
with errata lists and Supplement 1.

Alien plants of the British Isles
E. J. Clement & M. C. Foster. 1994. 616 pp. Lists 3,586 recorded non-native species (of which
885 are established), with English names, frequency of occurrence, status, areas of origin,

location of voucher specimens, references to published descriptions and illustrations, and
selected synonyms. Paperback. ISBN 0 901158 23 2.

Alien grasses of the British Isles
T. B. Ryves, E. J. Clement & M. C. Foster. 1996. 234 pp. A companion volume to the last,
listing over 700 non-native grasses; includes keys to bamboos and eight of the larger and more
difficult genera and 29 pp. of illustrations. Paperback. ISBN 0 901158 27 5.

Plant crib 1998
T. C. G. Rich & A. C. Jermy. 1998. 400 pp. An identification guide for some 325 ‘difficult’
taxonomic groups, with explanations, keys and illustrations of plant details. A4 paperback.
ISBN 0 901158 28 3.

British Red Data Books I Vascular plants
M. J. Wigginton, ed. 1999 (3rd ed.). 468 pp. Up-to-date information on 408 taxa (including 118
microspecies) regarded as threatened in Great Britain, with details of their habitats and
associated species, maps showing pre-1970, 1970-1987 and post-1987 records by 10-km
squares, and 1-km square frequency maps for 63 taxa. Hardback, published by JNCC,
Peterborough. ISBN 1 86107 451 4.

Scarce plants in Britain
A. Stewart, D. A. Pearman & C. D. Preston, comp. & ed. 1994. 518 pp. Accounts of 254
nationally scarce taxa (occurring in 16-100 10-km squares in Great Britain) and of 71 taxa
formerly thought to be so, with details of their habitats and associated species, reproductive
biology, changing British distribution and world range; updated distribution maps. Hardback,
published by JNCC, Peterborough. ISBN 1 873701 66 7.

Aquatic plants in Britain and Ireland
C. D. Preston & J. M. Croft. 1997. 365 pp. Summarises the distribution, habitat and
reproductive biology of 200 aquatic plants in 72 genera, with 200 distribution maps and 72
drawings. Hardback, published by Harley Books, Colchester. ISBN 0 946589 55 0.

Available from the official agents for BSBI publications, Summerfield Books (Jon & Sue Atkins),
Summerfield House, High Street, Brough, Kirkby Stephen, Cumbria CA17 4BX (Tel: 017683 41577.
Fax: 017683 41687. E-mail: bsbipubs @beeb.net).

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INSTRUCTIONS TO CONTRIBUTORS

Scope. Authors are invited to submit Papers and Notes concerning British and Irish vascular
plants, their taxonomy, biosystematics, ecology, distribution and conservation, as well as topics of
a more general or historical nature. Authors should consult the Hon. Receiving Editor for advice
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Submission of manuscripts

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Watsonia

August 2000 Volume twenty three Part two

Contents

BRIGGS, M. Presidential Address, 1999: The study of a local flora

Box, J. D. Mistletoe Viscum album L. (Loranthaceae) on Oaks in Britain

FOLEY, M. J. Y. A morphological comparison between some British Orobanche
species (Orobanchaceae) and their closely-related non-British a
from continental Europe: Orobanche reticulata Wallr. s. 1.

JONSELL, B., NORDAL, I. & ROBERTS, F. J. Viola rupestris and its hybrids in Beian
FOLEY, M. J. Y. & PORTER, M. S. Carex muricata L.
(Cyperaceae) - a review of its present status in Britain
OREDSSON, A. Choice of surveyor is vital to the Pane of flosighe. dines

studies

RICH, T. C. G. & SYDES, fo Rtecnniline andl te iechaes ei the Nationally Seatiee
plants Ajuga pyramidalis L. and Melampyrum sylvaticum L. : :

FOLEY, M. J. Y. Dactylorhiza incarnata (L.) So6 subsp. ochroleuca (Wiistnei ¢ ex
Boll) P. F. Hunt and Summerh. laces A comparison of British and
European plants a

RIcH, T. C. G. Conservation oF nena S Ddiodiversity Hienatnan dase
(Asteraceae), Welsh Hawkweed

RICH, T. C. G. & HOUSTON, L. Conservation of Brita S biodiversity Miocene
tavense (Asteraceae), Black Mountain Hawkweed

RANDALL, R. D. & RICH, T. C. G. Conservation of Britain’s biodiversity Res
trelleckensis (Rosaceae), Trelleck Bramble : :

USHER, M. B. The nativeness and non-nativeness of species :

MEIKLE, R. D. & ROBINSON, N. A. A new record for Salix; xX angusensis
(Salicaceae) Rech. f. from Ainsdale Sand Dunes National Nature Reserve, S.
Lancs. Vic. 39° ..:

HANDLEY, R. J. & DAvy, A. i Discovery of faale plants of Naja marina ra
(Hydrocharitaceae) in Britain ue son | oa ee ee

NOTES

Blunt, A. G. & Blunt, M. E. Corynephorus canescens (L.) P. Beauv. (Poaceae) at
Kinver Edge, Staffordshire: a re-assessment

Rich, T. C. G. A reanalysis of the Mistletoe (Viscum ibis | ‘Lemnihaeeas
survey data from the 1970s and 1990s _... —...

Rich, T. C. G., Jones, R. A. & Jebb, M. Three new Batish Sites fon Cares
depauperata With. (Cyperaceae) represented in the Irish National Herbarium,
Glasnevin oe

Dines, T. D. & Preston, C. D Fleoeiaris maniite ficeorercan in ‘Scotland

Rich, T. C. G. & Brown, N. Suaeda vera J. F. Gmel. (Chenopodiaceae), Shrubby
Sea-blite, present in Anglesey (v.c, 52), Wales

Piggott, C. D. A probably native and regenerating population of Tilia platyphyllos
Scop. In Bedfordshire -

Allen, D. E. Further British species of ueue L. (Reston in comhaed Fence

subsp. muricata

Allen, D. E. Rubus angusticuspis Sudre (Rosaceae) in Scilly
BOOK REVIEWS oe ae ee ae
OBITUARIES

REPORT

Published by the Botanical Society of the British Isles
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Typeset by D. K. & M. N. SANFORD
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257-267

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293—297

299-303

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323-326

327-330

331-334

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i

uae

“WATSONT A

Botanical Society of the British Isles

Volume 23 Part 3 February 2001
Editors: M. Briggs, D. L. Kelly, D. R. McKean,
D. A. Pearman, M.N. Sanford, D. A. simpson

Botanical Society of the British Isles

Patron: Her Majesty Queen Elizabeth the Queen Mother

Applications for membership should be addressed to the Hon. General Secretary,
c/o Department of Botany, The Natural History Museum, Cromwell Road, London,

SW7 5BD, from whom copies of the Society’s Prospectus may be obtained.

Officers for 2000—2001

President, Dr G. Halliday
Vice-Presidents, Mr M. Walpole, Mr D. J. McCosh, Prof. C. A. Stace
Acting Honorary General Secretary, Miss A. Burns

Honorary Treasurer, Mr M. E. Braithwaite

Editors of Watsonia

Papers and Notes, D. L. Kelly, D. R. McKean, M. N. Sanford*, D. A. Simpson
Book Reviews, D. A. Pearman

Obituaries, M. Briggs

*Receiving editor, to whom all MSS should be sent (see inside back cover).

© 2001 Botanical Society of the British Isles
The Society takes no responsibility for the views expressed by authors of Papers,
Notes, Book Reviews or Obituaries.

The cover illustration of Primula scotica Hook. (Scottish Primrose) was drawn by
Rosemary Wise.

Watsonia 23: 377—390 (2001) 6 iy

On the status of the genus Koeleria Pers. (Poaceae) in Britain

J. M. DIXON

Dept. of Environmental Science, University of Bradford, Bradffrd BD7 1DP

MAY 15 2001
LIBRARIES

Since 1980 there has been confusion over the number of species in the genus Koeleria found in Britain. Prior
to 1980, only two species - Koeleria macrantha (= cristata) and K. vallesiana - were reported; Flora
Europaea added K. glauca to these. Results of an extensive programme to clarify the status of the Koeleria
macrantha group (containing both K. macrantha and K. glauca) in Britain are presented here. Detailed
morphological measurements and observations were made on plant material collected from 51 sites
throughout Britain. The material was then grown under standard conditions; repeat measurements and
observations were then made. Chromosome determinations were also carried out on samples of all
populations. Whilst field specimens sometimes had morphological characteristics which might have identified
them as either K. macrantha or K. glauca, all plants grown under standard conditions in the glasshouse
converged in all their characteristics and were clearly referable to K. macrantha. The results are supported by
cytological evidence: all plants were tetraploid (K. macrantha is reported as 2n = 28 and K. glauca as 2n = 14
in Britain). The results strongly suggest that only two species of Koeleria - K. macrantha and K. vallesiana -
occur in Britain, and that confusion might have arisen as a result of the wide morphological variation inherent
in K. macrantha.

ABSTRACT

KEYWORDS: Koeleria macrantha, Koeleria glauca, morphological measurement, chromosome determination.

INTRODUCTION

During the latter part of the 19th century and early part of the 20th, the genus Koeleria in Britain
was divided into at least five species: K. vallesiana (Honck.) Gaudin, K. albescens DC., K. gracilis
(eugracilis) Pers., K. britannica Domin and K. pseudocristata Domin. There were nine varieties
and several forms distributed between them, together with the hybrid K. x mixta, considered to be
the hybrid between K. vallesiana and K. britannica (Howarth 1933).

Hubbard (1954) reduced the species to two (K. vallesiana and K. cristata), with a comment that
a coastal sand dune variant had been named K. albescens. However, he noted that the British plant
referred to as K. albescens was scarcely distinguishable from inland K. cristata, except perhaps by
its whitish panicles. A further note stated that K. britannica is a dwarf variant with short dense
panicles.

The delimitation of two species continued to be adopted in British Floras throughout the 1960s
and 1970s (Clapham et al. 1962, 1987; Keble-Martin 1965; Hubbard 1984). However, Humphries
(1980) described the genus Koeleria in Flora Europaea in which species 5—13 of the genus are
listed under the K. macrantha group. The author comments that the group is an imperfectly
understood complex, with much variation as a result of polyploidy and adaptation to a wide range
of ecological conditions. He goes on to state that many populations which have been given
specific, subspecific and varietal rank might well represent only ecotypes and variants of different
ploidy levels. As a result, the species in the K. macrantha group, which are based entirely on
morphology, might only represent minor taxa (Tutin ef al. 1980).

Within the K. macrantha group the species K. macrantha (Ledeb.) Schultes (a re-naming of the
older K. cristata) and K. glauca (Schrad.) DC. are both recorded as occurring in Britain, along
with K. vallesiana which in Britain is confined to a handful of sites in Somerset, and will not be
discussed further here.

British Floras and related publications from 1980 onwards have, for the most part, contained the
three species: K. vallesiana, K. macrantha and K. glauca, the last-named also belonging to the
macrantha group. Clapham et al. (1987) recorded the distribution of K. macrantha as “ in turf on

378 J. M. DIXON

well-drained, base-rich soils particularly on chalk and limestone. Locally common” and K. glauca
as “on sandy soils. Widely distributed throughout the British Isles”. Rose (1989) recorded K.
macrantha as “frequent to locally common on dry sandy or calcareous grassland, dunes” and K.
glauca as “common. On sandy soils, especially dunes”. Fitter & Fitter (1984) recorded the two
species as occurring in Britain but commented that the distribution of K. macrantha is not yet
distinguishable from that of K. glauca.

Some authors were rather more cautious: Grime ef al. (1988) merely noted that “recently K.
glauca has been separated from the more common taxon formerly called K. cristata’, Rodwell
(1992) only referred to K. macrantha, while Stace (1997) under the sub-heading for macrantha
noted that “despite several claims the diploid K. glauca has not been confirmed from BI’.

This paper offers a detailed examination of the status of the Koeleria macrantha group in
Britain.

MATERIALS AND METHODS

FIELD WORK

Observations and measurements were made on material belonging to the macrantha group from 51
sites throughout Britain, using the B.S.B.I. distribution map as a guide (Perring & Walters 1962).
The majority were coastal sites but a number of inland sites were also sampled. Site descriptions
are given in Appendix | and Fig. | indicates their approximate geographical location; at three of
the sites (numbers 5, 41 and 44 ) two or more different habitats were sampled. At each site
estimates were made of the average height of stands of vegetation containing K. macrantha (apart
from sites 1, 2 and 3), and an assessment was made of the status and habit of the species.
Accompanying species were also recorded but are not presented here. An estimate of soil depth
was made, and a soil sample collected from the top 10 cm for analysis. The soil was collected with
a Stainless steel trowel, after removing any surface vegetation and litter, and placed in self-sealing
polythene bags. Measurements of pH were made on fresh soil and the rest of the analyses,
comprising loss on ignition, extractable Ca, K, Mg and PO, - P and total organic nitrogen, were
carried out on air-dried soil: methods were according to Allen (1989). Soil analyses are presented
in Appendix 2.

Wherever possible about ten ramets of K. macrantha were collected from each site, far enough
apart to be separate genotypes, and an additional number of inflorescences were gathered from the
same population to ensure sufficient measurements were available. The ramets were wrapped in
damp newspaper and placed in polythene bags, as collection took place over several days.

LABORATORY AND GLASSHOUSE WORK

The length and width of the three longest leaves of eight selected ramets were measured and the
length of the three longest stems; their panicle lengths and widths (measured in the middle of the
panicle) were also measured. Where a ramet did not have three inflorescences, measurements were
included for that population from the extra ones collected. Thus for most populations a total of 24
measurements for each variable were obtained. Three populations could not be sampled
extensively and these are indicated by an asterisk in Appendix 1. In addition, for each population
six inflorescences were selected at random (from different ramets) and one or two spikelets were
removed from the middle of the panicle and measured to the nearest 0-5 mm. Observations were
also made on whether stem thickening was present, the shape of the glumes and lemmas, and the
degree of hairiness present on both leaves and floral parts. Hairiness was subjectively assessed as
glabrous, sparsely hairy, moderately hairy and densely hairy.

After the measurements had been made all the ramets were potted up in 9 x 9 cm plastic pots
containing John Innes No | compost and transferred to an unheated glasshouse. Each pot was
moved one position horizontally and laterally every three weeks to ensure any variations in
environmental conditions were evened out. The plants were watered as necessary, and in the
spring were given four feeds of seedling strength commercial fertilizer “Tomerite’ to encourage
flowering. .

Sampling had taken place during the summers of 1997 (northern Britain) and 1998 (southern
Britain). Each set of plants was grown in the glasshouse for approximately one year under standard
conditions and then harvested.

KOELERIA IN BRITAIN 379

| | | | |
: | H
| : | | |
| | |
i | |
: | !
O Km _ 100
Miles 100 |
| =
| |
|
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| | 35 |
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FIGURE |. Approximate locations of sites sampled for Koeleria.

Measurements were then made, where possible, on each population for the same set of variables
as had been made for the field samples. However, glasshouse populations from sites 15, 20 and 22
did not flower.

In addition, chromosome determinations were carried out for each population. During the spring
of 1998 and of 1999, when new roots were being produced, three pots were selected at random
from each population and about six new, healthy white root-tips were snipped off from those
emerging through the sides of the soil/root mass. These were fixed in 3:1 absolute ethanol / glacial
acetic acid fixative and kept in the refrigerator for periods not exceeding one week until they could
be processed. The root tips were washed in tap water and hydrolysed for 12 minutes at 65° C in
MHCI. This extended hydrolysis was found to be necessary to ensure efficient squashing. The root
tips were stained in Feulgen’s solution for 1—2 hours and then squashed in 45% acetic acid.

380 J. M. DIXON

From the literature K. macrantha in Britain is described as tetraploid (2n = 28) and K. glauca as
being diploid (2n = 14) (Clapham et al. 1987). Therefore, for the current study it was only
necessary to ascertain whether 14 or 28 chromosomes were present. In view of the quantity of
material being processed, to save time it was decided that any squashes with countable spreads of
20+ could safely be deemed to belong to the tetraploid and this method was adopted. For each
population two out of three plants sampled were considered acceptable if good counts were
obtained (not all plants had actively dividing roots); where necessary further plants were sampled.

RESULTS

MEASUREMENTS

These are presented in Appendix 3. Leaf length is highly plastic and mean leaf length was longer
in all the glasshouse plants with the exception of populations from site 39 (Broughton Bay), which
had almost the same mean lengths for field and glasshouse plants, and from site 4 (Nairn) where
the field population had a significantly longer (p = 0-05) mean leaf length (Nairn had the longest
mean leaf length of any field population). The increase was not significant in only five
populations - sites 5a (Branderburg), 6 (Portgordon), 21 (Allonby), 30 (Staxton) and 44a
(Sandwich Bay). These were all populations with mean field leaf lengths >11 cm. All the shortest
populations, that is with mean field leaf lengths < 6 cm, showed the greatest increase in leaf length
in the glasshouse populations and were all significantly (p = 0-001) longer, increasing to at least
twice the field length.

Leaf width is not very variable in K. macrantha, only ranging from 1—4 mm. However, all
populations increased their mean leaf width in the glasshouse, many of them significantly so. The
populations which had not increased significantly in leaf length also tended to be those not
showing significant increases in leaf width either.

Stem length was more variable but showed an increase in all glasshouse populations apart from
site 21 (Allonby) and site 24 (Coxhoe), with mean field lengths of 27-3 and 29-7 cm respectively.
These two populations had the longest mean field stem lengths.

Panicle length increased in most populations in the glasshouse; only one population had a mean
measurement less than that of the field population site 13 (Mull of Galloway), but this was not
significant. Some populations increased significantly in panicle length in the glasshouse while
others did not.

Panicle width also generally increased, but it is difficult to measure this with any degree of .
accuracy and it depends on the age of the panicle, which becomes more contracted as it matures.

Spikelet length, which in the field ranged from 3-5-6 mm, increased in all populations in the
glasshouse apart from site 46 (Camber) which remained the same. However, unlike the leaf length
measurements, there was no significant variation between the ratio of increase for any of the
populations. Whatever the mean field length the mean increase in spikelet length was around 1:1-2,
with the glasshouse spikelet lengths ranging from 4-5—7-5 mm.

OBSERVATIONS

Field populations varied considerably in their habit, with some very rigid, tufted populations such
as from sites 2 (Barra), 12 (Ayr), 17 (Auchenmalg Bay), 23 (Silecroft), 27 (Jenny Brown’s Point);
43 (Cheddar) and 44 (Sandwich Bay - foreshore). Some were very dwarf, prostrate populations
such as from sites 36 (Sizewell), 37 (Thorpeness Sluice) and 42 (Brean Down), some prostrate
“trailing” populations such as from sites 39 (Broughton Bay) and 40 (Rhossili Cliff) and some tall,
bushy populations as at sites 24 (Coxhoe) and 46 (Camber). These habit forms became much less
pronounced in the glasshouse, but populations from sites 2, 12, 17 and 23 remained tufted and
fairly rigid.

Although the field populations varied in leaf colour from light blue-green to glaucous to dark
green on the adaxial surface, and from mid-green to dark green on the abaxial surface, with coastal
populations being generally more glaucous than inland populations, after a year in the glasshouse
there was much closer correspondence in leaf colour, varying from blue-green to mid-green on the
adaxial surface and a uniform mid-green on the abaxial surface.

|

KOELERIA IN BRITAIN 381

Leaf hairiness is extremely variable, with plants from the same population being glabrous to
densely hairy and the degree of hairiness even varying on the same plant. Hairs can be minute and
downy, to long (1-5 mm) on both surfaces and on the margins. Generally leaves are less hairy
towards the tip, but in some populations, for example from site 24 (Coxhoe), the leaves were
densely hairy on both sides almost to the tip, while in others, for example from site 27 (Jenny
Brown’s Point) the leaves were almost entirely glabrous apart from a few hairs around the ligule,
with not even any marginal teeth. In most populations the abaxial surface is more hairy than the
adaxial. In general, leaves tend to be more hairy in southern populations than in northern, but
extremes have been found in both the north and the south of Britain.

None of the c. 1200 stems examined had any sign of thickening; some stems were very robust
but there were no bulbous swellings at the base.

All glasshouse inflorescences were greenish-purple until flowering, becoming silvery and losing
their greenness as the panicles closed, and finally straw-coloured as the panicles matured. Panicles
were variable in shape, and narrowly oblong, spiciform, pyramidal and some almost round
panicles were observed; these variations were maintained in cultivation. The majority of plants had
a moderately to densely hairy rachis, sometimes very fine and downy, sometimes with longer hairs
to 0-5 mm. In only five populations were plants found which had a glabrous to sparsely hairy
rachis - 13 (Ayr), 11 (Troon), 33 (Great Orme), 35 (Holme-next-the-sea) and 44 (Sandwich Bay -
fixed dunes).

Glumes varied from aristate to acuminate (most common) to mucronate; none were blunt when
flattened. Glumes were densely hairy with some hairs to 0-5 mm, to minutely hairy on the keel and
near the tip, to completely glabrous. Awned and very sharply-pointed glumes occurred more
frequently in northern populations, with mucronate glumes more frequent in southern populations.
Lemmas were also either aristate, acuminate (most common) or, rarely, mucronate. Lemmas also
varied in hairiness with some being glabrous and some minutely hairy on the keel and near the tip:
lemmas are never as densely hairy as glumes.

CHROMOSOME COUNTS
All populations were tetraploid with 2n = 28. No diploid plants were found.

TABLE 1. DIFFERENTIAL CHARACTERISTICS FOR KOELERIA MACRANTHA
AND K. GLAUCA

Characteristic Koeleria macrantha Koeleria glauca
1) Habit Tufted Rigid tufted
2) Stems Not thickened, 10-40 cm Robust and with bulbous thickening towards the
base, 10-45(—60) cm
3) Leaves Green or glaucous, flat or folded, Leaves of non-flowering shoots glaucous, more
sometimes slightly rough, up to 15cm x __ or less inrolled and rough to touch, at least above,
1-2 mm densely silvery, not more than 5 cm
4) Leaf sheaths Persistent, with parallel veins Persistent, breaking down into filaments when
old
5) Panicle 20—80(—100) mm x c. 10 mm. Narrowly 20-110 mm x c. 8 mm. Spiciform, interrupted
oblong, silvery-green purplish below; whitish/silvery; rachis densely puberulent
6) Spikelets 2—5(-6) mm 4-5 mm
7) Glumes Acuminate,aristate or mucronate green, = Blunt when flat, pale green or brown
brown or purplish with a silver hyaline
margin
8) Lemma Acuminate and glabrous Subobtuse or mucronate, puberulent, at least
towards the base
9) Chromosomes 2n = 28 (for Britain) 2n = 14 (for Britain)

Sources: Humphries (1980); Clapham et al. (1987).

382 J. M. DIXON

DISCUSSION

Taxonomic differences between K. macrantha and K. glauca are listed in Table 1.
If each character is examined in turn a comparison can be made between the samples collected
in the field and the glasshouse material grown under standard conditions:

1) A number of the field populations of K. macrantha had a tufted habit but with flexible leaves.
while several could be described as having a rigid tufted habit, which in four populations
remained so in the glasshouse.

2) Examination of the stem did not reveal any thickening or bulbous swelling in these four
populations nor indeed in any other population; not a single plant exhibited this characteristic.
Stem length is similar for the two species, but with K. glauca extending less commonly to 60
cm. Most of the K. macrantha populations, both in the field and in the glasshouse, did not
generally extend beyond 40 cm, but in three field populations and 19 glasshouse populations,
stems 44-57 cm long were recorded.

3) None of the plants collected from the field had densely silvery leaves; a few were rough enough
to be described as scabrid, but were not so in their glasshouse counterparts. Sixteen populations
had mean leaf lengths equal to or less than 5 cm, but in the glasshouse, in all these populations,
leaf length increased significantly, with minimal mean lengths of not less than 10 cm. Leaves of
Koeleria plants fold or inroll (K. macrantha does both) when subject to water stress - a
common occurrence on sand dunes and shallow soils over limestone, but are flat when not
subject to a water deficit: all glasshouse plants, watered freely, had flat leaves.

4) Observations on leaf sheath breakdown did not reveal the presence of filaments in either field or
glasshouse populations.

5) Panicles varied in size and shape as described in the results: none exceeded 9 cm either in the
field or in the glasshouse, and the rachis in the majority of populations varied from moderately
to densely hairy, only five populations had a sparsely hairy or glabrous rachis. These included
three small populations, an intermediate one and a tall population (based on leaf length).

6) Spikelet length for the two species is an overlapping range, with K. macrantha extending to 6
mm and K. glauca stated to be not less than 4 mm. In the field only two populations had
spikelets with measurements less than 4 mm, but these had increased to 5 mm in the glasshouse.
Nine populations contained spikelets of 6 mm or over in the field and in the glasshouse this had
increased to 41 populations containing spikelets measuring 6 mm or longer.

7) No plants had glumes that were blunt when flattened. Some had a well-developed hyaline
margin and others much less so. Colouring was variable, depending on the state of maturity.

8) Lemmas varied in shape and also in hairiness as described in the results, and at maturity the
thickened median rib was found to extend into the folded acute apex.

9) All plants were tetraploid.

Koeleria glauca is described as a plant of sand dunes and sandy soils, habitats which are
susceptible to drought and are often nutrient poor. Willis & Yemm (1961) investigated the nutrient
status of three types of dune soils in Devon, and reported that while these had sufficient minor
nutrients they were very deficient in nitrogen, phosphorus and often potassium. The vegetation of
these dune soils produced some forms of several of the common species which were very
depauperate, and which had even been recognised as dwarf varieties. Addition of the major
nutrients led to an increase in height’and weight, often two to three times greater than the control
plants. Ivimey-Cook & Proctor (1966) also commented that the sparse cover and dwarfing of
plants on calcareous grey dunes in the Burren, Co. Clare (v.c. H9) may reflect nitrogen,
phosphorus and potassium deficiencies in the skeletal soil.

In the present study, although no sites were deficient in calcium and magnesium (see Appendix
2), several sites, 9 (Balmedie), 11 (Troon), 12 (Ayr), 19 (Druridge Bay), 23 (Silecroft) and 36
(Sizewell), all from sand dunes or raised beaches are very low in nitrogen and all the plants are of
small stature. Site 36 is also deficient in phosphorus and potassium. Other short populations
occurred in very shallow soils, some were heavily grazed and some occurred on exposed cliffs,
usually in shallow soils. Plants in such stressed habitats are often dwarfed and very tufted relative

KOELERIA IN BRITAIN 383

to plants from more favourable habitats. If the stature of the short populations was a genetic
adaptation to such stressed habitats (as it is perhaps in K. glauca), then one would have expected
the ratio of increase in leaf length brought about by the more favourable conditions of the
glasshouse to be roughly the same regardless of the initial length of the leaf. Figure 2 shows that
this is not so and that the shorter the field population in leaf length the greater the ratio of increase
in the glasshouse, leading to the conclusion that the plants were suffering from environmental
stress in their original habitat, as were those investigated by Willis & Yemm (1961). This is borne
out by the fact that spikelet length, and thus the floral parts, which are generally much less plastic
than the leaf and stem characteristics, did not increase in the linear manner demonstrated by the
regression line in Fig. 2.

The results show that K. macrantha is very variable in many of its morphological characteristics,
and some plants collected in the field had a number of features which might lead them to being
identified as K. glauca, such as leaf length, spikelet length, inrolled leaves and indumentum.
However, after a year of growth under standard conditions all plants had converged in these (and
other) features and were unmistakably K. macrantha, an identification supported by the
cytological evidence of the constant tetraploid state. Wall (1988) investigated the role of B
chromosomes in K. macrantha and sampled extensively from 27 sites (mainly coastal) throughout
Britain, including two sites sampled by the present author. All plants that she examined were
tetraploid and clearly K. macrantha. Thus sampling in depth from 78 sites covering a wide range
of geographical locations and substrate types in Britain has yielded only tetraploid plants.

Wall also commented that plants with high numbers of B chromosomes (K. macrantha can have
up to 6) lack vigour and set fewer seeds. Thus it is possible that the shortest populations might be
those with high numbers of these accessory chromosomes; for example, in the present study
populations from sites 23 (Silecroft), 36 (Sizewell) and 41b (Porthcawl) all produced mean leaf
lengths of less than 11 cm in their glasshouse populations compared with the mean overall
glasshouse leaf length, for all populations, of 16-2 (+ 0-5) cm.

Ratio field to glasshouse leaf length
Uo

0 0.2 0.4 0.6 0.8 ] 122 1.4
Log) field leaf length (cm)

FIGURE 2. Log. mean field leaf length vs. ratio of field to glasshouse leaf length for sampled Koeleria
populations

384 J. M. DIXON

wih (\

FIGURE 3. Comparison of spikelets and spikelet parts of Koeleria macrantha and Koeleria glauca. Top row K.
glauca ; bottom row K. macrantha. G 1 = lower glume; G2 = upper glume; P = palea and pedicel; L = lemma.
Entire spikelets on left. Indumentum not shown.

Because many of the coastal sites are stressed habitats it is possible that some plants exhibiting
some characeristics of K. glauca may well have been misidentified from field specimens, as a
result of the inherent variation in K. macrantha, which is described in Flora Europaea (Tutin et al.
1980) as “ very variable in habit, leaf-size and indumentum”. Confusion may have arisen by the
inrolling of leaves by K. macrantha where water deficits occur.

To ascertain the degree of confusion which might arise in comparing K. macrantha and K.
glauca an examination was made of European material of K. glauca held in K. Measurements and
observations are summarised below.

Stems of K. glauca were mostly very sturdy but in only two specimens was there any slight
thickening which could be described as bulbous. Leaves of non-flowering shoots, which are
presented as an identifying characteristic in Flora Europaea as being not more than 5 cm,
surprisingly turned out to be in many cases considerably longer than this: mean leaf length was 6-1
(+ 0-2) cm, and the range for western Europe was 2-6—10-0 cm. Only about 37% of leaves had a
length less than 5-5 cm. Thus leaf length does not appear to be a strong differentiating feature for
the species. However, the maximum leaf length of 10 cm is considerably shorter than that recorded
by the author for British K. macrantha (24-8 cm). The majority of leaves were inrolled, but some
were folded and some flattened. However, this might have been an artefact of the preservation
process. Leaves of K. glauca were all “felted” with minute rough teeth, giving the silvery, scabrid
characteristic described for the species (see Table 1). This felting extends to the tip of the leaf and
is in contrast to K. macrantha where the leaves have longer, true hairs rather than teeth (which are
confined to the margins) and in most populations these do not extend to the tip. The leaf sheaths of
K. glauca break down into a network of filaments, in a similar manner to those of K. vallesiana, in
contrast to those of K. macrantha, where the leaf sheaths are persistent. =

Panicle lengths to 115 cm were measured and there is less variation in the shape of the panicles
in K. glauca compared with K. macrantha: 90% were spiciform with the rest exhibiting a slight
pyramidal shape. Koeleria glauca is characterised as having the panicles interrupted below and

KOELERIA IN BRITAIN 385

77% of the specimens examined were interrupted compared with only 14% in specimens of K.
macrantha (from a much larger sample) with interrupted panicles.

Spikelets of K. glauca ranged from 2-5—5-5 mm with a mean of 4-7 mm (median 4-0 mm) which
was very little different from those of K. macrantha for British specimens (range 3-5—6-0 mm,
mean 4-9 mm and median 4-8 mm). These figures do not agree with those quoted in Flora
Europaea (see Table 1).

Illustrations of the spikelets and the spikelet parts are provided in Fig. 3, and it can be seen that
the glumes of the two species are different, with those of K. macrantha being pointed and K.
glauca being blunt. The lemmas of K. macrantha are also more pointed than those of K. glauca
and the median rib is thicker and more rigid than in K. glauca, and extends into the point.

Some of the differential characteristics listed in Table 1, such as the rigid tufted habit of K.
glauca and panicle width, were not possible to determine from herbarium specimens. However,
these aspects notwithstanding, and in spite of the greater convergence of the two species than is
indicated in their descriptions in Flora Europaea, there is a marked separation of the two species
as far as the leaf sheath breakdown, leaf pubescence and floral characteristics are concerned,
together with the more slender and uniform spiciform shape of the panicle, which even in
herbarium specimens is more intensely silvery than those of K. macrantha.

Thus the presented measurements and observations provided in this paper do not support the
presence of K. glauca in Britain. In the most recent Floras, British authorities have more-or-less
discarded K. glauca from the British list (see Cope 1998); the current paper affirms this view.

ACKNOWLEDGMENTS

My thanks are due to Dr A. D. Headley for providing material from Barra, Rhum and Skye, to Dr
D. J. Hambler for drawing Fig. 3, and to Dr D. Comber, Dr W. H. G. Hale and Prof. M. R. D.
Seaward for help and advice.

REFERENCES

ALLEN, S., ed. (1989). Chemical analysis of ecological materials, 2nd ed. Blackwell Scientific Publications,
Oxford.

CLAPHAM, A. R., TUTIN, T. G. & Moore, D. M. (1987). Flora of the British Isles, 3rd ed. Cambridge
University Press, London.

CLAPHAM, A. R., TUTIN, T. G. & WARBURG, E. F. (1962). Flora of the British Isles, 2nd ed. Cambridge
University Press, London.

Cope, T. A. (1998). Koeleria, in RICH, T. C. G. & JERMY, A. C., eds. Plant crib 1998: B.S.B.I., London.

FITTER, A. & FITTER, R. (1984). Collins Guide to the grasses, sedges, rushes and ferns of Britain and
northern Europe. William Collins, Glasgow.

GRIME, J. P., HODGSON, J. G. & HUNT, R. (1988). Comparative plant ecology. Unwin Hyman, London.

HOWARTH, W. O. (1933). Notes on Koeleria Pers. Reports of the Botanical Exchange Club 10: 37-41.

HUBBARD, C. E. (1954). Grasses. Penguin Books, Harmondsworth.

HUBBARD, C. E. (1984). Grasses. 3rd ed. Penguin Books, Harmondsworth.

HUMPHRIES, C. J. (1980). Koeleria Pers., in TUTIN, T. G. et al., eds. Flora Europaea 5: 218-220. Cambridge
University Press, Cambridge.

IVIMEY-COoK, R. B. & PRocTorR, M. C. F. (1966). The plant communities of the Burren, Co. Clare.
Proceedings of the Royal Irish Academy 64B: 211-301.

KEBLE-MARTIN, W. (1965). The concise British flora in colour. Ebury Press & Michael Joseph, London.

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

RODWELL, J. S., ed. (1992). British plant communities. Vol. 3. Grasslands and montane communities.
Cambridge University Press, Cambridge.

ROSE, F. (1989). Grasses, sedges, rushes and ferns of the British Isles and north-western Europe. Viking,
London.

STACE, C. A. (1997). New Flora of the British Isles, 2nd ed. Cambridge University Press, Cambridge.

WALL, S. (1988). The role of the B chromosome in the genetic system of a tetraploid grass (Koeleria cristata
L.). Ph.D. thesis, University of Manchester.

WILLIS, A. J. & YEMM, E. W. (1961). Braunton Burrows: mineral nutrient status of the dune soils. Journal of
ecology 49: 377-390.

(Accepted July 2000)

386 J. M. DIXON

APPENDIX 1. SITE DETAILS FOR KOELERIA SAMPLING. NUMBERS REFER TO
APPROXIMATE LOCATION ON MAP HIG. 1

No Location NGR Alt. Slope Aspect

1* Nr. Dunvegan, Isle of Skye, Highland NG223549 3m flat :
Koeleria frequent on basalt; also occasional on sandy beach with broken coral

2* Eoligarry, Barra, Western Isles NL695054 lm flat -
Machair. Koeleria common to abundant in ungrazed areas

3. Fonchra, Rhum, Highland NG337003 3500s 2 Sa SE
Base of steep, vegetated cliff over basalt. Koeleria frequent

4 Nairn, Highland NJ866569 2m flat -
Pebbly track Koeleria dominant, also in longer vegetation at side of track

5 Nr Branderburgh, Grampian NJ204714 15m ZO? N
Dune-heathland with mix of calcicoles and calcifuges, also on rocks at edge of sea

6 Portgordon, Grampian NJ401645 3m flat .
Sandy soil over pebbles, Koeleria scattered but frequent

7 St. Combs, Grampian NK059630 15m 12° NE
Raised beach,vegetation to 40 cm high, Koeleria in isolated clumps

8 Cruden Bay, Grampian NK096358 15m 40° WSW
All plants scattered in pockets across face of granite cliff, Koeleria frequent

9 Balmedie, Fife NJ977182 15m 10° NNE
Short grazed turf, Koeleria common, plus longer ungrazed turf, with scattered Koeleria

10 Lunan Bay, Tayside NO691502 10m BD NW
Lightly grazed, fixed dunes, vegetation 3-6 cm. Koeleria scattered but frequent

11* Troon, Strathclyde NS324305 3m flat -
Fixed dunes, Koeleria scattered but frequent

12 Ayr, Strathclyde NS328205 10m 20° W
Edges of fixed dunes (top was mown), Koeleria very scattered

13. Dunure, Strathclyde NS254158 2m flat -
On rocks (hornfels) at top of beach in shallow pockets of soil, isolated plants

14 Girvan, Strathclyde NX160943 10m 90° N

In shallow pockets of soil on rocks (phyllite) at top of shore, also on flat tops of rocks, isolated plants, an
abundance of ants here

15 Labarax, Dumfries & Galloway NX972601 5m flat -
Koeleria scattered on fixed dunes, isolated plants also in rock crevices (slate)

16 Mull of Galloway, Dumfries & Galloway NX159307 50m ee E
On grassy slope and also on flat cliff top. Koeleria frequent but scattered

17 Auchenmalg Bay, Dumfries & Galloway NX243515 6m flat -
Very short, prostrate turf on track, vegetation 10-15 cm. Koeleria dominant here

18 Alnmouth, Northumberland NZ256115 3m flat -
Fixed dunes, vegetation 30-45 cm high. Koeleria frequent but scattered

19 Druridge Bay, Northumberland NZ292936 10m flat -
Fixed dunes, vegetation 10-15 cm high, grazed (horses). Koeleria frequent

20 Silloth, Cumbria NY090493 10m flat -
Fixed dunes, vegetation to 45 cm high. Koeleria frequent

21 Allonby, Cumbria NY080440 7m flat -
Partially fixed dunes,. Koeleria on top but not sides of dunes, vegetation to 45 cm

22 St. Bees, Cumbria NY957118 50m 5° E
Short, but ungrazed, turf on cliff top. Koeleria frequent to abundant

23 Silecroft, Cumbria SD123805 10m flat -
Top of bank of consolidated pebbles, short, ungrazed turf. Koeleria common, also on track, trampled and
very short

24 Coxhoe, Durham NZ326364 132m _- flat -
Dolomite quarry spoil heap. Koeleria in scattered small patches :

25. Far Arnside, Cumbria SD453760 12m 20° SW
Grassy slope over Carboniferous limestone, just above beach. Koeleria frequent in small patches

26 Thrang End, Yealand Redmanyne, Cumbria SD494766 70m 10° NNW

Carboniferous limestone pasture, sheep grazed. Koeleria frequent

* = only small samples available for measurements and observations

44

45

46

47

48

49

50

51

KOELERIA IN BRITAIN 387

Location NGR Alt. Slope Aspect

Jenny Brown's Point, Silverdale, Lancashire SD463734 5m 20" S
Sandy bank over Carboniferous limestone, sloping into sea. Koeleria scattered among rocks

Bishopdale, North Yorkshire SD943803 AL9m== 20° S
Grassy bank over Carboniferous limestone, ungrazed vegetation to 45 cm high. Koeleria frequent

Langcliffe, North Yorkshire SD833656 366m fiat .
Koeleria isolated but frequent in pockets of soil on clints of limestone pavement, and in shallow grikes

Staxton, North Yorkshire TAO11782 LOZ mse 5° E
Koeleria scattered among tall vegetation on overgrown path of disused chalk pit

Wormhill, Derbyshire SK107756 372m: 18: S
Isolated plants in pockets on Carboniferous limestone outcrop. Turf 3-12 cm high

Monyash, Derbyshire SK160661 274m fiat -
On top of Carboniferous limestone outcrop. Koeleria scattered in small clumps. Some also present on
more or less vertical rock face.

Great Ormes Head, Gwynedd SH780836 76m ise N

On Carboniferous limestone sea cliff, above tidal limits but within splash zone. Koeleria abundant, some
on vertical face nearer the sea

Little Ormes Head, Gwynedd SH8 13823 130m,” “45° W
Very shallow soil over Carboniferous limestone. Koeleria abundant

Holme-next-the-Sea, Norfolk TF708448 2m flat
Partially fixed dunes, long loose vegetation to 60 cm. Koeleria occasional and very scattered, musty at
edge of board-walk

Sizewell, Suffolk TM475628 3m flat :
Raised beach, just above sea level. Sparse, very short turf on almost pure sand. Koeleria scattered but
frequent

Thorpeness Sluice, Suffolk TM472593 240 flat -
On inland side of a shingle bar, shingle mixed with sand. Koeleria abundant

Aldeburgh, Suffolk TM465575 3m 6° NW
Sandy bank at side of car park. Koeleria dominant

Broughton Bay, West Glamorgan SN416928 30 m oy SE
Early fixed dunes, dense tall vegetation to 60 cm. Koeleria restricted to path edges where vegetation was
rather shorter

Rhossili Cliff, West Glamorgan SN405878 50m 25° N
Sloping cliff top, shallow soil over Carboniferous limestone. Koeleria scattered

Nr. Porthcawl, Mid Glamorgan ST804782 6m 5° NE
Fixed dunes, dense tall vegetation to 60 cm. Koleria very scattered, mostly on path edges. Also on
limestone outcrop nearby

Brean Down, Somerset ST286587 45m flat -
Dwarf population on exposed Carnoniferous limestone cliff top; very shallow soil. Koeleria abundant

Nr Cheddar, Somerset ST504517 245 m 15% S
Koeleria frequent on Carbonifeous limestone outcrop, occasional in pasture

Sandwich Bay, Kent TR364578 3m 10° WSW
Roadside bank, also on flat fixed dunes and on flat sandy/pebbly foreshore. Koeleria common on bank
and in dunes, more scattered on foreshore

St. Margaret's At Cliffe, Kent TR372447 80 m jae ESE
In short to moderately long vegetation (12-40 cm) on top of chalk cliff. Koeleria scattered in small
populations

Camber Golf Course, East Sussex TR946194 5m 25% ESE
Partially fixed dunes of deep sand. Koeleria on edges of banks, frequent

Nr. Beachy Head, East Sussex TQ592973 154m 90° WwW
Koeleria virtually limited to roadside vertical bank, not in the pasture behind it

Durlston Head, Dorset SZ025769 61m 22. S
Very shallow (c. 5 cm) soil over limestone. Very short (2—7 cm) grazed turf. Koeleria frequent but
scattered

Carbis Bay Golf Course, Cornwall ST294588 95m 12° W
Fixed dunes, Koeleria mainly on edges of banks and paths

Mullion Cove, Cornwall SW666106 61m 29° N
In short turf on top of cliff over serpentine. Koeleria scattered but frequent

Kynance Cove, Cornwall SW686135 64 m 10° SW

Koeleria abundant in very short turf over serpentine

388 J. M. DIXON

APPENDIX 2. SOIL ANALYSES FOR SITES SAMPLED FOR KOELERIA

Site No pH % Losson Camg 100g' Mg mg 100g1 Kmg100g-' PO,-Pmg_ Total N (%)
Ignition 100g-'

] 5:8 56-1 205 88 39 1-12 1-92
Z 6-4 5-5 292 4] ANG 1-12 0-23
3 6-2 30-8 198 66 ZS 0-88 0-84
+ 6-2 Res. 268 99 28 1-44 0-66
5a 5-9 6:8 205 52 14 0-56 0-02
5b 6-0 5-0 190 a 14 1-12 0-03
6 5:8 4-5 759 4] 16 0-90 0-20
yi 5-7 10-4 365 ik i) 0-92 0-39
8 6-1 17-1 348 119 34 0-92 0-56
9 6-0 =| 183 4] 16 0-90 0-10
10 5-9 4.] 198 40 ibs: 0-88 0-21
11 6-5 1:8 A Kos 38 14 12 0-06
2 6:0 1-7 190 4] 15 0-80 0-04
i 6-2 3-4 210 61 17 1-20 0-06
14 5-9 24-2 185 83 28 0-68 0-80
15 6-1 8-4 203 60 21 0-76 0-31
16 soil analysis not available

17 5:8 4-0 188 51 HIS 1-12 0-22
18 6-0 3-9 255 44 14 1-12 0-11
19 6:3 3-2 205 58 14 0-76 0-10
20 ao | 5-4 250 a2 Wy 0-60 0-21
21 6-0 7:8 360 56 18 0-70 0-24
Ze 6:5 8-4 150 60 24 0-88 0-27
23 6-4 5-4 210 iB 17 1-12 0-15
24 6:3 44 255 50 15 0-79 0-10
ZA 6:3 11-8 448 39 20 0-60 0-48
26 5-9 23-0 490 40 24 0-90 0-97
27 6:5 7:9 320 58 22 0-88 0-36
28 6:3 19-2 650 58 26 0-76 0-69
20 6-9 n/a 278 38 16 0-80 0-36
30 7-0 5-4 364 59 LF 0-68 0-47
a1 6:6 32°1 625 13 20 0-96 0-99
a2 6:6 11-9 330 14 2 2-80 0-57
33 6:2 17-2 550 56 25 0-56 0-33
34 6:3 7:6 383 39 19 0-68 0:27
35 5-5 1-7 69 13 5 0-48 1-00
36 5:8 0-3 56 16 5 0-12 0-10
3) 6-4 8-6 75 35 14 3-20 0-37
38 6-5 4-5 200 25 8 0-96 0-24
39 6-2 5-4 144 16 8 0-58 0-18
AO 6-4 16-4 460 55 40 0-46 0-75
Ala 6:6 6-6 356 25 11 0-48 0-25
Alb 6-3 32-9 _ 431 n/a 39 1-12 1-18
42 6-3 17-4 420 |W 2s) 39 1-00 0-77
43 6:0 17-7 363 20 13 0-48 0-61
44a 6-7 8-3 356 26 19 0-90 0-19
44b 6-8 4-0 119 24 8 iheesD 0-12
44c 6:8 ai 194 20 10 0-48 0-24
45 7-0 28-8 750 44 54 1-44 0-16
46 6-6 4.] Lol 23 9 0:90", 0-16
47 68 | 72 Zoi 2 11 0-94 0-26
48 7:0 16-7 500 100 54 1-00 0-59
49 7-0 7-4 a13 29 2) 0-46 0-29
50 6-1 31:3 150 388~ ». 50 1-58 1-05
51 6-0 35-2 106 200 46 0-31 0-88

5a = dune heathland; 5b = rocks; 41a = dunes; 41b = outcrop; 44a = roadside; 44b = foreshore: 44c = fixed
dunes

KOELERIA IN BRITAIN

389

APPENDIX 3. MEAN FIELD AND GLASSHOUSE MEASUREMENTS (+ SE) AND
SIGNIFICANT DIFFERENCES FOR KOELERIA SAMPLES

Site
1 Field
Glass.
2 Field
Glass.
3. Field
Glass.
4 Field
Glass.
5a Field
Glass.
5b Field
Glass.
6 Field
Glass.
7 Field
Glass.
8 Field
Glass.
9 Field
Glass
10 Field
Glass.
11 Field
Glass.
12 Field
Glass.
13 Field
Glass.
14 Field
Glass.
15 Field
Glass.
16 Field
Glass.
17 Field
Glass.
18 Field
Glass.
19 Field
Glass.
20. Field
Glass.
21 Field
Glass.
22 Field
Glass.
23 Field
Glass.
24 Field
Glass.
25 Field
Glass.
26 Field
Glass.

Leaf length
(cm)
5-7 (0-5)
19-8 (0-6) 3
6-0 (0-4)
16-4 (0-9) 3
8-5 (2:3)
16-5 (0-8) 3
16-3 (1-7)
14-7 (0-5) 1
5-9 (0-3)
16-5 (0-7) 3
12-7 (0-7)
14-0 (0-5) N
13-6 (0-8)
15-3 (0-5) N
11-9 (0-7)
23-8 (0-4) 3
9-2 (1:2)
15-5 (0-7) 2
4-5 (0-4)
16-5 (0-5) 3
7-1 (0-5)
18-0 (0-6) 3
3-9 (0-2)
16-5 (0-6) 3
3-6 (0-4)
18-5 (0-5) 3
10-8 (0-6)
22°1.(1-7) 3
7:5 (0-6)
11-1 (0-3) 3
3-3 (0-3)
16-2 (0-6) 3
8-6 (0-9)
17-6 (0-4) 3
4-6 (0-2)
13-9 (0-9) 3
8-1 (0-5)
21-1 (0-9) 3
3-9 (0-2)
20-7 (0-7) 3
11-7 (0-9)
16-4 (0-2) 3
13-7 (0-6)
15-3 (0-5) 1
5-5 (0:3)
15-3 (0-4) 3
3-8 (0-2)
10-0 (0-4) 3
15-1 (0-9)
25-8 (0-9) 3
8-4 (0-6)
17-9 (0-4) 3
4-9 (0-3)
17-7 (0-7) 3

Leaf width

(mm)
1-2 ©1)
2-9 (0-1) 3
1-5 (0-1)
P7T(O1)3
1-4 (0-1)
2-8 (0-1) 3
2.2 (0-1)
2.7 (0-2) N
m2)
2.4 (0-1) 1
1-8 (0-1)
22 (0.0)
2.0 (0-1)
2-5 (0-1) 1
1-9 (0-1)
2-6 (0-1) 3
2.3 (0-1)
2-9 (0-1) 3
1-6 (0-1)
2-4 (0-1) 3
1-8 (0-1)
2-5 (0-1) 3
1-6 (0-1)
21 (0:1)3
1-7 (0-1)
2301)
2-8 (0-1)
2.9 (0-2) N
1-9 (0-1)
2-4 (0:2) 2
1-8 (0-1)
D301)3
2.0 (0-1)
2-6 (0-1) 3
2.0 (0-1)
2-6 (0-1) 3
2.3 (0-1)
2.9 (0-1) 3
1-8 (0-1)
2-4 (0-1) 3
2.0 (0-1)
DIOLS
2-1 (0-1)
2-5 (0-1) 2
Dna)
3-1 (0-1) 3
2.1 (0-1)
2.7 (0-2) 1
2.0 (0-1)
2-5 (0-1) 3
2.3 (0-2)
3-0 (0-1) 3
1-5 (0-1)
2-5 (0-1) 3

Stem length

(cm)
9-7 (0-5)
24-3 (1-3) 3
18-2 (0-9)
33-5 (1-2) 3
22:6 (1-2)
32:1-°C1-3)3
PA eA Tp)
25-0 (1:0) N
10-9 (0-9)
20-8 (0-9) 3
17-4 (1-7)
19-1 (0-7) N
24-9 (1-7)
36-5 (1-5) 3
24-2 (1-2)
22-2 (0-7) N
18-6 (1-6)
30-7 (1-9) 3
17-5 (1-9)
28-0 (1-1) 3
9-6 (0-6)
31-0 (1-2) 3
3-9 (0-4)
25-4 (1-0) 3
11-4 (0-6)
24-9 (0-3) 3
17-0 (0-9)
23-4 (1-3) N
13-2 (1-3)
17-1 (1-0) 1
5-9 (0-7)
n/a
14-9 (2-5)
22-9 (1-6) 3
12-3 (1-2)
29-3 (0-9) 3
14-8 (0-6)
26-4 (1-9) 3
11-4 (0-7)
24-7 (0-2) 3
24-3 (1-5)
n/a
27-3 (0-9)
20-9 (0-9) 3

3-8 (0:3)
11-0 (0-8) 3
29-7 (2:9)
25-5 (0-8) N
15-5 (0-7)
33-9 (1-3) 3
21-4 (1:3)
37-6 (1-6) 3

(cm)
2:7 (0-2)
5-0 (0-3) 3
4.0 (0-1)
5-0 (0-2) 3
3-4 (0-2)
6-5 (0:2) 3
4.7 (0-3)
6-0 (0-2) 2
2-8 (0-2)
6-2 (0-1) 3
3-1 (0-2)
5-6 (0-2) 3
4-0 (0-2)
7-0 (0-2) 3
3-5 (0-2)
3-5 (0-1) N
3-1 (0-2)
S=1(0-1) 3
3-2 (0-2)
5-6 (0-3) 3
3-1 (0-3)
5-7 (0-2) 3
1-7 (0-2)
4-7 (0-2) 3
3-8 (0-3)
6-9 (0-1) 3
5-9 (0-4)
6-0 (0-3) N
3-0 (0-2)
4-1 (0-1) 3
2S (0-1)
n/a
3-6 (0-3)
3-3 (0-2) N
2-7 (0-2)
4-5 (0-1) 3
2-8 (0-2)
5-1 (0-2) 3
2-6 (0-1)
4-5 (0-2) 3
5-2 (0-2)
n/a
3-9 (0-5)
4-6 (0-2) N
3-4 (0-2)
n/a
2-3 (0-05)
4-7 (0-2) 3
5-4 (0-4)
7-1 (0-1) 3
3-6 (0-3)
7-0 (0-1) 3
3-5 (0-2)
5-6 (0-1) 3

Panicle length Panicle width

Spikelet

(mm) length (mm)
3-6 (0-3) 5-4 (0-1)
5-5 (0-4) 1 6-8 (0-2) 3
5-5 (0-6) 5-0 (0-3)
6-4 (0-2) N 6-8 (0-1) 3
4-3 (0-2) 5-2 (0-1)
6-3 (0-2) 3 6-3 (0-2) 3
4-0 (0-2) 5-1 (0-1)
4-5 (0-1) 1 5-8 (0-2) 3
3-7 (0-2) 4-8 (0-1)
5-5 (0-1) 3 5-2 (0-1) 1
3-3 (0-3) 4-8 (0-1)
5-0 (0-2) 2 5-9 (0-2) 3
3-9 (0-1) 4.7 (0-1)
5-2 (0-2) 3 6-9 (0-1) 3
4.9 (0-1) 5-2 (0-2)
5-2 (0-2) 3 6-0 (0-3) 1
4-5 (0-2) 5-0 (0-3)
5-8 (0-2) 3 5-6 (0:2) N
3-9 (0-2) 5-0 (0-1)
5-6 (0-3) 3 5-7 (0-1) -2
3-2 (0-2) 4-6 (0-2)
4-6 (0-2) 3 6-2 (0-2) 3
3-4 (0-2) 4-8 (0-1)
4.7 (0-2) 3 5-3 (0-1) 1
4-5 (0-2) 4.4 (0-2)
5-0 (0-3) N 5-6 (0-2) 3
7-1 (0-5) 5-5 (0-2)
4-6 (0-2) 3 5-8 (0-1) N
4-6 (0-2) 4-9(0-2)
5-1 (0-2) 2 5-5 (0-2) 1
4-1 (0-2) 4-3 (0-1)
n/a n/a
4-9 (0-3) 4.7 (0-2)
5-0 (0-1) N 6-1 (0-2) 3
4-8 (0-2) 5-3 (0-2)
5-8 (0-2) 2 5-4 (0-2) N
4-5 (0-2) 4-6 (0-2)
5-7 (0-2) 2 5-7 (0-2) 2
4-4 (0-2) 4-8 (0-2)
5-0 (0-2) 1 5-3 (0-2) N
4-5 (0-2) 5-2 (0-2)
n/a n/a
5-0 (0-3) 4.7 (0-1)
5-0 (0-2) N 5-2 (0-2 1
4-6 (0-2) 5-5 (0-2)
n/a n/a
4.2 (0-2) 5-3 (0-1)
4-4 (0-2) N 5-4 (0-2) N
4-8 (0-3) 4-8 (0-2)
4-1 (0-3) N 5-7-(0:2) 2
4-0 (0-2) 4.9 (0-2)
5-7 (0-4) 3 5-3 (0:2) N
n/a 4-7 (0-3)
4-5 (0-1) 5-8 (0-3) 2

n/a = not available 5a = dune heathland; 5b = rock.
Numbers in bold indicate levels of significance for t-tests between the field and glasshouse populations. N =
not significant, 1 = p at 0-05, 2 = p at 0-01 and 3 =p at 0-001

390

Site

27 Field
Glass.
28 Field
Glass.
29 Field
Glass.
30. —~ Field
Glass.
31 Field
Glass.
32 Field
Glass.
33 Field
Glass.
34 Field
Glass.
35. Field
Glass.
36 —- Field
Glass.
37 Field
Glass.
38 Field
Glass.
39 Field
Glass.
40 Field
Glass.
41a Field
Glass.
41b Field
Glass.
42 Field
Glass.
43 Field
Glass.
44a Field
Glass.
44b Field
Glass.
44c Field
Glass.
45 Field
Glass
46 Field
Glass.
47 Field
Glass.
48 Field
Glass.
49 Field
Glass.
50 Field
Glass.
51 Field
Glass.

Leaf length
(cm)

5-5 (0-7)
17-8 (0:6) 3
10-4 (1-1)
18-3 (0-4) 3
5-9 (0-3)
17-9 (0-9) 3
13-2 (0-7)
13-6 (1-1) N
6:3 (0-7)
13-7 (0-6) 3
6-6 (0-6)
14-2 (0-8) 3
4-4 (0-2)
18-4 (0-8) 3
4-4 (0-2)
19-6 (0-5) 3
9-6 (0-6)
22-9 (1-0) 3
4-6 (0-3)
10-0 (0-2) 3
5-0 (0-3)
13-3 (0-5) 3
7-0 (0-3)
13-5 (0-3) 3
12-1 (1-0)
12-0 (0-4) N
5-0 (0-3)
17-7 (0-8) 3
6-5 (0-3)
12-3 (0-3) 3
5-0 (0-3)
10-6 (0-4) 3
2-7 (0-2)
12-2 (0-3) 3
5-7 (0-3)
14-1 (0-7) 3
14-1 (0-6)
14-9 (0-3) N
13-6 (0-5)
16-7 (0:6) 3
11-0 (0-8)
15-4 (0-4) 3
9-4 (0-6)
18-1 (0-8) 3
9-2 (0-6)
19-0 (0-7) 3
8-8 (0-7)
17-5 (0-6) 3
4-6 (0-3)
13-7(0:5)3
8-4 (0-6)
16-2 (0-9) 3
9-1:(0-4)
13-8 (0-4) 3
6-9 (0-4)
11-9 (0-3) 3

Leaf width
(mm)

1-8 (0-1)
2-5 (0-05) 3
F001)
2-8 (0-1) 3
1-4 (0-1)
2-4 (0-05) 3
1-5 (0-1)
2:3 (0-1) 3
1-8 (0-1)
2101)2
1-9 (0-1)
2-0 (0-1) N
1-7 (0-1)
2-6 (0-1) 3
1-7 (0-1)
2-4 (0-1) 3
1-9 (0-1)
23 O4)2
1-8 (0-1)
2:3 (0-1) 3
2-0 (0-1)
2-1 (0-1) N
2-1 (0-1)
2.3 (0-1) N
2-2 (0-1)
2-4 (0-1) 1
L001)
2-3 (0-1)3
1-8 (0-1)
2-4 (0-1) 3
1-7 (0-1)
2:3 (0-1) 3
1-5 (0-05)
2-0 (0-1) 3
1-5 (0-1)
2-1 (0-1) 3
2-3 (0-1)
2.5 (0-1) N
2-4 (0-1)
2-6 (0-1) 1
2-0 (0-1)
2-4 (0-1) 3
2-0 (0-1)
2-5 (0-1) 3
2-1 (0-4)
2-5 (0-1) N
1-8 (0-)
2-4 (0-1) 3
1-5 (0-1)
2-3 (0-1) 3
1-8 (0-1)
27 O13
2101)
2.3 (0-2) N
2-1 (0-1)
2-6 (0-1) 3

J. M. DIXON

Stem length
(cm)

20103)
26-9 (1-7) 3
175 23)
274A03)3
16-2 (1-6)
29.6 (1-1) 3
22.5 (1-2)
25-9 (1-8) N
10-2 (1-1)
20-5 (2:6) 2
17-7 (1-1)
20-8:(1-2)3
8-5 (0-8)
23-6 (1-2) 3
8.5 (0-8)
30-9 (1-7) 3
17-5 (1-2)
34-4 (2-4) 3
11-8 (0-8)
28-0 (1-1) 3
12-0 (0-9)
44-1 (1-9) 3
19-3(1-0)
39-4 (1-8) 3
13-8 (0-6)
19-6 (1-1) 2
7-9 (0-9)
22-3 (1-4) 3
14-7 (0:5)
25-1 (1-4) 3
10-4 (0:6)
28-0 (0-9) 3
8-2 (0-8)
25-4 (0-9) 3
21713)
42-9 (2-1)3
29-9 (2:8)
30-9 (0-8) N
26-4 (2:2)
41-4 (0-8) 3
30-4 (1-4)
39-1 (1-0) 3
20-3 (1-0)
32-8 (2:6) 3
28-1 (0-9)
28-7 (1-6) N
14-8 (0-7)
23-5 (0-2) 3
10-8 (1-0)
25-2 (1-9) 3
15-5 (0-8)
16-4 (1-1) N
19-7-1-0)
20-8 (0-8) 3
10-8 (0-7)
14-5 (0-4) 3

Panicle
length (cm)

3-5 (0-3)
6:20-1)3
3-9 (0-2)
6-2 (0-2) 3
3-1 (0-2)
6-0 (0-1) 3
3-1 (0-2)
5-5 (0-2) 3
2-5 (0-2)
4-5 (0-3) 3
3-5 (0-2)
5-5 (0-2) 3
2-8 (0-3)
5-5 (0-1) 3
2-8 (0-3)
5-4 (0-2) 3
4-1 (0-2)
6-8 (0-1) 3
3.2 (0-2)
5-5 (0-2) 3
2-8 (0-1)
6-9 (0:2) 3
4-1 (0-1)
6:5 (0:2) 3
3-9 (0-2)
4-4 (0-2) N
2.3 (0-1)
5-0 (0-2) 3
3.3 (0-1)
5-8 (0-3) 3
2-9 (0:3)
4-6 (0-2) 3
2.2 (0-1)
5:7 (0-2) 3
3-5 (0-2)
6-2 (0:2) 3
5-9 (0:3)
6:8 (0:2) N
4-8 (0-2)
6-6 (0-13
5-4 (0:3)
6-9 (0-2) 3
4:3 (0-2)
6-6 (0:3) 3
5-6 (0:3)
5-6 (0-1) N
3-1 (0-2)
5.3 (0-2) 3
2.7 (0-2)
5-4 (0-2) 3
3.3 (0-2)
4-1 (0-1) 2
4-1 (0-2)
5.0 (0-1) 2
3.3 (0-2)
4-4 (0-1) 3

41a = dunes; 41b = outcrop; 44a = roadside; 44b = foreshore; 44c fixed dunes
Numbers in bold indicate levels of significance for t-tests between the field and glasshouse populations. N =
not significant, 1 = p at 0-05, 2 = p at 0-01 and 3 = p at 0-001

Panicle width

(mm)

n/a

4-6 (0-2)
3-8 (0-2)
4-4 (0-2) N
3-4 (0-2)
5-0 (0-2) 3
3-7 (0-1)
4-9 (0-2) 3
3-6 (0-2)
4-6 (0-3) 2
3-6 (0-2)
4-2 (0-2) 1
4-2 (0-2)
4-9 (0-2) 2
4-2 (0-2)
4-4 (0-2) N
3-3 (0-1)
3-9 (0-2) 2
2-9 (0-1)
4-6 (0-1) 3
2-9 (0-1)
4-9 (0-2) 3
3-8 (0-2)
4-7 (0-2) 1
3-7 (0-2)
5-7 (0-2) 3
3-1 (0-2)
6-0 (0-7) 3
3-9 (0-2)
5-8 (0-3) 3
3-4 (0-3)
4-7 (0-1) 3
2-9 (0-1)
4-2 (0-1 )3
3-3 (0-1)
3-9 (0-3) 1
3-9 (0-1)
5-1 (0-2) 3
3-8 (0-2)
4-7 (0-1) 3
3-7 (0-2)
4-8 (0-2) 2
3-9 (0-2)
4-5 (0-3) N
4-1 (0-1)
4-6 (0-2) N
3-3 (0-1)
4-4 (0-1) 3
3-0 (0-2)
4-5 (0-3) 3
4-1 (0-2)
5-6 (0-3) 3
4-3 (0-2)
5-4 (0-2) 2
4-9 (0-2)
6-1 (0-3 )2

Spikelet
length (mm)

4-9 (0-2)
5-8 (0-2) 3
5-1 (0-2)
5-6 (0-2) N
4-8 (0-2)
5-4 (0-1) 2
4-8 (0-1)
n/a

4-3 (0-2)
5-9 (0-3) 2
4-8 (0-3)
5-6 (0-1) 1
4-8 (0-2)
6-1 (0-2) 3
5-2 (0-2)
5-8 (0-2) N
5-2 (0-2)
6-0 (0-1) 3
4-4 (0-2)
5-3 (0-1) 2
4-2 (0-1)
5-2 (0-2) 2
4-3 (0-2)
5-7 (0-3) 2
4-8 (0-1)
5-1 (0-2) N
4-8 (0-1)
5:3: (0-1). 4
4-9 (0-1)
6-4 (0-1) 3
4-3 (0-1)
5-3 (0:2) N
4-3 (0-1)
5-3 (0-2) 2
4-8 (0-2)
5-4 (0-2) 1
4-5 (0-2)
5-4(0-1)N
4-7 (0-2)
5-0 (0-1) 2
4-9 (0-1)
5:3 (0-2) N
4-9 (0-2)
5-4(0-1)N
5-3 (0-3)
5-3 (0-3) N
4-3 (0-3)
5-0 (0:2) N
4-8 (0-2)
5:7 (0-1) 1
4-8 (0-2)
5-3 (0-2) N
4-8 (0-1)
5-9 (0-3) 2
5-0 (0-2)
5-6 (0-1) 1

Watsonia 23: 391-400 (2001) 391

The decline of Spartina alterniflora (Poaceae) in the British Isles
L. C. MASKELL and A. F. RAYBOULD

Centre for Ecology and Hydrology, Winfrith Technology Centre, Dorchester, Dorset DT2 8ZD

ABSTRACT

Spartina alterniflora was introduced from the U.S.A. in the 1820s. It is one of the parents of the fertile
amphidiploid Spartina anglica and is an important resource for the study of polyploid speciation. Throughout
the 20th century the distribution of S. alterniflora has declined and it is currently found at only one site, in
Southampton Water. This study compares present and past distributions and discusses reasons for its decline,
both throughout the harbour and at the one remaining site. Industrial and marine developments have resulted
in loss of habitat as well as causing changes in tidal action, sediment deposition and increased pollution. These
have all contributed to the decline of Spartina alterniflora. Competition with Spartina anglica has also played
a role.

KEYWORDS: decline, distribution, polyploid speciation Spartina anglica.
INTRODUCTION

Spartina alterniflora Lois. is a perennial grass native to saltmarshes of the eastern seaboard of
North America. Around 1820 it was unintentionally introduced to mudflats on the River Itchen,
Hampshire (the earliest confirmed record of S. alterniflora is a herbarium specimen collected by
Borrer from the Itchen in 1829), from where it spread throughout Southampton Water (Marchant
1967). In some areas, S. alterniflora grew beside S. maritima (Curtis) Fernald, a plant of Old
World saltmarshes, which is now extinct in Southampton Water (Gray et al. 1999). Fertilisation of
S. alterniflora by S. maritima pollen (Ferris et al. 1997) produced a sterile hybrid now named
Spartina x townsendii Groves & J. Groves. The first record of this plant is from Hythe in 1870
(Groves & Groves 1880). Chromosome doubling in S$. x townsendii formed Spartina anglica
(Marchant 1968; Raybould er al. 1991a), a fertile amphidiploid species.

Spartina anglica spread naturally along the south coast of England, the Isle of Wight and the
north coast of France by seed and by rhizome fragments (Oliver 1925). It was also introduced into
many saltmarshes in Britain and overseas for coastal defence and land reclamation (Ranwell
1967). Its rapid spread, and in the south of England a rapid decline, raised concerns about changes
to shipping channels and effects on populations of wading birds and other species occupying
intertidal mudflats (Gray et al. 1991; Raybould 1998).

In addition to economic and nature conservation relevance, S. anglica has immense scientific
importance. Because the species formed so recently, and because both its parental species are
extant, it provides a superb opportunity to study the ecological and genetic consequences of
polyploid speciation, a key process driving plant evolution. However, throughout the 20th century
S. alterniflora has declined in Southampton Water. It is extinct in the River Itchen and River
Hamble, and since the early 1960s the only remaining population of Spartina alterniflora in the
British Isles has been at Bury Marsh, Marchwood, Southampton Water. The map in Fig. | shows
the location of the River Hamble, River Itchen and the Marchwood site. This population is the last
remnant of the S. alterniflora material involved in the origin of S. anglica and is therefore a unique
resource to evolutionary biologists.

THE HISTORY OF SPARTINA ALTERNIFLORA IN THE BRITISH ISLES

Marchant (1967) gives a detailed account of the spread of S. alterniflora. At the height of its
distribution, S. alterniflora grew as a more-or-less continuous sward on the east shore of the
Itchen. There were also two smaller patches on the west shore of the Itchen. In Southampton

392 L. C. MASKELL AND A. F. RAYBOULD

River Itchen

6
0 Marchwood site

0

FIGURE |. Map of southern Britain with inset of Southampton water showing the locations of the River Itchen,
R. Hamble and the last remaining site for Spartina alterniflora; Bury Farm, Marchwood.

Water/River Test upstream of the Itchen, S$. alterniflora occurred as several discontinuous swards
from Hythe Marina to Cracknore Hard, and as a large sward on the Bury Farm marshes as far
upstream as Eling Churchyard. The species also occurred on the eastern shore of the Test from
Redbridge to Millbrook. Downstream of the Itchen, S. alterniflora grew at Hythe in a large patch
about half a mile south of the pier. S. alterniflora was also common in the Hamble at Burlesdon,
Lincegrove, Hamble-le-Rice and Warsash and occurred as far downstream as Hillhead at the
mouth of the Meon.

It is likely that S. alterniflora spread through Southampton Water by vegetative reproduction
only. Bromfield (1836) commented that S. alterniflora in the Itchen “seldom perfects its fruit” and
Marchant (1968) states that “British” S. alterniflora always “fails to set seed”. This is also the case
in New Zealand (Partridge 1987). This may be due to low pollen fertility (Marchant 1968). It is
possible that the introduced S. alterniflora is a single clone (Raybould et al. 1991b) and because
some clones of S. alterniflora are self-incompatible (Daehler 1998) even if viable pollen is
produced, self-incompatibility may prevent any seed set in “British” S. alterniflora.

Recession of S. alterniflora began around 1920 (Marchant & Goodman 1969) and by the 1930s
it was extinct in the Hamble and at Hythe. By 1924, the populations at Eling and Redbridge to
Millbrook were extinct. A small population at Dibden was destroyed by reclamation in 1963; this
left the Bury Farm marshes at Marchwood as the final site for the S. alterniflora material involved
in the evolution of S. anglica.

The main objectives of this work were to map the current distribution of S. alterniflora on the
Bury Farm marshes and to survey areas in Southampton Water and the Rivers Test, Itchen and
Hamble where S. alterniflora once occurred to assess why the populations had declined.

DECLINE OF SPARTINA ALTERNIFLORA 393

SS yo IN, Transect 1
NY Cara “.\, Transect 2
Nee /\ Transect 3
aye
NS MS only Spartina altemifiora
Ao Mixed; Spartina alterniflora and S.anglica

oy = <4 9 uo 200 Meters —
v4 = ge mee
vi

FIGURE 2: The distribution of Spartina alterniflora at Marchwood, Bury marsh, Southampton Water.

MATERIALS AND METHODS

BURY MARSH: MAPPING SPARTINA ALTERNIFLORA

Patches of Spartina alterniflora were identified from the morphological characters in Marchant
(1964) and Hubbard (1984). Each patch was mapped using a Magellan promark X hand held GPS
unit, taking measurements of dimensions and recording associated vegetation. The data were
imported to ARCVIEW to create a digital map of the marsh. This survey was carried out in
October/November 1999.

BURY MARSH: PHYSICAL NICHE AND ASSOCIATED VEGETATION
The topography of the marsh was surveyed in October/November 1999 using a Pentax Total
Station electronic theodolyte. Transects were set up across the marsh (Fig. 2). Transect 1 was
placed in the middle of the marsh and passed through several patches of Spartina alterniflora.
Transect 2 was established to the western side of the marsh and crossed a particularly low-lying
patch of S. alterniflora which lies at the back of the marsh near to an equally low-lying patch of S.
anglica. Transect 3 was placed to the eastern side of the marsh where there is far less S.
alterniflora present.

Measurements were taken at each point where the vegetation or topography changed
significantly and these have been plotted onto profile diagrams. The plant species found have been
related to the height (Ordnance Datum) at which they were found.

SURVEY OF PREVIOUS SITES
In addition to the above, the sites around Southampton water where Spartina alterniflora had been
recorded previously were visited to assess possible reasons for its loss.

RESULTS

BURY MARSH: MAPPING SPARTINA ALTERNIFLORA
Figure 2 shows the distribution of S. alterniflora on the Bury Farm Marsh and the extent of the

394 L. C. MASKELL AND A. F. RAYBOULD

taj eee a “

oa. t a Ui

i =

FIGURE 3a. Spartina alterniflora at Bury Marsh, Marchwood. This photo shows the topography of the marsh;
there is a large creek system.

FIGURE 3b. Spartina alterniflora growing at the front of the marsh, which is being visibly eroded.

DECLINE OF SPARTINA ALTERNIFLORA 395

marsh in relation to neighbouring coastal features. Figure 3a shows a view across the marsh with
S. alterniflora in the foreground and a part of the extensive creek system in the background.

Only two patches appear to be completely mixed stands of S. alterniflora and S. anglica,
although five other patches have S. anglica beginning to invade at the edges. The “mixed” and
“pure” patches are marked in Fig. 2. It is noticeable that S. alterniflora is now absent from areas
described as “mixed” by Marchant (1964), indicating that S. anglica has replaced it.

Atriplex portulacoides L., Limonium vulgare Miller, Aster tripolium L. and Triglochin
maritimum L. are all present in the patches. Atriplex portulacoides and Puccinellia maritima
(Huds.) Parl. are particularly abundant. Only one stand appears to be purely S. alterniflora with no
other species and this is found at the front of the marsh to the north west of the map. This stand
can be seen in Fig. 3b. The area in which the S. alterniflora is growing 1s visibly eroding.

Most patches of S. alterniflora are on high parts of the marsh in areas with high species
diversity. The species tends not to occur right on the edges of deep creeks where Atriplex
portulacoides grows in pure stands. It is also found growing in low-lying areas of the marsh.

The map in Fig. 4 shows the distribution of S. alterniflora at the site in 1964. This map has been
created using information in Marchant’s thesis (1964). The distribution shown in Marchant’s map
has been superimposed onto the shape of the marsh as it is today. The marsh has changed in shape
and extent since the original map so this new version is a pictorial representation rather than a
distribution map. It does, however, enable comparisons to be made between 1964 and the present.
The abundance of S. alterniflora has decreased. In 1964, the eastern side of the marsh contained
several mixed stands of S. anglica and S. alterniflora. These were restricted to the back of the
marsh, while the front of the marsh was dominated by S. anglica. Now there are only a few small
patches of S. alterniflora on the eastern side of the marsh and two of these are still mixed stands
today. Spartina alterniflora is concentrated on the west side of the marsh. In 1964, it was

7 Spartina alterniflora only
M07 2 iNiixed: Spartina anelica and S.akemificea

FIGURE 4. Map based on Marchant’s 1964 map showing the distributions of Spartina alterniflora, S. anglica
and mixed swards.

396 L. C. MASKELL AND A. F. RAYBOULD

distributed throughout this part but has definitely contracted. Marchant (1964) describes some of
the stands as degenerate. He noted that although the largest pure stands of S. alterniflora grew in
the low-lying areas, these swards were less vigorous and had lower pollen fertility. On the higher
sites, although other plant species are present, the individuals of S. alterniflora appear to be much
healthier.

BURY MARSH: PHYSICAL NICHE AND ASSOCIATED VEGETATION

The three marsh profiles are quite similar and can be seen in Figs. 5a, 5b and 5c. The height at the
top of the marsh was between 2-2-5 m OD. The raised parts of the marsh were all c. 1-5 m OD and
the creeks c. 0-5 m OD. Puccinellia maritima tended to be found on higher elevations around
1-5 m OD or above and consequently was more abundant towards the back of the marsh. There
appeared to be no particular pattern to the distribution of S. alterniflora. In Transect 1
S. alterniflora was found from 1-58—1-6 m OD. It occurs in Transect 2 from 1-1—1-59 m OD with
S. anglica and at 1-67 m OD in Transect 3 with S. anglica. Spartina anglica was found from
1-2-1-87 m OD in Transect 1, from 1-2—1-59 m OD in Transect 2 and from 1-42—1-88 m OD in
Transect 3.

SURVEY OF PREVIOUS SITES

Many of the sites where S. alterniflora was recorded in the past (before 1910) have now been
developed for docks and port facilities such as the river Itchen and the area between Hythe and
Cracknore Hard. The river Test has also been developed, the eastern shore now consists of docks
and port facilities whilst the Marchwood site on the western shore is quite isolated.

S. alterniflora used to be found along a stretch of Southampton Water east of Weston point.
However, the shoreline here is now bare mud and shingle; the S. alterniflora habitat has
completely disappeared, presumably due to erosion. There is a similar situation at the Hook (near
to the College of Maritime Studies, on the shore near Newtown (SU4905) from where S.
alterniflora was last recorded in 1907. There is no longer any salt marsh here and the substrate is
composed of shingle and sand rather than mud. All that remains of the marsh is a small patch of S.
anglica.

S. alterniflora was common in the Hamble at Burlesdon, Lincegrove, Hamble-le-Rice and
Warsash. There has been less development of the river Hamble than the river Itchen and there are

Height OD (m)

25 -
Festuca Mixed species,
rubra, Puccinellia od
Aster maritima, Spartina

tripolium Spergularia media, alterniflora
Aster tripolium, ii

‘i some Spartina

anglica f
Bare mud at the bottom of the Atriplex portulacoides at the edges of creeks,
| creeks, Atriplex portulacoides ae Aster tripolium, Puccinellia maritima on the
_on the edges. top, mud at the bottom of creeks.
ib:
ge
0.5 [-
; bed
0) 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160

Distance (m)

DECLINE OF SPARTINA ALTERNIFLORA 397

Height OD (m)

25 : :
Spartina anglica,
Puccinellia
maritima, Aster ;
tripolium, | Spartina
Limonium vulgare | : alterniflora,

Puccinellia maritima, : Spartina
2 Spartina alterniflora, : anglica,

Spartina anglica and :Puccinellia

Atriplex : maritima and : ; ;

portulacoides : Aster tripolium Spartina alterniflora and
i i bare mud

Atriplex
portulacoides
and bare mud

Atriplex portulacoides
and Spartina anglica

0.5

0 10 20 30 40 50 60 70 80 90 100
Distance (m)

Height OD (m)

2.2 ; :
Spartina anglica,
Puccinellia _ Spartina anglica,
maritima, Aster : Puccinellia
B | tripolium, ; eta | maritima, Aster
Atriplex Spartina anglica, tripo lium,
portulacoides, Spartina alterniflora Atriplex
Puccinellia i | id
1.8 _maritima, Aster 'P ORUIaCOlaes,
:tripolium,
Spartina anglica,
1.6 : 2 P ae
:Puccinellia maritima,
: Aster tripolium, Atriplex
:portulacoides,
1.4
Puccinellia
maritima,
Aster j
12 | tripolium, aaa vet
Triglochin i j
maritimum, Creek, Bare mud
1 |Spergularia
media
0.8
0.6
5c
0 20 40 60 80 100 120 140 160 180 200 220

Distance (m)

FIGURES 5 a., b., & c. Topographic profiles of the transects which were established at Bury Marsh,
Marchwood. The most abundant plant species have been added to the profile.

398

L. C. MASKELL AND A. F. RAYBOULD

Height OD (m)
3.5
Juncus | Spartina anglica, ae i
effusus, | Puccinellia maritima, | Spartina anglica,
Holcus _ Spergularia media, Mud in creek: Puccinellia maritima,
3 |lanatus, _ Salicornia spp. and Atriplex Atriplex _
Festuca portulacoides portulacoides
VP"? Holcus pheidee : Spartina
25 lanatus, Spartina anglica, anglica
Puccinellia | Puccinellia maritima, | age
maritima Atriplex
portulacoides
2 Salicornia spp.
Puccinellia
maritima,
" Spergularia
media,
Festuca ‘Bare mud |
rubra, NY
Salicornia | an)
SPP... | 2 : _ | Bare mud

(0) 10 20 30 40 50 60 70 80 90 100 110 120 130 140 150 160
Distance (m)

FIGURE 6. The topographic profile of the transect established at Hacketts marsh on the River Hamble.

still areas of mature marsh at Burlesdon (Lincegrove and Hacketts), that have been given protected
status. S. alterniflora was last recorded from these marshes pre-1910. The reasons for decline of
S. alterniflora here are unclear. Unlike the river Itchen it is not due to loss of habitat through
development. We established a transect on Hacketts marsh to compare the topographic profile and
vegetation with Bury Farm Marsh, Marchwood. This profile can be seen in Fig. 6. The top of the
marsh is around 2 m OD in height. There is a flat area containing a lot of Puccinellia maritima
which has been quite heavily grazed between 1-5 m OD and 2 m OD. The bottom of the creek is
approximately 1-7 m OD. The other raised marsh areas are at a height of 1-5 m OD and contain
small amounts of Atriplex portulacoides, Salicornia spp. and Spartina anglica. S. anglica is found
throughout the marsh, growing between heights of 0-99-1-91m OD. At the front it grows on top of
cliffs approximately | m high. It is not found on the low-lying bare mud at the front of the marsh
and does not form as dense stands as it does at Marchwood. Puccinellia maritima is abundant
particularly at the back of the marsh. Other marsh species include Aster tripolium, Limonium
vulgare, Atriplex portulacoides, Juncus maritimus Lam., Salicornia spp. and Triglochin
maritimum.

DISCUSSION

REASONS FOR THE DECLINE OF SPARTINA ALTERNIFLORA

As well as the direct impacts of industrial and marine developments such as loss of saltmarsh
habitat there are indirect impacts such as changes in wave action and tidal currents, changes in
sediment deposition and pollution which have implications for the whole of the estuary system,
including the existing site at Marchwood. There has been much erosion of saltmarshes and many
areas where S. alterniflora used to grow are now low-lying mud or shingle, the saltmarsh having
completely disappeared.

DECLINE OF SPARTINA ALTERNIFLORA 399

It is not clear why S. alterniflera has persisted at Marchwood, while becoming extinct on
saltmarshes in the Hamble. Marchant (1964) suggested that Marchwood is unique in its uneven
topography; however, this was not based on topographical levelling and our study suggests that
Hacketts Marsh is not dissimilar.

At Marchwood, the S. alterniflora population has declined significantly since 1964. This is
probably due to an interaction of factors. Spartina anglica has played a role, as it has completely
replaced S. alterniflora in areas where mixed swards once occurred. The degree of tidal inundation
is considered to be a dominant factor influencing plants in the intertidal zone; it affects a number
of factors, including degree of anoxia, salinity and build-up of chemicals. Determination of the
level above Ordnance Datum at which a species occurs should give some idea of the ecological
conditions that it prefers or can tolerate and suggest factors that limit its distribution. The
topographic levelling showed little difference in height above Ordnance Datum between the upper
and lower limits of the two. However, competition between the two species is not the only factor
in the decline of S. alterniflora on uneroded marsh at Marchwood. Marchant (1964) noted that
pure stands of S. alterniflora were showing signs of decline even though S. anglica was not
present.

Pollution may have affected vegetation generally by weakening the plants. Marchant (1964)
suggested that reclamation work downstream altered tidal movement and scour in the estuary
which reduced the size of the marsh and deepened the channel. The regular dredging of the deep-
water channel may be important. The soft mud at the surface of the marsh may have crept laterally
into the deep channel and lowered the level of the overlying marsh. The dredgings are taken to the
reclamation area so the supply of water-borne silt has been reduced, which causes the marsh
accretion rate to be reduced. The lowering of the marsh increases waterlogging and tidal
immersion of sites. In low-lying waterlogged sites the sediment that is deposited is also more
easily removed by re-suspension than at sites of higher elevation where it may have a chance to
dry out and become part of the marsh surface so the process is cyclical and self-enforcing (Reed &
Cahoon 1992). Finally, there may be an age-related decline in vigour because there is no sexual
reproduction of S. alterniflora in Britain. Clones of S. anglica decline in vigour with age (e.g.
Thompson et al. 1991) and it is likely that a similar process occurs in S. alterniflora. Marchant
(1967) noted that herbarium specimens of S. alterniflora collected in the 19th century are larger
than plants growing at Marchwood in the 1960s.

CONCLUSIONS

The loss and decline of Spartina alterniflora from many of the sites in Southampton water appears
to be due to changes in land use. This includes direct impacts from construction as well as indirect
effects from changes in tidal action, channel topography and increased pollution. Spartina anglica
may have out-competed S. alterniflora in other areas. At some of the sites such as Hacketts and
Lincegrove marsh in the Hamble it is difficult to determine an obvious impact and loss may be due
to a combination of factors. We consider the best strategy for conservation of S. alterniflora to be
preservation of its habitat at Marchwood. Periodic monitoring of the site would determine whether
erosion at the front of the marsh is continuing and whether S. alterniflora populations are being
invaded by other species. If the dieback of S. alterniflora on the remaining sections of marsh since
1964 is due to the loss of sediment, it would perhaps be appropriate to consider whether the marsh
can be protected. Future research should examine the physical processes taking place at this site
and determine the rate of erosion, size of particles and types of sediment which are involved in
marsh construction. Possible actions for protection may include deposition of dredging spoil which
is already being removed from the deepwater channel nearby. The low-lying areas at the western
end of the marsh might be most suitable for deposition. Control of S. anglica may also be needed.

ACKNOWLEDGMENTS

This work was commissioned and supported by Ian Pearson from English Nature. We would like
to thank Duncan Hornby and Kathy Hodder for help and advice on GIS.

400 L. C. MASKELL AND A. F. RAYBOULD
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Southampton.

MARCHANT, C. (1968). Evolution in Spartina (Gramineae). I. Chromosomes, basic relationships and the
problem of the S. x townsendii agg. Botanical journal of the Linnean Society 60: 381-409.

MARCHANT, C. J. & GOODMAN, P. J. (1969). Spartina alterniflora Loisel. Biological Flora of the British Isles.
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(Accepted August 2000)

Watsonia 23: 401-411 (2001) 401

Conservation of Britain’s biodiversity: Carex depauperata With.
(Cyperaceae), Starved Wood-Sedge

iC. G- RICH

Dept. Biodiversity and Systematic Biology, National Museum & Gallery, Cardiff CF 10 3NP

and
C.R. BIRKINSHAW

Missouri Botanical Garden, BP 3391, Antananarivo 101, Madagascar

ABSTRACT

Fifteen years conservation work on the rare, statutorily protected species Carex depauperata With.
(Cyperaceae), Starved Wood-sedge in Britain are summarised. In the British Isles it has been recorded
historically from 14 sites in v.cc. 6, 9/11, 16, 17, 52, 83, 90 and H5. About 60 plants are currently known in
two native sites in England and one in Ireland, and it has been transplanted to two more sites. It is widespread
in Europe and Asia though rare in many countries, with the greatest frequency of records in France and Spain.
It is a distinct species showing little variation. It is a polycarpic perennial that usually grows in dry, deciduous
woodlands on a range of soil types, and is often associated with tracks. It is a woodland gap plant that
responds to opening of woodland canopies through coppicing and disturbance, and declines if management is
neglected and shade increases. Periodic (c. every five years) disturbance of its woodland habitat by coppicing
or scrub clearance is recommended as management for conservation.

KEYWORDS: Rare species, habitat management, critically endangered species, distribution, ecology.

INTRODUCTION

Carex depauperata With. (Cyperaceae), Starved Wood-sedge, is a very rare plant in the British
Isles. It is currently known from two sites in Britain and one in Ireland, all with small populations.
In Britain it is listed as Critically Endangered (the highest I.U.C.N. threat level), and is protected
by Schedule 8 of the Wildlife and Countryside Act 1981 (Wigginton 1999). In the Republic of
Ireland it is listed on the Flora (Protection) Order, 1999. It is also reputed to be rare and scattered
across much of its European range, and often occurs in tiny populations. It is extinct in Germany
and Luxembourg, and is described as rare in Belgium, Corsica, Italy, Switzerland and Turkey
(Blab et al. 1984; Delvosalle et al. 1969; G. Aymonin, pers. comm., 1989; Landolt 1989; Nilsson
1985).

Although apparently once reduced to a single plant and on the verge of extinction, British
populations of C. depauperata are now recovering and increasing through a combination of
restocking, research, appropriate habitat management and pure chance. In this paper, we
summarise the data from about 15 years detailed work so that other conservation projects may
benefit from our experience. Full details are given in Birkinshaw (1990, 1991), Rich (1992, 1994a,
b), Rich & Fairbrother (1994, 1995, 1996) and McDonnell (1997).

DISTRIBUTION AND CONSERVATION IN THE BRITISH ISLES

Data have been abstracted from the literature, herbaria (ABD, BM, BRISTM, BTN, CGE, CLE,
DBN, DEE, E, GLAM, K, LIV, LTR, NUW, OXF, PTH, RNG, SLBI and TCD), field survey
and correspondence with botanists.

402 T. C. G. RICH AND C. R. BIRKINSHAW

1. Axbridge, North Somerset (v.c. 6)

Plants were first found in a wood between Axbridge and Cheddar in June 1860 by T. B. Flower
(BM) but the locality was later reported to have been destroyed (White 1912). In 1911 H. W.
Pugsley re-discovered it by a lane in the same area (but possibly in a different site) occurring
thinly on an open grassy bank over 100 yards long (White 1912). H. S. Thompson found more,
with an abundance of fruiting spikes, in 1915 (E).

The site has since been visited many times and its changes in population are well documented. A
photograph taken by J. E. Lousley in 1937 shows C. depauperata on an open, flowery bank
without scrub or hedges (RNG). The site gradually became overgrown, and by the late 1950s one
last plant remained on a shaded bend of the lane which was continually being damaged by farm
vehicles (R. M. Harley, pers. comm., 1981). The number of plants increased slightly during the
next two decades, and in 1974 R. S. Cropper found six plants. Concerned for their conservation, he
began to monitor them annually and initiated restocking, possibly single-handedly maintaining the
species in the wild in Britain for the next decade.

Cropper’s detailed monitoring notes were kindly provided to Rich & Fairbrother (1994), and are
briefly summarised, with unpublished notes by R. A. Jarman, as follows. By 1977, the six plants
were reduced to two due to damage from tractors, and there was no further change until 1981.
There was light clearance of Rubus around the plants in the winters of 1977/78 and 1978/79, and
the hedge opposite the plants was cut back in winter 1978/79. In October 1981, several plants
grown from a cutting taken a few years earlier were re-introduced. Between 1982 and 1990,
between four and six plants were counted annually, and in the winter of 1991/1992 or 1992/1993 a
second transplant took place, with nine plants being present in 1994. The hedge on the east side of
the lane was cut again in 1990.

In 1993 following the success at Ockford Wood (see below), Plantlife became involved in
management of the site in consultation with English Nature and Somerset Wildlife Trust. In 1994,
the large evergreen Quercus ilex which was shading the plants was cut down and a carpet of
Hedera helix and invasive Prunus spinosa suckers removed along a 40 m stretch. One year later
the average diameter of the plants had increased by c. 50% and plants were much more floriferous.
C. Hancock noted heavy grazing of inflorescences, so a small deer track was blocked resulting in a
ten-fold increase in the number of inflorescences in 1996. By 1999, there were at least 55 plants
with many seedlings with several dense patches (one patch c. 7 x 1 m dominated by C.
depauperata alone), and a new plant occurred about 50 m further down the lane.

2. Leigh Woods, North Somerset (v.c. 6)

A single immature specimen was gathered 1886 by H. S. Thompson, but was not identified until a
few years later by A. Bennett, by which time Thompson could not remember the exact place he
had gathered it (BRISTM; White 1912). This is still the only record for a very well-recorded
National Nature Reserve.

3. Templecombe, North Somerset (v.c. 6)
Murray (1896) reported it from Templecombe but it has not been seen again and no further
information is available.

4. Cranborne to Damerham, Dorset (v.c. 9)/South Hants (v.c. 11)

A herbarium specimen collected in the 1920s by H. H. Haines from hedgebanks between
Cranborne and Damerham was discovered in LTR in 1984. The area concerned has been searched
without success, though the exact site is not known (Pearman 1994).

5. Charlton Wood, Greenwich, West Kent (v.c. 16)

Carex depauperata was described as new to science from Charlton Wood after being found by Mr
Woodward in 1787. It was last recorded there by R. R. Hutchinson in 1830 (SLBI). This area has
now been built over.

6. Effingham chalk pit, Surrey (v.c. 17)
“Two or three tufts only” were first seen by W. W. Reeves in 1874 (Reeves 1874) and last
recorded in 1881 by F. W. Ward (RNG). This is presumed to be the same site as “Bookham”.

CONSERVATION OF CAREX DEPAUPERATA 403

7. Ockford Wood (Westbrook Copse), Godalming, Surrey (v.c. 17)

Plants were first found at Godalming by W. Borrer in 1807 (E), though the exact site is not known.
It was first reported by J. D. Salmon at “Westbrook Woods” in 1843 (CGE) and was well-
documented until its demise in the 1970s.

E. C. Wallace photographed the plant on an open flowery bank in 1936 (RNG). In the 1940s, the
plant was locally frequent in the wood, along the track side, on the bank, at the top of the bank and
on the slopes above (F. Rose, pers. comm., 1992). J. E. Lousley saw “five fine plants by lane” in
1949 (RNG). R. W. David and O. Polunin saw five plants “one plant in gutter, two on bank above,
and two more just into the wood at the top of the bank and 10 yards downhill” in 1961. By 1970,
one plant remained on the edge of a precarious bank which Mrs J. E. Smith and W. E. Warren took
measures to shore up. F. Rose and E. C. Wallace last saw one vegetative plant inside the wood in
1972. This plant seems to have disappeared from the site about this time under a landslide (J. E.
Smith, pers. comm., 1993).

J. E. Lousley and Mrs J. E. Smith searched all the known sites in 1973 without success (Lousley
1976) and R. W. David failed to find the plant in 1974. In 1975 the Surrey Flora Committee again
searched all the old localities and placed a request for information with a picture of the sedge in a
local paper (one Carex sylvatica was the only response). Thorough searches by Mrs J. E. Smith
with other botanists in 1986 and 1987 of many suitable areas around Godalming failed to find any
plants. C. Birkinshaw failed to find it in 1989.

It was thus something of a surprise when one flowering plant was found in 1992 by F. Rose, P.
Marren and T. C. G. Rich in exactly the same spot as where it was last seen. During the Great
Storm of 1987, a large branch had been blown down from a Tilia x vulgaris tree directly above the
old site which had opened up the canopy and probably stimulated buried seeds to germinate (Rich
1994b). The plant had remains of inflorescences from 1991, suggesting that it was in at least its
third year.

In November 1992, the canopy and hedge were thinned and the Hedera helix and Rubus
fruticosus in the ground layer removed taking care not to damage the plant. A local botanist Mrs
A. Fairbrother was recruited to keep an eye on the plant and keep the surrounding vegetation
down, which has proved very successful for growth of the plant to the present day.

8. Frith Hill, Godalming, Surrey (v.c. 17)
Plants were found in c.1842 by J. D. Salmon (Brewer 1863) and by many botanists up to 1891.

9. Charterhouse (Milton Wood), Godalming, Surrey (v.c. 17)

This site was first reported in c. 1843 as “copse near Hurtmore Cottages” by J. D. Salmon (Brewer
1863). Lousley (1976) recorded that he last saw it in 1938 and that it disappeared during the war,
probably after timber extraction.

10. Farnham, Surrey (v.c. 17)
There is a specimen in DBN collected by W. Mclvor from “near Farnham” in August 1846 (Rich
et al. 2000).

11. Porth-y-felin, Holyhead, Anglesey (v.c. 52)

In 1936 A. G. and W. Holder discovered C. depauperata on rocks below the sea wall at Porth-y-
felin, Holyhead (LIV; correspondence in NMW). In 1967 W. E. Hughes visited the site using the
Holder’s directions and found one plant. This site appears to have been erroneously reported as on
the edge of a quarry by Ellis (1983) which has misled many botanists.

Searches in 1998 and 1999 by T. C. G. Rich, S. Ellis and R. A. Jones of the exact site have
failed to refind the plant. Aerial photographs taken by the Luftwaffe in 1940-41 show the area
around the rock to have changed little, though there is now an invasion of Fallopia japonica and a
small concrete ramp has been constructed. Clearance and disturbance might regenerate the plant
from the seedbank.

12. Bonaly, Edinburgh (v.c. 83)

A specimen from “Wood above Bonelly” (assumed to be Bonaly) was collected by W. R. McNab
probably sometime between 1860 and 1869 (DBN; Rich et al. 2000). This area, now largely a
Regional Park, was searched by Mr C. Dixon without success in 1999 (pers. comm., D. McKean,
1999).

404 T. C. G. RICH AND C. R. BIRKINSHAW

13. Forfar (v.c. 90)

There are two records which require confirmation, but which could refer to the same site. Hooker
(1821) cited “woods near Forfar, G. Don”, which was accepted by Gardiner (1848) but rejected as
doubtful by Ingram & Noltie (1981) as the origin could have been Don’s garden (the source of
several other confusing records). Another specimen from “Forfarshire” collected by T. McFarlane
(ex herb. Rev. H. G. Carroll, DBN) appears to have been collected independently suggesting the
species has indeed occurred in the county (Rich et al. 2000).

14. Killavullen, East Cork (v.c. H5)

Plants new to Ireland were discovered in scrubby open woodland on a rocky Carboniferous
limestone outcrop by T. O’Mahony in 1973 (O’Mahony 1976), and a total of twelve plants were
found on more detailed searches in 1974 and 1975. In 1984, these original plants were found to
have gone extinct, but a new sub-population of six plants was found to the west. Two further sub-
populations with two and three plants respectively were found in 1992. By July 1994, the sub-
population of two plants had gone (despite being large vigorous plants in 1992), but the other two
sub-populations survived though fluctuating in numbers.

The site has a dense field layer of Hedera helix through which there are many small animal
tracks with which the plant is associated. Some limited canopy clearance work took place in 1982.
In 1989, although the canopy was beginning to close, 20 plants were recorded. The plant has since
declined and by 1998 the population had declined to five small tussocks of which only three were
fruiting (O’Mahony 1999). The site has been monitored regularly, and a full account of this site is
in preparation by T. O’ Mahony.

Errors or probable errors

A specimen in DBN labelled “Leith Hill, Surrey, 30 June 1890, per J. Leitch” may be a
corruption of Frith Hill (Rich et al. 2000).

There is a second Frith Hill in Surrey at SU9058. It appears that the Godalming Frith Hill
records have been erroneously plotted in SU95 in Perring & Walters (1990) and Wigginton (1999)
(A. Lockton, pers. comm., 2000); all records are assumed to belong to the one at Godalming
(SU9744).

There is a specimen from Ranworth, Norfolk collected by W. Curnow in 1878 in E whose status
is currently in doubt. The soils in this area are predominantly coarse loams or unsuitable wet fen
land. There is currently no ancient woodland of more than 2 ha size in the parish, and the woods
that are present are predominantly semi-natural woods managed for pheasant shooting, though
there is some hazel coppice and oak woodland on the valley sides (R. Handley, R. Ellis, pers.
comms., 2000). Curnow was an acute observer of natural history and specialised in lower plants,
and made some outstanding collections (Davey 1909), but he seems to have botanised
predominantly, though not exclusively, in Cornwall. Letters he wrote in 1876 indicate he was
trying to obtain sedges and grasses for his collection from other botanists, and it may be that the
specimen is of cultivated origin and Curnow confused the original locality (R. J. Murphy, pers.
comm., 2000).

It has thus been recorded from 14 sites in v.cc. 6, 9/11, 16, 17, 52, 83, 90 and HS. A distribution
map summarising the records is given in Fig. 1. It has a somewhat disjunct distribution in the
British Isles, characterised by a few long-known and persistent sites contrasting with the nearly
half which are represented by one historical herbarium specimen only. In 1999 it was only known
as a native in two sites in England and one in Ireland. Population counts for the last ten years for
these sites are summarised in Table 1. |

TRANSPLANT SITES

1. Cheddar Wood

On 30 October 1977 C. E. D. Smith and R. A. Jarman, as part of a project with the Somerset Trust
for Nature Conservation and Bristol Conservation Corps, transplanted 15 plants from Bristol and
Cambridge University Botanic Gardens (both grown from seed collected at Axbridge in 1965 by
D. E. Coombe) into each of two 10 m x 10 m quadrats, one cleared of ground, understory and
overhanging vegetation, and the other under closed overgrown canopy nearby. The plants were
reported heavily grazed soon after they had been planted. On 22 July 1979, all 15 plants had

CONSERVATION OF CAREX DEPAUPERATA 405

FIGURE 1. Distribution of Carex depauperata in the British Isles. ® 1990+. O pre-1990. ? exact location
uncertain. x error or probable error.

406 T. C. G. RICH AND C. R. BIRKINSHAW

TABLE 1. NUMBER OF MATURE PLANTS AT CAREX DEPAUPERATA NATIVE SITES

1990-199]
Site Godalming Axbridge Cork
1989 0 6 20
1990 i 4 -
1991 | 5 -
1992 1 6 ifs)
1993 3 - 17.
1994 3 14+ 7,
1995 3 14+ 10
1996 3 15+ uf
0d + 18+ i
1998 + 51+ 5
1999 + 55+ -

Data for Godalming by extrapolation, counts for Axbridge include transplants at native site only;
data for Cork courtesy of T. O’ Mahony.

survived in the cleared plot, but only five small vegetative plants remained in the overgrown plot.
One plant, probably a survivor from the original transplants was seen in the clearing in the mid-
1980s (R. Corns, pers. comm., 1995), but the exact location of the other plot is not now known. No
plants were found by C. Birkinshaw in 1988 or 1989, or by T. C. G. Rich in 1993 and 1994 but,
surprisingly, two plants were found in the clearing in 1995 by T. C. G. Rich, four in 1996, and nine
in 1999,

2. Edge of Cheddar Wood

In April 1988, 100 divisions of C. depauperata from Cambridge University Botanic Garden
(origin as above) were planted inside and outside an exclosure in ungrazed pasture/woodland edge
on the edge of Cheddar Wood to investigate the light, grazing and disturbance requirements.
Eighteen months later all plants were still alive, though varying in performance. Plants subject to
grazing were less fecund than those protected from grazing, plants placed in disturbed vegetation
were much larger than those in undisturbed vegetation suggesting a benefit from reduced
competition, and optimum fecundity occurred on the woodland edge (Birkinshaw 1990). By
October 1991, 86 plants were still alive (the losses mostly due to competition from Hedera), and
the main trends observed by Birkinshaw had continued (Edgington 1991). By 1993 plants in the
enclosure were shaded by dense Rubus fruticosus, and fewer plants survived within it than outside.
The fence was removed in 1998. At least 17 plants survived in 1999, mainly those planted outside
the original enclosure indicating that competition affected survival.

WORLD DISTRIBUTION

There is little recent information available on its distribution or frequency in Europe. Data were
compiled from herbarium specimens and literature to give a basic assessment of its distribution.
Information was abstracted from non-British specimens at the following herbaria: BM, COI, FI.
K, L, LTR, LIV, MA, NMW, OXF; P and RNG. Information was also abstracted from relevant
Floras held in the library at the National Museum of Wales, although the Flora holdings are patchy
and incomplete.

The European records traced are mapped in Fig. 2. It has been recorded from Albania, Belgium,
Bulgaria, Corsica, Croatia, England, France, Germany, Greece, Hungary, Ireland, Italy,
Luxembourg, Romania, Scotland, Sicily, Spain, Switzerland and Wales. In addition, it has been
recorded from Iran, Iraq, Turkey and the former U.S.S.R. (Caucasus, Ukraine, Pamir, Tien-Shan)
(Nilsson 1985), but the records are too fragmented and unlocalised to merit plotting. The records
traced show it is widely scattered across Europe and near Asia, though these are likely to be biased
towards western European sources. France and Spain appear to be its main strongholds, and it is
certainly still widespread in the latter.

CONSERVATION OF CAREX DEPAUPERATA 407

FIGURE 2. European distribution of Carex depauperata compiled from herbarium specimens and the literature
(all dates).

ECOLOGY

LIFE CYCLE

Carex depauperata is a distinctive sedge, instantly recognisable across its range by its few flowers
and large utricles. Vegetatively it is less distinct, and can be similar to C. sylvatica when young,
but usually differs in forming dense clumps with more erect leaves which droop at the tips to form
a “fountain”. The red coloration of the basal sheaths is distinctive when present; it is often more
pronounced on herbarium specimens than in live material. Vegetative plants can be very difficult
to find, and young plants are difficult to distinguish from other Carex species. No hybrids are
known.

Material from the two British, one Irish and one French site (the latter originally collected in
southern central France by P. A. Harmes) was cultivated in an open herbaceous border on clay
soils in a garden in Surrey by Mrs J. E. Smith between 1995 and 1999. After two years in
cultivation, no significant morphological differences could be detected between the plants.
Similarly, examination of a range of herbarium material from the western part of its range shows
that it varies little morphologically, except a little in size due to local growth conditions. Davies
(1956) reported a haploid chromosome number of n = 22 from a plant from Axbridge.

Carex depauperata is a polycarpic hemicryptophyte. The diameter of clumps and the number of
inflorescences can increase or decrease between years depending on growth conditions. Leaves are
usually green though the winter. In cultivation and in the wild plants have been observed to flower
in their second or third year. Although some plants in cultivation in Britain have lived for at least
25 years, wild plants may have shorter lives, e.g. in the Cork population the maximum life-span
recorded was 13 years and many individuals survived for only 2-5 years (T. O’Mahony, pers.
comm., 2000).

408 T. C. G. RICH AND C. R. BIRKINSHAW

Large tussocks in well-lit sites may produce up to 200 inflorescences, each consisting of a
terminal male spike and 1—4(—5) female spikes. Heavily shaded plants may not flower. Flowering
begins in April and usually continues until June. The inflorescences are protandrous and wind-
pollinated. Isolated plants have been observed to set fruit, and it is thus presumably self-
compatible. Observations at Cambridge Botanic Gardens found an average of 4-96 + 0-8 (s.d.)
nutlets per spike (n = 40), one of the lowest figures for a British sedge outside the Subgenus
Primocarex. The nutlets are large (c. 4-5 x 2-1 mm) and heavy (average weight air-dry nutlets
excluding utricles 0-0099 mg, n = 125). Fruit set is usually high; it sets abundant fertile seed in
both extant British sites and many ripe fruits have been observed on herbarium specimens. Fruiting
stems tend to droop around the plant late in the summer onto the soil, and may persist with fruits
intact through the winter. It is usually possible to find at least some of the previous year’s
inflorescences at the next flowering.

The method of seed dispersal is unknown. Wind dispersal is unlikely because the nutlets are
heavy and lack adaptations normally associated with this method of dispersal (e.g. wings).
However, it is possible that the nutlets have evolved for dispersal by graminivores such as rodents
(that could collect and cache the nutlets, and then sometimes fail to exploit these caches; for
examples of species dispersed by rodents see Janzen 1983), or by herbivores that could swallow
the nutlets when eating the foliage and at least sometimes pass them in a viable condition (for
discussion of this method of dispersal see Janzen 1984). In addition, the nutlets will certainly
sometimes be dispersed by the surface flow of water following rain. Failing all else, the long
infructescence culm (up to 1-5 m) allows nutlets to be deposited some distance from the parent
plant.

Fresh nutlets may take a long time to germinate. Experiments showed samples took 18-29
months after sowing to germinate, suggesting an after-ripening requirement like many other sedges
(Birkinshaw 1990; Jermy er al. 1982). Dormancy appears to be partly controlled by the pericarp;
30% germination (n = 20) occurred within 3—5 months when the pericarp was nicked. Germination
is also almost certainly controlled by environmental conditions such as light.

Seeds may remain dormant for long periods. This has been demonstrated experimentally by
Birkinshaw (1999), who showed that at least some seeds in samples buried deeply in the soil
germinated when brought to the surface after an interval of nine years. Observations of the
Godalming site suggest dormancy of nearly 20 years between the last plant being seen and the new
one re-appearing.

HABITATS

Birkinshaw (1991) summarised its habitat in Britain. [t usually occurs in dry, deciduous woodland,
often in south-facing sites, but also occurs along the base of an old hedge in Somerset, and has
been recorded from scrubby rocks by the sea on Anglesey. It is often associated with rocky
woodland in Europe, but not exclusively so.

Carex depauperata will prosper in conditions of full sunlight (provided that it is not out-
competed by more vigorous heliophytes) to semi-shade. It grows well in full sunlight in the “Rare
Plants Flower Bed” outside the English Nature Headquarters in Peterborough, and in gardens
elsewhere. It is tolerant of shade, but plants may become moribund and cease to flower in deep
shade. However, this sedge is probably best considered as a woodland gap plant adapted to the
cycle of gap formation followed by secondary succession that is a feature of natural forests. In
particular, it is hypothesised that the seeds of C. depauperata in the soil seed bank are stimulated
to germinate by environmental changes associated with the formation of a gap in the forest; these
seedlings grow rapidly to maturity in the conditions of high illumination and low competition
within the gap and replenish the seed bank with new seeds; then as woody plants colonise the gap
and close the forest canopy once again, the sedge declines and finally disappears leaving seeds
waiting for the formation of a new gap, maybe several decades in the future. In support of this
hypothesis is the long dormancy of C. depauperata seeds and various observations showing that
the plant increases in abundance following coppicing or localised felling (woodland management
operations that simulate gap formation; e.g. Lousley 1976; pers. comm., T. O’ Mahony 1989). At
Fontainebleau, it was formerly plentiful in woods near the River Seine but the population declined
to a handful of plants until coincidentally the French Forestry Department felled the Fagus trees
and churned up the soil during forestry operations, after which hundreds of plants re-appeared (F.
Rose, pers. comm., 1992).

CONSERVATION OF CAREX DEPAUPERATA 409

The soils associated with C. depauperata range from acid podzols to calcareous brown earths
(full soil analyses from five sites are given in Birkinshaw 1990). In Somerset it occurs on a
calcareous brown earth (pH 7-4), in Surrey on weakly calcareous sandy soils, in Cork on a Mull
rendzina (pH 6-2-6-6), at Fontainebleau, France it occurs on a podzol (pH 3-44-5), at Chantilly,
France on sandy brown earth (pH 5-8—7-2) and at Ohrid, Yugoslavia on a clay-rich terra fusca. It is
also reported from schistes in France. Soil type per se does therefore not seem to be critical
provided they are free-draining (Birkinshaw 1991), though oddly Hegi (1980) also records it
associated with damper soils with trickles of fresh water and notes a preference for nutrient-rich
soils in central Europe.

The populations at Axbridge, Godalming and Cork occur in rather patchy W8d Fraxinus
excelsior - Acer campestre - Mercurialis perennis woodland Hedera helix sub-community of the
national vegetation classification (Rodwell 1991). The site at Anglesey was probably in W24
Rubus fruticosus - Holcus lanatus underscrub. At Fontainebleau is occurs in managed beech
woods equivalent to the NVC community W15 Fagus sylvatica - Deschampsia flexuosa woodland
and at Chantilly in woodland equivalent to the W8 Fraxinus excelsior - Acer campestre -
Mercurialis perennis woodland, Primula vulgaris - Glechoma hederacea sub-community. In
general these equate to the order Fagetalia and it is typically recorded from such woodlands and in
Quercus pubescens woodland of the order Quercetalia pubescenti-petraeae through most of
Europe (Hess et al. 1967; Oberdorfer 1979; Hegi 1980; Ellenberg 1988). In Spain it occurs in
Pinus pinaster woodland, Quercus pyrenaica woodland, Castanea woodland and Corylus avellana
scrub (data from herbarium labels in MA).

It is often associated with tracks ranging in size from small animal tracks to rough vehicle tracks
(Birkinshaw 1991). This may be due to slightly enhanced light levels and its ability to tolerate
crushing (it can tolerate even quite heavy trampling and even some crushing by vehicles) which
competitive heliophytes may not be able to do.

Observations at all three extant sites indicate grazing damage, especially to the inflorescences,
presumably by rabbits, deer and/or horses. M. and C. Kitchen have observed deer eating plants in
their garden in Gloucestershire (pers. comm., 1994). The exclosure experiment on the edge of
Cheddar Wood showed plants subject to grazing by rabbits and deer were significantly less fecund
than those protected from grazing (Birkinshaw 1990). It is tolerant of at least light grazing, but the
number of inflorescences and consequently seed production may be significantly reduced.

In Britain it is a lowland plant, being recorded from + sea level on Anglesey to c. 50 m at
Axbridge and Godalming. In Europe it also occurs from near sea level in the Aegean (R. M.
Harley, pers. comm., 1981) to at least 500-950 m in Spain and 1260 m in the mountains of central
Europe (Hegi 1980).

CONSERVATION

Carex depauperata is a fully protected species under Schedule 8 of the Wildlife and Countryside
Act 1981. The Cheddar Wood site is part of a S.S.S.I. but the Ockford Wood site is not. In Ireland
it is protected under the Flora (Protection) Order, 1999. Plantlife monitor the species annually, and
have prepared a species action plan (Davis 1999).

Of the 14 sites from where it has disappeared or declined, there is no information on the causes
of the decline for ten. The main cause of decline at three others appears to be lack of appropriate
woodland management and increase in shade, with one recorded as probably being lost due to
forestry operations. The main current threats to the plant are lack of appropriate management
which may result in either dense shade or competition from other species, damage by vehicles,
potentially too much grazing, and, at Godalming, landslides which have caused temporary
extinction in the past. Whilst there are many specimens in herbaria, collecting by botanists has
probably been only a very minor part of the decline, and has not been responsible for extinction at
any sites as far as we are aware. Further visits to known historic sites may result in it being
rediscovered, as at Godalming.

Both native populations at Godalming and Axbridge have increased following conservation
management of opening up the canopy and/or coppicing with disturbance of the ground flora. Such
management, repeated perhaps every five years or so, is recommended for all sites, depending on

410 T. C. G. RICH AND C. R. BIRKINSHAW

growth of the canopy and competing species. Where populations are critically small, more regular
visits to clear competing vegetation around individual tussocks may be required and proved very
successful at Godalming. Some protection from heavy grazing may be required at some sites.

The potential for restoration of old sites in England from the seed bank has been investigated by
McDonnell (1997), who concluded that there was insufficient detailed historic data to merit
disturbance to regenerate it from the seedbank at any sites, and proposed reintroductions at Milton
Wood, Godalming and Effingham. The recently rediscovered locality details suggests that
restoration of the Anglesey site would be worthwhile.

Material from Somerset, Surrey and Cork is held in cultivation in several botanic gardens and
private gardens, and seeds from both English sites have been deposited in the Millennium Seed
Bank. It grows readily in cultivation and increases from seed in at least some gardens, contrary to
the statement by White (1912).

ACKNOWLEDGMENTS

We are grateful to David Allen, Ms J. R. Andrews, Richard Brinklow, Arthur Chater, Bob Corns,
P. Cuccuini, Dick David, Mike Edgington, John Edmondson, Bob Ellis, Sally Ellis, Anne
Fairbrother, Lynne Farrell, Ro Fitzgerald, Sam Hallett, Cif Hancock, Richard Handley, Stephen
Hewitt, W. E. Hughes, George Hutchinson, Matthew Jebb, Andy Jones, Stephen Jury, Mark
Kitchen, Gerald Legge, Peter Marren, Jim McGregor, Andy McVeigh, Douglas McKean, Rose
Murphy, Henry Noltie, Donald Pigott, Francis Rose, Alan Showler, Joyce Smith, Peter Stovold,
Sybil Tucker, Keith Watson, the Royal Botanic Gardens Wakehurst Place for help and
information, and especially Tony O’ Mahony for details of the Cork population and Liz McDonnell
for help at Axbridge.

We would like to thanks the Keepers of the herbaria for access to specimens and libraries. The
work was funded in various stages by the British Ecological Society, English Nature’s Species
Recovery programme, Nature Conservancy Council, National Museum & Gallery of Wales and
Plantlife.

REFERENCES

BIRKINSHAW, C. R. (1990). The ecology of Carex depauperata and its reinforcement at Cheddar Wood,
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(Accepted June 2000)

*

po

Watsonia 23: 413-419 (2000) 413

A morphological comparison between some British Orobanche
species (Orobanchaceae) and their closely-related, non-British
counterparts from continental Europe:

Orobanche rapum-genistae Thuill. s.1.

MEN Ye FOLEY

Department of Biological Sciences, Institute of Environmental and Natural Sciences, University of
Lancaster, Lancaster, LAl 4YQ

ABSTRACT

A morphological comparison has been made between British plants of O. rapum-genistae and plants referable
to subsp. rigens from Corsica, which is endemic to that part of the Mediterranean. Characters which best
separate the two taxa are indicated and it is concluded that the latter is more appropriately placed at species
level as O. rigens Loisel.

KEYWORDS: Orobanche rapum-genistae subsp. rigens, Corsica, taxonomy.
INTRODUCTION

BACKGROUND

Orobanche rapum-genistae sensu lato comprises a small but widespread group of western
European taxa thought to be distributed eastwards as far as Germany and Italy (limit at Val
Vestino, south Tirol (Gilli 1965)) and southwards to Spain and Portugal (Beck 1926-28); it is also
recorded for Algeria (Battandier 1890) and other north African countries, although its presence in
some may be doubtful. The current northern limit is in Scotland.

Fournier (1937) recognised three taxa at subspecific level. These were the type (subsp. rapum-
genistae), subsp. benthamii (Timb.-Lagr.) P. Fourn., differing from the former by its extremely
long bracts and a distinctly bilobed upper lip to the corolla, which had been recorded from Spain,
Portugal and northern Italy, and subsp. rigens (Loisel.) P. Fourn., apparently similar to the type but
almost glabrous, from Corsica and Sardinia and the associated small off-shore islands. It is not
definitely known whether the distributional ranges of the latter two taxa overlap with subsp.
rapum-genistae. More recent treatments (Chater & Webb 1972; Pignatti 1982) have supported
Fournier's approach.

The host plants are usually leguminous shrubs (Fabaceae), especially the genera Cytisus, Ulex
and Genista, but with subsp. rigens apparently restricted to species of Genista (Chater & Webb
1972; Camarda 1981; Pignatti 1982).

Throughout most of its European range O. rapum-genistae is represented by subsp. rapum-
genistae. In the British Isles this occurs southwards from southern Scotland and is widespread but
local, having substantially declined during the past century (Foley 1994) and receded into a
predominantly south-westerly distribution pattern, but with substantial populations remaining in
Wales; it also occurs in south-east Ireland. In the British Isles the host is invariably either Cytisus
scoparius (L.) Link or Ulex europaeus L., although there is an old record from Kent on Genista
tinctoria L. The procumbent, maritime subspecies of C. scoparius (subsp. maritimus) is parasitised
on sea cliffs in the Channel Islands (Rumsey & Jury 1991).

In France, from where the type (O. rapum-genistae Thuill.) was originally described, it is widely
distributed up to subalpine levels (Rouy & Foucaud 1909) and records for Switzerland are also
mainly from montane and (rarely) subalpine localities (Hess et al. 1972). In Germany it is
restricted to the west, known from the Rheinland and Schwarzwald eastwards to Westphalia (Gilli
1965); it is also present (as subsp. rapum-genistae) in Belgium and Holland (Chater & Webb

414 WE... ¥; FOLEY.

1972). In Italy, Pignatti (1982) gave all three subspecies: the type, widespread throughout; subsp.
benthamii from the alpine foothills of Piedmont and Lombardy; and subsp. rigens from Corsica,
Sardinia and Sicily. Also for Corsica, Gamisans & Jeanmonod (1993) recorded all three taxa but
the presence of two of them (the type and subsp. benthamii) is doubtful, whilst Camarda (1981)
also gave localities for subsp. rigens (as O. rigens) in Sardinia. Both the type and subsp. benthamii
are known from the Iberian peninsula (Willkomm 1893; Merino 1906; Coutinho 1913; Cadevall
1932; de Lima 1947; Franco 1984). For Algeria only the type was recorded by Battandier (1890)
and Quezel & Santa (1963) although there is a record for subsp. benthamii by Beck (1930). O.
rapum-genistae Ss. l. is apparently present in Morocco (Greuter et al. 1989) but doubtful in Tunisia
(Bonnet & Barratte 1896).

BASIC TAXA RELATED TO OQ. RAPUM-GENISTAE AND SUBSEQUENT TREATMENTS

O. rapum-genistae was described from near Paris, France (Thuillier 1799) where it parasitised
Cytisus scoparius and flowered in June: “caule crasso, conspicue pubente et angulato: spica
longissima: confertiflora, bracteis calycibusque villosissimis: corollis rufescentibus, brevi-
tubulatis’. The description is barely adequate, although the reddish (but shortly tubular) corollas
give some indication of its appearance. The type specimen is possibly in LE.

O. rigens was first described from Corsica (Loiseleur 1807) from specimens seen in Richard’s
herbarium “caule simplicissimo glabro, squamis lanceolatis rigentibus imbricatis, corollis
quadrifis, staminibus inferne nudis, stigmate 2-lobo, lobis distantibus, stylo glaberrimo, simillima
praedenti [O. major L.], sed discrepare videtur squamis rigentibus subpungentibus, et glabritie
omnium parte, exceptis bractaeis quae leviter pubescunt. Flores rubiginosi ut planta ipsa’.
According to this description, one of the main characters relates to the more or less glabrous nature
of the stem, style and lower part of the filaments - and indeed of most parts other than the lightly
pubescent bracts. The imbricate, lanceolate scale-leaves are rigid, the stigma lobes apparently well
separated and the flowers and the whole plant are of a reddish coloration. No specimen was
mentioned by Loiseleur but his description indicates that it was based on Corsican material
collected by Richard and retained in the latter’s herbarium; however such potential type material
appears now to be lost or destroyed. Loiseleur’s herbarium at Avignon (AV) also contains no
relevant material collected by Richard (Camarda 1981) but the Paris herbarium (P) has a Corsican
specimen gathered by Soleirol in 1826 and presumably studied by Loiseleur “au mont d’Oro prés
des neiges eternelles”. This has been proposed as the neotype for O. rigens (Camarda 1981).

O. benthamii Timb.-Lagr. was based upon the earlier O. rapum var. bracteosa which was
described by Reuter (1847) as “spicd valde elongata, bracteis flores superantibus apice comos@”’.
Reuter’s diagnosis referred to plants collected by Bentham at Collioure in southern France and by
Moris in the subalpine region of Piedmont, Italy. There is a fuller description of the same plant
collected at les Basses Corbiéres hills, near Durban, southern France as O. benthamii (Timbal-
Lagrave 1874). Again the bract character is referred to: “bractées.....dépassant de beaucoup les
fleurs, ce qui rend l’inflorescence chevelue” - i.e. making the inflorescence “hairy”. The corolla is
stated to be very large, bright red - is the stem - and with lobes with fimbriate teeth. The filaments
are glabrous and inserted at the base of the corolla and the stigma is white or yellow.

Beck (1930) placed all these taxa in his Grex Arcuatae, members of which were characterised as
follows. The corollas were broad and more or less ventricose above the point of filament
attachment and of similar internal and external colour, their dorsal line was strongly curved over
the whole length, the margins of the corolla lobes were glandular-ciliate, the filaments inserted
very low near the corolla base and the stigma lobes yellow. Within the Arcuatae he recognised
three species: | O. rapum-genistae, O. rigens and O. anatolica Reuter (1847) - the last is non-
European, being from the Caucasus and Iran.

As regards O. rapum-genistae itself, Beck (1930) recognised two varieties in addition to the
type. One of these was var. bracteosa (= O. benthamii Timb.-Lagr.) described above, the other,
var. insolita, was based upon Guimaraes’ description of O. insolita from Bussaco, Portugal
(Guimaraes 1904) where it apparently parasitised Eryngium campestre. This is the usual host for
O. amethystea and it is possible that O. insolita may be a variant of this, although filament, bract
and inflorescence characters and the presence of abundant white, glandular hairs (Guimaraes 1904)
are consistent with O. rapum-genistae s. l. Unfortunately, no material of O. insolita that is known
to have been seen by Guimaraes has been located.

OROBANCHE RAPUM-GENISTAE

415

TABLE 1. COMPARISON OF MORPHOLOGICAL CHARACTERS OF POPULATIONS OF
OROBANCHE RIGENS AND O. RAPUM-GENISTAE

CORSICA
Locality Tassenita valley
Asco
Number 10
Date 10.6.1997
Species rigens
Grid Ref./Coordinates 42 26'N; 8 58'E
Plant height 213-5 + 16-7
Stem width 10-1 + 1-0
pubescence 0-8 +0
colour pink-mauve -
yellow
Inflorescence length 133-0 + 9-8
density jy c= 3) 05
Leaf width 6:9 + 0-4
pubescence 0-8+0
shape ovate-tapering
colour pink-slt. mauve
Bract length 16-7 + 0-7
shape ovate-lanceolate
pubescence 1-5+0-1
colour darkish mauve
Corolla length 23:2 + 0-6
height 9-1+0-3
approximate shape subcampanulate
attitude 1-1+0-1
dorsal line 2:5+0-1
pubescence 1-:0+0
colour pink-mauve
Upper lip divided 0-5 + 0-2
Lower lip ciliate 0-2+0
denticulate 0
shape of lobes 0
relative size 10+0
(central to lateral)
Bracteole 0
Calyx length 13-6 + 0-3
segment division 1-8 + 0-4
pubescence 14+0-1
shape of teeth elongated
colour mauve
Filaments insertion 2:9+0-1
height
pilosity above 0
pilosity below 0
Anthers pilose 0
Stigma lobes separation Daly)
colour pale yellow
Apparent Genista
host salzmannii var.
lobelioides

CORSICA BRITAIN BRITAIN BRITAIN
Haut Asco Haydon Bridge Cyttir Mawr Amisfield
Asco Northumberland Anglesey Dumfries
i 10 10 10
10.6.1997 126.1993 5.61993 12 & 19.6.1993
rigens rapum-genistae rapum-genistae rapum-genistae
42 26'N; 8 58'E NY8565 SH5774 NY0083
165 £13-9 415-5 + 18-7 360 + 14-9 3799+ 20:5
8-9 + 0-5 10-8 + 0-3 11-6+0-3 13-14 0-8
0-9+0 Ian 1:7+0-1 1-8+0-1
pink. pink-yellow, pink-yellow __ pink-pale brown _ pale brown-pink
yellow
105-7 + 10-1 224-8 + 18-0 195-7 + 15-8 175-0 + 14-8
1-:8+0-1 1-3+0-1 1:9+0-1 1:-6+0-1
6-8 + 0-4 6-7 + 0-2 6-7 + 0-2 8-0 + 0-4
0:9+0 0-9 + 0-1 1-0+0 2 20:1
ovate-lanceolate linear-lanceolate triangular- tapering
lanceolate
pink-yellow brown medium brown fawn-dark brown
18-4 + 0-8 27-6 + 0-5 21:9: 11 25-1 + 1-0
ovate-lanceolate lanceolate lanceolate lanceolate
14+0-1 2:0+0 1-6+0-1 1:8+0-1
pink-mauve pale brown pink-brown dark brown
21-6 + 0-6 24-4 + 0-3 222205 25:6 + 0-6
7-71-05 10-0 + 0-3 10-4 + 0-4 10-3 + 0-2
subcampanulate + infundibuliform + infundibuliform + infundibuliform
1:0+0 1-3 +0°1 1-6+0-1 1-4+0-1
24+0-1 2:2 + 0-1 29 + 0-1 2-6 + 0-1
0-8 + 0-1 1540-1 1:2+0-1 1-4+0-1
pink-mauve, pink-fawn pink-pale fawn pink-fawn
pink-yellow
0 0 0-1+0-1 0
0-1+0-1 10+0 0-3 + 0-1 0-7+0:1
0 0 0 0
0 0-8 + 0- 0-8 + 0-1 0-8 + 0-1
10+0 1-4 + 0- 1-3+0-1 1:3+0-1
0 0 0 0
13-0 + 0-9 15-0 + 0-5 13-0 + 0-4 15-8 + 0-6
1040 1:0+0 1-4+0-2 12+0-1
1-2+0-1 2:0+0 1:2+0-1 1:7+0-1
triangular- tapering + acuminate tapering
lanceolate
mauve-pink pale fawn pale pink-fawn pale fawn
26+ 0-1 3-4+0-2 2:0+0:1 3-8 + 0-2
0 Oe 0 0-9 + 0-1 1-0+0
0 0 0 0
0 0 0 0-1+0
2:0+0 3-022,0 2:9+0-1 2:9 +0-1
pale yellow bright yellow bright yellow bright yellow

Genista
salzmannii var.
lobelioides

Cytisus scoparius Ulex europaeus Cytisus scoparius

Mean values and standard errors (in mm)

416 M. J. Y¥. FOLEY

Beck (1930) maintained O. rigens as a separate species from O. rapum-genistae on the basis of
the former’s more or less glabrous corolla, filaments and style. In contrast, Fournier (1937) in his
French Flora placed all the principal European Arcuatae taxa at subspecific level under O. rapum-
genistae. Such a treatment has been maintained by several later workers (Chater & Webb 1972:
Pignatti 1982) but modified by others. For example O. rigens was ranked at species level by both
Camarda (1981) and Uhlich ef al. (1995) despite having much earlier been considered a variety by
Fiori (1926). In other instances some of the taxa have been simply omitted or ignored completely.

METHODS

Three British populations of subsp. rapum-genistae (two parasitic upon Cyfisus scoparius and one
upon Ulex europaeus) were morphometrically compared with two Corsican populations of subsp.
rigens parasitic upon Genista salzmannii var. lobelioides (see Table 1 for locality details).
Characters measured or assessed and the methods used were as described in a prior paper (Foley
2000). Mean values and standard errors of the relevant characters are summarised in Table 1 and
the quantifiable numerical elements of these data have been subjected to PCA (Figure 1) in a
similar manner to that previously decribed (Foley op. cit.). By this technique, each specimen
measured is located in a multidimensional array, in which the number of dimensions is equal to the
number of characters measured, and where the most similar specimens are placed closest together.
This technique extracts the axes of greatest variation from the multidimensional space, thereby
simplifying the space to a few (usually 2 or 3) dimensions, so allowing the location of each
specimen to be visualised.

RESULTS AND DISCUSSUION

In Figure 1 it can be seen that the Corsican plants are clearly separated from the British on the PC1
axis. Within the British plants there is only slight overlap between plants at Cyttir Mawr, Anglesey
(on Ulex europaeus) and those of the other two populations on Cytisus scoparius (at Amisfield and
Haydon Bridge). Whether this difference is significant, possibly due to the influence of the
different hosts on the plant’s morphology, is open to question but this could additionally be
investigated by cultivation experiments.

When the characters of the two taxa (subsp. rapum-genistae and subsp. rigens) are compared
(Table 1), it can be seen where the principal differences lie. It is true that there is quite a close
similarity in many of them but relatively intangible morphological differences, readily apparent in
living plants in the field, are reflected in at least some of these.

Plants of subsp. rigens appear to be less robust than those of subsp. rapum-genistae (overall
height, inflorescence length, stem width etc.) - but this may in part be due to their occurrence at a
higher average altitude. In many plants the inflorescence occupies a higher proportion of the
overall height of the plant and often reaches close to ground level. Leaf and bract shape is
distinctly ovate (rather than lanceolate) and the corolla is subcampanulate rather than ventricose-
infundibuliform as in subsp. rapum-genistae (the latter is also somewhat carinate dorsally), with
the lobes of the lower lip much more rounded and of equal size. Corolla colour (and general plant
coloration - stem, leaves, calyces) differs, being markedly mauve-pink in subsp. rigens rather than
pink-fawn. Perhaps the clearest character difference lies in filament pubescence. Whilst the
filaments are inserted at similar heights, they are glabrous throughout their entire length in subsp.
rigens whereas those of subsp. rapum-genistae are very distinctly glandular-pilose in their upper
portion. The style and ovary are also distinctly pilose, being provided with colourless, few-celled,
glandular hairs; in contrast subsp. rigens is glabrous in this respect. Stigma lobe colour is also
much paler in this taxon and the stigma lobes are noticeably less separated.

Another character difference which is very apparent is the overall level of pubescence, this being
very much reduced in subsp. rigens compared to O. rapum-genistae. Since subsp. rigens appears
to be a much more montane-subalpine plant than subsp. rapum-genistae, it might be expected that
if the former were.merely a close variant of the latter then the pubescence, as in other montane
plants, would be much greater (not less) in the higher altitude taxon.

OROBANCHE RAPUM-GENISTAE 417

6 5 -4 3 2 = 0 1 2 3 4 5 6

. rigens (Tassineta, Stranciacone, Massif du Cintu, Corsica)

. rigens (Haut Asco, Massif du Cintu, Corsica)

. rapum-genistae (Amisfield, Dumfries-shire, Scotland)

. rapum-genistae (Cyttir Mawr, Anglesey, Wales)

. rapum-genistae (Haydon Bridge, Northumberland, England)

Qoeta
S49 S495

FIGURE 1. First two axes of Principal Components Analysis for individual plants of Orobanche rigens and O.
rapum-genistae. PC1 & PC2 jointly account for 43-6% of the total variance.

CONCLUSIONS

It is clear that whilst both subsp. rapum-genistae and subsp. rigens are quite similar, they also
possess several distinctive characters. In contrast to subsp. rapum-genistae, the following
characters are diagnostic for subsp. rigens: the very low level of overall pubescence, the
differering corolla shape and coloration, the more ovate leaf and bract shape, the glabrous
filaments and the differing stigma lobe characters. In the absence of evidence of intermediates
between the two, it is concluded that these differences are sufficient for subsp. rigens to be
separated at species level as Orobanche rigens Loisel. Other work, based mainly upon herbarium
specimens of Sardinian O. rigens (Camarda 1981), reached a similar conclusion; both those
findings and the present ones contrast with the approach taken by most other workers subsequent
to Fournier (1937), who have maintained O. rigens at subspecific level.

The situation regarding subsp. benthamii is less easy to resolve as it is a little-known taxon of
limited and relatively unknown distribution; it is not recorded for Britain. During this work no
living plants and only three herbarium specimens (out of many hundreds of this group examined)
could reasonably be referred to this taxon and even those differed from subsp. rapum-genistae
only in their significantly longer bracts. Other differences indicated in Chater & Webb (1972), e.g.
an erect, distinctly bilobed upper corolla lip in subsp. benthamii, were not evident in these
specimens. It is concluded that the latter should be considered to be merely a form of O. rapum-
genistae.

418 M. J. Y. FOLEY

TAXONOMIC DESCRIPTION

OROBANCHE RAPUM-GENISTAE THUILL.

(syn.: O. major L. pro parte; O. palatina Schultz; O. rapum var. bracteosa Reuter; O. bracteosa
(Reuter) Nyman; O. elatior Horn; O. foetida Duby, O. sarothamnophyta St Lager, O. benthamii
Timb.-Lagr., O. rapum-genistae subsp. benthamii (Timb.-Lagr.) P. Fourn.)

Described from near Paris, France. Type possibly in LE.

Typically 250-450 mm tall, with a quite dense, long, many-flowered inflorescence. Stem: 8—15
mm wide immediately below the inflorescence, distinctly glandular-pubescent, pink-yellow to
pink-brown. Leaves: maximum width 6-8 mm, + lanceolate, fawn-brown. Bracts: 20-28 mm long,
linear-lanceolate, light brown, distinctly glandular-pubescent. Calyx: 13-16 mm long, segments
fairly equally divided, tapering, pale pink-fawn, glandular-pubescent. Corolla: 22-26 mm long,
sub-erect, + infundibuliform, ventricose, somewhat carinate dorsally, pink-fawn, glandular-
pubescent, dorsal line distinctly curved; upper lip + entire, margins of lower lip somewhat ciliate,
not denticulate, rather acute, the central lobe often larger than the laterals. Filaments: inserted 24
mm above the corolla base, glabrous below, distinctly glandular-hairy above. Stigma lobes: deep
yellow, well separated. Parasitic mainly upon species of Ulex and Cyfisus.

This is a plant of scrub-invaded areas such as rocky hillocks, field margins and headlands,
roadside verges and other habitats typical of its hosts. Within Britain, observations have suggested
a possible west-east cline in host preference with Ulex europeaus apparently being preferred in the
west (especially Wales) and Cytisus scoparius the preferred host further east. Flowers April—July.
The species is apparently widespread but often very local in western Europe extending north to
southern Scotland and east to the Italian south Tirol.

The long-bracted form (syn.: O. rapum-genistae var. bracteosa Reuter; O. benthamii Timb.-
Lagr.) appears to differ from the above only in its exceptionally long bracts. Suggested differences
in corolla lip characters have not been confirmed. It is apparently of very limited occurrence and
recorded only very locally in southern France, Italy, parts of the western Mediterranean and the
Iberian peninsula.

OROBANCHE RIGENS LOISEL.
(syn.: O. condensata Moris pro parte; O. thyrsoidea Moris; O. rapum var. glabrescens Gren. &
Godr., O. rapum-genistae subsp. rigens (Loisel.) P. Fourn.)

Described from Corsica. Type thought to be lost (Richard herb.), neotype: “au mont d’Oro
[Corsica] prés des neiges eternelles, 1826, leg. Soleirol” (P!).

Typically 150-230 mm tall, with a fairly dense, many-flowered inflorescence which sometimes
extends to most of the length of the spike. Stem: 8-10 mm wide immediately below the
inflorescence, slightly glandular-pubescent to nearly glabrous, pink-yellow to somewhat mauve.
Leaves: maximum width 6—7 mm, usually + ovate, pink-yellow to pink-mauve. Bracts: 16-20 mm
long, + ovate, pink-mauve, somewhat pubescent. Calyx: 13-14 mm long, segments fairly equally
divided, teeth elongated, triangular-lanceolate, mauve-pink, somewhat glandular-pubescent.
Corolla: 21-24 mm long, erect to suberect, + subcampanulate, not noticeably carinate dorsally,
pink-mauve sometimes tinged yellowish, only slightly glandular-pubescent, dorsal line distinctly
curved. Upper lip sometimes noticeably divided, margins of lower lip sometimes slightly ciliate,
not denticulate, rounded, lobes of + equal size. Filaments: inserted 2—3 mm above the corolla base,
glabrous throughout their length. Stigma lobes: pale yellow, only moderately separated. Parasitic
upon species of Genista, in Corsica usually only on Genista salzmannii DC. var. lobelioides (or
occasionally var. salzmannii) and in Sardinia on Genista corsica. There is a suspicion that
Anthyllis hermanniae may also be parasitised (D. Jeanmonod, pers. comm. 1996).

In Corsica, this is a plant of montane-subalpine habitats mainly found in the Thymus-Genistetum
association of dwarf spiny shrubs at an altitudinal range of (600—) 900—c.2000 m, but in Sardinia it
occurs down to sea-level (Camarda 1981). Flowers (March—) May—June (—July at high altitudes). It
is endemic to Corsica and Sardinia and the associated off-shore islands. It is widespread but local
in suitable habitats but may be under-recorded. Records for Sicily are probably errors for other
taxa.

Although both O. rapum-genistae and its long-bracted form (var. bractosa) have been recorded
for Corsica, it is possible that these are errors and that O. rigens is the only member of the group
present.

OROBANCHE RAPUM-GENISTAE 419
ACKNOWLEDGMENTS

I am most grateful to the trustees of the Welch Bequest Fund of the Botanical Society of the
British Isles and to the Botanical Research Fund (Kew) for financial assistance during this project.
I am also grateful to D. Jeanmonod (Geneva) and I. Guyot (Bastia) for providing me with
information regarding Corsican localities for O. rigens, and to all those vice-county recorders who
have similarly helped with information on O. rapum-genistae in Britain. I also wish to thank the
curators of the following herbaria for their loan of specimens: B, BM, E, LIV, MA, P, SEV.

REFERENCES

BATTANDIER, J-A. (1890). Orobanche in BATTANDIER, J-A. & TRABUT, L. Flore de l’Algérie, 4: 659-664.
F. Savy, Alger & Adolphe Jourdan, Paris.

BECK, G. (1926-28). Orobanche in Die Pflanzenareale, | Reihe, Heft 7, Karte 64. Gustav Fischer, Jena.

BECK, G. (1930). Orobanchaceae, in ENGLER, A., ed. Das Pflanzenreich IV. 261: 1-348. Willhelm
Engelmann, Berlin.

BONNET, E. & BARRATTE, G. (1896). Exploration scientifique de la Tunisie. Imprimerie nationale, Paris.

CADEVALL, J (1932). Flora de Catalunya 4: 301. Palau de Generalitat, Barcelona.

CAMARDA, I. (1981). Le piante endemiche della Sardegna, 106-109. Bollettino della Societa Sarda di Scienze
Naturali 21: 373-382.

CHATER, A. O. & WEBB, D. A. (1972), Orobanche in Tutin, T. G. et al., eds. Flora Europaea 3: 286-293.
Cambridge University Press, Cambridge.

COUTINHO, A. X. P. (1913). Flora de Portugal, 565-571. Aillaud, Lisboa.

DE LIMA, A.P. (1947). Flora Portuguesa, 496-500. Imprensa Moderna, Porto.

Fiori, A. (1969). Nuova flora analitica d'Italia, 2: 389. Edizioni Agricole, Bologna.

FOLEY, M. J. Y. (1994). O. rapum-genistae, in STEWART, A. et al., eds. Scarce Plants in Britain.
Peterborough.

FOLEY, M. J. Y. (2000). A morphological comparison between some British Orobanche species
(Orobanchaceae) and their closely-related non-British counterparts from continental Europe: Orobanche
reticulata Wallr. s. 1. Watsonia 23: 257-267.

FOURNIER, P. (1937). Les Quatre Flores de la France, 796. Paul Lechevalier, Paris.

FRANCO, J. do’A. (1984). Nova flora de Portugal, 2. Sociedade Astoria, Lisboa.

GAMISANS, J & JEANMONOD, D. (1993). Catalogue des plantes vasculaires de la Corse, 11-258.
Conservatoire et Jardins botaniques de la ville de Geneve, Geneva.

GILLI, A. (1965). Orobanche, in HEGI, G., ed. Illustrierte Flora von Mittel-Europa, 6(1): 470-505. Carl
Hanser Verlag. Miinchen.

GREUTER, W, BURDET, H. M. & LONG, G. (1989). Med-Checklist, 4: 256-265. Conservatoire et Jardins
botaniques de la ville de Geneve. Geneva.

GUIMARAES, J. d’A. (1904). Monographia das Orobanchaceas Portuguezas. Broteria, 3: 5-207, Lisboa.

HEss, H. E., LANDHOLT, E. & HIRZEL, R. (1972). Flora der Schweiz und angrenzender Gebiete. Bd. 3: 250—
260. Birkhatiser Verlag, Basel & Stuttgart.

LOISELEUR, J. L. A. (1807). Flora gallica, 2: 384-385. Lutetiae, Paris.

MERINO, R. P. B. (1906). Flora Descriptiva é Ilustrada de Galicia, 3: 42-52. Tipografia Galaica, Santiago.

PIGNATTI, S. (1982). Flora d'Italia, 1-3. Edagricole. Bologna.

QUEZEL, P. & SANTA, S. (1963). Nouvelle Flore de l’Algérie et des Régions Désertiques Méridionales, 2:
852-859. Centre national de la Recherche Scientifique, Paris.

REUTER, G. F. (1847). Orobanchacaeae in DE CANDOLLE, A. P., ed. Prodromus Systematis Naturalis Regni
vegetabilis 11: 1-45. Victoris Masson, Paris. ;

Rouy, G. & FOUCAUD, J. (1909). Flore de France, XI. Les Fils d’Emile Deyrolle, Paris.

RUMSEY, F. J. & Jury, S. L. (1991). An account of Orobanche L. in Britain and Ireland. Watsonia 18: 257-
Day,

THUILLIER, J. L. (1799). Flore des environs de Paris, 2nd ed., 317. Gazette nationale, Paris.

TIMBAL-LAGRAVE, P. M. E. (1874). Mémoires. Academie des sciences, inscriptions et belles-lettres de
Toulouse, 7(6): 643-644.

UHLICH, H., PUSCH, J. & BARTHEL, K-J. (1995). Die Sommerwurzarten Europas: Gattung Orobanche, 1-235.
Die Neue Brehm-Bucherei Bd. 618, Westaerp Wissenschaften. Magdeburg.

WILKOMM, M. (1893). Supplementum Prodromi Florae Hispanicae, 185-191. Sumptibus E. Schweizerbart,
Stuttgart.

(Accepted July 1999)

. rT.

ye

Watsonia 23: 421-435 (2001) 42]

Brambles (Rubus L. sect. Rubus and sect. Corylifolii Lindley,
Rosaceae) of the Channel Islands

Dees ALLEN

Lesney Cottage, Middle Road, Winchester, Hampshire, SO22 5EJ

ABSTRACT

The history of the investigation of the predominantly apomictic genus Rubus L. in the Channel Islands (Les
Iles Anglo-Normandes) is outlined and the records made to date are listed and critically assessed. Of species
currently admitted to the British Isles list, 39 inclusive of two naturalised horticultural taxa are considered
reliably recorded: 26 from Jersey, 23 from Guernsey, 14 from Sark and 11 from Alderney. Though these totals
roughly correlate with island size, they are each made up of different species in substantial part. They include
one apparent endemic, R. cordatifolius (Rogers ex Riddelsd.) D. Allen, shared by Guernsey and Sark. Many
unnamed entities, however, also occur, one or two in each main island relatively widely, but the group has
been too little investigated on the French mainland to tell how far any extend there sufficiently to merit being
described should they not prove to belong to existing taxa. Dispersal and other factors determining the
presence or influencing the frequency of certain species in the islands are also examined.

KEYWORDS: apomictic species, Britain, France, Normandy, Brittany, distribution, bird dispersal.

INTRODUCTION

By virtue of their being Crown possessions in British home waters the Channel Islands - Les Iles
Anglo-Normandes to the French (to whom one group of islets belong) — have traditionally been
treated as part of their territorial responsibility by students of the British-cum-Irish flora. From the
strictly geographical point of view, though, it is to Continental Europe that they manifestly belong,
as confirmed by the Precambrian geology they share with Brittany and the nearest part of
Normandy as well as by a general floristic character broadly different from that on the other side
of the English Channel. That very degree of difference has tended to deter specialists in major
critical groups such as Rubus, who have been conscious of the extra dimension of difficulty these
islands are likely to present without some reasonable measure of familiarity with the
representatives of those groups in the nearby parts of the mainland. The Rubus flora is particularly
richly diverse, as it chances, in the Cotentin Peninsula of Normandy immediately to the east (Allen
1996) and only small portions of that, or of the whole of Brittany for that matter, have been
investigated even cursorily by batologists until the last twenty years. Though knowledge of that
wider context is still very incomplete, the islands themselves have by now been sufficiently well
worked to justify putting together up-to-date lists of the taxa that seem to have been reliably
recorded from each of the largest and to examine how far the presence or absence of some of the
members of this predominantly apomictic group has extra insights to provide about the floristic
affinities of the individual main islands — especially to one another — and of the archipelago as a
whole.

PREVIOUS WORK

Study of the islands’ Rubus flora dates from Babington’s pioneer exploration of 1837-8. Though
that was before he had begun to specialise in the group, by then he was already sufficiently
intrigued by its complexity to collect the odd specimen and seek the assistance of Borrer in naming
them. Not surprisingly, though, given the primitive state of Rubus taxonomy at that period, their
joint efforts proved on the whole inconclusive (Babington 1839); disappointingly, too, not all of
the specimens appear to have survived. There is equally slight surviving evidence in herbaria of a

422 D. E. ALLEN

visit made by H. C. Watson to Guernsey in 1852 and of one by A. Ley to both there and Jersey in
1885, while two species listed by Marquand (1892) from Guernsey were under names that cannot
be referred with confidence to any recognised today.

The islands were thus effectively still virgin territory batologically when W. Moyle Rogers, the
then leading British specialist, devoted a visit of almost a fortnight in 1897 expressly to closing
what had come to seem a glaring gap in his knowledge of the distribution in the British Isles of the
various taxa he was about to recognise in his forthcoming revision of the group (Rogers 1900).
Accompanied by his son, F. A. Rogers, who did some collecting as well, he concentrated on Jersey
and Guernsey, forgoing Alderney and managing a few hours only on Sark. In the subsequent
published reports (Rogers & Rogers 1898; Derrick 1898; Rogers 1899) he was to claim, with
evident relief, that the vast majority” of the forms encountered were “practically identical” with
ones with which he was familiar in Britain. The specimens collected, however, now mostly in BM,
provide notably less support for that conclusion than he had persuaded himself was the case, for
those that are of entities to which a name cannot be put today almost match in number the 16 taxa
in his list that he proves to have identified with British ones correctly. Had he written, rather, that
most of the widespread taxa encountered were also British, that would have been near the truth —
and is perhaps what he intended to convey.

Considering how many more of Britain’s more widespread Rubus entities had yet to be
discriminated and describea at that period, the extent to which Rogers mastered the group in the
islands on that single visit is nonetheless impressive. Unfortunately, for half a century after his
death there were not to be any further Rubus specialists, British or French, with direct experience
of the islands and attempts at determining material from there had a poor record of success by
comparison. When W. C. R. Watson was invited in 1930 to pronounce upon random specimens
G. C. Druce had collected on various visits, his touch was so uncertain that not only are one or two
of his determinations patently wide of the mark, but in two cases he changed his mind in later
years - but not for the better. He would really have been wiser not to have hazarded names at all,
for Druce’s material consists in the main of scarcely enlightening scraps. The mere handful now to
be found in OXF are evidently by no means the full number that Druce amassed, for some records
of his based at least partly on determinations by Rogers were published by him at an earlier date
(Druce 1907).

Most of Druce’s specimens are undated, but one from Jersey in 1923 shows that his collecting
just overlapped with the period of activity there of Fr. Louis-Arséne, whose herbarium, now at
JSY, contains a number of Rubus species allegedly from various named localities in that island. As
shown by Le Sueur (1982), however, the stated provenance of all Louis-Arséne’s botanical
specimens is suspect and it would be unsafe to do other than ignore the examples of Rubus in their
entirety.

RECENT WORK

Around 1970 a limited resumption of collecting took place in connection with the new Flora of
Guernsey then in preparation. Material of a few plants met with by D. McClintock and Lady A.
Brewis on different occasions was submitted to B. A. Miles for determination, with mixed success,
before being deposited in the herbarium of the Société Guernesiaise in St. Peter Port (STP) and the
few records resulting published in the eventual book (McClintock 1975).

Conscious of the inadequacy of the treatment of the group in that work, McClintock prevailed
upon the present writer to pay a week’s visit to Guernsey in July 1978 with a view to a more
intensive survey. The opportunity was taken to spend a day and a half in Sark as well and three
days in Jersey after that, with sufficiently productive results to warrant the publishing of new lists
for all three islands (Allen 1981; Le Sueur 1984).

Those lists have subsequently been considerably modified and extended in the light of four
further visits by the writer to Jersey and Guernsey for a week each (including a day on Herm) in
1990; to Jersey and Sark, similarly for a week each, in 1992; to Alderney for four days in 1993;
and finally to Jersey again for a week, with a day’s return to Sark, in 1994 (the updated Sark list,
without localities, was subsequently published by Marsden (1995)). All material collected has
been deposited in BM except for a few specimens donated to JSY and STP.

BRAMBLES OF THE CHANNEL ISLANDS 423

As a result Alderney, Sark and Herm can be claimed to have been investigated more or less
exhaustively; Jersey and Guernsey, however, have been covered only incompletely — possibly
important omissions in Jersey are Les Grands Vaux and parts of the north coast - and those larger
two have undoubtedly more to yield up.

ISLAND LISTS

The records for each of the main islands are considered below in turn, in the geographical
sequence of these from south to north. As no non-British taxa have as yet been detected, the
species order in each list is that in Kent (1992), to which reference should be made for sectional
and series names.

Many specimens cannot be matched at present with any entity known from the rest of the British
Isles or mainland Europe whether described or not. Unfortunately, authentic material of many taxa
recorded from northern France is not in any British herbarium and the location of type specimens
is all too often unknown. By no means all of those, however, would now be rated worthy of
taxonomic recognition, for names have frequently been conferred in the past on the unique
morphotypes (or “individua”’) that the peculiar reproductive system found in the Rubus fruticosus
ageregate not uncommonly gives rise to, or on entities with a distribution too narrowly local to
have any significance geographically. In each of the main islands, however, there are one or two
“innominati’ that are relatively widespread, some at least of which may qualify for description
should they turn out to occur widely outside the Channel Islands as well. Details of these select
few have been thought worth appending at the end of the respective lists.

Now that UTT-gridded Ordnance Survey maps have become available in recent years for each
of the four largest islands, it is at last possible to pinpoint Channel Islands records with six-figure
references, and these are provided below in appropriate instances.

Numbers cited for some of Rogers’ specimens are not his own (he did not use collection
numbers) but those of Herb. Barton & Riddelsdell, now in BM, into which were incorporated
many specimens of particular interest extracted from Rogers’ herbarium by Riddelsdell after it
passed into his possession.

JERSEY

The most southerly of the four main islands and the one with the warmest climate, Jersey is also
much the largest, its 115 km* making it almost twice the size of Guernsey. Though 24 km from the
mainland, not nearly as close as Alderney and Sark, it has much the shallowest intervening sea, a
principal reason for thinking that it was the last of the four to be severed from Normandy (Sinel
1912). While it resembles Guernsey in gradually sloping one way, it does so in the opposite
direction from north to south. The north and south-west coasts are fringed with steep cliffs with a
narrow belt of heath on top which widens in places into distinct commons and is one of the two
habitats in the island particularly productive of Rubus diversity. The other is the series of long
wooded valleys running down the centre of the island to the south coast. The lengthiest of these,
the St. Lawrence Valley (since the building of two large reservoirs better known as Waterworks
Valley), preserves at its north tip, close to the museum at Hamptonne, a fragment of rocky
woodland with an exceptional assemblage of unusual brambles, almost all of them so far
unmatched with any known from Britain or France. This is the one piece of ground encountered in
the whole of the Channel Islands that most gives the impression of a portion of Normandy
surviving little changed from the period before it was cut off by the sea.

R. briggsianus (Rogers) Rogers Recorded by Rogers from one spot near Waterworks Valley and
by Druce from La Corbiére. In neither case does a specimen seem to have been kept. Still not
certainly known on the European mainland, the species may yet survive in the coastal heath.

R. couchii Rilstone ex D. E. Allen Common and widespread in the north-east corner, rising to
abundance in Rozel Woods. A specimen of Druce’s in OXF from Flicquet Bay (where the
species is prominent today) was erroneously referred by Watson (Druce 1931) to “R. cariensis”
sensu Rogers (1.e. R. altiarcuatus W. C. Barton & Ridd.).

R. laciniatus Hayne & Willd. Mont du Vallet, L’Etacq, since the 1950s (Le Sueur 1984).

424 D. E. ALLEN

[R. lindleianus Lees Druce (1907), who was familiar with this common English species, claimed

to have seen it “in several places” on his recent visit, leaving it ambiguous whether Rogers had
confirmed his identification. None can be found in OXF today, no sign of the species has been
discovered in Jersey since and it appears to be absent from north-west France. Presumably one
or more other white-flowered, eglandular brambles were mistaken for it.]

[R. macrophyllus Weihe & Nees Rogers may have been correct in recording this from a deep lane

on the west side of Bouley Bay, but his specimen (BM) is not satisfactory enough for certainty.
Though occasional in woods in central Normandy, this species has only once been recorded in
the Cotentin Peninsula. |

. questieri Lef. & P. J. Mull. Common to abundant in several wooded tracts, but absent from

large areas. Rogers believed he saw this only in the Vallée des Vaux, where Ley had collected it
earlier, but there are specimens of his in BM (4900, 7459) from two other localities which he
failed to recognise, suggesting that his concept of the species was too narrow.

. cardiophyllus Lef. & P. J. Mull. Rare, as in Normandy, and nowhere more than a patch. On

roadsides in La Ville des Quennevais and Waterworks Valley but on wood margins in the
Mourier Valley and two places in St. Peter’s Valley. Earlier records are dubious: the surviving
specimen (BM, 8093) of what Rogers took to be a small-leaved form of this, at Rozel and Pont
Marquet, is of some other bramble, while Druce’s material from Le Portelet and St. Aubin’s
(Druce 1931) was so determined by Rogers in an aggregate sense only.

. davisii D. E. Allen Open heath south of Gros Nez, extending thence along the cliff-top towards

Plémont.

. dumnoniensis Bab. Rogers recorded a supposedly strong form of this “especially abundant at

Gorey Bay”, but his specimen (K) is of some quite different (but unknown) bramble. Another
specimen of his from Rozel (BM) he also incorrectly referred to this species. His claim to have
seen it in abundance in Waterworks Valley must accordingly be treated with reserve; though
certainly present, it is no more than occasional there today at best. Apart from a bush in the next
valley to the west (on the wooded margin of La Rue de I|’Aleval) and several on the upper
common at Le Ouisné it has been noted otherwise only on the north coast, at Le Marionneux (a
colony), Le Jardin d’ Olivet and La Chrétienne.

. polyanthemus Lindb. Occasional. Mostly among bracken on the north coast heaths, but in

woodland in the Vallée de Bellosanne.

. prolongatus Boulay & Letendre Rare. Several patches at top of Waterworks Valley and a bush

on a wood margin near Tesson Mill in St. Peter’s Valley. Rogers’ specimen (BM) of the “R.
micans” he listed from Anne Port proves to be this and probably his Gorey record of that
belongs here too.

[R. rhombifolius Weihe ex Boenn. “South of England incurvatus”’, the name by which Rogers

mostly understood this species, was recorded by him from St. Aubin’s Bay. No specimen has
been traced and the determination seems unlikely to have been correct. ]

[R. rubritinctus W. C. R. Watson Very rare. Two patches on lane at rear of Le Jardin d’Olivet.

R.

Recorded by Watson (1958) on unknown evidence and almost certainly in error. ]

sprengelii Weihe Very local. Apart from a find by Druce in 1908 vaguely localised as “St.
Helier” (OXF), where it must surely have been obliterated by building, all records are from the
north-east of the island, where it is plentiful especially around Bouley Bay.

. armeniacus Focke “Himalayan Giant” An increasingly widespread escape, though in colony

strength as yet only in La Rue de I’ Aleval. This cultivar, so very extensively grown in English
gardens since the Second World War, has found little or no favour in France and is
conspicuously absent from Normandy and Brittany.

. lamburnensis Rilstone Specimens (BM, 8043-4) of a bramble found by Rogers “in good

quantity” in Waterworks Valley match well Cornish material of this species in BM
authenticated by Rilstone himself. These Cornish specimens were initially labelled by him as R.
macrostemon Focke, to which Focke had independently referred, rather hesitantly, Rogers’
Jersey specimens earlier.

. ulmifolius Schott Very common.
. adscitus Genev. Very local. Abundant in wood along east bank of Queen’s Valley Reservoir;

also a patch or:two in St. Peter’s Valley and its Rue de I’ Aleval branch. A specimen of Druce’s
from Gros Nez (OXF), misdetermined by Focke as the Anjou species R. andegavensis Bouvet,
was initially referred to R. adscitus by Watson (Druce 1931), a determination, however, he later
withdrew; its identity remains unestablished.

a if eo = eee

BRAMBLES OF THE CHANNEL ISLANDS 425

. boraeanus Genev. Very common and (as Rogers noted) generally distributed, including on the

fixed dunes of Les Quennevais.

. caesarius D. E. Allen Locally abundant. Especially plentiful in the coastal districts, on banks,

roadsides and open heathland, but scarce in the central valleys. This is clearly the “abundant”
bramble which Rogers passed as the species now known as R. vestitus Weihe, although, as he
admitted, “usually untypical”.

. corbieri Boulay ex Corbiére Very rare. Single bushes at Portinfer and Bouley Bay.
. leightonii Lees ex Leighton Locally common at several points on the north coast, very thinly

scattered elsewhere.

. percrispus D. E. Allen & R. D. Randall Extinct? Old quarry, Le Jardin d’Olivet, one clump

1978, gone by 1990.

. dentatifolius (Briggs) W. C. R. Watson Locally plentiful on wooded slopes around Bouley

Hay; otherwise seen only in St. Peter’s Valley (including a colony in its Rue de |’ Aleval
branch) and upper part of Waterworks Valley. There is, however, a specimen in BM collected
on the south-west cliffs of La Moye by R. P. Murray in 1900, misdetermined as R.
dumnoniensis.

[R. adenanthoides Newton Listed for Jersey by Watson (1958) under the name mostly applied by

R.

him to this species, R. pseudadenanthus W. C. R. Watson. As none of Druce’s material
received this name from him, it appears he was repeating Focke’s determination of one of
Rogers’ specimens as R. adenanthus Boulay & Gillot. This was a very handsome” plant found
in some quantity in a very sunny spot in Gorey Bay. Rogers subsequently provided a
description of it under that name in his Handbook (Rogers 1900: 53), indicating that it was a
very hairy, considerably glandular-aciculate plant intermediate between R. radula Weihe ex
Boenn. and R. vestitus. At least one specimen was formerly in his herbarium, but none can now
be traced. ]

bloxamii (Bab.) Lees Rare. Except for a population in oakwood at the upper end of the Vallée
des Vaux seen only on the north coast cliffs: at intervals from Plémont to Gros Nez and on the
east side of Bouley Bay.

[R. echinatus Lindley A record of this from heathland by La Téte d’Ane (Le Sueur 1984) is

R.

R.

suspect: a similar-looking unnamed bramble was all that could be found in 1994 at this
locality. |

peninsulae Rilstone Common to abundant in and near most stretches of woodland. A species
not described till 1950 and known elsewhere only in Devon, Cornwall and one wood on the
northernmost part of the Brittany coast, this was initially queried by Rogers as R. oigoclados P.
J. Mull. & Lef. (9169, 9170) or R. micans Godr. (7458) but eventually (Rogers 1900: 78)
regarded by him as a form allied to R. botryeros (Focke ex Rogers) Rogers.

. conjungens (Bab.) Rogers Rogers’ record from Bouley Bay of the species then known as “R.

corylifolius var. cyclophyllus” is confirmed by his specimen in BM. This widespread British
bramble has as yet been detected on the European mainland only in Belgium. It is frequent near
the coast in Devon and Cornwall.

nemorosus Hayne & Willd. Very local. Bouley Bay district and in and between the central
valleys, but nowhere more than a patch.

[R. pruinosus Arrh. Rogers’ sole record of this (Bouley Bay) must be treated with reserve in the

R.

absence of a specimen. |

transmarinus D. E. Allen Correctly identified by Rogers as the member of sect. Corylifolii
Lindley familiar to him round Bournemouth as “R. dumetorum var. ferox’, this is occasional,
rather than “generally distributed”. Since the species was described (Allen 1994) it has proved
to occur widely in Normandy and plentiful in the district of Brittany between St-Brieuc and St-
Malo.

Innominati. Two unnamed brambles occur quite widely. One of these, as yet unmatched with any
known outside the Channel Islands, is a member of ser. Rhamnifolii (Bab.) Focke with pink
flowers and many long, subulate rachis prickles. This is found mainly in the Bouley Bay - Rozel
area and the central valleys (up to colony strength beside the All in St Peter’s Valley at
WV605518), was twice collected by Rogers (BM, 10082, 10083) and has been detected in
Guernsey also, on a hedgebank near St. Saviour Reservoir.

426 DE. ALLEN

Unlike that, a robust member of ser. Mucronati (Focke) H. E. Weber has been encountered on
the mainland immediately to the east: round a small wood on the D42 | km north-east of St-
Maurice-en-Cotentin, near Barneville. This has been noted in Jersey in three different areas:
Waterworks Valley, in some quantity north of Millbrook Reservoir and again east of Hamptonne;
Vallée des Vaux at WV651514; and Val de La Mare Reservoir, by the dam. In the third of these
the population is wholly white-flowered, whereas elsewhere only pink petals have been seen.

In and near an oak-holly wood near Trois Bois (Coétil de la Qualité nature reserve, WV623525)
are patches of another robust bramble with large pink flowers, often red-based styles and incised
leaflets, which, despite a sometimes amply glandular rachis and slightly aciculate stem, seems best
in ser. Sylvatici (P. J. Mull.) Focke. This has also been collected in the Cotentin Peninsula, but in
more distant parts, and is common in woods in the South Hams district of S. Devon, v.c. 3, where
it is well-known to R. W. Gould (as “the Andrews Wood Bramble’) and was collected by Briggs
from 1872.

GUERNSEY

Covering 63 km*, Guernsey has much less ground conducive to Rubus diversity than Jersey,
lacking wooded valleys at all comparable in scale and length and surviving expanses of suitable
heathland of a similar size and number. The flat, rather exposed northern one-third is largely bereft
of interest for the batologist, in marked contrast to the south, where plantations, massively
augmented by self-sown trees, clothe the cliffs and extend some way inland in places. That some
natural scrub may have escaped on these cliffs the otherwise wholesale removal of the island’s
original tree cover is suggested by the present-day restriction to those of such characteristic ancient
woodland indicators as Euphorbia amygdaloides L., Ruscus aculeatus L. and Luzula sylvatica
(Huds.) Gaudin (McClintock 1975: 19). That might account for the presence on them too of
several Rubus species known nowhere else in the island — though these might equally well have
been brought by frugivorous migrant birds, for which the fringe of trees and shrubs along the cliff-
tops, especially its profusion of bullace, Prunus domestica subsp. insititia (L.) Bonnier & Layens
(which fruits heavily in some years at least (N. Jee, pers. comm., 1990)), must constitute a
favourite landfall.

R. briggsianus Seen by Rogers in several places (and “in considerable quantity’, according to a
postcard he sent to E. F.Linton in 1914, now in BM), including St. Peter, Petit Bot Bay, Cobo
and quarries between St. Sampson and L’ Ancresse, from the last two of which specimens of his
are in BM. Extensive draining of boggy ground since may have rendered it extinct.

R. laciniatus Several recent records for this naturalised escape are cited by McClintock (1975).
Rogers met with it only in Petit Bot Bay quarry.

[R. oxyanchus Sudre Cited for the Channel Islands by Edees & Newton (1988) as a result of the
patch of R. daveyi, q.v., having initially been determined as this. ]

R. plymensis (Focke) Edees & Newton Common in plantations round Belvedere House, Fort
George.

R. questieri Very local. Rogers said he had searched for this in Guernsey in vain, but as in Jersey
his over-narrow conception of the species is shown by his having collected it under other
names: his “R. villicaulis’ (STP) and “R. rhombifolius’ (BM, 7211; K), both from Fermain,
where it is abundant in woodland today, have proved to be this. Scattered bushes extend to
plantations on the south edge of St. Peter Port and occur in one or two other spots as well.

R. cardiophyllus Rare. Of the two stations recorded by Rogers, Fermain Bay yielded just one bush
in 1978 while Petit Bot Bay may have been a loose localisation of a colony that grows among
gorse on the cliff-top 0-5 km to the south. A bush has also been noted at Jerbourg.

R. cordatifolius (Rogers ex Ridd.) D. E. Allen Supposed by Rogers at the time of his visit to be
merely a luxuriant state of what he understood as R. dumnoniensis, this very distinct bramble
still hangs on at Petit Bot Bay, the source of his specimen (BM, 9823) that now constitutes the
lectotype. Elsewhere in the south it has been noted in coastal scrub at Icart and in Silbe Nature
Reserve in St. Peter; its presence, however, also on the north edge of St. Peter Port, in two spots
east of Le Hougue des Quartiers, suggests it could turn out to be fairly widespread. It is still not
known in any of the other islands apart from Sark or for certain anywhere outside them. A
Belgian bramble that has been passing there under the names of “R. nemoralis” and “R.
argenteus subsp. incarnatus var. leventii’” (H. Vannerom, pers. comm., 1993) has been

BRAMBLES OF THE CHANNEL ISLANDS 427

identified with it by Vannerom (1998), but the one specimen of that seen by the writer, though
very similar, lacks the pecuhar inflorescence structure of the Channel Islands plant; the
resemblance may therefore be fortuitous.

R. daveyi Rilstone One patch in a bullace thicket beside the cliff-top lane midway between
Portelet Bay and Les Fontenelles at WV300745 and a further bush nearby among gorse.
Hitherto believed endemic to Cornwall and a few places in Devon.

[R. dumnoniensis Rocquaine Bay was identified by Rogers (1899) as among the “one or two other
localities” apart from Petit Bot Bay (his locality for what was to prove R. cordatifolius) in
which he claimed in the initial paper to have seen this. Though its presence in the other three
main islands makes it a likely Guernsey species, the lack of later records and the over-broad
interpretation of the taxon by Rogers, revealed by material from Jersey and elsewhere
misdetermined as this species, make it necessary to treat his claim with reserve in the absence
of any specimen. ]

[R. herefordensis Sudre The basis for a claim by Watson (1958) to have seen a specimen of this
from St. Andrew has not been ascertained. The species as now understood is unknown outside
Hereford, v.c. 36.]

R. polyanthemus Local, but markedly more plentiful than in the other islands. Particularly large
populations in the south-west corner, as in Silbe Nature Reserve and round Pleinmont.

R. prolongatus Rare. Recorded from six localities in the south, half of them on the coast, but in
quantity only on Petit Bot cliffs.

[R. riddelsdellii Rilstone A specimen in STP collected by McClintock west of Chateau des Marais

was thought by Miles to be this, but the material is so poor that any determination would seem

unwise. ]

rubritinctus Listed by Watson (1958) on unknown evidence. Rogers remarked on its absence.

sprengelii Collected by Rogers at Fermain Bay (BM, 6073), but perhaps now extinct.

armeniacus “Himalayan Giant” As in Jersey, becoming a widespread escape.

pydarensis Rilstone Very local or overlooked. Common in hedges about 1 km inland in the

district south of Les Arquets, St. Peter (conf. L. J. Margetts). Specimens in BM collected by

Rogers at Fermain Bay (4898) and La Valette, referred by him to an aggregate taxon he called

“R. argentatus’, also turn out to be this species, hitherto believed endemic to Cornwall and a

few places in Devon.

R. ulmifolius Very common.

[R. adscitus Believed to have been seen in 1978 in Silbe Nature Reserve, but requires
confirmation].

R. boraeanus Occasional. Noted more especially in Castel and St. Andrew. Surprisingly, Rogers
appears not to have seen this. Of the only two localities given in the original paper (Rogers &
Rogers 1898), one is omitted from his 1899 paper, suggestive of second thoughts, while the
specimen from the other locality (BM, 10258) has proved to be a misdetermination. Possibly
the species has colonised the island subsequently to his visit.

[R. conspersus W. C. R. Watson Listed by Watson (1958) on the strength of his determination of
one of Rogers’ specimens in BM from L’ Ancresse Common as this in 1951. The specimen,
however, is R. corbieri.]

R. corbieri Mainly in the north, where it is frequent on hedgebanks and abundant in old quarries,
having probably colonised Guernsey from Sark subsequent to that island’s sixteenth-century
resettlement or, alternatively, direct from the Cotentin Peninsula beyond.

R. leucostachys Sm. A colony by the cliff path on the south side of Fermain Bay.

[R. vestitus This was the species, very common in much of England, to which Rogers and his
contemporaries generally misapplied the name R. leucostachys. In the supplementary list
(Rogers 1899) he provided for Guernsey that appears with “generally distributed” and “very
variable” against it. Presumably one or more of the island’s still-unnamed brambles passed as
this - as was certainly the case with specimens of Druce’s in OXF and Rogers’ own (BM,
10081) from St. Andrew. ]

R. coombensis Rilstone One bush in shade on the cliff path south of Le Vallon (WV330754).
When discovered in 1990, this was believed endemic to south-west England, but it has since
been recognised as a bramble local in quantity round Cherbourg.

ae ete

428 D. E. ALLEN

[R. moylei W. C. Barton & Ridd. Listed by Watson (1958) on unknown evidence. In his
supplementary list Rogers (1899) included the name R. lejeunei var. ericetorum, by which he
normally meant this species, but gave no locality. It is unlikely that Watson saw that paper,
however. |

R. norvicensis A. L. Bull & Edees Very local. Plentiful in plantations and shrubberies across the
south-east corner, from above Moulin Huet Bay and Sausmarez Manor to the south edges of St.
Peter Port. This species is otherwise known only in parts of England east of a line from
Salisbury to The Wash and along one valley in south-west Wales. The compact nature of its
Guernsey distribution and prevalence there in shady habitats comparatively recent in origin
both suggest a colonist in the process of primary spread, perhaps following accidental
introduction with nursery shrubs (as seemingly in some of its English stations). If so, that
introduction must have been over a century ago, for this turns out to be the identity of one of the
two brambles collected by Rogers’ son at Fermain Bay (BM) which, after some hesitation, his
father concluded was R. rudis Weihe “type or near it” (Rogers & Rogers 1898; Rogers 1899) —
an unlikely species to occur in the Channel Islands, however, on geographical grounds.

R. percrispus_ This is the identity of the other Fermain “R. rudis” collected by F. A. Rogers, the
specimen of which is in this case in K; it was initially determined by his father as the bramble
now known as R. micans Godr. A depauperate example of probably this same species was
collected on the cliff path above Petit Port in 1978. Recently described, like the previous
species. R. percrispus is otherwise known outside the Channel Islands as yet only in parts of
southern England and a single locality each in west Wales and Ulster.

R. dentatifolius Very local. Shares old quarries in Vale with R. corbieri and still as abundant in

one as when Rogers found it there. Most of the few other records are from the south coast.

. leyanus Rogers Rue du Banquet, near Pleinmont, one patch in hedge of the northernmost bend.

. bloxamii Locally common; much more plentiful than in Jersey. Mainly in the southern half of

the island but conspicuously absent from the self-sown woodland at Fermain. A specimen in
CGE collected by Babington at Fort George is labelled “R. koehleri var. lejeunei”’.

[R. radula Recorded from Fermain Bay by Rogers (1899) in his supplementary list, but no
specimen has been traced. A specimen in STP from Mount Durant determined as this by Miles
(McClintock 1975) has turned out to be R. norvicensis. |

[R. conjungens “Fairly frequent” according to Rogers, but there have been no later finds and his
statement must be treated with reserve in the absence of a confirming specimen. ]

R. transmarinus Rogers’ initial description of the bramble he knew as “R. dumetorum var. ferox”
as generally distributed was modified in his supplementary list to “frequent”. It appears,
however, to be thinly scattered at best, mainly in the vicinity of the south-east coast.

poe)

Innominati. Of numerous unnamed brambles, one is particularly widespread, a member of ser.
Vestiti (Focke) Focke with deep pink flowers, red styles and aculeate sepals, this is common in
woodland in the centre of the island, especially along the Fauxquets Valley. Seemingly the same
plant occurs in Jersey, in a wood by Gargate Mill Farm (WV604518) in St. Peter’s Valley.

SARK

The smallest of the four main islands and unique among them in being entirely a lofty plateau,
which gives it the appearance of an aircraft-carrier, Sark was probably a stretch of treeless heath
before its first substantial colonisation in the sixteenth century. Like the very similar [le de Bréhat
off the north coast of Brittany, it is essentially two islands, connected merely by a narrow isthmus;
a detached portion of the larger one, the now strictly private Brecqhou, lies just 200 m off the west
coast. High cliffs all round permit access to the sea in only a few, very restricted places. Protected
from Atlantic gales to some extent by Guernsey, 13 km to the west, it presents a comparatively
lush contrast to Alderney, as remarked upon by Druce (1907) after visiting the two for the first
time in quick succession. Much less ground than in the other main islands has escaped cultivation
and such of the little heath cover that remains is best represented by Eperquerie Common, at the
north tip.

R. altiarcuatus W. C. Barton & Ridd. In an isolated clump of tall gorse by La Valette
(WV474764) and several further bushes along the cliff-top to the north-west. The sole Channel
Islands locality for this widespread species of south-west Britain, also in the far west of both
Ireland and Brittany.

BRAMBLES OF THE CHANNEL ISLANDS 429

R. cordatifolius Scattered among gorse south-east of the Pilcher Monument (WV455754) — which
is by no means “near the Dixcart Hotel”, where Rogers recorded this in quantity (specimen in
BM). A search of the hotel’s vicinity has proved fruitless, but the area is no doubt much
changed since 1897.

dumnoniensis Several patches in a rough field west of D’Icart (WV466756) and one bush near

the top of Eperquerie Common.

nemoralis P. .J. Mull. In four widely separate places in Great Sark and one in Little Sark,

mostly as single bushes and nowhere more than a patch.

polyanthemus A heath relic. One or two bushes at Eperquerie Landing; also collected on a cliff

on Brecghou by Mcclintock in 1968 (STP).

prolongatus Several patches in the south-west of Little Sark and a bush in a gorse hedge just

below D’Icart.

rubritinctus Several patches at different levels on Greve de La Ville cliffs.

ulmifolius Very common.

corbieri Very common. A conspicuous feature of virtually every hedgebank, this probably

post-dates their construction, having spread across from the north-west of the Cotentin

Peninsula, where it occurs in comparable profusion locally. Rogers at first referred it to the

group of brambles to which he applied the name R. schlechtendalii (Derrick 1898), but by the

time he published his records (Rogers & Rogers 1898) had decided it best regarded as a strong
and highly glandular form of R. macrophylloides Genev., a species described from Angers.

Subsequent collectors were to have specimens referred also to R. adscitus or R. boraeanus

before the plant’s true identity was finally established c. 1990 (Allen 1992). Patches of apparent

hybrids with R. ulmifolius have been noted in two places.

R. leucostachys One bush on cliff-top north-west of La Valette and a clump by the cliff path above
Dixcart Bay. The latter may be much the same spot as the “furze brake near Dixcart Lane” from
which Rogers recorded in a supplementary list (Derrick 1898) a bramble under this name. By
this he normally understood R. vestitus, which is absent from the Channel Islands, or, in Jersey,
as R. caesarius.

R. dentatifolius Frequent in rough ground. Probably general in the pre-resettlement era.

R. conjungens An unlocalised Sark specimen in CGE collected by Babington in 1838 and
successively determined by him as “R. rhamnifolius” (as published in Babington 1839) and “R.
thyrsoideus” is in fact this, as first recognised by Miles.

R. nemorosus Frequent; much more plentiful than in the other islands. In places in open heath (in
which it was doubtless prevalent formerly).

R. transmarinus Occasional, mainly around the south-east coast. Perhaps in part the bramble
determined by E. F. Linton under the aggregate name of “R. corylifolius” which Hurst (1903)
claimed to be abundant on Brecghou.

Bo re Se ae Re aU ae

Innominati. Sark has two relatively widespread brambles that are also known outside the island but
so far unnamed. One, a pink-flowered member of ser. Sylvatici with strongly hairy anthers and
reminiscent of small forms of R. gratus Focke, occurs in some quantity on the cliff-top north-west
of La Valette and in two places on Eperquerie Common and also as single clumps on Harbour Hill
(WV476758) and by D’Icart. Since its original discovery (“Sa36” in Allen 1981) it has been found
in the third-largest of the Isles of Scilly (Allen 1997) and on a wood margin in the south-east
corner of Hampshire, v.c. 11. This distribution suggests it may occur in Brittany too. The other is a
member of ser. Radula (Focke) Focke recalling R. longithyrsiger Lees ex Focke but with pink
flowers, pale styles and hairy anthers. In addition to a colony at Eperquerie Landing this occurs on
the east cliff-tops between La Valette and Gréve de la Ville as well as by D’Icart (WV466755); it
is also in Jersey, dotted around Dannemarche Reservoir in Waterworks Valley.

Also worthy of mention is a very distinct member of ser. Discolores (P. J. Mull.) Focke with
hairy anthers, pink filaments, pink-based styles and mucronate terminal leaflets with a lengthy
mucro. Patches of this occur in Sark on the cliff-top at Gréve de la Yule and in the gorse heath
near the Pilcher Monument, in Alderney (where A. Newton has also seen it) in the old quarry on
Val Longis (WA581076) and on the coast west of Cherbourg near Landemer.

430 DU EALEEN

HERM
Covering 2-5 hectares, the largest of Guernsey’s satellites, privately-owned but publicly accessible,
Herm is an object lesson in the sharply disproportionate reduction in species diversity that can
occur in the case of islands that are below a certain size — even in the case of plants that are bird-
dispersed. In startling contrast to its neighbour some 5 km to the east, hardly any of Guernsey’s
numerous Rubus species are to be found, not even most of the commonest. Apart from the
predictably abundant R. ulmifolius, indeed, and a patch by the landing of a member of the “R.
dumetorum” aggregate, there appears to be only one other — and that not a Guernsey one, so far as
is known at present. Resembling R. orbus W. C. R. Watson of south-west England and belonging
like that to ser. Vestiti, this is common in the south half in ground with a modicum of shelter.
Herm’s own even smaller satellite, the more strictly private Jethou, had several bramble
specimens collected from it by Babington. Borrer, to whom he sent one, referred it doubtfully to R.
leucostachys (Babington 1839); but Miles, who discovered it in CGE in 1965, thought it was
indeed that species, in the original sense of Smith; though that has since been established as
occurring in both Guernsey and Sark. Babington’s material is unfortunately too scanty and
immature to be given a confident determination.

ALDERNEY
The northernmost of the main islands, 13-5 km west of Cap de la Hague and thus almost twice as
close to the mainland as Jersey, Alderney differs from the other main three in resembling an
inverted saucer, the one town standing on a hill in the centre. As in Guernsey, the northern third is
low-lying and only the more southerly part fringed by cliffs. Though its cultivation dates from a
much earlier period than Sark’s, it still has extensive tracts of rough grassland and heath, of which
those of La Giffoine are the most rewarding for the batologist. Introduced trees form dense
thickets in places, but rarely shelter brambles of interest. The most exposed of the islands to the
winds of the English Channel, Alderney’s Rubus flora mainly owes such distinctiveness as it has
to its consequently more oceanic climate.

R. viridescens (Rogers) T. A. W. Davis One patch among roadside gorse above Fort Tourgis just
south of Roc a |’Epine.

R. dumnoniensis Local; patches on the coast here and there in the western one-third. Relatively
more plentiful than in the other islands (though it seems to be missing from the coastline round
Cherbourg).

R. iricus Rogers In plenty with R. viridescens. Locally abundant on La Giffoine southwards from
east of Fort Clonque; also one bush in the Valley Gardens in the middle of St. Anne.

R. polyanthemus A patch in gorse heath on top of Essex Hill.

R. prolongatus Widespread and locally abundant on the south-west heaths; also abundant round
the television tower on Les Rochers and in some quantity on Essex Hill. Markedly more
plentiful than in the other islands.

R. armeniacus “Himalayan Giant” Escaping from a garden in St. Anne.

R. ulmifolius Very common.

[R. winteri P. J. Mull. ex Focke Listed by Watson 1958) following his determination of one of
Druce’s specimens in OXF as this, in place of Rogers’ “apparently R. leucostachys’. It is
neither of those, however, but may be the same as an unnamed Guernsey bramble of
hedgebanks in St. Andrew. |

[R. adscitus Another of Druce’s specimens in OXF was considered by Watson at one time to be
this (Druce 1932), but it appears to be merely a form of R. ulmifolius.]

R. boraeanus Common. -

R. corbieri Very thinly scattered, achieving quantity only in three particularly sheltered locations:
Barrack Master’s Lane (“Essex Glen”), Vau du Saou and the top of Le Val Reuters.

R. dentatifolius Common and locally abundant on the southern heaths and cliffs southwards from
Fort Tourgis in the west and Le Longue Pierre in the south; elsewhere rare or absent.

R. tuberculatus Bab. Common, taking the place of R. transmarinus.

Innominati. Given its greater proximity to the mainland, Alderney has a proportionately higher
share of brambles that cannot be matched with any known in Britain. One of these is widespread
on the neighbouring Cotentin Peninsula, a member of ser. Discolores resembling R. rossensis
Newton, this occurs at the north tip near the lighthouse and in two places on the south coast. A
large patch of another member of ser. Discolores on the upper margin of Barrack Master’s Lane at
WA591077 appears to correspond to a poorly-understood bramble of northern France that has

BRAMBLES OF THE CHANNEL ISLANDS 43]

passed under the illegitimate name of R. propinquus P. J. Mull. A very distinct, white flowered
member of ser. Sylvatici with many tiny rachis prickles and long, acuminate leaflets is also
widespread, with clumps at intervals along the south-east coast and several patches on La Giffoine.

DISCUSSION

Altogether, 39 entities currently accorded taxonomic recognition as species, including two
naturalised horticultural ones, are accepted as reliably on record from one or more of the islands.
Of these, 26 are on record from Jersey, 23 from Guernsey, 14 from Sark and 11 from Alderney.
Though these totals roughly correlate with island size, the diminution is by no means a straight
reflection of increasing failure to colonise on the part of certain species. Instead, as the Table
below demonstrates, there are marked discrepancies between the four main islands in how the
totals are made up:-

Jersey Guernsey Sark Alderney
R. briggsianus 0 0
R. couchii 3
R. laciniatus 1 py)
R. plymensis 1
R. questieri 3 1
R. viridescens l
R. altiarcuatus 1
R. cardiophyllus ]
R. cordatifolius 1 l
R. daveyi ]
R. davisii ]
R. dumnoniensis 2 1 2
R. iricus 3
R. nemoralis 1
R. polyanthemus 2 2 1 1
R. prolongatus l 2 1 3
R. rubritinctus l ]
R. sprengelii | 0
R. armeniacus 2 2 1
R. lamburnensis 1
R. pydarensis 1
R. ulmifolius 3 3 3 3
R. adscitus 1
R. boraeanus 3 2 3
R. caesarius 3
R. corbieri 1 pp yy
R. leucostachys ] 1
R. coombensis 1
R. leightonii 3
R. norvicensis 1
R. percrispus 0 0
R. dentatifolius D 2) 2 3
R. leyanus 1
R. bloxamii 1 3
R. peninsulae 3
R. conjungens 0 0
R. nemorosus 1 2
R. transmarinus 2 2 ye
R. tuberculatus 3

O - extinct ?, 1 - rare to very local, 2 - widely scattered, 3 - locally to very common

432 D. E. ALLEN

All of these species occur in Great Britain also - though in the case of R. iricus only in an island
group far offshore - with the single exception of R. cordatifolius, which is apparently endemic to
Guernsey and Sark. It is possible, however, that the seeming identity of R. caesarius with a
bramble locally plentiful in one small district of S. Hants., v.c. 11, between Southampton and
Portsmouth (Allen 1994), is the result of a fortuitous overlap in characters, in which case that
would be a second apparent endemic (for despite the abundance of that species in Jersey no sign of
it has so far been found in the nearer parts of the mainland, very surprisingly). Though no
Continental taxa unknown in Britain have yet been detected in the islands, at least one or two of
the many brambles to which no name can be put at present will almost certainly prove to come
into that category once the taxonomy of the Rubus flora of north-west France has been more fully
worked out. Until only two or three centuries ago, however, R. corbieri may have belonged in that
category, for in the south-east corner of Dorset, v.c. 9, the sole area in which this conspicuous
species is known on the other side of the English Channel, it has all the appearance of a
comparatively recent colonist, though probably present well before the first specimen was
collected, in 1892 (Allen 1992).

Of the 37 species apart from R. caesarius that the islands share with Britain, eight are not yet
known to occur on the European mainland. These are R. briggsianus, R. daveyi, R. davisii, R.
iricus, R. lamburnensis, R. pydarensis, R. norvicensis and R. percrispus. R. norvicensis, as already
intimated, appears to owe its presence to accidental introduction with nursery plants at some
comparatively recent date, while R. percrispus, despite a wide scatter of localities across southern
England, has had too brief a taxonomic history for its presence or absence on the European
mainland to have had time to be adequately tested as yet. The rest of the eight, though, are all
strongly western species whose ranges could well extend at least to Brittany. Nevertheless,
extensive sampling of coastal areas of both Brittany and western Normandy in recent years has not
only failed to reveal those, but has turned up instead other species with a similar range on the other
side of the English Channel: R. hastiformis W. C. R. Watson, for example, and R. aequalidens
Newton along the Céte de Granit Rose, the former in abundance, and R. rilstonei W. C. Barton &
Ridd. in no less abundance on Cap Fréhel, west of Dinard - each of them seemingly no less likely
candidates for the Channel Islands flora.

More perplexingly, there are several brambles abundant in the adjacent parts of France that
appear to be absent from the Channel Islands. The most conspicuous of these absentees is R.
venetorum D. E. Allen, which is plentiful, commonly in profusion, almost throughout Brittany,
including on the Ile de Bréhat, to Jerseys south-west. Almost as odd is the failure of R.
thyrsigeriformis (Sudre) D. E. Allen to reach at least the northern islands. It has found its way
from the north of the Cotentin Peninsula to a wide part of S. Hants., v.c. 11 (in which it gives the
appearance, like R. corbieri, of being a comparatively recent invader still in the course of primary
spread). Similarly, there is no sign of an unnamed member of ser. Sy/vatici that grows in profusion
on the heaths around Cherbourg. Why has R. questieri, not noticeably more plentiful on the
mainland than R. venetorum, been able to colonise Jersey and Guernsey and yet none of those?

It may be that the transmarine movements of the relevant species of frugivorous birds have
persistently been from other directions than from the mainland for the most part, in particular from
the north-west. It certainly looks suggestive that two species otherwise confined to Britain and
Ireland and especially plentiful in south-west England, R. leyanus and R. plymensis, are both
known from just a single locality in the Channel Islands (and in the westernmost of those,
Guernsey) and from just a single mainland locality too, in the Cotentin Peninsula to the east — as if
they were legacies of one and the same fly-line. R. couchii has (or rather had, for at Cherbourg it is
probably extinct now) a similar distribution pattern, though in its case the population in Jersey is a
very large one and in Britain the species appears confined to a solitary district in south-east
Cornwall, v.c. 2. Even more suggestively, R. daveyi, a common species of Cornwall which extends
thinly into Devon, has as its only Channel Islands site a bullace thicket on top of a Guernsey cliff,
precisely the kind of landfall likely to be made by a migrant coming in from the open sea to the
west. A species of warbler, most probably a Blackcap, Sylvia atricapilla, is likely to have been
responsible in such a case, for that is known to feed on blackberries (Snow & Snow 1988: 61) and
there have been recoveries in Guernsey of individuals ringed in Dorset. Moreover, unlike members
of the thrush family, the birds generally thought to be the principal disseminators of Rubus species.
Blackcaps and other warblers are commonly seen on the Guernsey cliffs (N. Jee, M. Austin, pers.
comm., 1990).

BRAMBLES OF THE CHANNEL ISLANDS 433

Of all the British species that have turned out to be in the Channel Islands R. iricus was the least
expected. At the time of its discovery in Alderney it was known elsewhere only along the Atlantic
seaboard of Ireland, where it had long been presumed endemic and in parts of which it is very
common. Subsequently it has proved to occur in abundance in the Isles of Scilly as well, which
may thus have served as a half-way stepping-stone without which such a great distance seems
much less likely to have been bridged by a bird with viable Rubus seed in its intestine (Allen
1997). The oceanic character of both Alderney and Scilly strongly suggests that the bird
responsible for the original transmission between the two was a sea-going species, perhaps a
member of the gull family. The possibility that gulls may disseminate Rubus species does not
appear to have been entertained previously, but the recent find of a bush of R. daveyi in the middle
of a gullery on Gugh in the Isles of Scilly (Allen 1997) lends that idea some credence. Certainly,
there is evidence that gulls feed on the fruits of other plants: Solanum nigrum L. has been noted in
quantity in gulleries on the Calf of Man, v.c. 71 (Allen 1984) and S. dulcamara L. in the crevices
of cliffs alongside nests on the off-shore islands of Pembroke, v.c. 45 (Gillham 1956: 446), while
Snow & Snow (1988: 167) cite records from Central Europe of different gull species eating the
fruits of Vaccinium and Empetrum spp., Hippophae rhamnoides L. and even the blackberry-sized
fruits of cherries (Prunus spp.), which flocks of Black-headed Gulls pluck from the trees in flight -
an apparently recently-developed habit.

The presence so far to the east of species characteristic of , or otherwise confined to, the extreme
south-west of England is explained by the similar climate of the two regions. Although the
Cotentin Peninsula of Normandy lies on the same longitude as Bournemouth and the Isle of Wight,
its western half is so exposed to maritime influence in general and to the Gulf Stream in particular
that its floristic affinity is rather with Cornwall and Devon (Allen 1996). That climate is shared
with the Cotentin’s offshore islands, which can be expected to experience an added measure of
oceanicity by virtue of their insular character. That essential homogeneity of the western Cotentin
and at least the more northern of the Channel Islands is well shown by the distribution of R.
corbieri, a robust, shaggy species clearly exceptionally well adapted to existence in such a wet and
windswept region. Apparently relatively recent in origin, with R. bloxamii patently as one
ancestor, it would seem to have spread aggressively at one period in the not-too-distant past,
spilling across from the Cotentin on to each of the four main islands and colonising the northern
three in proportion to the amount of shelter they offer.

Though the most characteristic of its species, R. corbieri is not alone in being seemingly well-
suited to the region’s relatively high oceanicity. R. iricus is a similarly robust and very hairy
bramble, with presumably many centuries, if not millennia of adaptation to the climate of the west
coast of Ireland. It can be no accident that Alderney is the only one of the Channel Islands in
which it has become established - and in considerable quantity. Besides R. corbieri the archipelago
is noticeably rich in other members of ser. Vestiti, a chief character of which is a dense
indumentum: they include R. boraeanus, one of the commonest species of Jersey and Alderney
(though absent from Sark), R. caesarius, similarly prevalent in much of Jersey, the unnamed
bramble plentiful in woodland in the centre of Guernsey and the unnamed one that dominates the
Rubus flora of Herm jointly with R. ulmifolius. Members of this series are likewise prominent in
the Cotentin Peninsula.

It is presumably for the same reason, that it can better cope with oceanic conditions, that R.
prolongatus is distinctly more widespread in Sark and especially in Alderney than in Guernsey and
Jersey, while the less hairy R. polyanthemus occurs in exactly the opposite island order of
frequency. In just the same way R. prolongatus occurs on four of the isles of Scilly and in local
abundance on the largest, whereas R. polyanthemus is known from only two and is very rare in
both (Allen 1997). The difference between this pair in apparent oceanicity tolerance, though,
would seem to obtain only at these latitudes, where R. polyanthemus, altogether a more northern
species extending to Scandinavia, is at the southern limit of its range and thus presumably at a
competitive disadvantage: on the west coast of Ireland it is R. polyanthemus that is locally
common and R. prolongatus that is rare.

That a preference for the relatively moister and milder climate of those parts of western Europe
more especially subject to the influence of the Gulf Stream is not necessarily synonymous with a
tolerance of oceanic conditions is particularly well exemplified by the distribution pattern of R.
rubritinctus. Rubus specialists accustomed to the prevalence of that species in Devon and

434 D. E. ALLEN

Cornwall (except the westernmost part) have difficulty in believing that in the Isles of Scilly it is
represented by no more than a bush or two. No less puzzling at first sight is its wide scatter around
Southampton Water yet total absence from the Isle of Wight, at most a mere 8 km offshore to the
south (Allen 1990) - and despite the fact that this is a bramble especially dependent on birds for its
dispersal, to judge from its marked tendency to occur as solitary bushes. The explanation for the
second of these anomalies is that the Isle of Wight shields Southampton Water from the open sea.
A slender plant with long ascending inflorescence branches and for the most part only sparsely
hairy, R. rubritinctus is seemingly ill-adapted to oceanic rigours, and an overriding requirement for
shelter from those largely excludes it from both the Channel Islands and the Cotentin Peninsula,
just as it does from Scilly. Significantly, of the only two sites known for it in the Channel Islands,
Jersey’s is on the inland margin of a coastal common and Sark’s is the only sheltered cove on the
side that faces the mainland. The only area in that region where it thrives in quantity is the
southernmost stretch of the Gulf of St-Malo coastline east of St-Brieuc, an area comparable to the
Lancashire side of Morecambe Bay in England in the extent to which it constitutes an isolated
“oasis” for this protection-demanding species.

Another distribution that merits some final comment is that of R. leightonii, which Jersey alone
of the islands possesses and that in considerable plenty. Together with R. bloxamii this has a
relatively narrow corridor of abundance down the centre of England from Staffordshire to the
Bournemouth district, to either side of which the two species diminish steeply. Though both ranges
continue southwards across the sea into Normandy , they lose their tightly corseted character there
and the species lose their closely parallel occurrence as well. Once across the English Channel
R. leightonii becomes noticeably the more warmth-demanding and intolerant of oceanic
conditions, extending apparently further south (to beyond the Loire) and much further east than
R. bloxamii and, unlike that, missing Guernsey and all but the south end of the Cotentin Peninsula.
Its prevalence in Jersey, however, is repeated along the southern coast of the Gulf of St-Malo,
from which R. bloxamii is seemingly debarred climatically - as both species are from the main
mass of Brittany to the west.

These examples illustrate how differences between the islands in the Rubus species they possess
or lack can in some cases be explained by scrutinising the wider ranges of those species for
illuminating anomalies. They also serve to underline the substantial size of the addition to the
phytogeographer’s armoury that the study of a critical group such as this can bring, by reason both
of its considerable species diversity even in an archipelago of relatively small isiands and of the
sensitivity of those individual species to subtle gradations in atmospheric moisture.

ACKNOWLEDGMENTS

A grant from the Appleyard Fund of the Linnean Society of London made it possible to extend to
Alderney the fieldwork on which this paper is based. A grant from the B.S.B.I Welch Bequest
also made it possible to replace the greater part of one year’s collection after its loss in transit. Iam
most grateful for both.

I would also like to thank David McClintock for introducing me to the Channel Islands and both
him and Cherry Lihou for their assistance in the field on my initial visit, the late Frances Le Sueur,
Patience Ryan and Marcia Marsden for hospitality and assistance in Jersey, Guernsey and Sark
respectively, La Société Jersiaise and La Société Guernesiaise for access to their herbaria (and the
latter also for the subsequent loan of specimens), Nigel Jee and Margaret Austin for information
on Guernsey bird movements and ringing recoveries, Chris Preston for guidance on map grids, and
Ray Gould, Len Margetts, Herman Vannerom, Giinter Matzke-Hajek and above all Alan Newton
for assistance with determinations. McClintock and Gould were additionally kind enough to
comment on this paper in draft.

REFERENCES

ALLEN, D. E. (1981). Brambles of Guernsey and Sark. Société Guernesiaise report & transactions 1980 20:
609-615. :
ALLEN, D. E. (1984). Flora of the Isle of Man. Manx Museum and National Trust, Douglas.

BRAMBLES OF THE CHANNEL ISLANDS 435

ALLEN, D. E. (1989). A neglected bramble of Guernsey. Watsonia 17: 436-437.

ALLEN, D. E. (1990). The Rubus flora of the Isle of Wight. Watsonia 18: 21-31.

ALLEN, D. E. (1992). Rubus corbieri Boulay (Rosaceae) in the British Isles. Watsonia 19: 131-133.

ALLEN, D. E. (1994). Four new Anglo-Sarnian species of Rubus L. (Rosaceae). Watsonia 20: 143-151.

ALLEN, D. E. (1996). British species of Rubus L. (Rosaceae) in the Cotentin Peninsula of Normandy.

Watsonia 21: 129-133.

ALLEN, D. E. (1997). Rubus L. (Rosaceae) in the Isles of Scilly: a revised list. Watsonia 21: 355-358.

BABINGTON, C. C. (1839). Primitiae florae Sarnicae. Privately published, London.

DERRICK, G. T. (1898). Additions to the Sark flora. Guernsey Society of Natural Science & Local Research
report & transactions 1897 3: 166-169.

DRUCE, G. C. (1907). Notes on the flora of the Channel Islands. Journal of botany 42: 395-402, 419-428.

DRUCE, G. C. (1931). New county and other records. Botanical Society & Exchange Club of the British Isles
report for 1930 9: 331-379.

DRUCE, G. C. (1932). New county and other records. Botanical Society & Exchange Club of the British Isles
report for 1931 9: 632-679.

EDEES, E .S. & NEWTON, A. (1988). Brambles of the British Isles. Ray Society, London.

GILLHAM, M. E. (1956). Ecology of the Pembrokeshire islands, 3. Manuring by the colonial seabirds and
mammals, with a note on seed distribution by gulls. Journal of ecology 44: 429-454.

Hurst, C. P. (1903). The flora of the island of Brechou. Guernsey Society of Natural Science & Local
Research report & transactions 1902 4: 1-14.

KENT, D. H. (1992). List of vascular plants of the British Isles. B.S.B.1., London.

LE SUEUR, F. (1982). The Jersey herbarium of Frére Louis-Arséne. Watsonia 14: 167-176.

LE SUEUR, F. (1984). Flora of Jersey. Société Jersiaise, Jersey.

MCCLINTOCK, D. (1975). The wild flowers of Guernsey. Collins, London.

MARQUAND, E. D. (1892). The flora of Guernsey. Guernsey Society of Natural Science & Local Research
report & transactions 189] 2: 129-151.

MARSDEN, M. H. (1995). A new check list of flowering plants and ferns wild on Sark and its islets. Société
Guernesiaise report & transactions 1994 23: 754-783.

ROGERS, F. A. & ROGERS, W. M. (1898). On the Rubi and Rosae of the Channel Islands. Journal of botany
63: 13-15.

ROGERS, W. M. (1899). The Rubi of Guernsey and Sark. Guernsey Society of Natural Science & Local
Research report & transactions 1898 3: 296-297.

ROGERS, W. M. (1900). Handbook of the British Rubi. Duckworth, London.

SINEL, J. (1912). The geology of Jersey, with special reference to its stratigraphy and relation to the
continental coast. J. T. Bigwood, Jersey.

SNOW, B. & SNow, D. (1988). Birds and berries. A study of an ecological interaction. Poyser, Calton.

VANNEROM, H. (1998). Wijzigingen aan de Rubus-inventaris (Sect. Corylifolii Lindl. en Sect. Rubus) in de
derde editie van de ‘Flora van Belgie’. Dumortiera 70-71: 45-49.

WATSON, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland. Cambridge University Press,
Cambridge.

(Accepted June 2000)

w

1
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Watsonia 23: 437-442 (2001) 437

Conservation of Britain’s biodiversity: Rubus dasycoccus
(Rosaceae), Thick-berried Bramble

R. D. RANDALL and T. C. G. RICH

Dept. of Biodiversity and Systematic Biology, National Museum & Gallery, Cardiff CF 10 3NP

ABSTRACT

Rubus dasycoccus W. C. R. Watson is a rare British endemic species. Field and herbarium studies indicate at
least 14 sites in three 10-km squares in v.cc. 34, 35 and 41, and many populations have fewer than ten plants.
It is a perennial which reproduces by seed and appears to colonise new habitats readily. It occurs in a broad
range of habitats and soils, but probably grows best in marginal habitats such as woodland edge and unkempt
hedgerows on deeper brown earths. The main potential threats appear to be from inappropriate forestry and
regular mowing of hedgerows or road verges, but its scattered distribution with broad ecological requirements
indicate few significant threats to its long-term survival.

KEYWORDS: Endemic, rare species, England, Wales.

INTRODUCTION

Rubus dasycoccus W. C. R. Watson, Thick-berried Bramble (from the Latin name dasy - thick,
coccus - berry), is a rare British endemic species with a very localised distribution centred on
Tintern and Trelleck in v.c. 35 Monmouth, with out-lying colonies at St Briavel’s and Bigsweir in
v.c. 34 West Gloucester and Rudry, Rhydygwern and Llanfedw in v.c. 41 Glamorgan. Along with
a number of other rare brambles, it has been included in the 3rd edition of the Vascular Plant Red
Data Book (Wigginton 1999), but as yet little is known about its detailed distribution or ecology.
In 1998 the information available about it was collated and combined with a field survey to
establish its current status and determine its needs for conservation. A summary of the work is
given below; full details are given in Randall & Rich (1999).

In many respects R. dasycoccus is similar to R. trelleckensis Edees & Newton, Trelleck Bramble
which grows in the same area (Randall & Rich 2000), but it has broader ecological tolerances, a
wider distribution and would appear to be less threatened.

TAXONOMY

Rubus dasycoccus would probably not have received a name if it had not been distributed in the
Set of British Rubi as supposedly representative of R. amplificatus Lees, a more widespread and
superficially similar species. It was first collected near Tintern by A. Ley in 1891 and distributed
through the Botanical Exchange Club as “R. montanus Wirtgen?”, where W. O. Focke suggested it
might be a “var. of R. amplificatus Lees?” (Rogers 1892). Material collected in 1892 was then
distributed in 1893 as No. 33 of the Set of British Rubi, labelled “R. macrophyllus Weihe & Nees
var. amplificatus Lees”. Watson (1931) pointed out the characters which distinguished the Trelleck
plant from R. amplificatus Lees and described it as R. lasiocarpus. Unfortunately this name was
already in use and he was forced to allocate it a new name, R. dasycoccus (Watson 1933).

R. dasycoccus 1s a distinct species and fits fairly well into its current position in Series Sylvatici,
but there are a number of other species in Series Sylvatici and Series Rhamnifolii which resemble it
to some degree. A guide to superficially similar species in South Wales and adjacent counties
(v.cc. 34-36, 41-43) is given by Randall & Rich (1999). Distinctive characters of R. dasycoccus
are the obovate leaflets with long cuspidate, often curved tip, cylindrical panicle, long patent
sepals, rather pale pink petals, white filaments, green styles and the densely hairy carpels.

No hybrids of R. dasycoccus are currently known, but in shaded habitats the fruits of
R. dasycoccus are often poorly formed so plants could be easily confused with hybrids.

438 R. D. RANDALL AND T. C. G. RICH

DISTRIBUTION

Locality and habitat information was abstracted from the literature, and from herbarium sheets at
the Natural History Museum (BM) and the National Museum of Wales (NMW) which between
them include much Botanical Exchange Club material and the herbaria of W. M. Rogers, H. J.
Riddelsdell and E. S. Edees (cf. Sewell et al. 2000). Because R. dasycoccus has been confused
with R. amplificatus Lees, R. albionis W. C. R. Watson, R. questieri Lef. & P. J. Mull. and
probably other species, earlier records cannot be accepted without voucher specimens.

HISTORIC LOCALITIES

V.C. 34, WEST GLOUCESTER
Brockweir, near Tintern (BM). Bigsweir, 1894, Herb. W. A. Shoolbred, det. B. A. Miles (Newton
1986; specimen not located at BM or in herb. Shoolbred at NWW).

V.C. 35, MONMOUTH

Trelleck (BM). Beacon Hill, Trelleck (NMW). Trelleck to Tintern (BM, NMW),. Trelleck to
Tintern (BM). Hedgerow between Tintern and Trelleck (NMW). Trelleck Hill, near Tintern
(“Set No. 33”, BM, NMW). Bottom of Trelleck Hill, Tintern (Shade-grown specimen, “Set No.
33”, BM). Hedgerow, Catbrook Lane, near Tintern (NMW). Hedgerow, lower end of Catbrook
Lane, near Tintern (BM). Catbrook road, near Tintern (BM). Open space in Wye Wood, Mon.
(NMW). Wooded hill, Barbadoes hill, Tintern (NMW). Near Tintern (BM, NMW).

V.C. 41, GLAMORGAN
Forestry track, Coed Cefn-pwll-du (NMW).

FIELD SURVEY 1998

The historic records indicate that R. dasycoccus is widely distributed in the Tintern and Trelleck
area, and these and other suitable sites were consequently checked during the field survey. The
survey was carried out between mid-July and early-September 1998, the best time being mid-July
to early-August when plants were flowering.

When estimating population sizes of Rubus dasycoccus it was not possible to give exact counts
as plants spread vegetatively and were often densely entangled. For linear populations it was
assumed that if the plant was found in two consecutive lengths of 2 m that one plant was found in
each; in some cases this may result in an underestimate of the population size but it is unlikely that
a serious overestimate has been made. Where the plant was found to colonize larger areas, as at
Upper Hales Wood, one plant per 2 m x 2 m was assumed; this is also likely to be an
underestimate but when a blanket cover occurs it is impossible to be certain where one plant ends
and another begins. Populations were estimated using the following log scale where exact counts
could not be made: A = 1-9 plants, B = 10-99 plants, C = 100-999 plants and D = 1000 or more
plants. It should be stressed that all counts are crude estimates and should only be used to indicate
the relative sizes of the populations.

V.C. 34, WEST GLOUCESTER
The Nedge, St Briavel’s (SO554039), one non-flowering plant was found on the path through
woodland (A).

V.C. 35, MONMOUTH

Hale Wood; plants occurred in four areas of this wood which is conifer or mixed plantation at
different stages of maturity. At Upper Hale Wood (SO512012), a population in excess of 2500
plants was found dominating the ground flora of the Picea plantation (D). In another section of
the Plantation (SO517017) on a gentle slope near “The Oaks”, fewer than ten plants were seen
scattered along the forestry tracks (A). At Bardadoes Hill (SO522007) more than 100 plants
were present along track sides (C). On the minor road between Yew Tree House, Whitelye
(SO513015) and Botany Bay (SO525019) several patches occurred scattered in hedges and
along verges and walls (B). Botany Bay to Coed Beddick (SO525021 to SO526019), about 30

CONSERVATION OF RUBUS DASYCOCCUS 439

Vt tpl
a

3 a

FIGURE |. Current distribution of Rubus dasycoccus (¢).

scattered plants were counted along a 300 m section of track through the plantation (B).
Catbrook Lane (SO518025 to SO519024), scattered along 200 m of the public bridleway
through the plantation off Catbrook Lane (B). Bargain Wood (SO520033), three clumps were
found at the bottom of the slope on level ground in typical heathland community regenerating
following tree felling and replanting (A). Roadside, Cicelyford to Cleddon (SO504038 to
$0513038), scattered colonies were found along the roadside and in an adjacent forestry track
(B). Parkhurst Rocks (SO501031), one small colony of about five patches was seen by the
footpath through the ancient Quercus petraea woodland (Park House Wood S.S.S.I.) (A).
Beacon Hill, Trelleck (SO512054), scattered patches occurred along the tracks and in recently
cleared plantations (B). Beacon Hill - North Slope (SO513057), abundant on the lower forestry
tracks but rare or absent from the upper heathy areas (B). Vicar’s Allotment, Trelleck
(SO512060), thriving populations occurred adjacent to the footpath on the open heathland
which until recently has been forestry plantation (B). Trelleck Hill near New House
(SO504072), one young, non-flowering plant was found along c. 200 m of track through a
dense plantation (A).

V.C. 41, GLAMORGAN

Llyn Hir/Coed Cefn-pwll-du (ST2086, ST2087, ST2187), up to 100 plants were present along
rides and track sides through the plantation (B). Morphologically the plants differed from those
in the Wye Valley with the leaflets here being mostly quite long and narrow and the stems and
petioles generally more hairy, and with variable development of felting under the leaves. Some
plants showed the typical broad-based, slightly curved reddish prickles but others had narrower
straighter prickles and a darker stem. Cefn Mably Woods (ST225842), a vigorous colony was
found along about 100 m of the drive to Cefn Mably Hospital (A), exhibiting the same variation
as noted for Llyn Hir above. Rudry Common (ST192868), four bushes were found towards the
top of the common where the bracken was not dense (A).

440 R. D. RANDALL AND T. C. G. RICH

OTHER LOCATIONS SEARCHED

A number of other locations were searched without success including v.c. 34 Plantation opposite
Bearse Farm, St Briavel’s (SO573052), Hudnalls, St Briavel’s (SO5403) and St Briavel’s
Common (SO5402), v.c. 35 Church Hill Common, Penallt, (SO5210) and New Wood, Fedw
(ST4998 NE, ST5098 NW), and v.c. 41 Michaelston-y-Fedw (ST2484), Pensylvania (ST2585) and
Pen-y-lan (ST2584).

Rubus dasycoccus has thus been found in at least 14 sites in three 10-km squares in v.cc. 34, 35
and 41, and it may be more widespread. It was refound in all the historic localities with the
exception of the West Gloucester sites whose exact location is not now known (and where it may
still be present), and was found in several new localities. Many of the populations are small with
fewer than ten plants, but some are also quite large. The current distribution map is given in Fig. 1.
It appears that R. dasycoccus is a significant, if uncommon, element of the “Archenfield” regional
flora (cf. Newton 1980).

ERRONEOUS AND UNCONFIRMED RECORDS

Watson (1931) suggested that Ley had collected R. dasycoccus at Aconbury Wood, v.c. 36
Hereford, in 1888 but this has not been confirmed and is not repeated in his subsequent handbook
(Watson 1958). Watson (1958) listed R. dasycoccus for v.c. 46 Cardigan but the source has not
been traced and an error is suspected.

Watson (1958) stated that it was recorded from Vierset-Barse in Belgium under the name
R. dumnoniensis var. amplificatus (Lees) Sudre, presumably in Sudre’s account of Belgian
brambles (Sudre 1928-1929). There is a record for R. pyramidalis Kaltenb. subsp. amplificatus
Lees in Legrain (1958), where he mentions it recorded for Vierset-Barse. This, no doubt, refers to
the same plant, since in this work R. dumnoniensis is also relegated to a subspecies of
R. pyramidalis. An error has been assumed.

ECOLOGY

LIFE CYCLE

Rubus dasycoccus is, like all blackberries, a more or less evergreen perennial with some leaves
often remaining until the new year. Panicles arise from buds in the axils of the previous year’s
leaves as in other species and vegetative branches sometimes grow from the axils of the stronger
stems in the first season. The plants are fairly bushy and so it seems likely that individual stems
sometimes live longer than the usual two years.

The flowering time is from mid-July to late August (exceptionally into September). Fruits were
beginning to ripen at the end of August 1998 while the last flowers were still opening, but on many
plants the seed set was rather uneven; whether this was due to environmental or other factors is not
known.

Dispersal of seed by birds and mammals probably occurs, the normal mode of dispersal for
brambles. Vegetative spread by adventitious roots from the ends of the arching stems was noted
but only in a few places did this result in extensive colonies developing, as at Upper Hale Wood
where it has become the dominant ground cover. The occurrence of single vegetative plants in new
sites suggests it has the potential to colonise further areas where the environment is suitable.

HABITATS, VEGETATION AND SOILS
Rubus dasycoccus was found in a broad range of habitats, including woodland, conifer plantations,
woodland rides and tracks, hedges, heathland, roadside banks and verges. An attempt was made to
relate the vegetation types of some of the sites to the National Vegetation Classification (Rodwell
et al.1991 et seq.), but its occurrence in recently disturbed plantations, edge habitats and road sides
has made this quite difficult (Table 1). The original vegetation types of the sites it now inhabits
may have been W16 Quercus - Betula - Deschampsia woodland and W10 Quercus - Pteridium -
Rubus community), and the H8e Calluna - Ulex gallii heath.

Marginal habitats like woodland edge and unkempt hedgerows probably had the best-grown
plants. The least: healthy plants occurred in the dry, exposed areas on the more acidic soils, and
plants at Bargain Wood were suffering from Felt disease, which may indicate that conditions were

CONSERVATION OF RUBUS DASYCOCCUS 44]

TABLE 1. PROBABLE NATIONAL VEGETATION CLASSIFICATION TYPES AT SITES OF

RUBUS DASYCOCCUS
Site National Vegetation Classification types
Botany Bay W25 Rubus - Pteridium scrub in hedge over-topped by Salix caprea and S. cinerea,

woodland ride edges in W14 Fagus - Rubus woodland, W8 Fraxinus-Acer-
Mercurialis woodland hedge.

Bargain Wood H8e Calluna - Ulex gallii heath, Vaccinium sub-community developing in 5-year old
cleared and replanted conifer plantation.

Cicelyford to Cleddon, W23 Ulex-Rubus scrub grading into W16 Quercus - Betula - Deschampsia woodland

roadside along roadside wall.

Beacon Hill Open H8e Calluna - Ulex gallii heath, Vaccinium sub-community in recently cleared
conifer plantation, and abundant in W16 Quercus - Betula - Deschampsia
woodland on disturbed edge of possibly original W10 Quercus - Pteridium -
Rubus woodland.

Vicars Allotment H8e Calluna - Ulex gallii heath, Vaccinium sub-community developing in 5-year old
cleared and replanted conifer plantation.

Llyn Hir W24 Rubus - Holcus underscrub and W25 Rubus - Pteridium underscrub.

Cefn Mably Woods W24 Rubus - Holcus underscrub.

unfavourable. Plants growing under sunny but more sheltered and moister conditions were
generally vigorous but a reasonable amount of shade was tolerated at Upper Hale Wood and
Parkhurst Rocks.

It is distributed on a range of soils, tolerating poor, dry heath soils but growing best on deeper
brown earths (Table 2). With the exception of Llyn Hir where plants grow on Dolomitic limestone,
all colonies grew in situations where the underlying geology involved sandstones.

CONSERVATION REQUIREMENTS

Rubus dasycoccus is certainly a rare plant. It is not protected under Schedule 8 of the Wildlife and
Countryside Act 1981, but is listed in the Vascular Plant Red Data Book (Wigginton 1999) which
should draw attention to its rarity, and this should be sufficient to ensure its survival. It occurs on
two S.S.S.I.s (at Parkhurst Rocks and Cleddon Bog), though populations at both sites are small. If
S.S.S.I.s need to be designated for this species, the largest site at Hale Wood and the type locality
at Trelleck Hill are the best candidates.

Given its widely scattered distribution with broad ecological requirements in a range of habitats,
there appear to be relatively few significant immediate threats to its survival overall. The main
potential threats to individual colonies appear to be change of land-use from forestry to farmland
or recreation, weed killing during forestry operations, long periods of deep shade (e.g. colonisation

TABLE 2. SOILS COLLECTED FROM THE RUBUS DASYCOCCUS ROOTING ZONES

Site Soils

Upper Hale Wood, Plantation: a gritty brown organic soil, pH 3-1. Recently disturbed ground: a gritty
Whitelye dark-brown soil some organic matter and pebbles, pH 7-5.

Botany Bay A fine, light-brown sandy soil, pH 7-0.

Bargain Wood A fine grey-brown heathy soil with lots of organic matter, pH 4-0.

Beacon Hill Top: A fine brown heathy soil with organic matter, pH 5-0. North side: A gritty light

brown soil, pH 5-0. Track edge: A red-brown, gritty stony soil, pH 7-9.

Vicars Allotment A blackish organic soil with large quartz sand grains, pH 3-4.

Llyn Hir A reddish brown earth with many limestone fragments on track side, pHs 7:5 and 7-7.

Cefn Mably Woods A red brown earth, pH 6-8.

pH was measured in a 50:50 deionised water: soil slurry.

442 R. D. RANDALL AND T. C. G. RICH

by Rhododendron, lack of clearance of conifers), and introduction of regular close cropping of
paths, hedgerows and roadside scrub, or destruction of hedgerows due to road-widening or other
schemes.

The most favourable forestry management would be a regime similar to that at Beacon Hill,
where trees have been cropped in rotation with adjacent areas being felled several years apart
allowing colonisation into recently felled areas. For woodland that is being maintained as an
amenity or a nature reserve, natural regeneration after cropping followed by selective thinning to
produce small clearings and glades would be beneficial. For hedgerows and other marginal
situations annual trimming should be avoided as it is detrimental as two or three years are required
for stems to reach the flowering and fruiting stage. Where populations are significant, footpaths
could be cleared on alternate sides in alternate years, or perhaps on both sides once every 3-4
years.

A small number of fruits from four plants at Beacon Hill (2 populations) and Whitelye have
been deposited in the Millennium Seed Bank at the Royal Botanic Gardens, Wakehurst Place for
long-term storage; larger samples from a wider range of plants would be worthwhile collecting. No
plants are known to be held in cultivation.

ACKNOWLEDGMENTS

This work was jointly funded by Countryside Council for Wales and National Museum & Gallery,
Cardiff as part of a joint project into conservation and ecology of critical species. The Royal
Botanic Gardens, Kew, contributed towards collecting blackberries for the Millennium Seed Bank.

We would like to thank Andy Jones and Lindi Rich of CCW for their help, and Alan Orange and
Jessica Carey for help with field work. We would like to thank the Keeper of BM for access to the
herbarium.

REFERENCES

EDEES, E. S. & NEWTON, A. (1988). Brambles of the British Isles. Ray Society, London.

LEGRAIN, J. (1958). Rubus L., in ROBYNS, W., ed. Flore Générale de Belgique 3: 69. Bruxelles.

NEWTON, A. (1980). Progress in British Rubus studies. Watsonia 13: 35-40.

NEWTON, A. (1986). Rubus L., in HOLLAND, S. C. ed. Supplement to the Flora of Gloucestershire. Grenfell
Publications, Bristol.

RANDALL, R. D. & RICH, T. C. G. (1999). Distribution and conservation of Rubus dasycoccus, Thick-berried
Bramble. Countryside Council for Wales and National Museum of Wales, Cardiff.

RANDALL, R. D. & RICH, T. C. G. (2000). Conservation of Britain’s biodiversity: Rubus trelleckensis
(Rosaceae), Trelleck Bramble. Watsonia 23: 317-322.

ROGERS, W. M. (1892). R. montanus Wirtg.? Rep. Botl. Exch. Club Br. Isles 1: 329.

ROGERS, W. M. (1900). Handbook of British Rubi. London.

SEWELL, R. M., EVANS, M., RANDALL, R. D. & RICH, T. C. G. (2000). Catalogue of Rubus specimens in the
Welsh National Herbarium (NMW), National Museum & Gallery, Cardiff. National Museums &
Galleries of Wales, Cardiff.

WATSON, W. C. R. (1931). Bramble Notes, 1930. Rep. Botl. Exch. Club Br. Isles 9: 434-435.

WATSON, W.C. R. (1933). R. dasycoccus nom. nov. Rep. Botl. Exch. Club Br. Isles 10: 21.

WATSON, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland. Cambridge University Press,
Cambridge.

WIGGINTON, M. J. (1999). British Red Data Books: Vascular Plants, ed. 3. J.N.C.C., Peterborough.

(Accepted June 2000)

Watsonia 23: 443-453 (2001) 443

Notes

NITELLA GRACILIS (SMITH) AGARDH, AN ELUSIVE CHAROPHYTE NEW TO
CARDIGANSHIRE (V.C. 46)

The Slender Stonewort (Nitella gracilis) is, globally, a very widespread species, recorded from
every continent except Antarctica. At least in Europe, however, it is very sparsely scattered, with
only a few localities recorded in each country. This is also true for Britain and Ireland where it has
been recorded from just a few, widely scattered sites in Sussex, Essex, Cornwall, Gwynedd,
Cumbria, Ayrshire, Sutherland and Co. Wicklow, and not in some of these for over 50 years
(Stewart in litt.) It is consequently listed in the Stoneworts British and Irish Red Data Book
(Stewart & Church 1992), and as a Priority Species in the UK Biodiversity Action Plan (UKBAP;
Anon 1995).

The picture has, however, been complicated by a number of misidentifications, particularly for
the similar — but more robust — N. mucronata (A. Braun) Miq. var. gracillima J. Groves & Bull.-
Webst.; records from Hampshire, Greater London, Norfolk, Perth, and Co. Dublin are now thought
to be doubtful or in error. This has confused the view of these species’ ecology, since Nitella
mucronata var. gracillima grows in mesotrophic to eutrophic waters whilst N. gracilis appears to
be restricted to nutrient-poor water bodies, particularly in the uplands (cf. Stewart & Church
£9):

Most records of Nitella gracilis have been from chance captures by grapnels or rare jetsam
specimens after stormy weather and many of its sites have only yielded one or two specimens —
although this could be due to collecting difficulties, seasonal growth or transient colonisation.
Even so, there are good indications that the species has also disappeared from some sites (Stewart
in litt.) For all these reasons there is little published information about the ecology of N. gracilis in
Britain and Ireland and this short note aims to add to the current understanding, and to encourage
further surveys of likely habitats.

LLYN GYNON (V.C. 46)

At an altitude of 425 m in the Elenydd uplands of Ceredigion and straddling two hectads (SN76—
86), Llyn Gynon is probably the richest lake in Cards (v.c. 46). Floating Water-plantain (Luronium
natans) 1s locally abundant (Monteith, 1995), as has been Spring Quillwort (/soetes echinospora)
in the past, and there is a small colony of Six-stamened Waterwort (Elatine hexandra) towards the
north-western inflow. In 1964 Brian Seddon also found Pillwort (Pilularia globulifera) here at a
depth of about 45-60 cm, by grapnelling from a boat, but it has not been refound since, despite
diligent grapnel hauls from shore and boat, searches by wading, and searches of driftline debris (A.
Chater, pers. comm.).

Each of these species is nationally scarce; Pilularia and Luronium are priorities in the UKBAP
and placed in the IUCN category “Vulnerable” in Europe and Luronium has statutory protection in
national and international law (Schedule 8 of the Wildlife and Countryside Act; EC Habitats and
Species Directive, Annexes IIb & IIv). As a consequence, the Countryside Council for Wales
undertook a sample survey of the western side of the lake, on 25 September 1998, to map the
distribution of important plant species and to search for deep-water colonies of Pilularia. The
methodology followed from earlier, largely unpublished surveys for scarce aquatic macrophytes
(Lomas et al. 1998; Scott 1996 & 1998), and included drysuits, snorkels and sub-aqua equipment.

The survey gave a detailed description of the distribution of Luronium and Elatine and of their
associated species, but did not find any Pilularia or Isoetes echinospora. Between 1964 and 1989
I. echinospora was found abundantly in Llyn Gynon and it seems from these and other
observations (A. Chater, pers. comm.) that populations of the species may be subject to sudden and
rather mysterious fluctuations. Where Isoetes lacustris dominance declined (at about 2-5 m depth)
and with occasional plants of Myriophyllum alternifolium and Callitriche hamulata, vegetative L.
natans (forma submersum) became the most frequent deep-water species (Fig. 1). Individual

444

NOTES Watsonia 23 (2001)

KEY TO SPECIES

Nitella gracilis ly
\Z Juncus bulbosus

Isoetes lacustris

a Fontinalis antipyretica
5 va Ae Littorella uniflora
Luronium natans ue oy Lobelia dortmanna
ole ees Subularia aquatica
Batrachospermum sp.

L. natans (shallow water)

Myriophyllum alternifolium f Porifera sp. (indet.)

“Ke, Ranunculus flammula

@ ss Stone

Callitriche hamulata

FIGURE 1. Llyn Gynon submerged vegetation. Diagrammatic section (western shore).

NOTES Watsonia 23 (2001) 445

rosettes of L. natans with leaves 20-30 cm in length were scattered in deep, soft silt at a frequency
of 4-10 plants per square metre.

It was this association which yielded occasional plants of a Nitella sp., subsequently determined
by Arthur Chater & N.F.S as N. gracilis. Five plants were noted by R.A.J. in a sample of
approximately 30 m of “fringe” vegetation along the western shore of Llyn Gynon, at c. 3-5 m
depth. Individual plants were c. 30-40 cm across, domed in profile and, compared to the
associated species, notably free of silt deposition. Of the branches from two plants collected for
identification, both turned out to have well-formed, orange-coloured oogonia, and both broke up
substantially within 24 hours of collection, although the tougher, peat-stained, lower branches on
one plant seemed more resilient.

The location of N. gracilis in Llyn Gynon at 3-5 m depth on the western shore is distinctly
sheltered, but it is not possible to generalise from this without further survey. Similarly, the
fertility of plants so late in the year, their stained lower branches and absence of silt cover could all
represent seasonal growth patterns (even, perhaps, perennation) but confirmation of this would
require at least another visit. Other sources (e.g. John et al. 1982) have noted the value of
charophytes as invertebrate habitat, and N. gracilis might be locally significant in some
oligotrophic, acidic lakes for this reason. What can be concluded with confidence, however, is that
surface-based methods of sampling (such as grapnel, grabs and bathyscopes) stand a low chance of
detecting delicate species growing at such low density and at this depth and distance from the
shore (see also Wade & Bowles 1981). This might also explain why attempts to relocate the plant
by N.F.S. at its two other Welsh sites in Snowdonia (Llyn Dwythwch and Llynau Mymbyr) in
October 1998 by shore grapnel surveys were unsuccessful (Stewart 1999). Nitella gracilis is likely
to be an overlooked species of nutrient-poor, acidic lakes in Britain and Ireland and it will be
located with certainty only by the use of trained divers.

ACKNOWLEDGMENTS

We should like to thank Mike Bailey, Alan Hale, Duncan Smith and Gabrielle Wyn for their help
with the fieldwork, and Arthur Chater for his overall contribution and his comments on an earlier
draft of this note.

REFERENCES

ANON (1995). Biodiversity: the UK Steering Group Report. Volume 2: Action plans. H.M.S.O., London.

JOHN, D. M. , CHAMP, W. S. T. & Moore, J. A. (1982). The changing status of Characeae in four marl lakes
in the Irish Midlands. Journal of life sciences of the Royal Dublin Society 4: 47-71.

LOMAS, E., TEEARU, T. & ROWLANDS, A. (1998). Survey to determine the status of Luronium natans (L.) Raf.
in a sample of six mid-Wales lakes. Unpublished report to the Countryside Council for Wales, Bangor.

MONTEITH, D., ed. (1995). Integrated classification and assessment of lakes: Phase II — Final report.
Unpublished report to the Countryside Council for Wales, Bangor.

SCOTT, S. (1996). Action for Pillwort, Pilularia globulifera. Unpublished report to Scottish Natural Heritage /
Plantlife, London.

ScoTT, S. (1998). Partnership project for Action Plan work on Pillwort, Pilularia globulifera, 1997-2000.
Annual Report 1997-8. Unpublished report to Scottish Natural Heritage / Plantlife, London.

STEWART, N. F. & CHURCH, J. M. (1992). Red data books of Britain & Ireland: Stoneworts. Joint Nature
Conservation Committee, Peterborough.

STEWART, N. F. (1999). Welsh Stoneworts Survey 1998. Unpublished report to the Countryside Council for
Wales, Aberystwyth.

WADE, P. M. & BOWLES, F. (1981). A comparison of the efficiency of freshwater macrophyte surveys carried
out from underwater with those from the shore or a boat. Progress in underwater science 6: 7-11.

R. A. JONES
Countryside Council for Wales, Plas Gogerddan, Aberystwyth, Ceredigion SY23 3EE

N. F. STEWART
Kingfishers, Cholwell, Posbury, Crediton, Devon, EX17 3QF

446 NOTES Watsonia 23 (2001)

COLONISATION BY COCHLEARIA DANICA L. ALONG TRUNK ROADS IN CENTRAL
SCOTLAND FROM 1996 TO 2000

The spread of halophytes along motorways and trunk roads in England and Wales in the 1980s and
early 1990s was widely observed (Scott & Davison 1985; Scott 1985; Leach & Rich 1989; Leach
1994). The factors causing and influencing this spread have been much debated (Leach 1990: Scott
1990; Roper 1994), but it is generally held that coastal roads were colonised first and the
halophytes were then dispersed by vehicular traffic along roadside habitats made suitable by road-
salting (Scott & Davison 1985). For Cochlearia danica (Danish scurvygrass), rates of advance of
10-20 km yr" were deduced by Leach (1994) from observations on first colonisation.

In Scotland C. danica became established on trunk roads much later than in England. The first
record according to Leach (1994) was made in 1993 when plants were observed on the A74 in
Dumfriesshire (v.c. 72). By this year much of the English motorway system had been colonised,
and the species was absent inland in only six English vice-counties (Wight, E. Kent, E. Norfolk,
Salop, Derbys, S.E. Yorks) (Leach 1994). As Cochlearia danica was frequent along the M6
through Cumberland (v.c. 70) in the early 1990s, it is very likely that the Dumfriesshire colony
resulted from northward spread, rather than being a fresh colonisation from the nearby Solway
coast. Similarly, the colonies of C. danica observed on the Al in Berwickshire (v.c. 81) in 1993
(Braithwaite 1997) probably arose from dispersal northwards from colonies along the Al in
Northumberland (v.c.c. 67 and 68).

From the mid 1990s onwards, I have made 4—6 journeys each spring between N.W. England and
N.E. Scotland using the M74/A74/M73/A80/M80/M9/A9/A90 roads up to Stonehaven (Fig. 1). In
1995 and 1996 C. danica became increasingly obvious on the M74 and A74, and I reported what
appeared to be its northern limit in 1997 near Hamilton (Welch & Welch 1998), there being a

Stonehaven

Laurencekirk @."
¢

50 km

oe
Dunblane ve

t
Stirling je

e?®
ef
e
e

FIGURE 1. Map of Central Scotland, showing the trunk roads monitored (dashed) and other nearby trunk roads
(dotted).

NOTES Watsonia 23 (2001)

447

sizeable patch on the M74 central reservation by the Atlas Works close to Junction 9 (Table 1).
From 1997 onwards I have carefully looked for Cochlearia along the whole route north of
Junction 9 of the M74, but driving at an average speed of 60-70 mph it is inevitable that small
plants and non-flowering colonies will have been missed. In 1997, 1999 and 2000 I also drove
along the M90 between Perth and the Forth Bridge (Fig. 1), and in 2000 I made a single traverse of
the M876/M$9 triangle east of Stirling Services and the M8 from Newbridge to Newhouse.

TABLE 1. LOCATIONS AND SIZE OF COLONIES OF COCHLEARIA

DANICA ON CENTRAL SCOTTISH TRUNK ROADS

Vice

Road _ Place Grid square county 1997 1998 [O99 2000
A90__Laurencekirk NO77 91 4 ptchs” - ¢. 350 plis” 51 plts

in60m* in57m*
A90 Dundee Ring Road NO33 90 X X X 1 patch
A90 1kmELongforgan NO33 89 Xx Ks X 1 patch
A990 1kmWLongforgan NO22 89 x 1 patch 1 patch 1 patch

(20m long) (35m long)
A90 _ nr Inchture Jn NO22 89 X x x 2 ptchs
A90 _— nr Inchmichael Jn NO22 89 X X 1 patch 3 ptchs
A90 nr Leetown Jn NO22 89 % x X 1 patch
A90__ nr Inchyra Jn NO12 89 X few ptchs x X
M90 around J9 NO11 88 x . sev. ptchs _c. 250 m
M90 1kmNJn4 NT19 85 x - x few ptchs”
M90 lkmS Jn3 NT18 85 x x 1 patch
A90 ‘just N of Forth Bridge NT18 85 X - sev. ptchs _ sev. ptchs
A9 nr Findo Gask Jn NOO1 88 X X few ptchs few ptchs
A9 nr B9141 Jn NN91 88 Xx X few ptchs few ptchs
A9 Blackford bypass NN90 87 X x X 2 ptchs
A9 3 km W Blackford NN80 87 x X 1 patch 1 patch

(20 m long)
A9 5 km W Blackford NN80 87 x x X 2 ptchs
M9 =- Jn 11-Jn10 NS79 87 m8 X few ptchs occas.
M9 ~—=- Jn10—Jn9 NS78,79 86 x x frequent frequent
M9 = Jn9-Jn7 NS88 86 - - - occas.
M876 Jn8—Jn5 NS88 86 . - - occas.
M876 Jn7—W of Jn3 NS88 86 - - - occas.
M876 _ near Jn3 NS98 86 - . . few ptchs
M80 = _-Jn9-Jn5 NS88 86 x X frequent frequent
M/A80 Jn5—Jn3 NS77 86 X few ptchs _—_ frequent frequent
M73 = Jn3-Jn1 NS76,77 ie, x frequent occas. occas.
M8 near Jn2 NN17 83 . - - frequent
M8 ~~ W of Jn2-Jn3A NN06,07 84 . . - frequent
M8 = Jn3A—Harthill Services NS96 84 - - occas.
M74 Jn4-Jn5 NS66;75,76 77 X Xx few ptchs _—_ few ptchs
M74 Jn5-Jn8 NS74,75 Woh, x few ptchs occas. frequent
M74 Jn8—Jn9 NS74 Tal Xx few ptchs _ occas. occas.
M74 — around Jn9 NS74 77 few ptchs occas. frequent frequent
Y= on verge or hard shoulder; other colonies are on the central reservation; - = not searched; x = not seen;

Jn = Junction; ptchs = patches; frequent = c. 10% of central reservation occupied; occas. = c. 1% of central
reservation occupied; sev. patches = >5 patches observed; few patches = 2-5 patches observed.

448 NOTES Watsonia 23 (2001)

The direct route between Junction 9 of the M74 and Laurencekirk on the A90 is 225 km long,
and C. danica colonised most of this between 1996 and 2000 (Table 1). In 1998 and 1999
Cochlearia was much more frequent along the southernmost 80 km of this route up to Dunblane
than on the next 80 km from Dunblane to Dundee, and its abundance on the M73 in 1998
suggested it had been overlooked there in 1997. However, Macpherson & Macpherson (1999),
recording C. danica along Lanarkshire (v.c. 77) roads on a 1-km square basis, also made many
new records in 1998, three of their total of 17 records being from the M73 and ten from the M74.

Important factors affecting the presence and visibility of C. danica along roads are the laying of
gravel, the spraying of herbicides, and competition from dense grass swards. Fresh gravel laid
along the M73 central reservation in 1996 or 1997 probably contributed to the abundance of
Cochlearia there in 1998, whereas its apparent depletion in 1999 (Table 1) may have been related
to grass and coarse plants spreading. Herbicide treatment along most of the route across central
Scotland up to Dundee in spring 1999, which became obvious in mid May, much reduced grasses
and weeds, but had only short-term impact on C. danica judging from its distribution and
frequency in spring 2000.

It has been suggested that C. danica is actually introduced to inland roads in the gravel and hard
core used in their construction (Coombe 1994; Lansdown & Pankhurst 1995), and this is a possible
explanation for the phenomenal increase of the species in central Scotland. A pinky-red gravel has
been placed in the central reservation on many sections of the M73, the A80, the M80, the M876,
the M9 and the M90, and possibly this gravel contained Cochlearia seeds. However, I doubt this
explanation because there are long sections of red gravel with no Cochlearia plants visible
especially between Stirling and Dundee (Fig. 1). Also the colony on the M90 near the Inchyra
junction (Table 1) was lost when a dressing of red gravel was laid in summer 1998 after the
erection of a crash barrier.

The Laurencekirk colony of C. danica on the A90 at the far north of the monitored roads
perhaps did not arise from colonisation produced by vehicles sweeping seeds along the
carriageways. The four small patches observed in 1997 actually occurred on the nearside verges of
the two carriageways about 3 km apart, and could have resulted from introduction in road-making
materials besides transport on vehicles. Similarly, the colony of C. danica observed on a roadside
at Flemington (v.c. 77) in 1989 (Macpherson & Macpherson 1999) would seem not to be part of
the trunk-road invasion.

The potential rate of increase of C. danica was shown at the Laurencekirk colony between 1997
and 1999 (Table 1). Two patches of c. 400 cm’ on the southbound verge in 1997 had become c.
350 scattered plants in 1999 over a 60 m section of verge extending 0-3 m from the road kerb.
Some disturbance appears to have occurred, as some stiff red clay was exposed in spring 1999,
perhaps from skimming during grass-cutting in the previous summers. However, drain digging
during 1999 c. 4 m from the kerb, and intermittent parking, compacted the verge soil and damaged
the plants, and by spring 2000 the colony was reduced to 51 plants along a 57 m length 0-2 m
from the road kerb. Also, the two patches observed on the northbound verge could not be refound
in 1999 or 2000, nor any plants of Cochlearia in the 200 m of verge around their position.

The rate of spread of C. danica across central Scotland, calculated on the basis of strong highly
visual patches at the north ends of fairly continuous colonisation (Atlas Works on M74 in 1997,
J10 on M9 in 1999), is 40 km yr'!. This is considerably greater than Leach*s estimate of 10-20 km
yr'', and perhaps the 1997 start point was actually further north than the Atlas Works. This would
in turn mean an advance north up the A74/M74 from 1993 to 1997 at more than 20 km yr’.

Why C. danica was so slow to colonise Scottish motorways and trunk roads is puzzling. The
suggestion that less salt was being used in Scotland than northern England in the 1980s (Scott
1985; Scott & Davison 1985) is unlikely, given the climate; the success of C. danica in the last
few years indicates that roadside conditions had become very suitable by the 1990s. I believe that
for successful colonisation the quantity of seeds being dispersed has to be great, which needs both
large fruiting colonies and much traffic moving seeds along, and also the receiving habitat has to
be extensive and fairly continuous. Probably other local establishments, like the Laurencekirk
colony, have occurred unnoticed in Scotland, and have not produced enough seed for nearby
suitable habitat to be colonised before becoming extinct due to grass competition, road works,
herbicide treatment, etc. The great build-up of C. danica on the trunk roads leading into Scotland
by the early 1990s, the increased traffic density compared to the 1980s, and the greater extent of

NOTES Watsonia 23 (2001) 449

suitable habitat in the central reservation (bare gravel now being preferred to grass because
maintenance is easier), have combined to produce sufficient seed dispersal to create the present
spectacular carpets of pale pink flowers across central Scotland in spring.

ACKNOWLEDGMENTS

My wife Muriel has helped in the spotting and recording of roadside colonies. Peter Macpherson
has kindly given me details of his Lanarkshire observations.

REFERENCES

BRAITHWAITE, M. E. (1997). Maritime plants on roads in the Scottish Borders. B.S.B.J. Scottish newsletter 19:
15.

COOMBE, D. E. (1994). “Maritime” plants of roads in Cambridgeshire (v.c. 29). Nature in Cambridgeshire 36:
37-60.

LANSDOWN, R. & PANKHURST, T. (1995). Coastal gravel as a source of Cochlearia danica on inland
roadsides. BSBI news 69: 17-18.

LEACH, S, J. (1990). Cochlearia danica on inland roadsides. BSBI news 56: 20-21.

LEACH, S. J. (1994). Cochlearia danica on inland roadsides — an update. BSBI news 65: 12-13.

LEACH, S. J. & RICH, T. (1989). Scurvy-grasses take to the road. BSBI news 52: 15-16.

MACPHERSON, P. & MACPHERSON, A. C. (1999). Danish scurvygrass — another seaside-roadside plant in
Lanarkshire. B.S.B.I. Scottish newsletter 21: 11.

ROPER, P. (1994). Motorway scurvygrass. BSBI news 65: 13.

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

ScoTT, N. E. (1990). Cochlearia danica on inland roadsides — source of seed? BSBI news 56: 11-12.

ScoTT, N. E. & DAVISON, A. W. (1985). The distribution and ecology of coastal species on roadsides.
Vegetatio 62: 433-440.

WELCH, D. & WELCH, M. J. (1998). Colonisation by Cochlearia officinalis L. and other halophytes on the
Aberdeen — Montrose main road in North-east Scotland. Watsonia 22: 190-193.

D. WELCH
East Fernbank, Woodside Road, Banchory, Kincardineshire AB31 5XL

ALCHEMILLA GLAUCESCENS WALLR. IN V.C. 81 BERWICKSHIRE

Alchemilla glaucescens has its headquarters in Britain on the limestone of the Ingleborough district
of Yorkshire but has small disjunct populations in Northern Scotland and Ireland. It is a Red Data
Book species. Two populations have now been found in Berwickshire.

LAMBERTON

On 18 September 1999 M.E.B. was studying Rosa in scrub on the steep sea braes at Hilton Bay
NT9659 within the Lamberton Coast S.S.S.I. An unexpected open area was reached which carries
a limestone flora with several species of note. An interesting Alchemilla was collected in fruit,
suspected to be A. glaucescens. The site was revisited by M.E.B. with P. S. Lusby on 16 October
1999 and again by M.E.B. on 1 May 2000 when the Alchemilla was in full flower. P. S. Lusby and
Dr S. M. Walters confirmed the determination from the autumn plants, but with mild reservations.
The spring growth allowed fresh specimens to be presented to P. S. Lusby whose reservations
were dispelled.

The Alchemilla is closely associated with Sanguisorba minor subsp. minor, Primula veris and
Leontodon saxatilis. The grassland is also notable for large populations of Carlina vulgaris and
Catapodium rigidum. Other associates include Anthyllis vulneraria, Carex flacca, Leontodon
hispidus, Linum catharticum, Ononis repens and Viola hirta. The Alchemilla certainly seems to be
native, though, with the railway adjacent, introduction cannot be wholly discounted. The
population is estimated at 500 plants.

The site has gradual erosion that keeps the vegetation open to some degree. The substratum of
Hilton Bay is Carboniferous limestone and sandstone but there are also igneous porphyritic dykes
near the site (Greig 1988).

450 NOTES Watsonia 23 (2001)

Although the bay is secluded and not overgrazed there is one major threat. The very erosion that
the existence of the vegetation community depends on is a threat to the main North-South railway
nearby. British Rail (now Railtrack) have fairly recently carried out works to halt the erosion as
much as possible. Concrete structures have been erected at the edge of the beach and plastic mesh
has been dug into the slopes above. The concrete has holes in it and these have colonised
splendidly with such species as Carlina and Catapodium. However, the area with the mesh is now
covered by quite coarse vegetation and if any of the rarities were formerly present there they have
probably been lost. It should be noted that large areas of the braes are not strongly calciferous and
are covered naturally by coarse vegetation - indeed that is partly why these open communities have
not been discovered before - it is not obvious that this is the most interesting part of the braes.

Adjacent coastline was searched for the plant on 27 September 1999, 16 October 1999 and
1 May 2000 but no further colonies have been found.

CHIRNSIDE

Alchemilla glaucescens had earlier been discovered by M.E.B. in 1982 in a disused railway cutting
near Oldcastles, Chirnside, NT8558. It was described by him in ‘The Botanist in Berwickshire’,
(Braithwaite & Long 1990), as growing “on ballast” and treated as an introduction. This statement
is not accurate. The site was visited by M.E.B., P. F. Braithwaite and L. Gaskell on 6 June 1998.
The Alchemilla grows on a bank that appears to be part of the original hillside and to have been
preserved by being enclosed within the former railway fence. The cutting as such is below. The
plants are not growing on ballast and there is no evidence of ballast being used in the drainage of
that particular section of cutting. There is no reference to ballast in M.E.B.’s field notes in 1982.
M.E.B. had found interesting communities on ballast-drained railway cuttings in Roxburghshire
(Braithwaite 1976).

The Alchemilla is associated with depauperate base-rich grassland and is closely associated with
Trifolium medium. Other associates are Crepis capillaris, Fragaria vesca, Luzula multiflora,
Primula vulgaris, Succisa pratensis and Vicia sativa subsp. nigra. Linum catharticum and
Trisetum flavescens were recorded in 1982. This site must now be reconsidered. The Alchemilla is
in all probability native here also.

The Alchemilla population is healthy with about 100 plants, the determination of which has been
checked by Dr S. M. Walters in 1982, and P. S. Lusby in 1999.

The substratum is “unusually coarse-grained red sandstone” of the Upper Old Red Sandstone
(Greig 1988).

The vegetation is kept open by cattle grazing, which appears to be beneficial. Nevertheless the
site is so small that the future of the Alchemilla is most doubtful.

DISCUSSION

The locally scarce limestone species with which the Alchemilla is associated at Hilton Bay can all
be presumed to have reached the Scottish Borders by a fairly clear-cut path from the limestone
areas of Northumberland where they are more frequent (Swan 1993). Some of them have other
stations in the Scottish Borders on the Silurian and on the Old Red Sandstone where there are
lime-rich pockets. There is no reason why the Alchemilla should not have colonised by a similar
route but, if so, no ready explanation can be offered as to why it is not found between
Berwickshire and the Ingleborough area today. It seems therefore as likely that it is a chance
introduction in the distant past, perhaps by bird-borne seed from Scandinavia, as the site is very
much on a bird migration route. However, Tofieldia pusilla was found near Berwick by John Ray
in 1671 (Ray 1677), probably within two km of the Hilton Bay site, and it is possible that the
former moorland of this area held, in limestone pockets, refugia for a specialised post-glacial
community that included the Alchemilla and Tofieldia.

REFERENCES

BRAITHWAITE, M. E. (1976) A Railway Flora of Teviotdale. Hawick
BRAITHWAITE, M. E. & LONG, D. G. (1990) The Botanist in Berwickshire. The Berwickshire Naturalists Club,
Buccleuch Printers, Hawick.

NOTES Watsonia 23 (2001) 451

GREIG, D. C. (1988) Geology of the Eyemouth District. British Geological Survey, H.M.S.O. London.

Ray, J. (1677) Catalogus Plantarum Angliae (2nd ed.). London.

SWAN, G. A. (1993) Flora of Northumberland. The Natural History Society of Northumberland, Newcastle
upon Tyne.

M. E. BRAITHWAITE
Clarilaw, Hawick, Roxburghshire, TD9 SPT

GERANIUM ROBERTIANUM L. VAR. MARITIMUM (GERANIACEAE):
AN EARLIER AUTHOR

For a century and a half the authorship of this name for the prostrate and often glabrous coastal
ecotype of G. robertianum has been attributed to Babington, who first published it in the third
edition (1851) of his Manual of British botany. It has, however, been overlooked that the same
epithet was validly published for the very same plant by Cooper (1834), whose authorship has
priority.

Cooper cited only the one locality: Cockbush Common, Shoreham (W. Sussex, v.c. 13). There is
a specimen of the plant in CGE collected at Cockbush Common the previous year by G. E. Smith,
by whom and, even more, Borrer (the two leading Sussex botanists of that period) the
inexperienced Cooper was provided with most of the localised records that featured in his list — a
fact which Cooper concealed from the public, to the lasting indignation of Borrer (1851; Allen
1979). It is thus probable that the name was a manuscript one already in use by one or other of
those two. Babington may well have been apprised of it later by Borrer, who may by then have
forgotten (or wanted to forget) that it had appeared in print already.

Baker (1956) considered that the rather distinctive plant at Shoreham (which was also the type
locality of Babington’s taxon) represented just one extreme in a cline of hairiness, allowing all the
prostrate plants of shingle beaches to be grouped together under a single name. Because these have
a wide geographical distribution and he had detected in cultivation a physiological difference in
addition to a loose cluster of external characters, he felt justified in promoting the variety to
subspecific rank. In this he has been followed by subsequent authors, though it might perhaps be
questioned whether an ecotype with, as pointed out by Stace (1991), only one constant external
character really merits such an elevated rank.

REFERENCES

ALLEN, D. E. (1979). The plagiarisms of Thomas Henry Cooper. Journal of the Society for the Bibliography
of Natural History 9: 275-279.

BAKER, H. G. (1956). Geranium purpureum Vill. and G. robertianum L. in the British flora. Il. Geranium
robertianum. Watsonia 3: 270-279.

BORRER, W. (1851). Botanical memoranda. Botanical gazette 3: 98.

CoopER, T. H. (1834). The botany of the county of Sussex. Sussex Press, Lewes. (Preprint of Cooper in
HORSFIELD, T. W. (1835). The history, antiquities and topography of the county of Sussex, 2: 5-22).

STACE, C. A. (1991). New Flora of the British Isles. Cambridge University Press, Cambridge.

DD. E: ALLEN
Lesney Cottage, Middle Road, Winchester, Hampshire, SO22 5EJ

452 NOTES Watsonia 23 (2001)
EPILOBIUM x KITCHENERI McKEAN (ONAGRACEAE) IN EAST CORNWALL

The hybrid between Epilobium pedunculare A. Cunn. and E. montanum L. has hitherto been
reported only as a single plant found in West Perth (v.c. 87) in 1996, from which it was named as
E. x kitcheneri by McKean (1999). This is the sole report from the British Isles of a hybrid
involving the alien Rockery Willowherb E. pedunculare from New Zealand, which is only locally
established in the wild in the British Isles, whereas the Broad-leaved Willowherb E. montanum is a
widespread native piant that forms hybrids with numerous congeners (Stace 1997; Kitchener &
McKean 1998).

In 1996 Mary and Tony Atkinson discovered that E. pedunculare was naturalised in abundance
alongside tracks through Hancock’s Wood, a coniferous plantation on the northern side of the
Glynn Valley, East Cornwall (v.c. 2) (Murphy 1998). During 1999 we searched independently for
possible hybrid Epilobium in the vicinity of the naturalised E. pedunculare in the Glynn Valley.
These searches resulted in the discovery of two plants of E. x kitcheneri, providing the second and
third records for this hybrid and the first reports of it outside Scotland.

The first find of the hybrid in East Cornwall was made on 9 July 1999 by D.T.H., in woodland
north-east of Trago Mills, grid reference SX182648; specimen in herb. D.T.H. The single plant
was growing on the rocky edge of a track in a coniferous plantation, close to E. ciliatum Raf., E.
montanum, E. obscurum Schreb. and E. pedunculare, the last of these species being naturalised
along c. 150 m of the track edges. The hybrid had two main stems, the longest of them 28 cm tall
and stiffly erect above its procumbent base, the other stem 27 cm tall with two shorter branches,
both stem and branches being erect throughout. The specimen accords well with the Scottish type
material of E. x kitcheneri in having leaf shape midway between the orbicular leaves of E.
pedunculare and the ovate leaves of E. montanum, the influence of E. pedunculare apparent in
strongly toothed leaf margins and the leaves bronzed on the underside, and the influence of E.
montanum apparent in the presence of a shallowly four-lobed stigma.

The second find of the hybrid in East Cornwall was made a month later, at Hancock’s Wood just
over | km away (SX171649). It was found by Ian Bennallick in company with Mary and Tony
Atkinson and G.D.K. The hybrid plant was also found at the edge of a woodland track with the
same Epilobium associates as the first find. It consisted of three closely associated, separately
rooted stems, 41 cm, 35 cm and 20 cm long. These may have been separate plants or the results of
layering of one original plant which may have overwintered. Subsequent cultivation by G.D.K. has
demonstrated that overwintering can take place, with rooting from the nodes.

The north side of the Glynn Valley in East Cornwall apparently has the most extensive
naturalised colonies of E. pedunculare in the British Isles. The species extends in quantity along
more than a kilometre of forestry tracks in Hancock’s Wood and Cross Plantation. The means by
which it was originally introduced there is uncertain, although the adjoining woodlands have some
exotic plantings, including a bamboo. There is evidence that forestry operations have contributed
to its subsequent spread along tracks.

In the Glynn Valley E. pedunculare mostly grows in damp, shaded situations alongside
woodland tracks, including ditches and moist banks shaded by coniferous trees. In August 1999 it
was noticed that it was often dried up and perished where it was growing in the more open parts of
tracks in Hancock’s Wood and Cross Plantation. This preference for moist shaded sites shown by
E. pedunculare in Cornwall corresponds to its predilection for moist shaded banks in Nothofagus
forests in New Zealand (Raven & Raven 1976).

Although E. pedunculare is extensively naturalised in the Glynn Valley and it often grows
alongside native Epilobium species there, a thorough search revealed only one other putative, but
unidentifiable, hybrid with E. pedunculare parentage. The species grow intermixed on track edges
with partial shade, but the damp and shade favoured by E. pedunculare are circumstances which
have been found to delay opening of the flowers in Epilobium (Brockie 1959). This increases the
likelihood of self-fertilisation, and therefore diminishes the opportunities for hybridisation.

ACKNOWLEDGMENTS

We are grateful to Mary and Tony Atkinson for information on the whereabouts of Epilobium

NOTES Watsonia 23 (2001) 453

pedunculare in East Cornwall. They also participated, along with Ian Bennallick, in a search for
Epilobium hybrids that resulted in the second find of E. x kitcheneri in Cornwall.

REFERENCES

BROCKIE, W.B. (1959). Breeding systems in New Zealand species of Epilobium L. Transactions of the Royal
Society of New Zealand 87: 189-194.

KITCHENER, G. D. & MCKEAN, D. R. (1998). Hybrids of Epilobium brunnescens (Cockayne) Raven &
Engelhorn (Onagraceae) and their occurrence in the British Isles. Watsonia 22: 49-60.

MCKEAN, D. R. (1999). A new Epilobium hybrid from Scotland, FE. pedunculare A. Cunn. x E. montanum L.
Watsonia 22: 417-419.

MurPHY, R. J. (1998). Plant notes and records. Botanical Cornwall 8: 47-86.

RAVEN, P. H. & RAVEN, T. E. (1976). The Genus Epilobium (Onagraceae) in Australasia: a systematic and
evolutionary study. New Zealand Department of Scientific and Industrial Research Bulletin 216,
Christchurch, New Zealand.

STACE, C. A. (1997). New Flora of the British Isles. 2nd ed. Cambridge University Press, Cambridge.

D. T. HOLYOAK
& Edward Street, Tuckingmill, Camborne, Cornwall, TR14 SPA

G. D. KITCHENER
Crown Villa, Otford Lane, Halstead, Sevenoaks, Kent, TN14 7EA

i
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-
‘

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Watsonia 23: 455461 (2001) 455

Book Reviews

Ireland. A Natural History. David Cabot. Pp. 512, with over 230 illustrations. Harper Collins,
London, 1999. Hbk £34-99. ISBN 000-—220079-1. Pbk £17.99. ISBN 000—220080-5.

This book has been much needed. Ireland, always in the news, is a source of intense fascination to
outsiders, not least naturalists and botanists. This splendid addition to the New Naturalist Library,
not so much introduction as comprehensive synthesis, is a fact-packed guide to the natural
wonders of our smaller, enigmatic western partner. A friend, quizzing me about Ireland, observed
dryly “They tell me it is very green”. Yes, for here is an Atlantic land of water — soft rainy days,
lakes, rivers, streams, boglands, beaches and lush pastures. Conversely some of Ireland’s richest
habitats are the (relatively) dry islands of glacial eskers, sand-dunes, cliffs and, above all,
limestone pavements.

Natural history in Ireland is all about the ecology of animals and plants at the furthest north-
western fringe of Europe, made the more exciting by millennia of isolation. For the visitor, or a
naturalised denizen like David Cabot, Ireland is irresistible, not least for plentiful opportunities to
compare and contrast with Britain. The author’s Preface, which tells how he was instantly grabbed
by the sight of unfamiliar hooded crows, and much else beside, has a familiar ring (in my case it
was Brent geese feeding near my tatty Dublin lodgings). Cabot is primarily a zoologist but his
interests range far and wide.

The book starts from a historical perspective, taking us from the Christian saints and scholars of
the Dark Ages through bemused 17th century English visitors and invaders, the confident and
learned world of 18th century Georgian Dublin and the 19th—early 20th century heyday of Irish
natural history, to more recently departed polymath giants like Robert Lloyd Praeger (1865-1953)
and the B.S.B.I.’s own David Allardice Webb (1912-95). The author notes the remarkable upsurge
in Irish plant and animal recording during the last 25 years, although he could perhaps have taken
a littke more account of recent floristic literature and so many exciting discoveries such as
Trifolium occidentale. He then talks us skilfully through the biological history and chapters on the
vegetation, flora and fauna: mountains and uplands, peatlands, lakes, rivers, the limestone
pavements and turloughs, the (sadly depleted) woodlands, the farm landscape,and the coastline and
Surrounding seas.

The book covers many well-known botanical areas, notably the legendary Burren landscape and
the Killarney oakwoods, and equally fascinating but less publicised treasures such as the Scragh
Bog fenland in Westmeath, the species-rich mineral flush in Bellacorrick Bog in West Mayo,
Uragh Wood in Co. Kerry (with its Kerry spotted slugs), the precipitous arctic-alpine plant
communities of Benbulbin, Co. Sligo, and the extensive sand-dunes and machair of the north-
western coasts. The text also covers several of those individual Irish plants such as Strawberry
Tree, Arbutus unedo; Cottonweed, Otanthus maritimus (on the Co. Wexford coast) or the special
heaths of Connemara, that are absent — or largely so — from Britain, and so fascinate natives and
visitors. The chapter on mountains looks at the conundrum whereby Ireland has a relatively poor
mountain flora, but one of huge botanical interest, with arctic-alpines frequently descending to sea-
level.

The last chapter is devoted to conservation, for a long time haphazard in Ireland but now given
strength by EU law. Appendices at the end of the book list, with brief notes on interest and
location, the nature reserves in the Republic of Ireland and Northern Ireland. Although the two
political regions of the island have by necessity had separate statutory bodies for conservation, it is
to the credit of the natural history community that it has never established any cross-border or
sectarian division.

Here is a fine example of the high quality scientific natural history writing that is at the heart of
the best New Naturalist volumes. For the British botanist keen to know Ireland, I can think of few
better prescriptions than to read and digest Cabot’s book, browse through the classic works of
Praeger and, while you are at it, take out a subscription for Irish Naturalists’ Journal to keep the
story updated.

J. R. AKEROYD

456 BOOK REVIEWS

Flora Nordica. Volume I. Edited by B. Jonsell. Pp. xxii + 344. The Bergius Foundation, the Royal
Swedish Academy of Sciences, Stockholm. 2000. €60. ISBN 97 7190 033 0.

The Flora Nordica Project was initiated in 1987 to produce “‘a comprehensive, diagnostic Flora of
high scientific standard covering the vascular plants”, some 4,600 altogether, of Denmark, Finland,
Iceland, Norway and Sweden, including the Faeroes, Jan Mayen, Bear Island and Spitsbergen. The
chairman and editor-in-chief is Bengt Jonsell, well known in the B.S.B.I. and one of our Honorary
Members. The first of the expected eight or so volumes is now published, covering Lycopodiaceae
to Polygonaceae in Flora Europaea order. All but one of the 31 authors are from the Nordic
countries. A good flora, and this is clearly a very good one, can only be properly assessed after
years of constant use, so rather than writing a review I will attempt to indicate how useful this
book is likely to be to botanists in Britain and Ireland.

Flora Nordica is written in English. It gives practical and concise dichotomous keys, plenty of
synonyms which are in most cases adequate for our purposes, vernacular names where these are
already available in the Nordic languages, and indication of types for all names based on Nordic
material. The good and clear descriptions are about as long as those in Sell & Murrell’s Flora of
Great Britain and Ireland. Distribution and habitat are given in considerable detail, and most
species are mapped by provinces. There are paragraphs on Biology, Variation, Hybridization and
Similar Taxa, and there are numerous line drawings, mostly of diagnostic features. Chromosome
numbers are included, with the origin of those based on Nordic material being given. Subspecies
are given full treatment, and varieties are included though these do not get much of a showing in
the present volume. An outstanding feature is that all hybrids are described or at least diagnosed.
Casuals are treated in reduced typeface and format depending on their frequency, and this leads to
a rather fussy appearance and, for British readers, a sometimes annoying inconsistency of
descriptive information. Flora Nordica thus provides a great deal more information, both in kind
and in depth, than any modern British Flora.

Volume | turns out to be very applicable to the British flora. In the families covered, 200 out of
the 270 species in Stace are also in Flora Nordica, 1.e. 74%. (Conversely 200 out of the 288
species in Flora Nordica are also in Stace, i.e. 70%.) The accounts of the two largest families
serve us particularly well, 90% of the Stace species in the Salicaceae being covered, and 81% in
the Polygonaceae. In both ferns and conifers 69% are covered. In Salix, Flora Nordica describes
no fewer than 111 hybrids, while Stace has 67 and describes only 15. Probably no other foreign
Flora is for us so relevant and worth studying. For its descriptive treatment of hybrids alone it is
unique, and in the paragraphs on biology, habitat and variation in particular there is a vast amount

of information otherwise available, if at all, only widely dispersed in the literature. Although they

are working from opposite ends of the classification and direct comparison is impossible, on the
evidence of the available volumes Flora Nordica and Sell & Murrell provide similar amounts of
description and infraspecific taxonomy, while the former provides a great deal more other
information.

The taxonomy is generally familiar, and where unfamiliar it is usually enlightening. For
example, Polygonum aviculare has six subspecies, including our P. boreale, P. rurivagum and P.
arenastrum, surely a more reasonable solution than trying to separate them absolutely as species.
On the other hand, the subspecies of Rumex crispus are not recognised, and it is suggested that part
of the variation may be due to introgression with other species. In the very thorough account of
Salix, S. fragilis is very narrowly defined, with the material with hairs on the buds and young
leaves being ascribed to S. x rubens, the hybrid with S. alba. (In one of the few conspicuous errors
in the book, S x rubens, though included in the vegetative key, is omitted from the key to material
with female flowers.) Quite extensive essays are devoted to some of the major taxonomic
problems, such as in Betula pubescens and B. tortuosa, Rumex acetosella and Salix repens. There
are numerous instances where one is stimulated to apply characters and ideas to our own plants. Is
it really true that anther-length provides an absolute distinction between Rumex acetosella subsp.
acetosella (1-2—1-5 mm) and subsp. angiocarpus (0-8-1 mm)? If so, it would be very useful in this
dioecious species, but the occurrence of mixed populations makes it difficult to check with
certainty.

As Flora Europaea neared completion in the mid 1970s, a group of British botanists under
David Valentine began planning a “Critical Flora of Great Britain and Ireland”. This envisaged a

ee

BOOK REVIEWS 457

work very much along the lines of Flora Nordica, but it never got beyond the planning stage,
partly for lack of funding. Much of the momentum went instead into the Floras by Stace on the one
hand and Sell & Murrell on the other, with Clapham, Tutin and Moore occupying the middle
ground. We must congratulate the Royal Swedish Academy of Sciences for supporting such an
uncompromisingly thorough international project as Flora Nordica through the Bergius
Foundation, and our Nordic colleagues for bringing this first volume to completion. Anyone
seriously interested in the British flora will find it both stimulating and very useful, and will be
consumed by envy.

A. O. CHATER

William Turner. Libellus de re herbaria novus 1538. Edited with a translation into English by Mats
Rydén, Hans Helander & Kerstin Olsson. Acta Societatis Litterarum Humaniorum Regiae
Upsaliensis 50 (ISSN 0280-0918). Pp. 146. Swedish Science Press, P.O. Box 118, SE-751 04
Uppsala, Sweden. 1999. Paperback with dustcover, SEK150 (+ postage and handling). ISBN 91-
630—8620-4.

“The chief purpose of the present volume,” begins the preface to this simple but scholarly and
elegantly produced book, “is to make the whole of William Turner’s Libellus de re herbaria novus
(1538), the first scientific botanical treatise published in England, available in English, with
historical, textual, linguistic and botanical commentaries.” For this, the team who produced it
deserves the gratitude of anyone interested in the history of British botany, and particularly of our
plant names. William Stearn’s lavish edition of Libellus and The names of herbes (1548),
published by the Ray Society in 1965, provided facsimiles of the two texts and revised versions of
indexes identifying Turner’s plants, originally published by B. Daydon Jackson and James Britten
over a hundred years ago, but no full translation of Turner’s Latin text.

William Turner, described by Charles Raven as “the true pioneer of natural history in England”,
was born at Morpeth in Northumberland around 1510, became a student and then a Fellow of
Pembroke Hall (now College), Cambridge, and died in London in 1568. He is best known for his
great Herball (1551-1568), written after several years of self-imposed exile on the Continent for
religious reasons from 1540 had widened his experience; but his pioneering work, Libellus,
published before he left Cambridge, reveals him as first and foremost a botanist (with a primary
interest in the naming and description of plants) rather than a herbalist.

This new edition of Libellus has a 13-page introduction, which succinctly discusses Turner as
botanist and plant-name scholar; the significance, sources and contents of his first botanical book;
the English, scientific and officinal plant names in it; Turner’s Latin; previous editions of the
book; and the editorial principles of the present one. We learn that the quarto volume of 20 pages
is organised, in roughly alphabetical order, in 148 sections in the sequence of the scientific names,
describes or mentions some 175 plant taxa, the great majority flowering plants (native and non-
native), and includes some 255 English names (or forms); of these “no fewer than 44% are used as
standard or Flora names today (form and spelling disregarded), with same or different botanical
reference” and some 15% are, the editors believe, “first attested there”.

There follows a facsimile of the only known extant copy of Libellus, in the British Library.
Though the facsimile in the Ray Society edition is cleaner, this one, despite some quite serious
show-through from the reverse sides of the pages, creases and other blemishes often removed in
the former, is actually clearer where the printing is faint or smudged. It also provides a more
authentic representation of the original. Next come a transcript and translation, set out on facing
pages. The latter, derived from a “basic translation ... carried out as a term paper in Latin” by Ms
Kerstin Olsson of the Department of Classical Philology, Uppsala University, is unstilted but
faithful to Turner’s Latin. [Indeed, having compared the translations of the preface and the entry
under “NARCISSVS” with those of Charles Raven reproduced in the Ray Society edition, I tend
to favour the new ones.] This section is followed by a helpful commentary on selected phrases, set
out under headings identical to those in the transcript and translation. Together, these two sections
occupy 69 pages. One of my few criticisms is that, when one is looking something up, it is difficult
S see at a glance which of the two one has opened: running headings would have been helpful

efe.

458 BOOK REVIEWS

Much has been made of the unwieldy length of pre-Linnaean plant names, but, as the
introduction to this book points out, there are “no polynomial phrase-names in Turner’s works”:
indeed “one-term terminology” (e.g. “ACORVM” for Iris pseudacorus) is the rule, occasionally
with a synonym, while 9% of the headings are binomials and more of these occur within the text; a
few (e.g. “Plantago maior” and “rumex acetosus’’) are “still with us as standard names, unaltered
or only slightly modified”.

The book ends with three tables (the first with six pages of notes), a brief index of persons and
places, and five pages of bibliography. The first table, in four columns, gives Turner’s English and
scientific names with their modern equivalents, the second Turner’s scientific names with their
modern equivalents, and the third the modern scientific names and the corresponding English
names in Libellus. These are approximately equivalent to Tables C, A and B respectively on
pp. 61-73 of the Ray Society volume, based on those of Jackson (1877 and 1878), and reveal few
changes in the identification of Turner’s taxa.

I strongly recommend this excellent little book (or libellus!).

P. H. OSWALD

Flora of North America north of Mexico. Volume 22 Magnoliophyta: Alismatidae, Arecidae,
Commelinidae (in part), and Zingiberaceae. Flora of North America Editorial Committee. Pp.
Xxiil + 352. Oxford University Press, New York & Oxford. 2000. £65-00. ISBN 0—19-513729-9
(y..22),

This is the fourth volume of the Flora of North America to be published. It provides accounts of
the monocot families from Butomaceae to Marantaceae in the Cronquist system, excluding the
families Cyperaceae and Poaceae which are reserved for later volumes. Many of the 30 families
included are widespread in the North Temperate zone, and have numerous representatives in the
British Isles. The largest of the Temperate families are Juncaceae (118 species), Potamogetonaceae
(37) and Alismataceae (34). However, a substantial minority of the families covered are primarily
Tropical, with many of their North American species restricted to the southern fringes of the
continent. These include Commelinaceae (51 species), Arecaceae, perhaps more familiarly as
Palmae, (29), and the slightly more widespread Xyridaceae (21).

The format of this Flora has been outlined in the reviews of two earlier volumes in Watsonia 21:
141 (1996) and 22: 195-6 (1998). In addition to the basic features common to most Floras, it
contains distribution maps, some biological, ecological and ethnobotanical information and very
useful literature references. (Perhaps mindful of the ever-litigious nature of American society, the
editors warn readers that they do not encourage or recommend the reported folk remedies or
culinary practices!). Botanists accustomed to Stace’s New Flora will notice relatively few
differences in the treatment of species which are common to the British Isles and North America.
Generic concepts are usually similar, although Potamogeton Subgenus Coleogeton is separated as
the genus Stuckenia. At lower taxonomic ranks there are occasional divergences in nomenclature
(e.g. Potamogeton oblongus rather than P. polygonifolius) and in taxonomy (e.g. Juncus balticus is
treated as J. arcticus var. balticus, part of a “wide-ranging and obviously polymorphic complex”).
As European authors usually restrict their attention to European plants and North American
authors to the American flora, there are often unresolved questions about the relationships between
the two. Are the American Juncus arcticus var. alaskanus, Potamogeton strictifolius and
Sparganium eurycarpum synonymous with the European J. arcticus sens. str., P. rutilus and S.
erectum? These issues are sometimes discussed, sometimes ignored.

In the account of the Potamogetonacaeae by R. R. Haynes & C. Barre Hellquist, I was less
struck by divergences in taxonomy than by the differences in approach to the genus compared with
European authors. They “strongly recommend that no one collect specimens of Potamogetonaceae
that are lacking reproductive structures” and that “all specimens should be collected when in fruit”.
A herbarium built up on these lines would be a useful collection of voucher specimens, but would
provide no information on the over-wintering habit of species, or their first leaves or phyllodes:
even flower and pollen characters would be past their best and turions probably under-represented.
Species such as P. robbinsii (which “rarely flowers”) would rarely be collected, and sterile hybrids
would be overlooked. In fact, although the authors say that “hybridization is common among

BOOK REVIEWS 459

members of the genus” they provide very little information about hybrids, simply listing those
accepted by Hagstr6m in his 1916 monograph and by later authors. This comes as a severe shock
to anyone familiar with the history of Potamogeton studies, as Fernald in his 1932 revision of the
North American linear-leaved species famously criticised Hagstr6m for proposing “preposterous
hybrids (of hypothetical parents thousands of miles apart)”. It is a pity that the opportunity was not
taken to produce a revised treatment of the North American hybrids in this volume.

The account of Typha provides a contrast to that of Potamogeton, as C. Galen Smith draws on
the results of 40 years’ study of the genus in North America to provide a treatment in which
hybrids are keyed out and T. x glauca (a “noxious weed’’) is illustrated alongside its parents T.
angustifolia and T. latifolia.

Exchange of plant material between Europe and North America has been taking place ever since
the European discovery (or rediscovery) of the continent in 1492. This Flora provides details of
North American aliens which are to varying degrees naturalised in Britain, including Elodea
canadensis, E. nuttallii, Juncus tenuis, Lysichiton americanus, Sagittaria latifolia and S. rigida. In
the account of Elodea Haynes notes that in contrast to their behaviour in Europe, he knows “of no
instance in North America where Elodea nuttallii or E. canadensis is weedy”. Surprisingly, there
is nO mention in this account of the occurrence of hybrids between these two species in North
America, although the monograph by Cook & Urmi-KG6nig in which this possibility is suggested is
cited. There are many more European plants naturalised in North America than vice versa, and
some of the species included in this volume are Butomus umbellatus, Hydrocharis morsus-ranae,
Juncus capitatus(!), J. compressus, J. inflexus, Luzula pallidula and Potamogeton crispus.
Hydrilla verticillata has spread widely since it was first recorded in the U.S.A. in 1959; it is
described as “widely distributed in the Eastern hemisphere but it is uncertain as to where it is truly
native’. Could this species, like Aponogeton distachyos (S. Africa) and Egeria densa (S. America),
have been introduced to both Europe and North America?

Botanists who need to identify plant material from North America will clearly find this Flora
indispensable. I hope that I have written enough to show that those whose interest lies in the
taxonomy and distribution of the British and Irish flora will also find much of interest in it.

C. D. PRESTON

A flora of Tiree, Gunna and Coll. D. A. Pearman and C. D. Preston. Pp. 168. Published privately
by the authors. 2000. £9.00. ISBN 0—9538111-1-5.

After several recent floras which have left a visible bend in my bookshelves, it is a pleasure to
have a book which can be taken along on a trip without a second rucksack. And to use the book
will involve a trip, as Coll and Tiree, along with Colonsay, are the least accessible of the Argyll
islands and taking a car is an expensive luxury. In terms of pages per unit area, it is arguable that
this is actually a very large flora, as the islands are small, covering bits (sometimes very small bits)
of only eight hectads and this, at my reckoning, runs out at about 20 pages per hectad. The relative
value is even better if you work it out in tetrads.

Whatever currency you use, this excellent account of the flora of these islands is a good buy.
From a decision to produce a flora in 1996 to its publication has taken a mere four years and only
one dedicated field trip, and this during a period in which both authors were much involved in
other projects. Much of the usual process of balancing coverage of different areas and habitats and
the checking of records has clearly been cut short. As the authors acknowledge, it would have been
a difficult process from hundreds of miles away. I am glad that this was not regarded as an
obstacle, as I would much rather have this somewhat incomplete account than no flora at all.

As with all floras, this account has the sense of a “work in progress”, heightened by the
remoteness of the islands and the history of recording here. Throughout most of the Highlands and
Islands, resident botanists are (and have always been) very sparse and most recording is done by a
limited number of expeditions, usually in high summer with all the bias that this induces.
Fortunately, Coll and Tiree have proved an attractive target, both for itinerant botanists,
particularly Joan Clarke and friends, and institutional study, notably Heslop Harrison in the past
and Scottish Natural Heritage more recently, as the impressive bibliography shows. As a result,
Coll and Tiree are probably as well-recorded as much of mainland Argyll and, certainly, the plant

460 BOOK REVIEWS

communities on the islands have attracted more learned discourse. This book is a synthesis of the
results of these botanical visits and one of its strengths is that a lot of disparate data are drawn
together into a concise whole.

With such diverse sources of information there will always be a number of erroneous records
and these have been dealt with in a common-sense manner, as has the particular problem posed by
the stigma attached to the accounts of the flora by Heslop Harrison. The detective work this has
involved makes interesting reading. In fact, the whole of the introduction is excellent. The
descriptions of the main habitats are succinct and it is good to see NVC community categories
being used as a matter of course. The section on phytogeography may seem rather long for these
small islands but given their position on the edge of things, the association with Heslop Harrison
and his theory of per-glacial survival and Chris Preston’s recent work on geographical
relationships in plant distribution, this seems justified. The comparison with the nearby island of
Mull and the somewhat more distant Outer Isles is fascinating.

The layout is clear and straightforward with the comparatively long introductory section
followed by the species accounts and finally by a short account of the stoneworts by Nick Stewart
and ecological notes, with tetrad maps, on the Cyperaceae of Tiree by David Pearman. Other tetrad
maps are provided only for a selection of aquatic plants, probably reflecting the lack of precision
of older records, the main thrust of visits by the authors and a desire to save space. For most
species of interest, the frequent citing of localities in the text with map references more than makes
up for this. The species accounts are commendably concise but fuller accounts are given where
there is a story to be told or a question to be asked, be it about rarities like Eriocaulon aquaticum
or the more mundane Eriophorum vaginatum. Some accounts are rather enigmatic with Juncus
squarrosus “common on heathy ground” but “very poorly recorded”. To someone used to the
mainland Argyll flora, the rarity of plants like Digitalis purpurea, Eriophorum vaginatum and
Conopodium majus, abundant plants in main Argyll, comes as a surprise.

Do go and visit Coll and Tiree, they are wonderful islands, and do take this flora with you.
Almost every page of the species accounts has at least an implied query on which the curious
botanist could shed some light.

G. P. ROTHERO

Lichens. Oliver Gilbert. Pp. 214. The 86th volume in the New Naturalist Series. Harper Collins.
2000. Hbk £34-99. ISBN 0 00220081 8. Pbk £19.99. ISBN 0 00 2200821.

The high standard we have come to expect of this invaluable series is maintained or even excelled
in this latest volume. The student of the smaller fungi, like the bus passenger, has had a patient
wait (55 years since the first New Naturalist Volume) and now two books have turned up at once
with the publication of Ingram and Roberston’s “Plant Disease” as number 85 in 1999. In an age
when “biodiversity” is on everyone’s lips and in theory the otter or eagle now have equal status
with the rarest lichen or plant pathogen the publishers and editors of this series are to be
congratulated in bringing to the press two books which must further the appreciation and
understanding of this huge slice of our wildlife.

In his opening chapter Gilbert shows that lichens are of more than just scientific interest with a
review of the way they have impinged on our life. If you enjoy the look, feel and scent of Harris
tweed, relish the long lingering qualities of the finest perfumes or just want to test the pH of your
soil with litmus paper he describes the essential part lichens play.

You are then led through clear and succinct chapters on the nature of lichens, creatures that need
lichens, and air pollution and lichens, to the main part of the book. Here the important role that
lichens play in most of our major ecosystems is described, with chapters on woodland, acid rocks,
heaths and moors, chalk and limestone, the built environment of church and village, our mountain
areas, the rivers, lakes and coastal habitats. At a glance these pages seem liberally splattered with
Latin names (since few lichens are blessed with English names). The non-lichenologist should not
be put off. Read any paragraph and they trip easily by with Gilbert’s lively writing style almost
invariably linking them into some thought-provoking ecological process or ecological niche you
never thought of. These range from the effects of wind blasted ice and grit in subarctic heaths of
the Cairngorms to the “canine zone” of tree bases in suburbia and from crashed aircraft on St
Kilda, through old shoes at Girvan, to the drip zone under galvanised motorway crash barriers.

BOOK REVIEWS 461

Three appendices provide information on the use of lichens for dating objects, the biological
estimation of air pollution and the use of lichens as indicators of ecological continuity. There is
also an adequate glossary and extensive bibliography.

Misprints and errors are few and mostly inconsequential. Occasionally a now superseded lichen
name creeps back in and the reference to “Soya” sheep on page 237 should probably not be
proffered as evidence of a previously overlooked transgenic industry in western Scotland. The
high standard of paper, print and plates we have seen in some of the recent “New Naturalists” is
maintained here.

This book conveys the excitement of the recent rapid accumulation of knowledge of the part
lichens play in British ecosystems and identifies areas where the amateur can still make a
significant contribution. It handsomely realises the aspirations of the editors in “seeking to interest
the general reader by recapturing the enquiring spirit of the old naturalists”’.

If this spirit moves you to discover the lichen genus Rocella in Ireland the author offers a more
tangible spirit prize in the shape of a bottle of whiskey to the finder. Even without this inducement
I am certain this book will stimulate you to get out and look at lichens, towns, cities and
countryside in a new and exciting way.

R. G. WOODS

Alien Plants of Yorkshire. G. T. D. Wilmore. Pp. 316. Yorkshire Naturalists’ Union. 2000. Price
£15.00 (reduced price to members of the Yorkshire Naturalists’ Union £12.00) ISBN 0—9504093-
3-2.

It is perhaps surprising that in a small country with a flora that has been extensively botanised for
over 400 years, so little time and effort has been devoted to the study of alien plants. For most of
that period they have formed an increasingly substantial minority of the flora of the British Isles,
(over 40% of taxa at the last count) but have usually been treated with suspicion or disdain. Often
regarded as “second class citizens” (try telling that to people in areas of the country suffering from
the ravages of Fallopia japonica, Crassula helmsii or Rhododendron ponticum) they were at best
relegated to appendices in Floras or at worst completely ignored. Clive Stace’s New Flora of the
British Isles changed the position overnight, not only were botanists able to see what alien plants
were present they also had help in identifying them.

However, some areas of the Country did have a better record in studying the alien flora and this
excellent and fascinating book sets out the history of such studies in Yorkshire and catalogues the
present state of our knowledge.

Some 3,046 taxa in 131 families are included and details given include: Scientific and English
name; a brief statement on its status in Yorkshire, with Country or region of origin and list of
Yorkshire vice-counties in which it has been recorded. For most taxa there then follows a more
detailed list of records for each vice-county, but for about 90 common taxa there is a “thumbnail
sketch” of habitats and history. The comprehensiveness of the book is well illustrated by the
reviewer having penned a criticism on the omission of taxa alien to Yorkshire but native to Britain,
only to find that appendix 3 covers this subject very well. Other appendices deal with less well
documented taxa and taxa not included in Stace or the B.S.B.I.’s two “Alien” books, and a
bibliography and comprehensive index complete the book.

Misprints and “typos” are commendably rare; over half a dozen taxa are wrongly attributed with
illustrations in the index and page numbers are missing from the contents page. But this reviewer
does have an intense dislike of single lines of a paragraph at the top or bottom of a page., and there
are too many of these “orphans”.

However, these are no more than minor irritations that in no way detract from the excellence of
the book itself. Geoffrey Wilmore and the Yorkshire Naturalists’ Union are to be congratulated on
producing a book which deserves to be on the shelf of every botanist with an interest in the flora of
these islands and hopefully will serve as an inspiration to others to produce similar volumes for
other regions.

R&GoEREIS

Watsonia 23: 463-464 (2001) 463

Obituary

RAYMOND PAUL BOWMAN
(1931-1999)

It was with much sadness and some shock that naturalists (particularly, but not only, in
Hampshire) learnt of the sudden death of Paul (as he was known to us), on Calshot Spit on the
Hampshire coast in July 1999. He had gone there to count the estuary birds and to record the local
flora, and suffered a heart attack on this very hot day, apparently after a swim.

Paul was born in Hythe, Hampshire, on the east side of the New Forest, in 1931. His remarkable
mother Dorothy was born near Exbury on the south-east edge of the New Forest. When he was
three years old, the family (which came to include his four younger sisters, June, Dawn, Wendy
and Heather) moved to the Maybush district of north-west Southampton, but summer holidays
were nearly always spent at the caravan at Longdown, on the edge of the Forest, south east of
Ashurst. His mother describes how his schoolboy passion for wildlife blossomed at an early age.
Tragically, at the age of 15, an infection of meningitis destroyed his hearing. This disaster had a
profound influence on his lifestyle, and focused his interests even more on wildlife. Though best
known later as a botanist, he patiently and methodically studied the bird life with binoculars, as of
course he could not hear their calls, and became an outstanding ornithologist. He was a member of
the Southampton NHS and Hants Ornithological Society for over 40 years, and the Recorder of the
NHS for over 20. His skills as a field worker were outstanding - indeed quite exceptional - in the
effort he expended in verifying records.

Sadly, he never learnt to lip-read effectively (except within his own family) so although he was
by nature a most outgoing and sociable person, communication with other naturalists was not easy.
He could tell others all he wanted to say, but they had to write things down in reply.

He became a quite outstanding botanist with an encyclopaedic knowledge of the flora of South
Hampshire. As a member of the Botanical Society of the British Isles, his knowledge and
meticulous work led to him being appointed botanical Vice-County Recorder for South Hants
(v.c. 11), a post he held for 20 years until his death. When the work on the Flora of Hampshire
was revived after various misfortunes, he became one of the three co-authors of this book
(published eventually in 1996) with Lady Anne Brewis and myself; and the most casual inspection
of the pages of this book reveals how much work he did towards its completion; not least in the
detailed distribution maps, but in his patient and meticulous care in making records. I personally
think he did more work for the Flora of Hants than most of the rest of us combined! And all of an
incredibly high standard.

In this period he did not neglect his ornithological studies, and every October he travelled,
usually to the Scilly Isles, to take part in annual bird migration counts - incredibly without being
able to hear a single bird song.

Professionally he worked as a draughtsman and cartographer with the Ordnance Survey at
Southampton. It is clear that his professional work in this field was of the same outstanding quality
as his natural history work, and his work was recognised by the award of the British Empire
Medal. His cartographic skills were of great help in his botanical work, and he helped much with
the geographical and mapping aspects of the Flora of Hampshire. Unlike many botanists I know
(some quite eminent!), if Paul gave you a grid reference you knew you could rely on it implicitly.
He was a very good and patient photographer - some of his photos are included in the Hants Flora,
and are superb.

His ready smile and amiable, enthusiastic personality managed to shine through his severe
disability, particularly during the protracted (and sometimes heated!) arguments that developed in
committee during the gestation of the Flora. He took all these problems in his stride, with his
excellent sense of humour. As soon as he was given brief written notes of what was actually going
on in committee, he ‘cottoned-on’ instantly to the problems and was always very helpful.

He was a botanist of reputation well beyond Hampshire, and had extensive correspondence with
naturalists all over Britain.

464 OBITUARY

Apart from his work as a naturalist and, above all, as a botanist, he has permanent memorials not
only in the published Hants Flora, but in the horsetail hybrid, Equisetum x bowmanii C. N. Page
(E. sylvaticum L. x E. telmateia Ebrh.) named after him by the pteridologist Dr C. N. Page. He
found this hybrid, new to science, in 1986 in the northern part of the New Forest. It is still there,
but it has not yet been found outside the New Forest, (Page 1987 - Watsonia 17: 273-277).

I found Paul to be a lovable personality and a great naturalist whom we miss very much in
botanical circles in Hampshire.

FRANCIS ROSE

Watsonia 23: 465-467 (2001) 465

Report

ANNUAL GENERAL MEETING, 8 MAY 2000

The Annual General Meeting of the Society was held at the Ulster Museum, Belfast, at 10.45 a.m.
The President, Mrs M. Briggs took the chair 56 members were present.

Apologies for absence were read and the minutes of the 1999 Annual General Meeting
(published in Watsonia 23: 373-375) were circulated, approved as a correct record and signed by
the President.

REPORT OF COUNCIL

The report had been circulated to members and the President went through it commenting on the
main achievements of the Society during the past year, noting especially the good progress with
the Threatened Plants Database and Atlas 2000 and she thanked all who had contributed — the
Co-ordinators, the Atlas organiser, the inputters and all the members. She finally thanked every
one who had, voluntarily, worked so hard for the Society.

The Treasurer then presented the financial part of the Report. Summary accounts had been
circulated to members; full sets of accounts had been available on request by post and were now
available at the meeting. The Treasurer reported that the Society’s accounts were in good order
and that we had worked within our means. He pointed out that as Gift Aid had replaced Covenants,
the Society is able to recover tax paid on subscriptions on completion of a simple form which
would be circulated to all members; in response to a question, however, it was pointed out that this
could not be done if the subscriber was, as a professional, claiming the subscription against tax in
his business accounts.

The Membership Secretary had reported that the Society membership had declined, slightly,
over the past year but that this did not appear to be as a result of the increase in subscription.

The Treasurer proposed the adoption of these reports which was seconded by Mr E. F.
Greenwood. The President thanked the Treasurer for all his hard work, particularly the extra effort
involving the Charity Commissioners.

ADOPTION OF REVISED RULES

Copies of the proposed revised rules had been available by post from the Honorary General
Secretary and were now available at the meeting. The Treasurer stated that the changes were
purely administrative and were in response to advice from the Charity Commissioners:-

They emphasise compliance with Charity Law.

They emphasise best practice in financial control.

They strengthen the quorum requirements at meetings of Council.
They update details of permanent working committees.

Under a requirement of Charity Law, they remove the vote from junior members under the age of
18.

They clarify that family members have only one vote per family.

The adoption of the Revised Rules was proposed by Mrs M. Lindop and seconded by Mrs J.
Robertson and carried unanimously.

At this point, Mrs M. Briggs stood down as President and, as she did so, thanked all members for
their help and support during her presidency; in particular she thanked Mr Peter Thomson, Council
Minuting Secretary and Mrs Margaret Perring who was retiring after handling B.S.B.L
publications for 23 years; she welcomed Mr Jon and Mrs Sue Atkins of Summerfield Books who

466 REPORT

were taking over. Finally, she welcomed her successor as president, Dr Geoffrey Halliday, well
known to members for his work with Records Committee and for the acclaimed Flora of Cumbria.

ELECTION OF PRESIDENT

Dr G. Halliday was nominated by Council. He gave the Society’s thanks to Mrs M. Briggs M.B.E.
for all her good work on behalf of the Society — 2 years as President, preceded by 23 years as Hon.
General Secretary. He expressed his gratitude for his election and said that he was looking forward
to making his own contribution to the office of President. He was elected unanimously.

ELECTION OF VICE-PRESIDENT

Professor C. A. Stace was proposed by Miss A. Burns and seconded by Mrs M. Briggs. His
election was unanimous.

ELECTION OF ACTING HONORARY GENERAL SECRETARY

The President thanked Mr R. G. Ellis, the retiring Hon. Gen. Sec. for all his hard work in that
office and noted with gratitude that he was continuing as Editor of ‘News’. Mr Ellis thanked Mrs
Briggs and the Treasurer for their assistance and support and also recorded his thanks to Mr P. J.
Fry for his work as Assistant Secretary at the B.S.B.I. desk at the Natural History Museum. The
administration of the Society was under review, pending the outcome of the review, Miss A. Burns
is nominated by Council to serve as Acting Honorary General Secretary.

RE-ELECTION OF HONORARY TREASURER

Mr M. E. Braithwaite, nominated by Council was re-elected, proposed by Mr R. G. Ellis, seconded
by Mr A. O. Chater. The President thanked him very much for all his good work, particularly his
continuing efforts with the Charity Commissioners.

RE-APPOINTMENT OF EDITORS, INDEXERS AND REPRESENTATIVES OF COMMITTEES

The President proposed the re-appointment of the Honorary Editors, Indexers and Representatives
of Committees and thanked them very much for all their hard work, mentioning especially, Mr C.
R. Boon for indexing Watsonia.

At this point, the President also recorded his own thanks to Mr D. A. Pearman, Dr C. D. Preston,
Mrs J. M. Croft and Dr T. D. Dines for their enormous amount of work with the Atlas.

ELECTION OF COUNCIL MEMBERS

In accordance with Rule 12, nominations had been received for Miss R. Davis, Dr J. Edmondson
and Mr M. S. Porter. Profiles had been circulated and would be published. The election of these
new Council members was proposed by Mrs M. Briggs and seconded by Mr E. F. Greenwood;
their election was unanimous.

ELECTION OF HONORARY MEMBERS

The President stated that three members had been nominated for Honorary Membership of the
Society this year and that all three had made outstanding contributions to the Society in their
different ways. Sponsors for all three had produced appreciations of their respective candidates
and these were either given, or, if not able to be present, read to the meeting and would be

REPORT 467

published in ‘News’. Their election was proposed by Mr R. G. Ellis, seconded by Dr D. L. Kelly
and was carried unanimously.

The three new Honorary Members are, Dr F. E. Crackles M.B.E., MSc., sponsor Mrs M. Briggs;
Dr P. Macpherson F.R.C.P., F.R.C.R., sponsor Mr M. E. Braithwaite; Mr R. D. Meikle, sponsor
Dr E. F. Greenwood.

RE-ELECTION OF HONORARY INDEPENDENT EXAMINER

The President expressed the Society’s gratitude to Mr John Coats for his exemplary examination
of the Society’s accounts. His re-election as Honorary Examiner was proposed by Mr A. O. Chater
and seconded by Mr R. G. Ellis.

There was no other business and the meeting closed at 12.40 p.m.

AILSA BURNS

BOTANICAL SOCIETY
OF THE

BRITISH ISLES
(B.S.B.L.)

The B.S.B.I. traces its origin to the Botanical Society of London founded in 1836 and has a membership
of 2,850. It 1s the major source of information on the status and distribution of British and Irish
flowering plants and ferns. This information, which is gathered through a network of vice-county
recorders, is the basis for plant atlases and for publications on rare and scarce species and is vital to
botanical conservation. The Society published Atlas of the British flora in 1962 and revised editions and
is working towards a new atlas, to be completed in 2000. It organises plant distribution surveys,
publishes handbooks on difficult groups of plants and has a panel of referees available to members to
name problematic specimens. The B.S.B.I. arranges conferences and field meetings throughout the
British Isles and, occasionally, abroad. It welcomes as members all botanists, professional and amateur
alike.

Details of membership and any other information about the Society may be obtained from:

The Hon. General Secretary,
Botanical Society of the British Isles,
c/o Department of Botany,

The Natural History Museum,
Cromwell Road,

London, 5W7 S5BD.

BSBI Handbooks

Each Handbook deals in depth with one or more difficult groups of British and Irish plants.

No.1 Sedges of the British Isles
A.C. Jermy, A.O. Chater & R.W. David. Revised edition, 1982. 272 pp., with descriptions,
drawings and distribution maps for all 73 species of Carex. Paperback. ISBN 0 901158 05 4.

No.2 Umbellifers of the British Isles
T.G. Tutin. 1980. 200 pp., with descriptions and drawings of 73 species of Apiaceae
(Umbelliferae). Paperback. ISBN 0 901158 02 X.

No.3 Docks and knotweeds of the British Isles

J.E. Lousley & D.H. Kent. 1981. 208 pp., with descriptions and drawings of about 80 native and |

alien taxa of Polygonaceae. Paperback. ISBN 0 901158 04 6. Out of print. New edition with
distribution maps in preparation; orders recorded.

No.4 Willows and poplars of Great Britain and Ireland
R.D. Meikle. 1984. 200 pp., with descriptions and drawings of 65 species, subspecies, varieties
and hybrids of Salix and Populus. Paperback. ISBN 0 901158 07 0.

No.5 Charophytes of Great Britain and Ireland
J.A. Moore. 1986. 144 pp., with descriptions and drawings of 39 species and varieties of
Characeae and 17 distribution maps. Paperback. ISBN 0 901158 16 X.

No.6 Crucifers of Great Britain and Ireland
T.C.G. Rich. 1991. 344 pp., with descriptions of 148 taxa of Brassicaceae (Cruciferae), 129 of
them with drawings, and 60 distribution maps. Paperback. ISBN 0 901158 20 8.

No.7 Roses of Great Britain and Ireland
G.G. Graham & A.L. Primavesi. 1993. 208 pp., with descriptions and drawings of 13 native and
nine introduced taxa of Rosa, descriptions of 76 hybrids, and 33 maps. Paperback. ISBN
0 901158 22 4.

No.8 Pondweeds of Great Britain and Ireland
C.D. Preston. 1995. 352 pp., with descriptions and drawings of all 50 species and hybrids of
Potamogeton, Groenlandia and Ruppia, most of them with distribution maps: detailed
introductory material and bibliography. Paperback. ISBN 0 901158 24 0.

No.9 Dandelions of Great Britain and Ireland
A.A. Dudman & A.J. Richards. 1997. 344 pp., with descriptions of 235 species of Taraxacum,
most of them illustrated by silhouettes of herbarium specimens; drawings of bud involucres of
139 species and 178 distribution maps. Paperback. ISBN 0 901158 25 9.

No.10 Sea beans and nickar nuts
E. C. Nelson. 2000. 156 pp., with descriptions of nearly 60 exotic seeds and fruits found
stranded on beaches in north-western Europe (many illustrated by Wendy Walsh) and of the
mature plants (some with drawings by Alma Hathaway), accounts of their history and folklore,
growing instructions etc. Paperback. ISBN 0 901158 29 1.

Other publications

List of vascular plants of the British Isles
D.H. Kent. 1992. 400 pp. Nomenclature and sequence as in Clive Stace’s New Flora of the
British Isles (1991, 1997), with selected synonyms. Paperback. ISBN 0 901158 21 6. Supplied
with errata lists and Supplement 1.

Alien plants of the British Isles
E.J. Clement & M.C. Foster. 1994. 616 pp. Lists 3,586 recorded non-native species (of which
885 are established), with English names, frequency of occurrence, status, areas of origin,

location of voucher specimens, references to published descriptions and illustrations, and
selected synonyms. Paperback. ISBN 0 901158 23 2.

Alien grasses of the British Isles
T.B. Ryves, E.J. Clement & M.C. Foster. 1996. 234 pp. A companion volume to the last, listing
over 700 non-native grasses; includes keys to bamboos and eight of the larger and more difficult
genera and 29 pp. of illustrations. Paperback. ISBN 0 901158 27 5.

Plant crib 1998
T.C.G. Rich & A.C. Jermy. 1998. 400 pp. An identification guide for some 325 ‘difficult’

taxonomic groups, with explanations, keys and illustrations of plant details. A4 paperback.
ISBN 0 901158 28 3.

British Red Data Books 1 Vascular plants
M.J. Wigginton, ed. 1999 (3rd ed.). 468 pp. Up-to-date information on 408 taxa (including 118
microspecies) regarded as threatened in Great Britain, with details of their habitats and
associated species, maps showing pre-1970, 1970-1987 and post-1987 records by 10-km
squares, and 1l-km square frequency maps for 63 taxa. Hardback, published by JNCC,
Peterborough. ISBN | 86107 451 4.

Scarce plants in Britain
A. Stewart, D.A. Pearman & C.D. Preston, comp. & ed. 1994. 518 pp. Accounts of 254
nationally scarce taxa (occurring in 16-100 10-km squares in Great Britain) and of 71 taxa
formerly thought to be so, with details of their habitats and associated species, reproductive
biology, changing British distribution and world range; updated distribution maps. Hardback,
published by JNCC, Peterborough. ISBN 1 873701 667.

Aquatic plants in Britain and Ireland
C.D. Preston & J.M. Croft. 1997. 365 pp. Summarises the distribution, habitat and reproductive
biology of 200 aquatic plants in 72 genera, with 200 distribution maps and 72 drawings.
Hardback, published by Harley Books, Colchester. ISBN 0 946589 55 0.

Available from the official agents for BSBI publications, Summerfield Books (Jon & Sue Atkins),
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. East Kent

West Kent

. Surrey

. South Essex
. North Essex
. Hertfordshire

. Monmouthshire
. Glamorgan

. Breconshire

. Radnorshire

Carmarthenshire

. Dumfriesshire
. Kircudbrightshire

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. Ayrshire

Renfrewshire

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Selkirkshire
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West Lothian

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Mid Cork

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Co. Waterford
South Tipperary
Co. Limerick
Co. Clare

North Tipperary
Co. Kilkenny
Co. Wexford
Co. Carlow
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Bedfordshire 62.
Huntingdonshire 63.
Northamptonshire 64.
East Gloucestershire 65.
West Gloucestershire 66.
Herefordshire 67.
Worcestershire 68.
Warwickshire 69.
Staffordshire 69b.
Shropshire 70.
South Lincolnshire Tele
WALES
Pembrokeshire 50.
Cardiganshire ai
Montgomeryshire a2.
Merionethshire
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Stirlingshire 99.
West Perthshire 100.
Mid Perthshire 101.
East Perthshire 102.
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South Aberdeenshire 105.
North Aberdeenshire 106.
Banffshire 107.
Moray 108.
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South-east Galway H29.
West Galway H30.
North-east Galway H31.
Offaly H32.
Co. Kildare H33.
Co. Wicklow H34.
Co. Dublin H35.
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Co. Longford H38.
Co. Roscommon H39.
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Anglesey

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Clyde Isles
Kintyre

South Ebudes
Mid Ebudes
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West Sutherland
Caithness
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Orkney
Shetland

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Co. Cavan

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Fermanagh
East Donegal
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Watsonia

February 2001 Volume twenty three Part three

Contents

DIXON, J. M. On the status of the genus Koeleria Pers. (Poaceae) in Britain

MASKELL, L. C. & RAYBOULD, A. F. The decline of eee Coe
(Poaceae) in the British Isles

RICH, T. C. G. & BIRKINSHAW, C. R. ee of Britain’ S bioversy cues
depauperata With. (Cyperaceae), Starved Wood-Sedge

FOLEY, M. J. Y. A morphological comparison between some British Orieue
species (Orobanchaceae) and their closely-related, non-British aaa
from continental Europe: Orobanche rapum-genistae Thuill. s.1.

ALLEN, D. E. Brambles (Rubus L. sect. Rubus, and sect. ic: Lindley
Rosaceae) in the Channel Islands

RANDALL, R. D. & RICH, T. C. G. Conservation of Britain’s biodiversity ihe
dasycoccus (Rosaceae), Thick-berried Bramble ea Nee ae hee

NOTES

Jones, R. A. & Stewart, N. F. Nitella gracilis (Smith) ean an elusive
charophyte new to Cardiganshire (v.c. 46) a

Welch, D. Colonisation by Cochlearia danica L. song sink a in eed
Scotland from 1996 to 2000

Braithwaite, M. E. Alchemilla glaucescens Wallr. in v.c. 81 Benda

Allen, D. E. Geranium robertianum L. var. marinum: an earlier author ;
Holyoak, D. T. & Kitchener, G. D. planet x kitcheneri McKean (Onagracene)

in East Cornwall
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Botanical Society of the British Isles

Volume 23 Part 4 August 2001
Editors: M. Briggs, R. G. Ellis, D. L. Kelly, D. R. McKean,
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© 2001 Botanical Society of the British Isles

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The cover illustration of Euphorbia hyberna L. (Irish Spurge) was drawn by
Rosemary Wise.

Watsonia 23: 469-480 (2001) 469

What is Anthyllis vulneraria L. subsp. corbierei (Salmon &
Travis) Cullen (Fabaceae)?

te C.G. RICH

Dept. Biodiversity and Systematic Biology, National Museum & Gallery, Cardiff CF 10 3NP

ABSTRACT

Anthyllis vulneraria L. subsp. corbierei (Salmon & Travis) Cullen, originally distinguished by having
spreading hairs on the stems and petioles, is reputed to be a British endemic confined to a few localities in
Anglesey, Caithness, Cornwall and the Channel Islands. Examination of herbarium material indicates that
plants with spreading hairs occur widely around the British coast and occasionally inland, encompassing a
range of forms. Similar material with spreading hairs also occurs on the coast of northern France. Field work
at previously reported sites (including the type locality) showed that most populations were polymorphic with
respect to hair, corolla and calyx characters. Different hair types occurred in random combinations with
different corolla and calyx characters. The occurrence of spreading or appressed hairs is probably simply
polymorphic variation. I do not recognise subsp. corbierei either as an endemic or as a subspecies.

KEYWORDS: Britain, France, Red Data Book, polymorphism, infraspecific variation.

INTRODUCTION

Anthyllis vulneraria L. is taxonomically very complex and consists of numerous inter-grading
variants which are to some extent ecologically and geographically separated (Cullen 1976). Cullen
(1968, 1976, 1986) provided accounts of the infraspecific variation, in which three native and two
introduced subspecies were recognised in the British Isles. One of these, subsp. corbierei (Salmon
& Travis) Cullen, is reputed to be a rare plant endemic to Britain, and was included in the third
edition of the vascular plant Red Data Book (Wigginton 1999). However, Wigginton
acknowledged that the RDB account was based on little recent information, and he suggested
further investigation was required. This prompted R. A. Jones and T.C.G.R. to review the
information available about subsp. corbierei and to carry out field work to assess its current status.

The taxon was first described by Salmon & Travis (1917) as A. maritima Schweigg. var.
corbierei (coastal forms of Anthyllis were at the time ascribed by some botanists to A. maritima
Schweigg., now often referred to A. vulneraria subsp. vulneraria var. langei Jalas; Cullen 1986). It
was distinguished as having an abundance of spreading hairs on the stems and petioles (‘Caules
folia petiolique valde patento-villosae’) in contrast to typical A. vulneraria which has appressed
hairs. After studying herbarium material (including type specimens), cultivation experiments and
field work at South Stack, Cullen (1967) raised the variety to subspecific rank for Flora Europaea
(Cullen 1968). In his résumé of the A. vulneraria complex, Cullen reported subsp. corbierei from
only four localities, in Anglesey (stated as forming large, distinct, homogenous populations),
Cornwall and Sark, and suggested it could occur elsewhere (Cullen 1976).

Cullen (1986) provided a description, clearly based on his holotype in LIV: ‘Upright perennial.
Stems hirsute over their whole length. Leaves somewhat fleshy, concentrated in the lower part of
the stem. Calyx usually not red-tipped. Corolla clear yellow’. This description has formed the basis
of descriptions in subsequent floras (e.g. Stace 1997; Akeroyd 1998) except for the poor
description in Clapham et al. (1987) which was derived indirectly from Cullen’s Flora Europaea
account.

The taxonomic complexity of A. vulneraria forced me to a wider investigation of subsp.
corbierei from the literature, herbarium specimens, field work and limited cultivation experiments
to try to clarify which populations were that taxon and therefore which might require conservation.
The work has led to a different conclusion from Cullen about the most appropriate rank for the
taxon and its distribution, and consequently its priority for conservation. The work applies only to
A. vulneraria from the British Isles and northern France, and does not apply to other European
taxa with spreading hairs (see Cullen 1968, 1976).

470 TC, G-RICH
ANTHYLLIS VULNERARIA FORMS WITH SPREADING HAIRS IN THE BRITISH ISLES

As the taxon was originally distinguished on the basis of spreading hairs on the stems and petioles,
a review of such herbarium material was carried out. All material of A. vulneraria from the British
Isles was examined at BM, BRISTM, CGE (mostly seen by J. Cullen or J. R. Akeroyd), DBN, K
(mostly seen by J. Cullen, also Marsden-Jones & Turrill’s 1933a material), LIV (which now
includes Cullen’s research collection transferred from LIVU), NUW, OXF, RNG (mostly seen by
J. R. Akeroyd), TCD (mostly seen by J. R. Akeroyd) and material named as subsp. corbierei was
borrowed from E and LTR.

The herbarium specimens with spreading hairs showed a large range of variation, especially
with regard to corolla and calyx coloration. Notes on the specimens with most of the stem with
spreading hairs seen, and their differences from Cullen’s description of subsp. corbierei are given
in Appendix 1 with other records. The records are mapped in Figure 1 with selected silhouettes:
these show a large range of variation in general growth form, distribution of leaves on the stem
and in leaf shape.

The herbarium material showed significant variation in the density, coverage and coloration of
spreading hairs. Plants with spreading hairs right the way up the stem were uncommon, and many
specimens had appressed hairs on the top and/or lateral peduncles (including Cullen’s holotype).
Inland plants tended to be less densely hairy than coastal plants. In plants with spreading hairs, the
occurrence of red-tipped calyces was more frequent than green-tipped calyces; these character
states are simple polymorphic variation which is widespread in many populations in Britain, with
red-tipped calyces being dominant in most (but not all) populations. Variation in corolla colour
was also frequent, especially in Cornish material.

Spreading hairs occurred in material which had been determined by various botanists as subsp.
vulneraria or subsp. lapponica (Hyl.) Jalas, which are more or less recognisable subspecies, or as
intermediates with subsp. corbierei. Plants with red corollas and spreading hairs were often
ascribed to subsp. vulneraria var. coccinea L. but many could equally have been treated as subsp.
corbierei.

ANTHYLLIS VULNERARIA FORMS WITH SPREADING HAIRS IN NORTHERN FRANCE

It has been suggested that subsp. corbierei might occur on sea cliffs in the western part of France
(e.g. Cullen 1976, noting he had seen little material from this area). Corbiére (1894) had noted
variation in pubescence in Normandy similar to that described by Salmon & Travis (1917), but the
name he had applied (var. sericea Bréb.) could not be clearly related to Salmon and Travis’s taxon
from Brébison’s description. Couderce (1975) revised the French infraspecific taxa of Anthyllis
vulneraria, and cited subsp. corbierei as synonymous with his subsp. vulgaris (Koch) Willk. &
Lange var. maritima (Koch) Willk. His description included plants with a mixture of hair types,
including those with spreading hairs throughout the stem: ‘a pilosité généralement apprimé, mais
parfois hirsute dans la moitié inférieure ou méme sur toute leur longueur’ .

Specimens from the Musée National d’ Histoire Naturelle, Paris (P) and the Herbier de la Société
des Sciences Naturelles et Mathématiques de Cherbourg (CHE) were examined for material which
might be regarded as subsp. corbierei (these herbaria contain Corbiére’s collections, the former
having been transferred from Caen CN). Eight collections from the coast had dense spreading
hairs below and at least sparse spreading hairs above (Appendix 2). These are regarded as falling
within the variation noted in the British Isles (cf. above and Appendix 1). Corbiére’s annotations
on the sheets indicate that he regarded such plants with spreading hairs as var. sericea Bréb., as
surmised by Salmon & Travis (1917). Similarly, there are two more recent inland collections from
the Pas-de-Calais in the Centre Régionale de Phytosociologie/ Conservatoire Botanique National
de Bailleul herbarium.

ANTHYLLIS VULNERARIA SUBSP. CORBIERE]I 47]

FIGURE 1. Map showing distribution of forms with spreading hairs in Britain with silhouettes drawn from
selected specimens.

472 T.C.G_RICH

FIELD INVESTIGATION OF KNOWN POPULATIONS

The variation in herbarium material prompted a more detailed investigation of variation in the
field. Field work was carried out in four areas, concentrating on those from which Cullen had cited
material: Anglesey (including the type locality “grassy banks, cliffs near South Stack’), Cornwall
and the Channel Islands (the latter by M. Marsden, B. Ozanne, R. Rabey and R. Veall in 1999),
and also Dorset. At each site the phenotypes present were noted. The variation is summarised in
Table 1 with phenotypes coded as follows:

Corolla: R = red, Y = yellow, W = red standard with white keel, O = orangey-red. This includes
the most frequent phenotypes, though there is variation within them (cf. Marsden-Jones &
Turrill 1933a).

Stem and petiole hairs: A = appressed, S = spreading to ascending. The hairs on stems of some
plants can be spreading to ascending below but appressed above (cf above).

Calyx: G = whitish-green-tipped, T = red-tipped. These are the two variants, though the
development of the redness in the tips is variable, and may sometimes be almost purple. I have
not seen any plants with red or orangey corollas with green-tipped calyces.

Thus plants coded YAT have yellow corollas, appressed hairs and red-tipped calyces (typical
‘yulneraria’), those coded YSG have yellow corollas, spreading hairs and whitish-green-tipped
calyces (subsp. corbierei sensu Cullen), etc. Representative specimens have been deposited in
NMW.

ANGLESEY

Most populations investigated were polymorphic, containing two or more phenotypes (Table 1).
The composition of the populations differed between sites; for instance, populations from
Aberffraw to Porth Trecastell were dominated by plants with appressed hairs and red standards
with white keels (other forms being rare), whilst those at South Stack all had yellow corollas and a
mixture of hair types.

Plants equating to subsp. corbierei were found only at South Stack, Addoloy-Penrhosfeilw
chapel and Porth Trecastell. No plants were refound at Porth Nobla, Rhosneigr or Aberffraw,
though the exact original sites are not known and vegetative plants with spreading hairs were
found in the latter in 1993 (R. A. Jones, pers. obs.).

At the type locality at South Stack there was significant variation in the growth form. Some
plants in deep turf sheltered by rocks were erect, but others in more exposed positions were
decumbent (in cultivation, some variation in growth form is retained; photograph in NMW). The
leaves were somewhat fleshy (thick by comparison with inland plants), but the cauline leaves did
not appear to be noticeably concentrated in the lower part of the stem (this is also true of the type
specimens at LIV and BM). All plants seen had + uniformly yellow corollas. Four phenotypes
were present, differing in the combinations of spreading or appressed hairs and red-tipped or
green-tipped calyces, but otherwise identical. For plants with spreading hairs, there was some
variation in the density and distribution of hairs. Most plants had dense spreading hairs along the
length of the stem, though a few had sparse spreading hairs, and one was noted with spreading
hairs below and appressed hairs on the peduncles. There was also a little variation in the redness of
the calyces, with some plants with only the very tips red.

CORNWALL

Plants with spreading hairs equating to subsp. corbierei were found at five of the previously-
reported localities (Hayle Towans, Tubby’s Head, Perranporth, Porth Joke and Pentire Point East),
and at two new sites (Watergate Bay and Rocky Valley) (Table 1). No plants were refound at
Bedruthan Steps or Tintagel, but they could still be present. Plants with spreading hairs may be
widespread around the Cornish coast, but they are not present in every population.

The north Cornish Anthyllis populations are highly polymorphic, and probably the most variable
in Britain. In all localities, spreading hairs occurred with practically all other combinations of
corolla and calyx colour, and such plants could not otherwise be separated from the variation in
plants with appressed hairs. They were quite variable in growth form, probably related to
exposure.

ANTHYLLIS VULNERARIA SUBSP. CORBIEREI 473

TABLE 1. PHENOTYPES OF ANTHYLLIS PRESENT IN SELECTED SITES IN ANGLESEY,

CORNWALL AND DORSET
SITE PHENOTYPES
Anglesey (June 1999)
Tywyn Aberffraw (SH356688) WAT
Ynys Meibion, cliffs opposite (SH327686) RAT, WAT
Porth Trecastell, cliffs and roadside (SH3370 area) OAT, RAT, WAT, YAG, YAT, YST
Rhosneigr, roadside (SH319726) VAT
Rhosneigr, roadside (SH324739) YAT
Black Creek, Bodior, Rhoscolyn, rocks (SH294757) YAG, YAT
Gromlech, Rhoscolyn, cliffs and tracks (SH262764 area) OAT, RAT, WAT, YAT
Porth Diana, Trearddur (SH253782) WAT, YAT
Porth-y-post, Trearddur, roadside (SH243796) WAT, YAT
Addoloy-Penrhosfeilw chapel, roadside (SH226806) YAT, YST
Pen-y-bonc, Penrhosfeilw, roadside (SH217812) YAT, YAG
South Stack, cliffs (SH2082 area) YAT, YAG, YSG, YST
Porth Tywyn-mawr, rocks (SH285850) YAG, YAT
Ynys y Fydlyn, Llanfairynghornwy, cliffs (SH292917) RAT, WAT, YAT
Cornwall (June 2000)
Hayle Towans, fixed dune grassland (SW554385) OAT, RAT, WAT, YAG, YAT, YSG, YST
Porthtowan, cliffs (SW69248 1) OAT, RAT, WAT, YAG, YAT
Chapel Porth, cliffs (SW697495) OAT, RAT, WAT, YAG, YAT
Tubby’s Head, cliffs (SW698505) OAT, WAT, YAG, YAT, YSG, YST
Droskyn Point, Perranporth, cliffs (SW755545) OAT, WAT, YAG, YAT, YSG
Porth Joke, cliffs (SW772604) OAT, OST, WAT, YAG, YAT, YSG, YST
Pentire Point East, grassland (SW782615) OAT, OST, RAT, RST, WAT, WST, YAG, YAT,
YSG, YST
Watergate Bay, cliffs (SW841648) OAT, OST, RAT, WST, YAG, YAT, YSG, YST
Bedruthan Steps, cliffs (SW8469) OAT, RAT, WAT, YAG, YAT
Tintagel, cliffs (SX0589) OAT, RAT, WAT, YAG, YAT
Rocky Valley, cliffs (SX073896) OAT, YAG, YAT, YSG, YST
Dorset (July 2000)
South of Fortuneswell, Portland, old quarry (SY693731) YAG, YSG, YST
Easton, Portland, car park (SY696723) YAG, YAT, YSG, YST
Handfast Point, Ballard Down, grassland (SZ054825) YAG, YAT, YSG, YST

See text for coding of phenotypes

DORSET

Three sites were investigated, all of which had a mixture of phenotypes (Table 1). Plants equating
to subsp. corbierei were found in all three, though they had less dense spreading hairs than plants
in Anglesey and Cornwall, and all had appressed hairs above. Again, plants with spreading hairs
covered the same range of variation in corolla and calyx colour as those with appressed hairs.
Growth form also varied markedly between sites.

CHANNEL ISLANDS
Plants on Sark had spreading hairs, yellow corollas and red-tipped calyces (M. Marsden, R. Veall,
pers. comms., 1999). Plants with appressed hairs have also been recorded on the island (K; see
above). Similarly, on Guernsey the plants with spreading hairs had yellow corollas and red-tipped
calyces, and occurred in mixed populations with plants with appressed hairs (B. Ozanne, R. Rabey,
pers. comms., 1999).

The field work showed that most of the larger populations are polymorphic, and that plants with
spreading hairs were otherwise identical to those with appressed hairs and always grew with them.

474 TCG, RICH
VARIATION IN FREQUENCY OF PHENOTYPES WITHIN POPULATIONS

The variation of the corolla, calyx and hair characters prompted further analysis to see if plants
with spreading hairs were most commonly associated with yellow corollas and green-tipped
calyces (cf. Cullen’s 1986 description), or occurred in random combinations with all corolla and
calyx colours. The frequency of characters was scored for 100 plants along transects at four sites.

The numbers of plants with each combination of characters recorded in three transects at the
type locality in Anglesey are shown in Table 2 (all plants had yellow corollas). 27% of the plants
had spreading hairs (range on transects 24-32%), and 69% of the plants had green-tipped calyces
(range 66-74%). All phenotypes were scattered + randomly along the transects, though often
patchily. A y? analysis showed that the calyx and hair characters occur together in random
combinations independent of each other (y? = 0-42, p>0-1), and there is no particular association of
green-tipped calyces with spreading hairs (typical ‘corbierei’ ) or red-tipped calyces with appressed
hairs (typical ‘vulneraria’ ).

Single transects were recorded at Hayle Towans, Pentire Point East and Watergate Bay in
Cornwall (Table 3). The composition of the populations differed between sites, and the population
was especially variable at Pentire Point East. The percentages of plants with spreading hairs at
these sites were 14%, 28% and 15% respectively. Again, a y? analysis showed that the corolla,
calyx and hair characters occurred together in random combinations (y? = <0-001, p>0-9;
x? = 0:58, p>0-:5; 7? = 0-03, p>0-9 respectively).

TABLE 2. NUMBER OF PLANTS OF EACH PHENOTYPE RECORDED IN EACH OF
THREE TRANSECTS OF 100 PLANTS AT TYPE LOCALITY, SOUTH STACK, JUNE 1999

Phenotype NW facing SW facing SW facing

grassy slope grassy slope rocks Mean
YAG 49 3D 4 49-33
YAT (‘vulneraria ) De 19 24 23°33
YSG (‘corbiere?’) 7 19 23 19-66
YST 7 7 9 7:70

See text for coding of phenotypes

TABLE 3. NUMBER OF PLANTS OF EACH PHENOTYPE RECORDED IN TRANSECTS OF
100 PLANTS AT CORNISH SITES, JUNE 2000

Phenotype Hayle Towans Pentire Point East Watergate Bay
OAT l 18 9
OST 0 8 1
RAT 0 11 3
RST 0 Z 0
WAT 0 3 0
WST 0 4 2
YAG DS 6 58
YAT (‘vulneraria ) 60 34 16
YSG (‘corbierei’ ) 9 2 8
YST 5 12 6)

See text for coding of phenotypes

ANTHYLLIS VULNERARIA SUBSP. CORBIEREI 475
DISCUSSION

The herbarium material showed that plants with spreading hairs vary in morphology between sites
in Britain (cf. Figure 1) and France. The field investigation also showed that much variation occurs
within populations, in which plants with spreading hairs could not otherwise be separated from the
variation in those with appressed hairs. Whilst there were always some plants with spreading hairs
and yellow corollas which conformed to Cullen’s description of subsp. corbierei in the populations
investigated (allowing for the variation in calyx colour which I do not regard as of taxonomic
significance), there were other plants with spreading hairs which did not. The analysis of the
frequency of phenotypes within four populations showed that spreading hairs occurred in
apparently random combinations with corolla and calyx colour.

Cullen (1976) described the type population of subsp. corbierei at South Stack as homogeneous,
but the field work showed that this is not true for hair type, the essential character on which the
original variety was based. Indeed, Salmon & Travis (1917) had already noted that most of the
plants were typical with regard to hair-clothing (i.e. had appressed hairs).

Genetic experiments are required to establish the basis of inheritance of the spreading and
appressed hair characters, though the complex variation in pubescence seen suggests it is unlikely
to be a simple Mendelian inheritance controlled by one gene. Marsden-Jones & Turrill (1933a)
cultivated plants to investigate the genetic basis of variation (primarily corolla colour), but the only
observation reported on hair type was that three out of 20 plants cultivated from wild seed
collected from one plant at Par Harbour (R.21) had spreading hairs on the stems, the others having
appressed hairs. Three seedlings grown by T.C.G.R. from seed collected from a plant with
spreading hairs at South Stack also had spreading hairs showing that it can breed true.

I am thus unable to delimit a homogeneous taxon which can be equated with Cullen’s subsp.
corbierei, and which can be consistently distinguished from variation in other Anthyllis vulneraria
with which it always grows. My current interpretation of the hair characters in Britain is that they
are polymorphic, and that plants with spreading hairs do not merit recognition at subspecific rank.
Indeed, plants with spreading hairs occurred in collections which would otherwise be treated as
subsp. vulneraria or subsp. lapponica. What is interesting is that spreading hairs occur most
frequently in populations of the western Atlantic coast, indicating some ecological and
distributional integrity. There is little value in giving names to each random combination of
characters (ten names would be required to name the different combinations of hair, corolla and
calyx characters observed at Pentire Point East alone, cf. Table 3), and I follow Marsden-Jones &
Turrill’s (1933b) advice in refraining from making new combinations until the genetic basis has
been resolved. Meanwhile, I regard subsp. corbierei as synonymous with subsp. vulneraria and
suggest it should be removed from the Red Data Book (Wigginton 1999).

ACKNOWLEDGMENTS

This work was stimulated by Andy Jones who kept asking questions that were difficult to answer,
at least initially, and I am grateful for his prompting and assistance. I would like to thank the
Keepers of the herbaria for loan of or access to material, and Arthur Chater, John Edmondson,
Sally Ellis, Trevor Evans, Vera Gordon, Richard Gornall, Gwyll Lewis, Alex Lockton, Marcia
Marsden, David McClintock, Andy McVeigh, Bridget Ozanne, Alison Paul, Chris Preston, Rachel
Rabey, Benoit Toussaint, Roger Veall, Rosie White and Goronwy Wynne for assistance with field
work or specimens, Andrew Pang for drawing the silhouettes, and Jane Croft for details of records
at the Biological Records Centre. The field work in England was funded by the Threatened Plants
Database. The map in Figure | was plotted using DMAPW by Alan Morton.

REFERENCES

AKEROYD, J. R. (1998). Anthyllis vulneraria, in RICH, T. C. G. & JERMY, A. C. (1998). Plant Crib 1998. pp.
181-182. Botanical Society of the British Isles, London.

ALLEN, A. & HILTON, B. (1994). Flowers of Sark. Privately published, Barnstaple.

CORBIERE, L. (1894). Nouvelle Flore de Normandie. Caen.

476 TC-GaRICH

COUDERC, H. (1975). Contribution a la révision des taxons infraspécifiques de l’Anthyllis vulneraria L. en
France. Revue Générale de Botanique (Paris) 82: 93-118.

CULLEN, J. (1967). Anthyllis vulneraria L. Watsonia 6: 295.

CULLEN, J. (1968). Anthyllis L. in TUTIN, T. G. et al. eds., Flora Europaea 2: 177-182. Cambridge University
Press, Cambridge.

CULLEN, J. (1976). The Anthyllis vulneraria complex: a résumé. Notes from the Royal Botanic Garden
Edinburgh 35: 1-38.

CULLEN, J. (1986). Anthyllis in the British Isles. Notes from the Royal Botanic Garden Edinburgh 43: 277—
281.

JERMY, A. C. & CRABBE, J. A., eds., (1978). The island of Mull. British Museum (Natural History), London.

MARGETTS, L. J. & DAVID, R. W. (1981). A review of the Cornish flora. Institute of Cornish Studies, Redruth.

MARSDEN, M. (1994). A new check list of flowering plants and ferns wild on Sark. Report and Transactions
Société Guernesiaise 1994: 754-783.

MARSDEN-JONES, E. M. & TURRILL, W. B. (1933a). Studies in variation of Anthyllis vulneraria. Journal of
genetics 27: 261-285.

MARSDEN-JONES, E. M. & TURRILL, W. B. (1933b). Notes on the taxonomy of British material of Anthyllis
vulneraria. Journal of botany 71: 207-213.

MCCLINTOCK, D. (1975). The wild flowers of Guernsey. Collins, London.

ROBERTS, R. H. (1982). The flowering plants and ferns of Anglesey. National Museum of Wales, Cardiff.

SALMON, C. E. & TRAVIS, W. G. (1917). Notes from Carnarvon and Anglesey. Journal of botany 55: 320.

STACE, C. A. (1997). New Flora of the British Isles. 2nd ed. Cambridge University Press, Cambridge.

WIGGINTON, M. J. (1999). Anthyllis vulneraria ssp. corbierei (Salmon & Travis) Cullen (Fabaceae), in
WIGGINTON, M. J., ed., Red Data Books of Britain and Ireland 1: Vascular plants. 3rd ed. p. 40. Joint
Nature Conservation Committee, Peterborough.

(Accepted February 2001)

ANTHYLLIS VULNERARIA SUBSP. CORBIEREI AT7

APPENDIX 1. ANTHYLLIS VULNERARIA V ARIANTS IN THE BRITISH ISLES WITH
MOST OF THE STEM WITH SPREADING HAIRS

WEST CORNWALL, V.C. 1

Lizard, 4 June 1926, A. H. Wolley-Dod; plants with red calyces, some of which have spreading
hairs but the sheet is named by C. E. Salmon as var. coccinea and not var. corbierei (BM). Housel
Bay, Lizard, 6 June 1926, A. H. Wolley-Dod (BM). Caethillean Bay, Lizard, 2 June 1900, no
collector (BRISTM). Cadgwith, cliffs, 26 May 1900, no collector (BRISTM). St Agnes cliffs, 27
and 28 May 1913, F. Rilstone; red corollas, appressed hairs on peduncles (NMW). St Agnes cliffs,
19 June 1913, F. Rilstone (NMW). St Agnes Head, 28 May 1989, C. D. Preston/BSBI meeting;
very variable population, some plants of which det. as subsp. corbierei by J. R. Akeroyd, 1992
(CGE). Mullion, cliff top on serpentine, 11 June 1969, P. D. Sell, D. E. Coombe & S. Wegmiiller;
dense spreading hairs except top peduncle (CGE). Perranporth cliffs, 22 May 1952, T. J. Wallace;
mixture of phenotypes present in small area (LIV). Headland by Droskyn Castle Hotel,
Perranporth, 26 May 1952, D. A. Reid (K). Hayle Towans, 8 June 1933, A. A. Bullock, red-tipped
calyces, and both red and yellow corollas (K; Cullen 1976, cited incorrectly as Hazel Towans). It
was also reported from Bedruthan Steps, 1958, J. H. Chandler (Margetts & David 1981; no
material traced) and at Porth Joke and Pentire Point East in 1962 (Biological Records Centre; no
further details known).

EAST CORNWALL, V.C. 2
Tintagel, 23 May 1961, I. W. Evans (BRISTM).

SOUTH DEVON, V.C. 3
Salcombe Regis beach, June 1845, R. Cupnell; mixture of plants with spreading and appressed
hairs (TCD).

NORTH SOMERSET, V.C. 6

Railway embankment near Winscombe, 10 June 1933, H. S. Thompson; spreading hairs
throughout, though sparse above (LIV). Congresbury, May 1882, D. Fry; sparsely hairy
(BRISTM).

DORSET, V.C. 9

Portland Bill, 21 May 1938, herb. Dunston (BM). Old quarry south of Fortuneswell, Isle of
Portland, 16 May 1987, S. L. Jury & M. F. Watson no. 8195; four plants with red-tipped calyces,
det. J. A. Akeroyd, 1988 (RNG). Car park bank, Easton, Portland, 1999, R. White (specimens
shown to T. Rich, not retained). Ballard Down, Studland, 17 May 1959, W. S. Cattling; one plant
with red-tipped calyces ‘varying towards subsp. vulneraria...,’ det. J. R. Akeroyd, 1988 (RNG).

ISLE OF WIGHT, V.C. 10

Woody Point, 3 July 1925, R. Melville (NMW). Ventnor, June 1888, J. H. A. Stueart; red-tipped
calyx, spreading hairs all way up (LIV). Carisbrook Castle, 13 June 1827, no collector (CGE) and
1865, F. Stratton (OXF); material in other herbaria from this site is more variable.

KENT, V.C. 15 AND 16

Material with sparse spreading hairs, sometimes along the whole length of the stem, has usually
been determined as A. vulneraria subsp. vulneraria var. vulneraria forma polyphylloides Sagorski
in a number of herbaria (e.g. Durham Downs, 29 May 1948, J. E. Lousley; det. J. Cullen, CGE).

SURREY, V.C. 17
Field on chalk near Merstham, 28 April 1945, N. Y. Sandwith (BRISTM).

MONMOUTH, V.C. 35
Coastal bank, Sudbrook, June 2000, T. G. Evans (NMW).

WORCESTER, V.C. 37
Broadway, 8 June 1898, L. Rea; sparse spreading hairs throughout (BM).

478 T.CsG-RICH

GLAMORGAN, V.C. 41

Baldwin’s Tip, Crymlyn Burrows, 19 June 1939, J. A. Webb; corollas purplish, one of five
specimens with spreading hairs (NMW). Taffs Well, railway embankment, 21 June 1927, A. E.
Wade (NMW).

CARMARTHEN, V.C. 44
Machynys, urban waste ground, 23 June 1992, I. K. Morgan (NMW).

PEMBROKE, V.C. 45

Trevine, grassy cliff, 16 June 1963, T. A. W. Davis (BSBI field meeting) ‘calyx concolorous, one
plant with calyx-tipped reddish. Plants rather small, those with hairy stems a small proportion of a
large population..’; “shaggy form of subsp. vulneraria var. langei... J. Cullen 1963’; some plants
also with spreading hairs below only (LIV; NMW). St Anne’s Head, Dale, 5 June 1963, T. A. W.
Davis; peduncle with appressed hairs (LIV).

CARDIGAN, V.C. 46

Shaley cliffs above the sea west end of Cribach Bay, R. A. E. site, Aber-porth 5 June 1994, A. O.
Chater; calyces purple-tipped, plants with appressed hairs also present (NMW). Dry, shaley bank
by track above Cribach Bay, R.A.E. site, Aber-porth, 25 May 1997, A. O. Chater; calyx purple-
tipped (NMW). Dry shaley track side bank near west end of R. A. E. site, Aber-porth, 3 May
1997, A. O. Chater; calyx purple-tipped (NMW).

DENBIGH, V.C. 50
Bryn Euryn, Colwyn Bay, 1966, W. E. Hughes; appressed hairs on peduncles (NMW)

FLINT, V.C. 51
Prestatyn, 1923, Miss B. Allen; with plant with appressed hairs (herb. G. Wynne).

ANGLESEY, V.C. 52

Holyhead Island, Anglesey, June 1850, J. Ball; red corollas (E). Holyhead, Anglesey, 4 June 1859,
J. Ball; det. J. R. Akeroyd 1986 (E). South Stack, June 1916, W. G. Travis (syntype of var.
corbierei and holotype of subsp. corbierei LIV; syntype also at BM; with the type at LIV is
another specimens with the same details but with appressed hairs, cf. Salmon & Travis 1917).
Trearddur Bay, July 1970, A. Jones; mixed forms with spreading and appressed hairs (NMW).
South Stack, 28 July 1998, R. A. Jones; with plants with appressed hairs (NMW). Cliffs near
Holyhead, 3 June 1960, V. Gordon (exact site not noted, possibly c. ¥2 mile south of South Stack;
pers. comm. 1999); mixed gathering, one plant with red corollas and spreading hairs (NMW).
Coast near Aberffraw, 9 July 1927, A. Wilson; one specimen only with red corollas and spreading
hairs, sparsely so on peduncle (NMW). According to Cullen (1976) there are specimens he
collected from South Stack, 29 March 1958 and Rhosneigr, 22 March 1961 in Liverpool
University herbarium (LIVU); this collection was transferred to Liverpool Museum (LIV) but the
specimens could not be traced in 1999. A collection from Rhosneigr, 9 May 1961, J. M. Scott has
all plants with appressed hairs only and was det. by J. Cullen as intermediate between subsp.
corbierei and subsp. vulneraria (LIV).

MID PERTH, V.C. 88

There are several collections of plants with spreading hairs either all the way up or not including
the last peduncle from the Breadalbane Mountains which would otherwise be referable to subsp.
lapponica. (e.g. rock ledges, Creag na Caillich, 6 August 1933, J. E Lousley; RNG, and other
forms with spreading hairs all way up in OXF and NMW).

MID EBUDES, V.C. 103

Staffa, 2 July 1969, British Museum Mull Survey, no. 3706; ‘basically subsp. /apponica .. but also
reminiscent of subsp. corbierei, J. Cullen, 1971’ (BM; Jermy & Crabbe 1978). Ardalanish, Mull,
22 May 1971, A. G. Kenneth & A. McG. Stirling; noted both by J. Cullen and J. R. Akeroyd to be
subsp. lapponica but varying towards subsp. corbierei (BM).

WEST SUTHERLAND, V.C 108
Bettyhill, July 1951, F. Rose; variably hairy, spreading hairs sparse or absent on peduncle
(NMW).

ANTHYLLIS VULNERARIA SUBSP. CORBIEREI 479

CAITHNESS, V.C. 109

Duncansby Cliff, Caithness, May 1928, R. Meinertzhagen; a distinctive plant with lemon-coloured
corollas, and close to corbierei but intermediate with subsp. lapponica (BM). Sea cliffs east of
Port of Brims, 1 August 1988, T. C. G. Rich & G. M. Kay, det. J. R. Akeroyd, 1988 (collected as
subsp. lapponica; specimen not traced). Holborn Head, | July 1936, J. E. Lousley, det. as subsp.
vulneraria but varying towards corbierei, J. R. Akeroyd (RNG). Dunbeathe, riverbank near the
sea, 11 July 1968, M. McC. Webster; sparse patent hairs (CGE).

CHANNEL ISLANDS

Guernsey. Le Jaonnet, 31 May 1970, D. McClintock, det. J. Cullen (herb. Société Guernesiaise;
not seen T.C.G.R.). Le Jaonnet, base of steps, June 1999, B. Ozanne; all with spreading hairs,
calyx red-tipped (NMW). Le Jaonnet summit, 27 June 1999, R. Rabey; mixed population with
spreading and appressed hairs, all corollas yellow and calyx red-tipped (NMW). It was also
reported from La Bette Bay adjoining Le Jaonnet Bay by M. English, 1957 (McClintock 1975),
but was not refound in 1999 and no material has been traced.

Sark. Cliff at Creux harbour, 11 June 1929, F. Ballard & E. A. Golby; red-tipped calyces, and
some material with appressed hairs (K; Cullen 1976). Creux harbour, 14 May 1999, R. M. Veall;
two plants with red-tipped calyces (NMW). La Coupée, 25 May 1965, A. A. Bullock (K). La
Coupée, 4 June 1986, R. J. Murphy; one plant with red-tipped calyx (BM). La Coupée, 11 June
1999, Mrs M. H. Marsden; three plants with red-tipped calyces (NMW). According to some of the
literature, only subsp. corbierei is present on the island (Allen & Hilton 1994; Marsden 1994).

IRELAND

No material has been found in all the herbaria examined, though a very few plants with sparse
spreading hairs below have been seen (e.g. Killiney Hill, 1849, no collector, TCD). J. R. Akeroyd
has also not seen any plants with spreading hairs in the field (pers. comm., 2001).

480 T. C. G. RICH

APPENDIX 2. ANTHYLLIS VULNERARIA FORMS WITH SPREADING HAIRS IN
NORTHERN FRANCE

MANCHE

Jobourg et Herqueville, 4 Juillet 1883, herb. L. Corbiére (as var. sericea Breb.); mixed collection,
one with spreading hairs throughout and others with spreading hairs below (CHE). Herqueville,
falaises, 20 Juin 1886, herb. L. Corbiere (as var. sericea Breb.); spreading hairs throughout though
sparsely above (CHE).

CALVADOS

Cotes de Calvados, dans le falaises, 1838, A. Lenormand (P). Pentes des falaises calcaires du
rivage de la Manche a Fontenailles, 18 Juin 1890, Bertot; spreading hairs throughout (CHE, P).
Falaises maritime, Longues prés Bayeux, 8 Aout 1872, L. Corbiére (as var. sericea Breb.);
spreading hairs throughout though very sparsely above (CHE). Falais d’ Arromancher, Calvados,
Juillet 1846, E. Cosson, det. “forma inter subsp. vulgaris (Koch) Hayek et subsp. polyphylla (Kit.)
Cab.’ by J. Cullen, 1961 (P).

SIENE-MARITIME

Falaises de Fécamp, Siene-Inferieur, 12 Juin 1892, I. Thériot & L. Corbiére (as var. sericea Breb.);
sparse spreading hairs throughout (CHE, P) and I. Thériot; one plant with spreading hairs
throughout and others with spreading hairs below (CHE).

PAS-DE-CALAIS
Pernes, 19 July 1995, B. Destines (Herbarium CRP/CBNBL). Setques, 8 June 1995, B. Toussaint
(Herbarium CRP/CBNBL).

Watsonia 23: 481-491 (2001) 48]

The distinction between Cerastium tomentosum L. and
C. biebersteinii DC. (Caryophyllaceae), and their occurrence in
the wild in Britain

M. K. KHALAF and C. A. STACE

Department of Biology, University of Leicester, Leicester LEI 7RH

ABSTRACT

Differences between Cerastium tomentosum and C. biebersteinii are summarised. Due to the enormous
variation of the former in most readily observable characters, certain identification can be effected only by
using features of the morphology of the capsule teeth (not present in most material) and the anatomy of the
non-glandular hairs (a microscopic character). No material of C. biebersteinii has been seen from the British
Isles other than in a few botanic gardens, and there is no good evidence of its existence here either as a garden
ornamental or in the wild, nor of its hybridization with C. tomentosum. Hybrids between C. tomentosum and
C. arvense are frequent both in cultivation and in the wild.

KEYWORDS: Cerastium, morphology, chromosome numbers, hair anatomy, hybridization.

INTRODUCTION

Cerastium tomentosum L. and C. biebersteinii DC. belong to a group of up to twenty species that
can conveniently be termed the C. tomentosum group. This was recognised by Schischkin (1936)
as series Grandiflora (Borza) Schischk. in the following classification:

Subgenus Dichodon (Bartl.) Fenzl
Subgenus Eucerastium (Boiss.) Pax (= Cerastium)
Section Strephodon Ser.
Section Schizodon (Fenzl) Schischk.
Section Orthodon Ser. (= Cerastium)
Subsection Fugacia (Fenzl) Pax & K. Hoffm.
Subsection Perennia (Fenzl) Pax & K. Hoffm. (= Cerastium)
Series Lasiostemona (Fenzl) Schischk.
Series Leiostemona(Fenzl) Schischk.
Series Alpina (Borza) Schischk.
Series Arvensia (Hayek) Schischk. (= Cerastium)
Series Grandiflora (Borza) Schischk.

The names of four of the above taxa have to be corrected to Cerastium, as shown, since C. arvense
L. is the type of the genus.

None of these five series is well defined, however, and certain hybrids between species in the
last three can be synthesised with ease (Khalaf & Stace 2000). The C. tomentosum group was
defined by Buschmann (1938) by its perennial habit, presence of many non-flowering axillary
shoots, relatively large flowers and presence of (usually very dense) long white hairs. Although
these characters are not strictly diagnostic of the group, either individually or in combination, all
apply to C. tomentosum and C. biebersteinii, which can be considered among the core species of
this informal grouping. Subsection Perennia (= Cerastium), as redefined by Sdllner (1954) when
he removed series Leiostemona (which includes C. fontanum Baumg.), seems to be a well defined
taxon which is best not subdivided above the species level (Khalaf & Stace 2000).

Cerastium tomentosum itself is an extremely variable taxon, and there have been considerable
divergences of opinion about its delimitation. Moggi (1963) pointed out that at least six species

482 M. K. KHALAF AND C. A. STACE

had been segregated from it in Italy and Sicily alone. On the other hand the name has been used
for plants from Spain (e.g. Linnaeus 1753), the Alps (e.g. Nyman 1855), the Balkans (e.g.
Stojanoff & Stefanoff 1924), the Crimea (fide Schischkin 1936) and the Caucasus (e.g. Fenzl
1842). Today it is recognised as endemic to Italy and Sicily, where no other taxa from the C.
tomentosum group occur. Of all its variations, specimens from higher altitudes on Mt Etna, Sicily,
are the most distinct in having only sparse hairs, narrower leaves, shorter petals and capsules and
smaller seeds; these can be recognised as var. minus C. Presl (var. aetnaeum Janka).

The only other Linnaean species possibly in the C. tomentosum group, Cerastium repens L.
from France and Italy, is a name of uncertain application. No type specimen has been traced; the
specimen in LINN is not a type and is in our opinion referable to C. arvense, a conclusion
apparently behind De Candolle’s (1822) inclusion (with some doubt) of the name under the latter
species. It has also been used for the Italian species (C. tomentosum) by Buschmann (1938) and for
the Crimean species (C. biebersteinii, see below) by Marschall von Bieberstein (1808).

The first species recognised in the group from outside the Mediterranean area were C. repens M.
Bieb. (from Crimea) and C. argenteum M. Bieb. (from Tiflis, now Tbilisi in Georgia), described
by Marschall von Bieberstein (1808). The Crimean species was realised by De Candolle (1822) to
be different from C. repens L., so he renamed it C. biebersteinii. De Candolle’s material consisted
of cultivated plants originating from material collected by Steven in Crimea. Buschmann (1938)
noted “I have seen the plants which were collected by Steven in Tauria [Crimea] in 1821. He
called it C. repens and certainly it belongs to C. biebersteinii’. Schischkin (1936) stated that “C.
biebersteinii is an endemic Crimean plant, very widespread in the mountainous part of the Crimea.
It displays the closest relationship to C. szowitsii Boiss. (Eastern Transcaucasia) and C. argenteum
M. Bieb. (Central Transcaucasia) which are similarly white-tomentose but differ in some other
characters. Early investigators identified the Crimean plant with C. tomentosum, although this
species is confined to the central and southern parts of Italy. Cerastium biebersteinii does not
occur in a wild state outside the Crimea. The report by Taliev and Voinovskii (1902) referring to
the occurrence of this species in inundated meadows of Buguruslan County of Samara Province
[north of the Caspian Sea in Ukraine] is certainly wrong”. However, Buschmann (1938)
considered that C. biebersteinii does occur in Transcaucasia, since he favoured a wider taxonomic
concept of it, including two of Schischkin’s Caucasian species in his C. biebersteinii subsp.
transcaucasicum Buschm. According to Schischkin (1936) the only other member of the
C. tomentosum group to occur in Crimea is C. stevenii Schischk., which is a very different-looking
species and perhaps does not belong to this group at all. It is therefore likely that C. tomentosum
and C. biebersteinii each occur in widely separated areas neither of which supports any other
closely similar taxa.

C. tomentosum var. tomentosum has long been introduced in central and northern Europe as a
garden ornamental. From here it has escaped into the wild and in places has become extremely
well naturalised not only in marginal and disturbed ground but sometimes in natural habitats, such
as sandy grassland and maritime sand-dunes. Already by 1960 it was found scattered in the wild
all over the British Isles (Perring & Walters 1962). It has caused taxonomic problems in two ways.
Firstly, it has quite frequently hybridised (in eastern and south-eastern England and one part of
central Scotland) with the native C. arvense (belonging to a different series from C. tomentosum,
see Classification above) to produce plants that superficially resemble another member of the
C. tomentosum group, the Balkan C. decalvans Schlosser & Vuk., and which have been wrongly
so called. Hybrids between C. tomentosum and C. arvense have also been reported from Canada
(Morton 1975), Sweden (Karlsson 1997) and Luxemburg and Finland (Kaleva 1966), as well as in
several botanic gardens throughout Europe. Secondly, there are several reports of the introduction
to gardens of the Crimean C. biebersteinii DC., and the variability of C. tomentosum has often
been partly explained by determination of some naturalised plants as this or as various Balkan
species of the group. Clapham (1987) stated that C. biebersteinii “is also much grown in gardens”
and that “some cultivated material may well be of hybrid origin”. Walters (1989) claimed that both
C. biebersteinii and Caucasian taxa were introduced in the 19th century and that “complex
hybridization seems to have taken place in cultivated plants”.

As part of a study of the breeding relationships of subsection Perennia sensu Schischkin, we
attempted to describe precisely the differences between C. tomentosum and C. biebersteinii and to
investigate the occurrence in the wild of these two taxa in Britain.

CERASTIUM TOMENTOSUM & C. BIEBERSTEINII 483

Due to their wide geographical separation the two species are not often treated in the same Flora.
Fenzl (1842) covered both species, but his C. tomentosum from “provinciis caucasicis. Iberia”
must have been misapplied. Boissier (1867) stated that C. biebersteinii has a denser tomentum,
longer narrower sepals and capsule teeth with flat (not revolute) margins. Buschmann (1938)
separated C. biebersteinii from a group of species including C. tomentosum in his key (couplet 8):
capsule straight, strong-walled, with patent teeth flat on the margins in the former; capsule straight
or curved, soft-walled, with erect teeth revolute at margins in the latter. Jalas et al. (1964) agreed
about the narrower sepals and flat capsule-teeth in C. biebersteinii, and added shorter stems,
shorter and narrower leaves and erect (not patent, contrary to Buschmann) capsule teeth. Walters
(1989) included both species in the “C. tomentosum group” and did not attempt to distinguish
them. Kaleva (1966) is apparently the only person to have studied these species in a scientific and
comparative way. He listed eight differences: growth habit; leaf size and shape; petal shape;
capsule size; capsule wall anatomy; seed size; pollen grain size; and chromosome number.
However, he concluded that the revolution of the margins of the capsule teeth is not a good
discriminator (“+ flat” in both), and he did not mention capsule-teeth posture. Nevertheless his
photographs showed clearly more patent capsule teeth in C. tomentosum, in agreement with Jalas
et al. (1964) but not with Buschmann (1938).

MATERIALS AND METHODS

Living accessions were obtained as plants or seeds by collecting in the wild, from correspondents
in Continental Europe, and by the seed-exchange schemes. Frequently material from the last of
these sources was misidentified. Much herbarium material was also examined. Up to 25
measurements were made of each character for each accession. Unfortunately, our observations on
C. biebersteinii were restricted to two living accessions and about a dozen herbarium specimens,
few of which carried ripe capsules.

Seed accessions were germinated at 15—19°C in a 3:7:2 loam:peat:grit mixture under a light
regime of 16 hr light/8 hr dark. Germination occurred between 4 and 7 days; seedlings were potted
on and after 1-2 months were transferred to an unheated glasshouse where they were kept
permanently. Plants flowered much more profusely after vernalization over winter in a cold
glasshouse. Vouchers of all accessions at flowering are deposited in LTR.

Chromosome counts were made on root-tips obtained from fresh stem-cuttings after 10—15 days;
these possessed larger meristems and a higher mitotic index than root-tips gathered from mature
plants or from seedlings. They were pretreated in 50% saturated aqueous paradichlorbenzene,
washed in tap-water, fixed for 3 hr in ethanoic acid:ethanol (1:3), washed again and stored in 70%
IMS. After storage they were rewashed, hydrolysed in SN HCI at room temperature for 3—6 min,
washed in 70% ethanol, and transferred to a drop of 45% ethanoic acid on a slide. The
meristematic region was dissected from the rest of the tissue and transferred into a drop of 2%
aqueous aceto-orcein on another slide. The meristem was dissected apart and squashed in the usual
way and then gently heated over a spirit lamp to flatten the cells, and the coverslip was sealed with
rubber solution. Suitable spreads were photographed.

Thirty characters (Table 2) were measured by us for all the material at our disposal to see
whether any discrimination could be achieved. Means per plant, overall ranges or ranges of means
were used according to which we considered the most diagnostically useful.

RESULTS

CHROMOSOME NUMBERS

Table 1 shows that all our accessions from the wild in Britain of C. tomentosum var. tomentosum,
C. arvense and their hybrids had 2n = 72, representing the octoploid level, except that six of the
seventeen accessions of C. tomentosum had up to nine B-chromosomes. Octoploid counts are also
the commonest ones reported in the literature for C. tomentosum and C. arvense, but in addition
there are tetraploid and dodecaploid counts (2n = 36 and 108) for C. tomentosum, and tetraploid
and decaploid (2n = 36 and 90) for C. arvense (Khalaf & Stace 2000). However, the three previous
counts of C. arvense and one of C. tomentosum from Britain (Brett 1955, Blackburn & Morton

484

M. K. KHALAF AND C. A. STACE

TABLE 1. CHROMOSOME COUNTS OF WILD MATERIAL FROM THE BRITISH ISLES OF
CERASTIUM TOMENTOSUM VAR. TOMENTOSUM, C. ARVENSE AND THEIR HYBRID

Site Vice-county Grid Ref. Count
C. tomentosum var. tomentosum
CER 4 Gorebridge Midlothian NT353605 2n = 72 + 1-3B
CER 5 Blackford Quarry Midlothian, NT261702 2n = 72+ 5B
CER 206621 Shingle, Snettisham W. Norfolk TF648328 2n= 742
CER 25 Shingle, Snettisham W. Norfolk TF648333 2n= 12
CER 26 Shingle, Snettisham W. Norfolk TF648333 2n=72 + 2B
CER 31 Sand-dunes, Holme W. Norfolk TF702442 on = 72
CER 103 Near Dumbarton Rock Dunbarton NS37 Dnj=12
CER 104 Sea front, Crosby S. Lancashire SJ39 2ni= 12
CER 105 Behind beach, Southwold E. Suffolk TM507753 7A a
CER 106 Wall, Old Town, St Mary’s Scilly SV912013 2n=72+7B
CER 118 Wall-top, Llaniestyn Caernarvonshire SH282337 2n = 72 + 9B
CER 126 Near railway, Great Heck, York M.W. Yorkshire SE5920 2n= 72
CER 139, 140 & 141. Heathland, near Ipswich E. Suffolk T 14 A
CER 143 Hedgerow, Orrisdale Isle of Man S$€325930 2n = 72+ 9B
C. arvense
CER 27 Grassy roadside, near Snettisham W. Norfolk TF669336 272
CER 29 Sand-dunes, Holme W. Norfolk TF700442 n= 72
C. arvense x C. tomentosum
CER 6 Borthwick Bank Midlothian NT369603 n= 12
CER 19 Shingle, Snettisham W. Norfolk TF648328 2n= 72
CER 23 Shingle, Snettisham W. Norfolk TF648330 Pi 12
CER 24 Shingle, Snettisham W. Norfolk TF648333 on =z
CER 28 Sand-dunes, Holme W. Norfolk TF700442 2m= 2
CER 30 Sand-dunes, Holme W. Norfolk TF701442 hive

1957) have all been octoploids, as have the numerous counts we have made of British garden
material of C. tomentosum.

The other members of the C. tomentosum group vary from tetraploid (2n = 36) to 16-ploid (2n =
144), and several species are represented by more than one ploidy level (Khalaf & Stace 2000). C.
biebersteinii has been reported as tetraploid (one count of wild Crimean material, Kaleva 1966)
and as octoploid (three counts of botanic garden material); this species has been widely
misidentified and the true identity of the three cultivated octoploids requires confirmation. We
used two accessions of C. biebersteinii, one of wild South Crimean (“Tauria meridionalis”) and
the other of Miinchen botanic garden origin; both were tetraploids (2n = 36).

MORPHOLOGICAL AND ANATOMICAL CHARACTERS

The vast range of morphological features of wild Italian material of C. tomentosum var.
tomentosum strongly suggested to us that many of the characters of C. biebersteinii would be
covered by those of the former taxon, and that few of them would be likely to distinguish the two
species. The results shown in Table 2 can be classified as follows:

i - characters showing differences in the means, or in a good part of the ranges, between the two
species, but with overlap, viz. sepal length and breadth, pollen grain diameter, capsule length,
capsule shape, capsule wall rigidity, capsule tooth length and breadth, capsule tooth margins,
and seed length (also seed breadth, not listed in Table 2).

ii - characters apparently clearly discriminating the two species, viz. presence of rhizomes and
stolons, leaf thickness, filament length, anther length, ovary length, and seed weight.

iii - characters not discriminating between the two species, viz. 13 of the other 16 characters (the
three hair characters are covered below).

CERASTIUM TOMENTOSUM & C. BIEBERSTEINII 485

TABLE 2. DIAGNOSTIC CHARACTERS OF CERASTIUM TOMENTOSUM VAR.
TOMENTOSUM AND C. BIEBERSTEINII

|
|

Character C. tomentosum var. tomentosum C. biebersteinii
Height of flowering stems to tip of highest 200-450 mm 100-300 mm
flower*
Presence of rhizomes and stolons Both Neither
Length of uppermost internode of flowering stem, 13-3-120 mm 63-7-125 mm
i.e. internode between uppermost leaf-pair and
lowermost bract-pair
Mean length of next 3 lower internodes 7-2-57 mm 8-4-69-5 mm
Length and breadth of uppermost pair of leaves 749-7 x 17-8-42 x
on flowering shoot 1-5-8-5 mm 1-9-7 mm
Thickness of same leaves 0- 1-0-2 mm 0-2—0-3 mm
Length and breadth of lowest bracts 4-9.7 x 24 mm 5-9-5 x 2:54 mm
Length of scarious tip of lowest bracts 0-2-1-8 mm 0-5-0-9 mm
No. flowers per inflorescence 2-6-12-2 4-3-10-5
Pedicel length of terminal flower 12-7-42-8 mm 13-7-47 mm
Sepal length and breadth 4-9 x 2-3-4 mm 6-5-9-7 x 2-2-7 mm
Length of scarious tip of sepals 0-2-1-3 mm 0-5—1-1 mm
Petal length and breadth 11-18 x 5-10 mm 17-17:3 x 6-8-7 mm
Ratio petal cleft length/petal length* 0-2-0-3 0-2-0:3
Filament length* 4—5 mm 5-4-5-8 mm
Anther length* 1-1-1 mm 0-7—0-9 mm
Pollen grain diameter (tetraploid) 34-3-36-5 um 35-8-37 um
Pollen grain diameter (octoploid) 35-3—40-1 um -
Pollen grain diameter (dodecaploid) 43—45-5 um -
Ovary length* 2-2-5 mm 1-7-1-:9 mm
Style length* 3-6 mm 34-4 mm
Capsule length* 8-12 mm 10-15 mm
Capsule shape Straight or bent Straight
Capsule wall rigidity Pliable Pliable or rigid
Capsule tooth length, 0-7-1-3 x 1-1-5 x
and breadth (0-2 mm behind apex)* 0-1—0-3 mm 0-3-0-5 mm

Capsule tooth posture
Capsule tooth margins

Erect to patent
mostly revolute

Erect to patent
mostly + flat

Seed length (from hilum to furthest point)* 1-1-9 mm 1-5—2-2 mm
Seed weight per 100 seeds 33-75-42 mg 67—70-5 mg
Upper leaf-surface hair density (1) 5-62:8 4-426
Lower leaf-surface hair density (1) 6-6-5] 12-8-30-2

Length-ratio terminal/subterminal hair-cells

(3-7)4—19(19-8)**

(1)1-1-1-7(1-8)**

Ranges of means per plant, or (*) overall ranges, or (**) range of means with overall ranges in parentheses
(1) Number of hair-bases in circle of diameter of 250 um (0-05 mm”) at midpoint of uppermost pair of leaves

on flowering shoot

HAIR CHARACTERS

Both C. tomentosum var. tomentosum and C. biebersteinii possess only unbranched non-glandular
uniseriate-celled hairs. The great range in visible hair density in the former suggested that this
character would be of no value in distinguishing between the two taxa, and precise counts bear this

out (Table 2).

However, a good difference in hair anatomy was discovered between these two taxa (Table 2).
The uniseriate hairs consist of three to six (usually four or five) cells, of which the terminal one is
very long and the basal one or two (rarely three) are very short. The subterminal cell is much
shorter than the terminal cell in C. tomentosum var. tomentosum (and also in C. tomentosum var.
minus) but only slightly so in C. biebersteinii; the ratio between the two cells clearly separates the

486 M. K. KHALAF AND C. A. STACE

|
Z| \
C4 =

FIGURE 1. Representative leaf-hairs of Cerastium. A: C. tomentosum. B: C. biebersteinii.

two species in all cases (Table 2; Fig. 1). Of the other seven species of the C. tomentosum group
examined in this study, none agrees with C. tomentosum in this character, five (C. argenteum M.
Bieb., C. decalvans Schloss. & Vuk., C. gibraltaricum Boiss., C. lineare All., C. moesiacum Friv.)
agree with C. biebersteinii , and two (C. candidissimum Correns, C. grandiflorum Waldst. & Kit.)
differ from all the others in having branched hairs (of quite different form in the two species -
dendroid in C. candidissimum, T-shaped in C. grandiflorum). In addition some or all plants of
C. gibraltaricum, C. moesiacum and C. decalvans possess unbranched, uniseriate-celled glandular
hairs, which are also present in C. tomentosum var. minus.

CERASTIUM TOMENTOSUM & C. BIEBERSTEINII 487
DISCUSSION

Our chromosome counts lead us to believe that British wild and cultivated material of the above
three species mainly or entirely consists of tetraploid C. biebersteinii and octoploid C. tomentosum
and C. arvense, although of course some clones at other ploidy levels might exist as yet
undetected.

In general appearance C. biebersteinii is a shorter, less mat-forming, more sturdy plant than
C. tomentosum, with larger leaves, a denser, more matted tomentum and longer sepals. However,
the ranges of many characters of C. tomentosum are wider than those of C. biebersteinii and in
several cases completely overlap them. This is due not only to the much greater variation shown
by C. tomentosum but also to the relatively small number of specimens of C. biebersteinii that
were examined. In general our measurements, even the means, show wider ranges than those of
Kaleva (1966), presumably due to our greater sample size and possibly to the fact that our samples
of C. tomentosum included tetraploids and dodecaploids as well as octoploids. Because of this,
some characters previously claimed to separate the two species were found by us to be of virtually
no value, viz. leaf length and breadth, leaf-hair density, petal length and breadth, depth of petal
cleft and capsule tooth posture.

Clapham (1987) and Walters (1989) suggested that there has been hybridization between
C. tomentosum and C. biebersteinii in cultivation; their views are plausible, but we are unable to
substantiate them. We have found no evidence that any spontaneous hybrids between these two
species exist. In our experiments (Khalaf & Stace 2000) no capsules were produced from crosses
between tetraploid C. tomentosum and C. biebersteinii. In crosses using octoploid C. tomentosum
as the male parent some hybrid seeds were produced, of which 7:7% was viable, but no mature
hybrid plants could be obtained. But when octoploid C. tomentosum was used as the female parent
74% of the hybrid seed was viable and mature flowering hybrid plants were obtained. Moreover,
these were fertile to some degree, despite being hexaploids (2n = 54). Specimens have been
deposited in LTR. All the hybrids obtained in the C. tomentosum group resembled their female
parent more closely than their male parent. In the case of the C. tomentosum x C. biebersteinti
hybrids, the length ratio of the terminal/subterminal cells of the non-glandular hairs was well
within the range of that of C. tomentosum.

Hybrids between C. arvense and C. tomentosum occur in the wild in Britain as well as in botanic
and other gardens. They have no valid binomial. They are fertile octoploids and have the potential
of extensive and repeated back-crossing. Similar fertile hybrids were synthesised reciprocally by
us with ease. They can be distinguished from C. tomentosum by their greener appearance, with
much less dense non-glandular hairs, frequent possession of glandular hairs, and less sturdy
growth habit. These characters are shared by C. tomentosum var. minus, which might have such a
hybrid origin.

The 16 characters that show some promise as distinctions between C. biebersteinii and
C. tomentosum var. tomentosum are briefly discussed below.

Presence of rhizomes and stolons

Rhizomes and stolons are certainly more evident and far-reaching in C. tomentosum, but we
cannot be certain of their absence in C. biebersteinii. Field studies in Crimea are required to
elucidate this.

Thickness of uppermost pair of leaves on flowering shoot

According to our measurements this character separates the two species, but further sampling is
required to see whether there is some overlap. In any case this is not a character that is easy to
measure for identification purposes.

Sepal length and breadth

The sepals are usually longer and narrower in C. biebersteinii, but the ranges of both
measurements overlap. The length/breadth ratio provides a better discriminator, but there is still
overlap. Kaleva’s (1966) sepal length measurements were contained within ours and gave a clear
separation of the two species, but with a sample size of only 10.

488 M. K. KHALAF AND C. A. STACE

Filament, anther and ovary lengths

These all gave good separation in our material, but further sampling is desirable to ensure that
there is no overlap. The ovary must be measured at anthesis, before there is any chance of its
swelling after pollination.

Pollen grain diameter

The pollen of the tetraploid C. biebersteinii is similar in size to that of tetraploid and octoploid
C. tomentosum, but that of dodecaploid C. tomentosum is slightly larger. Kaleva’s (1966)
measurements were very close to ours.

Capsule length
Again, there is a difference, but with overlap. Kaleva’s (1966) measurements were very close to
ours.

Capsule shape
Whereas C. biebersteinii always has straight capsules, those of C. tomentosum can be straight or
bent/curved.

Capsule wall rigidity

The walls of the ripe empty capsules of C. ftomentosum are rather soft and pliable, whereas those of
C. biebersteinii are harder and more rigid, often fracturing when pressed with a needle. Kaleva
(1966) showed that this difference is based on the capsule wall anatomy; in C. biebersteinii the
lignified outer wall of the outer epidermal layer is at least twice as thick as in C. tomentosum.
However, we found that some capsules of C. biebersteinii had pliable walls, although this might be
due to the conditions under which the plants grew. Field studies in Crimea are required to check
this.

Capsule tooth length and breadth

The capsule teeth are generally longer and (especially) wider in C. biebersteinii, but there is some
overlap in length and ranges of breadths are contiguous. Breadth measurements are difficult to
compare as the point at which the measurement is taken is critical.

Capsule tooth margins

This is the most frequently cited diagnostic character, and we found it reliable (Fig. 2). The greater
revolution in C. tomentosum renders the apparent capsule tooth breadth more distinct from that of
C. biebersteinii than it would otherwise be. Kaleva (1966), however, found this character to be
unreliable, stating that both species possessed “+ flat” tooth margins. But his “C. tomentosum
strain B”, which he very plausibly concluded was C. tomentosum x C. arvense, had “revolute”
tooth margins. The capsule teeth in C. arvense are even more revolute at the margins than in
C. tomentosum, and it is therefore possible that Kaleva differed from us simply in his verbal
interpretation of the same structures.

Seed length
The ranges of the two species differ, but overlap. Kaleva’s (1966) measurements agree closely
with ours.

Seed weight
Seed weight gave a good separation between the two species; this is a combination of seed length,
breadth and thickness plus any possible anatomical differences.

Length ratio of terminal and subterminal hair-cells

A very clear separation was obtained between C. tomentosum and C. biebersteinii (along with all
other species in the group) (Fig. 1). The difference is so clear that after some experience it is not
necessary to make detailed measurements.

Chromosome number

Octoploid counts of C. biebersteinii need to be confirmed among wild collections; until then it 1s
safer to consider this species a tetraploid. C. tomentosum var. tomentosum is usually octoploid and
var. minus always so. Rarely tetraploid and dodecaploid populations of var. tomentosum are
reported; all our material of these two cytotypes was of botanic garden material, but according to
Favarger (1972) all three ploidy levels occur in the wild in the Italian Apennines.

CERASTIUM TOMENTOSUM & C. BIEBERSTEINII 489

FIGURE 2. Dehisced capsules of Cerastium. A: C. tomentosum. B: C. arvense. C: C. biebersteinit.
CONCLUSIONS

All the material of the C. tomentosum group that we have seen either from the wild or cultivated as
ornamentals in Britain has been referable to C. tomentosum var. tomentosum or to hybrids between
that and C. arvense. Despite the apparent absence of C. tomentosum var. minus and
C. biebersteinii, all three taxa are described here in order to allow future identification:

CERASTIUM TOMENTOSUM L. VAR. TOMENTOSUM

Plant mat-forming, with stolons and/or rhizomes, almost completely covered with simple non-
glandular hairs; length ratio of terminal to subterminal cell (3-7)4—19(19-8); glandular hairs absent;
flowering stems 20—45cm, their uppermost internode 13-3—120 (135) mm. Uppermost leaves on
flowering shoots linear to oblong- or elliptic-lanceolate, 7-49-7(54) x 1-5-8-5(19) mm, 0-1-0-2
mm thick, acuminate or acute to subobtuse at apex, greyish-green to greyish white, with hair
density on upper and lower surfaces (4)5—62-8(68) and (5)6-6—51(55) per 0-05 mm? respectively.
Inflorescence contracted to spreading, with (1)2-6—12-2(16) flowers; pedicel of terminal flower
(10)12-7-42-8(60) mm; lowest bracts 4—9-7(10) x 2-4 mm, with scarious tip (0-1)0-2—1-8(2) mm.
Sepals 4—9(9-2) x 2—3-4(4) mm, lanceolate to oblong- or ovate-lanceolate, with scarious tip 0-2—
1-3(1-5) mm, subacute to obtuse at apex. Petals (10-3)11—18(18-5) x (4-5)5—10(11) mm, with cleft
3-6 mm. Filaments (4)4-2—4-6(5) mm; anthers 1-1-1 mm; pollen grains (32-5)34-3-36-5(38) um in
tetraploid, (35)35-3-40-1(42) um in octoploid, and (42)43—45-5(50) um in dodecaploid. Ovary (2)
2-2—2-4(2:5) mm; styles (3)3-2—5-7(6) mm. Capsules 8—11(12) mm, straight to bent, with pliable
walls, with teeth (0-7)0-8—1-2(1-3) mm and (0-1)0-2-0-3 mm wide and with revolute lateral
margins. Seeds (1)1-19-1-6(1-9) mm. 2n = 36, 72, 108.

CERASTIUM TOMENTOSUM VAR. MINUS C. PRESL

Plant mat-forming, with stolons and/or rhizomes; flowering stems 5—20cm, their uppermost
internode (20)33-42-5(46) mm and with simple non-glandular hairs, simple glandular hairs, or
both. Uppermost leaves on flowering shoots linear to lanceolate, (8-3)9-30 x 1—3 mm, 0-2—0-3 mm
thick, acute to acuminate at apex, greyish-green to green, with simple non-glandular hairs only,
with hair density on upper and lower surfaces (4)4-8-13-4(16) and (5)5-3-6-8(18) per 0-05 mm?
respectively; length ratio of terminal to subterminal cell (6)6-5—15-8(16). Inflorescence contracted,
with (2)3-3-4-1(7) flowers; pedicel of terminal flower 10—19-3(23) mm; lowest bracts (4)4-5—7-3
(7-5) x 2-3 mm, with scarious tip (0-1)0-2-0-4(0-7) mm. Sepals (4:5)5—8 x 2-2-2(2-5) mm,
lanceolate to oblong-lanceolate, with scarious tip (0-2)0-3—1 mm, subacute to acute at apex. Petals
(8-9)9-11-7(12) x (5-3)6(7) mm, with cleft 3-4-1(4-5) mm. Filaments (3-8)4—4-1 mm; anthers 1—
1-1 mm; pollen grains (35)35-5—39-2(42) um. Ovary (1-8)1-9-2 mm; styles (3)3-1-3-5(4) mm.
Capsules 8—8-1(8-2) mm, straight to bent, with pliable walls, with teeth 1-1-1 mm long and 0-1—0-2
mm wide and with revolute lateral margins. Seeds (0-9)1-18—1-4(1-5) mm. 2n = 72.

490 M. K. KHALAF AND C. A. STACE

CERASTIUM BIEBERSTEINII DC.

Plant forming tussocks or small mats, without(?) stolons or rhizomes, almost completely covered
with simple non-glandular hairs; length ratio of terminal to subterminal cell (1)1-1—1-7(1-8);
glandular hairs absent; flowering stems 10-—30cm, their uppermost internode (53)63-7—125(160)
mm. Uppermost leaves on flowering shoots lanceolate to oblong-lanceolate, (14)17-8-42(50) x
(1-5)1-9-7 mm, 0-2—0-3 mm thick, acute to obtuse at apex, white to greyish white, with hair
density on upper and lower surfaces (4)4-4—26-8(29) and (10)12-8-30-2(32) per 0-05 mm?
respectively. Inflorescence contracted, with (3)4-3—10-5(12) flowers; pedicel of terminal flower
(13)13-7-47 mm; lowest bracts 5-9-5 x 2-5-4 mm, with scarious tip 0-5—0-9 mm. Sepals 6-5—9-7
(10) x 2—2-7(3) mm, lanceolate to oblong- or ovate-lanceolate, with scarious tip 0-5—1-1(1-2) mm,
acute to obtuse at apex. Petals (16-5)17—17-3(17-5) x (6-5)6-8-7(7-1) mm, with cleft (3-5)4(4-2)
mm. Filaments (5-4)5-5—5-7(5-8) mm; anthers (0-7)0-8—0-9 mm; pollen grains (32-5)35-8—37(37-8)
um. Ovary (1-7)1-8-1-9 mm; styles (3)3-1-4-1(4-4) mm. Capsules 10(10-1)-14(15) mm, straight,
with usually rigid walls, with teeth (1)1-2—1-4(1-5) mm and (0-3)0-4-0-5 mm wide and with + flat
lateral margins. Seeds (1-5)1-83-1-87(2-2) mm. 2n = 36, (?72).

IDENTIFICATION

Despite the enormous range of variation of C. tomentosum in many morphological characters, after
some familiarization with these two species it is possible to distinguish between them in the great
majority of cases using the following characters as clues: growth habit; leaf size, thickness and
indumentum; sepal size; filament, anther and ovary length; capsule, capsule-tooth and seed size,
and capsule wall rigidity. It is very difficult, however, to express these as opposing character-states
in a key, and a fully confident identification cannot be guaranteed using them alone. Study of a
much wider range of wild material of C. biebersteinii is required before we can be sure of the
diagnostic value of the above characters.

Notwithstanding the conclusions of Kaleva (1966), we have found the revolution of the lateral
margins of the capsule teeth to be absolutely reliable (Fig. 2). Unfortunately most herbarium
specimens do not carry ripe capsules, and it seems that these are not always produced. This is not
due, however, to self-incompatibility, as was surmised by Walters (1989), but to lack of self-
pollination in these highly protandrous but self-compatible plants (Khalaf & Stace 2000). The lack
of capsules does, of course, present a problem in certain identification. In such cases the hair
anatomy (Fig. 1) will provide an unequivocal answer, since this character always clearly
differentiates between the two species.

REFERENCES

BLACKBURN, K. B. & MorTON, J. K. (1957). The incidence of polyploidy in Caryophyllaceae of Britain and
Portugal. New Phytologist 56: 344-351.

BOISSIER, E. (1867). Cerastium, in Flora Orientalis 1: 712—730. Herbier Boissier, Geneva.

BRETT, O. E. (1955). Cytotaxonomy of the genus Cerastium, 1. Cytology. New Phytologist 54: 138-148.

BUSCHMANN, A. (1938). Uber einige ausdauernde Cerastium-Arten aus der Vermandtschaft des
C. tomentosum L. Feddes Repertorium 43: 118-143.

CLAPHAM, A. R. (1987). Cerastium, in CLAPHAM, A. R., TUTIN, T. G. & WARBURG, E. F. Flora of the British
Isles, 3rd ed., pp. 131-136. Cambridge University Press, Cambridge.

DE CANDOLLE, A. P. (1822). Sur les plantes rares ou nouvelles du jardin de botanique. Mémoires de la
société physique et d’histoire naturelle de Genéve |.

FAVARGER, C. (1972). Contribution a |’étude cytotaxonomique de la flore des Apennins, 2. Le groupe de
Cerastium tomentosum L. Saussurea 3: 65-71.

FENZL, E. (1842). Cerastium L., in LEDEBOUR, C., ed., Flora Rossica 1: 396-416. E. Schweizerbart, Stuttgart.

JALAS, J., SELL, P. D. & WHITEHEAD, F. H. (1964). Cerastium L., in TUTIN, T. G., HEYwWoop, V. H. et al.,
eds., Flora Europaea 1: 136-145. Cambridge University Press, Cambridge.

KALEVA, K. (1966). Biosystematic notes on Cerastium Biebersteinii DC. and some ornamental strains of
C. tomentosum L. (Caryophyllaceae). Annales Botanici Fennici 3: 100-109.

KARLSSON, T. (1997). Forteckning Over svenska karlvaxter. Svensk Botanisk Tidskrift 91: 241-560.

KHALAF, M. K. & STACE, C. A. (2000). Breeding systems and relationships of the Cerastium tomentosum
group. Preslia, Praha 72: 323-344.

LINNAEUS, C. (1753). Species Plantarum. Laurentius Salvius, Stockholm.

MARSCHALL VON BIEBERSTEIN, M. (1808). Flora Taurico-caucasica 1. Charkow.

CERASTIUM TOMENTOSUM & C. BIEBERSTEINII 49]

Moca, G. (1963). Considérations géographiques et systématiques sur la flore d’Italie du sud. Webbia
118: 65-72.

Morton, J. K. (1975). Cerastium L., in STACE, C. A., ed., Hybridization and the Flora of the British Isles,
pp. 171-174. Academic Press, London. =

NYMAN, C. F. (1855). Sylloge Florae Europaeae. N. M. Lindh, Orebro.

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

SCHISCHKIN, B. K. (1936). Cerastium L., in KOMAROV, V. L. & SCHISCHKIN, B. K., eds., Flora URSS 6:
430-466. Izdatel’stvo Akademii Nauk SSSR, Moscow & Leningrad.

SOLLNER, R. (1954). Recherches cytotaxonomiques sur le genre Cerastium. Bulletin de la Société Botanique
Suisse 64: 221-354.

STOJANOFF, N. A. & STEFANOFF, B. (1924). Flore de la Bulgarie. Minist. Agric. Bulg., Sofia.

WALTERS, S. M. (1989). Cerastium, in WALTERS, S. M. et al., eds., The European Garden Flora 3,
pp. 191-192. Cambridge University Press, Cambridge.

(Accepted March 2001)

'

—

Watsonia 23: 493-503 (2001) 493

The vegetation associated with Spiranthes romanzoffiana Cham.
(Orchidaceae), Irish Lady’s-tresses, on the Isle of Coll, Inner
Hebrides

S. A. HENDERSON

Dept. of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD

ABSTRACT

This paper details some environmental preferences of the scarce orchid Spiranthes romanzoffiana Cham. on
the Isle of Coll in the Inner Hebrides. Samples were categorised into four site types that form a continuum
rather than a series of discrete habitat types. High cover of Carex panicea L., and to a lesser extent Molinia
caerulea (L.) Moench and Juncus articulatus L., occurs throughout. The habitats appear to be influenced by
varying degrees of high water and low nutrient availability, as suggested by the disparate species. There is no
apparent difference between former and present habitat of the orchid on the island, but few conclusions can be
made from vegetation description alone. As it stands, the habitat as defined in this study is not uncommon, and
it may be that other factors, such as seed sterility or availability of mycorrhizal associate, are limiting the
species distribution.

KEYWORDS: Habitat preference, vegetation analysis.

INTRODUCTION

Spiranthes romanzoffiana Cham. (Orchidaceae), Irish Lady’s-tresses, is one of the few British
natives with an amphi-Atlantic distribution, found widespread in North America and limited to the
Western fringes of Europe (Preston & Hill (1997) included it under their “boreal-montane
element’). Indeed, within Europe it is almost entirely confined to the west of Scotland and south-
western, western and Northern Ireland. Other species that share a similar “asymmetrical” amphi-
Atlantic distribution include Potamogeton epihydrus Raft., Najas flexilis (Willd.) Rostkov. & W.
Schmidt and Eriocaulon aquaticum (Hill) Druce.

Although not discovered until 1810 in Ireland (Hackney 1992), it is widely accepted that
S. romanzoffiana is native (e.g. Heslop-Harrison 1953; Love & Love 1958; Stace 1997). Theories
explaining the amphi-Atlantic distribution include migration across a former land connection
between Europe and North America (Dahl 1963; Hulten 1963) followed by a contraction in the
European distribution during glaciation with survival in or near the present localities (see Coxon &
Waldren 1995), or long distance dispersal (Heslop-Harrison 1953; Perring 1965). One of the
outstanding features of North American elements in the British flora is that they are all aquatic,
marsh or lake margin plants, suggesting a similarity in propagule dispersal. While controversy
persists over the explanation for the uneven distribution of S. romanzoffiana, its similarity in
geography to other species and the difficulty in accounting for seed transport to such relatively
remote and more or less natural habitats by human means, are justifications of its native status (see
Webb 1985).

The main populations of $. romanzoffiana occur round the Galway-Mayo lakes area and Lough
Neagh in Ireland, and on Colonsay, Coll and the Outer Hebrides of Scotland, and it has one
outlying post in South Devon, discovered in 1957 (Anon. 1958). Stewart, Pearman & Preston
(1994) categorise the species as Scarce, found in only eighteen 10-km squares in Britain. As the
only widely recognised orchid species of Britain not to be found elsewhere in Europe, and with a
stronghold in Scotland, Scottish Natural Heritage has designated S$. romanzoffiana a priority
species for conservation. None of the Scottish localities recorded before 1981 now show the
orchid, although several new sites have since been discovered (UK Biodiversity Group 1999).
While little is known of the natural fluctuations of the populations, these results have increased
concern over the orchid’s status in the British Isles.

494 S. A. HENDERSON

S. romanzoffiana coexists with an agricultural system based on extensive grazing. The orchid is
supposedly sensitive to subtle changes in the type, timing and intensity of grazing (e.g.
Summerhayes 1951; Ferreira 1978; Horsman 1994). Traditionally, pastoral agriculture in the
Hebrides entails cattle grazing from September to May followed by a short period allowing for
plant growth and a subsequent harvest for hay. The land management is conducive to both survival
and fecundity of S. romanzoffiana, the two factors that determine population status. Cattle grazing
enables survival by keeping more competitive tussock-forming species low while the period free
from grazing coincides with the orchid’s flowering period, thus ensuring fecundity. In Coll, a
general trend in agricultural intensification is confounded by the heavy conservation priority
placed on the corncrake, Crex crex, a bird which has now disappeared from most other parts of
Britain (Stroud 1992). By encouraging “‘corncrake friendly” extensification of grazing on some
nesting areas, the RSPB has forced intensification in other areas. Ironically, the conservation
measure may pose a threat to populations of S$. romanzoffiana.

In North America typical habitats are moist: in bogs, marshes, meadows, salt flats, muskegs,
thickets, on sandy-gravelly beaches, but also occasionally on dry woods and dry open hillsides
(Luer 1975; Correll 1978; Case 1987; Smreciu & Currah 1989; Homoya 1993). Typically a
northern species, in the more southern parts of its range in America it is a mountain plant where it
occurs up to 3000 m asl and flowers until October (Correll 1978). In the British Isles, however, it
occurs within a narrower range of habitats, suggesting that here the populations are ecologically as
well as geographically marginal.

In Ireland it grows in damp meadows, close to rivers and lake-shores and in cut-over bogs
(Praeger 1934; Harron 1986; Hackney 1992). In Britain, the orchid is similarly described as
occurring on low-lying sites that are at least periodically irrigated and sometimes inundated
(Scottish Natural Heritage 1995). Horsman (1994) described Spiranthes romanzoffiana as having a
distinct habitat of Molinia caerulea carpet on old cattle-grazed lazy-beds. The habitats of S.
romanzoffiana have been surveyed on Coll, Colonsay (Scottish Natural Heritage 1995; Gulliver
1996) and Loch Sheil (Martin & Milnes 1993) using the National Vegetation Classification (NVC)
system (Rodwell 1991). All fall into the categories for mire, wet heath and rush pasture (Table 1).
Collectively, and on a national scale, these communities are not infrequent. However, as NVC
categories are based on the means of nation-wide data, they fail to describe the particular
idiosyncrasies of an actual plant community (see Legg 1992). Multivariate ordination and
classification techniques are preferable for the level of detail required in the study of rarities. For
example Prober & Austin (1991) used the ordination technique of correspondence analysis to
compare floristically sites where a rare species occurred with similar sites where it does not.

Bowles et al. (1993), Ratcliffe, Birks & Birks (1993), Vazquez & Norman (1995) and Read & ©

Tweedie (1996) all used numerical classification techniques to characterise the habitats of rare
species.

In order to investigate the floristic component of S. romanzoffiana’s habitat, a multivariate
analytical approach was used, with data from samples taken at two spatial scales. The two-scale

TABLE 1. NATIONAL VEGETATION CLASSIFICATION COMMUNITIES AND SUB-
COMMUNITIES IN WHICH SPIRANTHES ROMANZOFFIANA HAS BEEN FOUND

Code NVC Community and sub-community (Rodwell 1991)
M25 Molinia caerulea-Potentilla erecta mire

M25 a Erica tetralix sub-community

M25 b Anthoxanthum odoratum sub-community

M23 Juncus effusus-Galium palustre rush pasture

M23 a Juncus acutiflorus sub-community

M23 b Juncus effusus sub-community

Ml5a Scirpus caespitosus-Erica tetralix wet heath Carex panicea sub-community
M10 Carex dioica-Pinguicula vulgaris mire

M6 _ Carex echinata-Sphagnum recurvum/auriculatum mire
Mé6d Juncus acutiflorus sub-community

|
!
|
|
|

SPIRANTHES ROMANZOFFIANA ON COLL 495

approach enables a distinction to be made between the habitat preferences at the scale of the
individual plant and those of the typical mire community; habitat description is scale-dependent
due to spatial heterogeneity. Further, the identification of associated single species at a small scale
may facilitate fine-grained ecological insights into the nature of the rare species’ requirements.
The study was carried out on the Isle of Coll, in the Inner Hebrides, where the majority of the
Scottish S$. romanzoffiana populations are found, and where there is a relatively long history of
recorded individuals (Ferreira 1978; Horsman 1990, unpublished; Scottish Natural Heritage 1995).

METHODS

FIELD SURVEY

All 107 sites (numbering 16 locations, although these do not necessarily form discrete populations)
with accurate records of the orchid were visited, including two new sites found in 1996. Grid
references, sketch maps (Ferreira 1978) and photographs (Horsman 1990, unpublished; Scottish
Natural Heritage 1995) were used to relocate individuals or sites with previously recorded
individuals which were, in 1996, apparently absent. The locations were also thoroughly searched
for any unrecorded individuals. 1 x 1 m quadrats (““meso-scale samples’) were placed around the
orchid(s), including only visually homogenous areas of vegetation. The equivalent area was taken
where the homogenous vegetation did not fit to a 1 m*. Similarly, for those samples not containing
S. romanzoffiana, the quadrat was placed around the site of the absent orchid (this was made
possible by having relatively detailed and accurate records). Within each sample the number of
individuals of S$. romanzoffiana and the abundance of all species were recorded using the Domin
scale. Percentage bare ground was also estimated. 10 x 10 cm quadrats (“micro-scale samples’’)
were then placed around each individual of S. romanzoffiana found and presence/absence of all
species was recorded. This area was thought small enough to investigate the more specific
requirements of the individual orchid but large enough to incorporate other associated species.
Plant nomenclature follows Stace (1997) for vascular plants and Watson (1981) for bryophytes.

ANALYSIS

Ordination was used to summarise the sample data collected at both scales. Analysis of data used
detrended correspondence analysis (DCA) (Hill 1979a; Hill & Gauch 1980), a modification of
correspondence analysis (ter Braak 1987). Two-way indicator species analysis (TWINSPAN) (Hill
1979b; Gauch & Whittaker 1981) was used to classify the meso-scale samples. The computer
packages CANOCO (ter Braak 1988) and CANODRAW (ter Braak 1992) were used for DCA
analysis and ordination plots and VESPAN (Malloch 1995) for TWINSPAN analysis, raw data
tables and descriptive statistics.

RESULTS

MESO-SCALE SITE TYPES

Spiranthes romanzoffiana was present in 54 of the 107 meso-scale sites, (distributed over 11 of the
16 locations) and was absent in 53 of the sites (in 8 locations). In total 93 species were found.
Carex panicea and Molinia caerulea were constant occurring in 94% and 86% of samples
respectively, and 11 other species were frequent, occurring in 50% or more of samples: Juncus
articulatus, Carex nigra, Hydrocotyle vulgaris, Nardus stricta, Ranunculus flammula, Holcus
lanatus, Anagallis tenella, Succisa pratensis, Anthoxanthum odoratum, Carex echinata and
Eriophorum angustifolium.

The indicator species at divisions | and 2 of TWINSPAN are shown in Fig. 1. The first division
distinguishes samples containing Erica tetralix from samples with Holcus lanatus, Galium
palustre, Ranunculus flammula and Ranunculus acris. The characteristics of each of the four final
groups of division 2 are summarised in Table 2.

The sample ordination by DCA with the TWINSPAN groups superimposed (Fig. 2) shows that

496 S. A. HENDERSON

TABLE 2. MOST FREQUENT SPECIES AT DIVISION TWO OF TWINSPAN. PERCENTAGE
FREQUENCY IN 1M’? SAMPLES IS DENOTED IN PARENTHESES

Species present in 80% or more samples Species present in 50%-80% samples
Site type Al
Spiranthes romanzoffiana (70)
Carex nigra (100) Equisetum fluviatile (75)
Hydrocotyle vulgaris (100) Carex echinata (70)
Ranunculus flammula (100) Filipendula ulmaria (70)
Carex panicea (95) Holcus lanatus (70)
Juncus articulatus (95) Potentilla palustris (70)
Galium palustre (90) Mentha aquatica (65)
Caltha palustris (80) Calliergon cuspidatum (65)
Eriophorum angustifolium (80) Leontodon autumnalis (55)

Myosotis laxa (55)
Senecio aquatica (55)
Anthoxanthum odoratum (50)
Pedicularis palustre (50)

Site type A2
Spiranthes romanzoffiana (49)

Carex panicea (97) Hydrocotyle vulgaris (78)
Molinia caerulea (95) Ranunculus flammula (78)
Juncus articulatus (92) Carex nigra (76)
Holcus lanatus (89) Ranunculus acris (76)
Nardus stricta (81) Anthoxanthum odoratum (73)
Succisa pratensis (70)
Trifolium repens (62)
Potentilla erecta (58)
Agrostis canina (51)
Carex echinata (51)

Site type B1
Spiranthes romanzoffiana (50)

Carex panicea (100) Nardus stricta (77)
Molinia caerulea (95) Anagallis tenella (75)
Juncus articulatus (66)
Carex nigra (61)
Potentilla erecta (57)
Succisa pratensis (57)
Agrostis canina (55)
Hydrocotyle vulgaris (55)
Carex echinata (53)
Eriophorum angustifolium (53)
(note, Erica tetralix 39)

Site type B2
Spiranthes romanzoffiana (11)

Erica tetralix (100) Anagallis tenella (67)
Molinia caerulea (100) Carex panicea (67)
Carex hostiana (89) Selaginella selaginoides (56)
Schoenus nigricans (89) Scorpidium scorpioides (56) |
(note Drosera rotundifolia, Myrica gale and if
Narthecium ossifragum at 44) |

Those species characteristically present in high abundance are shown in bold. While subjective.
this aids interpretation.

SPIRANTHES ROMANZOFFIANA ON COLL 497

Divison |
A (357) B (50)
Holcus lanatus, Galium palustre, Erica tetralix
Ranunculus acris,
Ranunculus flammula
Divison 2

Al (20) A2 (37) B1 (41) B2 (9)
Caltha palustris, Nardus stricta without Schoenus nigricans Schoenus nigricans
Equisetum fluviatale,
Potentilla palustris

FIGURE 1. Indicator species of the TWINSPAN hierarchy for the first two divisions. The number of samples in
each category is shown in parentheses.

3

Axis 2

Axis 1

FIGURE 2. Sample ordination plot for the first two axes of Detrended Correspondence Analysis (DCA) of
meso-scale habitat data for Spiranthes romanzoffiana. TWINSPAN-defined groups are superimposed.

the TWINSPAN-defined site types are not readily separated, either at the first or the second
divisions. The first four axes extracted in DCA explain only 22.8% of the total variation in species
data, and the first two axes of the ordination (which are used in the ordination plot) explain only
17% of floristic variation between samples. There is thus a large proportion of variation that is not
explained by the ordination plot.

The sample ordination plot shown with S. romanzoffiana presence/absence data superimposed
(Fig. 3), shows for each group an ellipse within which, on average, 68% of the individuals will lie.
Considerable overlap is shown which indicates little pattern of differentiation between current and
former sites. The full set of samples were consequently analysed as potentially suitable.

MICRO-SCALE SITE TYPES

A total of 81 micro-scale samples were taken, all of which contained at least one individual of
S. romanzoffiana. A total of 68 species were recorded and the mean per sample was 9 (standard
error 0-29). Sample ordination by DCA showed that species composition reflects that of the larger
scale though inevitably fewer species are present. The proportion of samples within which the 13
most constant species were present is shown in Table 3. Carex panicea was the most associated

498 S. A. HENDERSON

TABLE 3. THE 13 MOST ASSOCIATED SPECIES WITH SPIRANTHES ROMANZOFFIANA
AT THE MICRO-SCALE, AS MEASURED BY THE PERCENTAGE OF THE 10 CM?
SAMPLES WITHIN WHICH THEY WERE PRESENT

Species % constancy
[Spiranthes romanzoffiana 100]
Carex panicea 84
Molinia caerulea 63
Carex nigra 44
Hydrocotyle vulgaris 43
Anagallis tenella 38
Nardus stricta 6s)
Eriophorum angustifolium Zi
Ranunculus flammula 2]
Carex echinata 26
Trifolium repens PG)
Holcus lanatus 21
Carex hostiana 20
Juncus articulatus 20

species at the micro-scale occurring in 84% of the samples, Molinia caerulea occurred in only
63% of the samples. No other species was present in more than 50% of samples. Juncus
articulatus, a species occurring with S. romanzoffiana in 76% of the meso-scale samples, is
present in only 20% of the samples at the micro-scale.

DISCUSSION

SAMPLING CRITIQUE
The accessibility of all the sites relocated opens to question the extent to which the orchid occurs
in relatively inaccessible areas. Known sites tend to occur close to roads - perhaps a sampling
artifact (as it is assumed that not all individuals in the population will have been recorded, perhaps
the unrecorded individuals grow in less accessible areas), indeed “road map recording” is a well
known phenomenon. Alternatively, the distribution may reflect true habitat preference for the
marginal land between dune and moor, generally known in the Western Isles as the
“blackland” (Hambrey 1986). The transitional zone between lime-rich and acid soils is not only
important for many species of plant but also tends to provide the most suitable terrain for roads.
Sampling was biased towards flowering spikes; vegetative individuals were normally only found
when searched for thoroughly in a small site where an individual was known to occur, or in short
open turf. Bias was additionally encountered by the relative ease of spotting plants (both
vegetative and flowering) in short vegetation. Sites within taller vegetation may thus be under-
represented. It is difficult to determine whether an individual is truly absent from a former site or
whether it has just been missed. The difficulty in distinguishing absences which are a result of
poor sampling from those which are genuine is a serious problem in the study of rarity in general
(McArdle & Gaston 1993: Gaston 1994) and accentuated here by the potential for S.
romanzoffiana to lie dormant underground (Gulliver et al. 2000). Finally, there is sample bias
towards those locations that have a longer history of recorded individuals; at the two new sites,
absences cannot be recorded for the obvious reason that there is no comparative data from
previous years.

THE HABITATS OF SPIRANTHES ROMANZOFFIANA

Horsman (1994) described Spiranthes romanzoffiana in Scotland as having a distinct habitat of
Molinia caerulea.carpet on old lazy beds grazed by cattle. M. caerulea is also listed as the closest
associate in 17 Scottish sites surveyed in 1995 (Scottish Natural Heritage 1995). The present

SPIRANTHES ROMANZOFFIANA ON COLL 499

3
: @ Present
: © Absent
p
N
&
x< =
< eee?
1 @ ®@
O 8
0 1 2 3 4

Axis 1

FIGURE 3. Ordination plot for all meso-scale samples for the first two axes of Detrended Correspondence
Analysis (DCA), with presence or absence of Spiranthes romanzoffiana in each sample superimposed. 68%
confidence ellipses are shown.

author, in agreement with Gulliver’s study on Colonsay (1996), found only one site on an old lazy
bed. Further, despite it being in general abundant, and the second most associated species with the
orchid at the micro-scale, it was found that in a number of samples, typically those of Site Type
Al (the habitat in which S. romanzoffiana was most frequent (Table 2)), M. caerulea was not
abundant or even present. All the NVC habitats previously described for the orchid include M.
caerulea as a typical species at intermediate to high frequency and abundance. However, to
compare the results directly to equivalent NVC categories would be inappropriate without
considering the effects of scale and pattern within the vegetation. NVC demands larger samples (2
x 2m or over) and treats minute mosaics as a single vegetation type (Rodwell 1991). The resulting
NVC classification is therefore at a coarser grain than that presented here, which describes the
floristics immediately surrounding the orchid. It would be similarly misleading to convert the
present data into NVC classes.

All site types have an abundance and constancy of the generally low-growing Carex panicea
and an abundance and frequency of Hydrocotyle vulgaris, Ranunculus flammula and/or Anagallis
tenella. The species are characteristic of unshaded, soligenous mire on peaty, mildly acidic soils
where growth of potential dominants is suppressed by low fertility and grazing pressure (Grime,
Hodgson & Hunt 1990). While the soil characteristics were not analysed, grazing was evident in
all sites. In these aspects, the habitats are similar.

However, TWINSPAN and ordination by DCA indicate differential abundances of the
dominants and distributions of more minor species between the four species groups. The habitats
are distinguished by these lesser species which have quite different ecological preferences and may
reflect differences in water and nutrient levels.

Site-type Al contains a species component tolerant of a degree of waterlogging and typical of
neutral/mildly acid conditions (Grime, Hodgson & Hunt 1990): Caltha palustris, Potentilla
palustris and Equisetum fluviatile define the habitat and are largely restricted to it. Other wetland
species present in these sites and less abundant or absent in others include Mentha aquatica,
Myosotis laxa, Filipendula ulmaria, Senecio aquaticus and Iris pseudacorus. Carex nigra, present
and abundant in all samples within this site type, has a wide habitat tolerance but is usually found
in mires with some degree of water movement or mineral enrichment (Jermy, Chater & David
1982). Hydrocotyle vulgaris is constant, and is a wetland species suggested to have an inability to
exploit sites which are waterlogged all year (Grime, Hodgson & Hunt 1990), while Ranunculus
flammula, also a constant, is adapted to both submergence and desiccation (Cook & Johnson

500 S. A. HENDERSON

1968). The evidence suggests that Al is composed of sites that are periodically flooded. Indeed,
samples of this type are primarily found around a periodically flooded pasture and a flat, poorly
drained area beside a lochan.

Al grades into site-type A2, a drier, slightly more acidic habitat, as indicated by the significant
contribution to cover of some graminoids less tolerant of inundation such as Nardus stricta,
Molinia caerulea and Holcus lanatus, but one which still contains an element of species of Al in
the wetter samples, such as Galium palustre, Senecio aquaticus and Filipendula ulmaria. Nardus
stricta, the TWINSPAN indicator species which distinguishes the habitat from A1, is a grass that
is characteristic of free-draining, acid soils (Grime, Hodgson & Hunt 1990). Species found almost
exclusively in this habitat, such as Cynosurus cristatus, a species susceptible to both drought and
waterlogging (Grime, Hodgson & Hunt 1990) and Achillea ptarmica, again suggest a habitat that
is moist but not waterlogged. Site type A2 thus appears to be of a wet, slightly acid graminoid and
herb-rich turf.

Site type A2 (and to a lesser extent Al) grades into site type B1. Species that are common to
both habitats A2 and B1 but not the others include frequent species such as Succisa pratensis,
Potentilla erecta, Carex echinata as well as less abundant species such as Plantago maritima, P.
lanceolata, Carex dioica and Euphrasia scottica. B1, however, contains elements typical of wet
acid heath (Grime, Hodgson & Hunt 1990; Rodwell 1991; Stace 1997), primarily shown by the
presence of the indicator species at division one, Erica tetralix, but also the other woody shrubs
Calluna vulgaris and, occasionally, Salix repens and Myrica gale. The low cover and presence of
these species in the samples indicates a habitat where their dominance is restricted to some extent.
Further, the abundance of these subshrubs is sufficiently low to allow a relatively high cover and
constancy of the creeping species Anagallis tenella, and the low-growing Drosera spp., typical of
unshaded and wet, nitrogen-poor sites of heaths and bogs (Stace 1997). The wet, acidic conditions
of the site type are further suggested by the presence of Sphagnum auriculatum and, more
frequently, S. recurvum. Eleocharis quinqueflora, characteristic of damp, peaty places and wet
flushes in the west of Scotland (Phillips 1994), also appears in this habitat. It seems therefore that
these habitats may comprise flush vegetation within more enclosed communities where
competition from low-growing shrubs would otherwise suppress the orchid.

The constancy of Erica tetralix and presence of species such as Myrica gale, Pinguicula
vulgaris, Selaginella selaginoides, Drosera spp., and Narthecium ossifragum render site type B2
similar to B1; the habitat still appears to be a flush in wet, nutrient-poor heath. However, B2 is
distinguished from B1 by the presence of the tuft-forming Schoenus nigricans, occurring in all
samples but one, and also the bryophyte Scorpidium scorpioides. The combined presence of these
species suggests a habitat of wet flushes where the water content is fairly base-rich (see Phillips
1994; Sparling 1968; Watson 1981). Such base and cation enrichment may result from flushed
shell-sand or directly from the bedrock (Rodwell 1991).

Carex panicea is the species most associated with S. romanzoffiana at both the large and small
scale. C. panicea is characteristically a wetland species, relatively low-growing, with a pH range
from 4-7-5 (Grime, Hodgson & Hunt 1991); in vitro seed germination experiments have revealed
that S. romanzoffiana prefers a soil pH from 4-7—5 (Arditti 1992). C. panicea is widespread
throughout the British Isles, occurring in all British vice-counties, and has a capacity to exploit
such diverse habitats as soligenous mire and limestone grassland (Grime, Hodgson & Hunt 1991)
as well as dwarf-shrub heaths (Jermy, Chater & David 1982). The occurrence of C. panicea in a
wide range of vegetation types outside those described here make it an unsuitable indicator species
for potential S. romanzoffiana habitat. Associated species are therefore not necessarily good
indicator species. However, the results suggest that, on Coll at least, habitats without C. panicea
are unlikely to support S. romanzoffiana.

The low association of Juncus articulatus with S. romanzoffiana at the micro-scale, despite the
presence of the rush in 76% of the samples at the meso-scale, may reflect an ecological preference
of the orchid. Grime, Hodgson & Hunt (1990) claim that J. articulatus is morphologically variable
(ranging in height from 20 mm to 600 mm), often depending on the level of grazing. It is possible
that S. romanzoffiana cannot grow in the shade that the tall-growing rush provides, but can survive
when J. articulatus is relatively short. Alternatively, the tufted nature of the rush may make
survival and growth of the orchid amongst its stems difficult.

SPIRANTHES ROMANZOFFIANA ON COLL 501
CONCLUSIONS

Overall vegetation description alone cannot provide a comprehensive summary of all habitat
variables. The structure of vegetation may be one of the most important conditions determining
habitat suitability and may be altered in a relatively short time by grazing. A period of inertia
before species composition responds would then render floristic measurements inadequate. This
may explain the apparent similarity in the current study of former and present habitats of S.
romanzoffiana. Vegetation description also fails to account for environmental variables that affect
only one species, in particular the requirement for orchid mycorrhizal infection before germination
(Wells 1981). It would therefore be interesting to examine the presence of the specific symbiotic
fungus in the soil in suitable and potentially suitable habitats for S$. romanzoffiana.

There are other reasons why an individual may appear absent from a potentially suitable site.
Dormant individuals will be recorded as absent unless destructive methods are used (hardly an
option for a scheduled species!). The related orchid S. spiralis can lie dormant for at least a year
before flowering (Wells 1967) and S. romanzoffiana has been documented to subsist for five years
underground before producing above-ground growth (Gulliver et al. 2000). The role of stochastic
events is also important; the processes of population extinction and colonisation may cause
suitable sites to become unoccupied, at least temporarily. For this reason, all habitats in which S.
romanzoffiana was known to occur, however infrequently, were included in the range of
vegetation types designated suitable.

These results provide a more accurate depiction of the immediate habitat of S$. romanzoffiana
than do the NVC descriptions (Rodwell 1991). They demonstrate the orchid’s ecological affinity
to C. panicea and avoidance of tussock-forming species at the micro-scale. They also show that
the orchid does not apparently have an affinity to M. caerulea, at least not at the spatial scale
examined here. However, the plant associations recognised in this study are not uncommon.
Gaston (1994) recognised two broad ranges of factors causing limited ranges and/or low
abundances: environmental limitations and colonisation abilities. The results here suggest that
factors other than the environment may be limiting the distribution of $. romanzoffiana. There is
evidence that the species is reproductively limited where it is found in the British Isles; Heslop-
Harrison (1953) found that, while it was an active colonist of cut-away bogs around Lough Neagh,
there were few years when it set seed. While flowering spikes are produced, it appears that
Scottish populations may produce fertile seed rarely (Gulliver 1997; Gulliver et al. 2000) and may
even reproduce entirely vegetatively (Horsman 1994; UK Biodiversity Group 1999). Whether or
not this accounts for the limited distribution of S$. romanzoffiana in Britain is not known; further
studies into the dispersal and colonisation abilities of S. romanzoffiana are therefore required.

ACKNOWLEDGMENTS

I am indebted to Neil Cowie at the R.S.P.B. for his suggestions and support, and to the farmers on
the Isle of Coll for their co-operation. Thanks also to Fred Rumsey, Bob Press and Richard
Bateman for valuable comments on earlier drafts of the manuscript, and to Nancy Garwood for
help with the graphics. The work was provided with financial support from Scottish Natural
Heritage.

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(Accepted November 2000)

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Watsonia 23: 505—516 (2001) 505

A re-assessment of the putative Carex flava agg. (Cyperaceae)
hybrids at Malham Tarn (v.c. 64): A morphometric analysis

N. BLACKSTOCK and P. A. ASHTON

Dept. of Natural and Applied Sciences, Edge Hill College of Higher Education, St. Helens Rd,
Ormskirk, Lancashire L39 4QP

ABSTRACT

Carex flava L., a rare species in the British Isles, is currently only recognised from one extant population. The
presence of three putative C. flava x C. viridula Michx. subsp. brachyrrhyncha (Celak.) B. Schmid hybrid
(= C. x alsatica Zahn.) populations suggests that it may have formerly been more widespread. Alternative
taxonomic treatments have been considered populations, including the possibility that they could be C. flava
subsp. jemtlandica (Palmgr.) P. D. Sell, a Fennoscandian taxon not previously recorded in this country. Uni-
and multivariate statistical analyses were used to elucidate the taxonomic affinity of the largest of these
populations. A new combination C. flava subsp. jemtlandica to C. virtdula Michx. subsp. brachyrrhyncha
(Celak.) B. Schmid var. jemtlandica (Palmgr.) Blackstock & P. A. Ashton, is made to bring the nomenclature
of this taxon in line with the current nomenclature of this group. The presence of a second extant population of
C. flava in the British Isles is also reported.

KEYWORDS: Carex viridula subsp. brachyrrhyncha var. jemtlandica, Principal Components analysis (PCA),
ANOVA, Discriminant Analysis (DA).

INTRODUCTION

In the British Isles Carex flava L. (Large Yellow Sedge) is currently only recognised from one
extant population and one historic record. In addition, populations generally considered to consist
of C. flava x C. viridula Michx subsp. brachyrrhyncha (Celak.) B. Schmid hybrids (= C. x
alsatica Zahn) have been recorded from three sites; the Tarn Moss, Malham (v.c. 64) (Shaw 1946),
Greywell Moors, Hants (v.c. 11) (Brewis et al. 1996) and by the River Corrib near Menlough,
north-east of Galway City (v.c. H17) (Perring 1970). C. flava is considered to have formerly been
extant at these localities but is now believed to be extinct, probably through hybridisation with
C. viridula. subsp. brachyrrhyncha (Jermy et al. 1982). C. flava is categorised in The British Red
Data Book: I. Vascular Plants (Taylor 1999) as vulnerable in Britain, but not threatened in
Europe. Populations are also found in Western and Eastern North America at latitudes similar to
those in Europe.

Although early Floras covering the various regions of the British Isles are littered with accounts
for C. flava s. I. (e.g. Goodenough 1792; Green 1933), the first authenticated specimen for C. flava
s. s. was collected by J. Dickinson in 1836, somewhere in the Ennerdale area of Cumbria (v.c. 70).
This population is now only known from one voucher specimen (LIV), although doubts as to the
authenticity of the site locality have been raised (Halliday 1997). Evidence of a population
previously unaccounted for in the British Floras comes from a herbarium specimen collected by
T. W. Edmondson from Hebden, W. Yorks (v.c. 64) in 1906. This population is also only known
from one voucher specimen (GH) which has been examined by W. J. Crins and has been
determined by him to be C. flava s. s. (Crins 1985).

The only currently recognised extant population of C. flava s. s. is at Roudsea Wood,
Haverthwaite, Cumbria (v.c. 69) being discovered by D. Lumb in 1913 (B.E.C. Rpt vol Il pt VI
p. 504). The Roudsea Wood population is well known and the number of plants there appears to be
relatively stable at around 1500-2000 presumed genets (Blackstock, unpublished data).

In 1946 G. A. Shaw reported C. flava from the Tarn Moss, Malham (v.c. 64). This population
was also referred to as C. flava s. s. by Davies (1953). However, doubts as to the true identity of
this population were raised by Jermy & Tutin (1968). They state that “the only typical material” of

506 N. BLACKSTOCK AND P. A. ASHTON

C. flava in Britain “is from N. Lancs” i.e. Roudsea Wood. Jermy ef al. (1982) state categorically
that “plants intermediate to C. flava and C. lepidocarpa Tausch. (= C. viridula subsp.
brachyrrhyncha) ...occur at Malham Tarn, Lough Corrib and N. Hants, where C. flava presumably
once occurred but is now extinct’. Stace (1997) reiterates this view by formally referring to these
populations as C. x alsatica.

A different approach to the putative C. x alsatica populations found at Malham Tarn, Lough
Corrib and Greywell Moors has been taken by Sell & Murrell (1996) who classify them as C. flava
subsp. jemtlandica (Palmgrn.) P. D. Sell (= C. jemtlandica Palmgrn.). This taxon is generally
regarded as a Fennoscandian endemic (Hedrén & Prentice 1996; Pykalé & Toivonen 1994) and
has not previously been recognised in the British Isles.

Clearly the Malham Tarn, Lough Corrib and Greywell Moors populations of yellow sedges have
been subject to a degree of taxonomic confusion. Davies (1953) considers them to be C. flava s. s.;
Jermy et al. (1982) consider them to be C. x alsatica, a C. flava x C. viridula subsp.
brachyrrhyncha hybrid; while Sell & Murrell (1996) treat them as a subspecies of C. flava
comparable with their classification of other C. flava subspecies.

Multivariate statistical analyses have been successfully used to clarify taxonomic problems in
Carex complexes e.g. sects. Vesicariae (O. F. Lang) Christ. (Shepherd 1975), Phyllostachys
(J. Carey) L. H. Bailey (Naczi et al. 1998) and Ceratocystis Dumort. (Crins 1985). Studies within
sect. Ceratocystis have utilised multivariate statistical methods such as Detrended Correspondence
Analysis (DCA) (Schmid 1980), Principal Components Analysis (PCA) (Crins 1985) and
Discriminant Analysis (DA) (Crins 1985) based upon herbarium specimens. The use of two of
these methods (PCA and DA) on living specimens of C. flava, C. viridula subsp. brachyrrhyncha,
the putative C. x alsatica from Malham Tarn and C. jemtlandica has been adopted here to
elucidate the taxonomic boundaries between these taxa. This study includes plants from North
America, the British Isles and continental Europe and seeks to place the British and Irish taxa into
a wider context, while clarifying the taxonomic status of the problematic Malham Tarn yellow
sedges.

MATERIALS AND METHODS

Specimens were collected in the field, either as ramets or seed, from clumps situated at least 3 m
apart from each other. Members of the C. flava agg. have a caespitose growth habit, so each
sample was presumed to represent an individual genet. The individual specimens were cultivated
under similar conditions at Edge Hill for 1-2 years until the plants had reached maturity.
Specimens of C. flava from Canada provided a valuable comparison, as these populations are
approximately 700 km away from the nearest C. viridula subsp. brachyrrhyncha population.
Voucher specimens are currently held at Edge Hill College of Higher Education and it is intended
that they will be deposited at LIV upon completion of the associated work. Details of the specimen
localities are given in Table 1. Four a priori groups (Table 1) were identified for analysis:
1. C. flava, (40 specimens, eleven populations); 2. The putative C. x alsatica from Malham Tarn
(MT) (ten specimens, one population); 3. C. viridula subsp. brachyrrhyncha (40 specimens, 13
populations) and 4. C. jemtlandica (ten specimens, four populations). All of the C. jemtlandica
sites were included in a study of C. lepidocarpa s. 1. (Hedrén & Prentice 1996). The specimens
collected from Dalbyn, near Ore, Dalarna, Sweden are from the same site as specimens collected
and photographed by Palmgren for his monograph of the C. flava agg. (Palmgren 1959). As no
single classification recognises all of these groups, they will be referred to as listed above until
their status is re-assessed later in this paper. The putative C. x alsatica from Greywell Moors is
only known from a single plant and this has not been seen in recent years. Therefore this
population has not been included in our analyses. In addition, it has not been possible to obtain
samples of the putative C. x alsatica from Ireland.

Multivariate analyses provide a preliminary re-assessment of the a priori classification and of the
phenetic relationships between the taxa. Eleven characters (Table 2) were included in the
morphometric analysis. All morphometric data were derived from living plants that were
producing a fertile culm with perigynia containing well-developed nutlets. This eliminated the
possibility that Fl-hybrids could be included in the data set and ensured that only complete data

CAREX FLAVA AGG. AT MALHAM TARN

507

TABLE 1. LOCALITIES OF THE POPULATIONS OF YELLOW SEDGES INCLUDED IN
THE PCA. TAXON REFERS TO THE A PRIORI CLASSIFICATION OF THE GROUPS

Number of
Species Location Country Location individuals Taxon
sampled
Nr. Campbellville, Halton Co., Canada 43°31’N 79°59°W 2 C. flava
Ontario
Nr. Orangeville, Peel Co. OntarioCanada 43°53’N 80°06’ W 1 C. flava
8 km NW Tranas, Smaland Sweden 58°06’N 14°58’E D) C. flava
Tannas, Harjedalen Sweden 62°40’N 12°41’E 2 C. flava
Sjugelmyren, S6rboda, Dalarna Sweden 61°07’N 15°13’E 1 C. flava
Lake Essen, Nr. Rattvik, Dalarna Sweden 60°57’N 15°13’E 4 C. flava
Vastana, Boda, Dalarna Sweden 60°59’N 15°12’E i C. flava
Romedenne Belgium 50°10’N 04°42’E 5 C. flava
Roudsea Wood, Cumbria England 54°14’N 03°08’ W 6 C. flava
Nittedal Hakadal Norway 60°05’N 10°50’°E 9 C. flava
K4arnten Austria 47°N 13°E 1 C. flava
Malham Tarn Fen, England 54°06’N 02°11’ W 10 MT
Mid-west Yorkshire
Stromyren, Boda, Dalarna Sweden 61°01’N 15°13’E 2 C. jemtlandica
Vadeermyran, Hammerdal, Sweden 62°52’N 15°16’E B) C. jemtlandica
Jamtland
Ljusnedal, Harjedalen Sweden 62°48’N 12°36’E 2 C. jemtlandica
Dalbyn,Ore, Dalarna Sweden 61°08’N 15°10’E 3 C. jemtlandica
Benestads Bakan, Tomelilla, Sweden 99°57 N13" 53; EF 4 C. viridula subsp. brachyrrhyncha
Skane
Lyngsjon Lake, Skane Sweden 55°48’N 13°53’E 4 C. viridula subsp. brachyrrhyncha
Hiiumaa Island Estonia » 59°10°N'22°30°E po C. viridula subsp. brachyrrhyncha
Locheres France 47°41’N 04°5S’E 1 C. viridula subsp. brachyrrhyncha
Crummack Farm, Inglebrough, England 54°08’N 02°21’W 1 C. viridula subsp. brachyrrhyncha
N. Yorkshire
Crummack Dale, Inglebrough, England 54°08’N 02°21’ W 4 C. viridula subsp. brachyrrhyncha
N. Yorkshire
Malham Tarn outflow, England 54°06’N 02°11’>W 6 C. viridula subsp. brachyrrhyncha
Mid-west Yorkshire
Woodbastwick, E. Norfolk England 52°41N 01°27’E 3 C. viridula subsp. brachyrrhyncha
Greywell Moor, Hants. England 51°15’N 00°58’ W 1 C. viridula subsp. brachyrrhyncha
Hawes Water, Lancashire England 54°11’N 02°48’ W 6 C. viridula subsp. brachyrrhyncha
Fearnan, Loch Tay, Scotland 56°47’N 03°48’ W 1 C. viridula subsp. brachyrrhyncha
Perth & Kinross
Glen Fender, Perth & Kinross Scotland 56°34’N 04°05’ W 5 C. viridula subsp. brachyrrhyncha
Prission, S-Tirol Austria 47°N 12°E 2 C. viridula subsp. brachyrrhyncha

sets were included for subsequent analysis. Measurements were taken using Rabone vernier
calipers graduated to 0-1 mm, with the exception of a coarse measurement (character VEGLEAF)
which was measured to the nearest mm using a standard 30 cm line ruler. Previous morphometric
studies, either within the C. flava agg. or Carex sub-genus Carex, have utilised a greater number of
morphological characters (e.g. Shepherd 1975; Schmid 1980; Crins 1985; Hedrén 1998; Naczi et
al. 1998). However, many of the characters included in the above studies are associated with the
position of the pistillate spikelets. Such spikelets are not necessarily equivalent within the C. flava
agg. Therefore, they and their associated characters have not been included in this study.
Characters BEAKL, PERLE and BRISTLE are all associated with the perigynia. Crins (1985)
and Rothrock et al. (1997) noted that there was greater variability in perigynia from the lower parts
of the pistillate spikelets. Therefore perigynia from the mid-section of the pistillate spikelets were
used for the measurements of all perigynia characters. All characters were then used in PCA to
summarise variation patterns within all of the taxa studied. Data were standardised for PCA so that

508 N. BLACKSTOCK AND P. A. ASHTON

TABLE 2. MORPHOLOGICAL CHARACTERS, WITH THEIR ABBREVIATIONS,
MEASURED FROM LIVING PLANT SPECIMENS OF THE CAREX FLAVA AGG. ALL
CHARACTERS WERE INCLUDED IN THE PCA AND ANOVA

Character Abbreviation
Vegetative leaf length/width ratio VEGLEAF
Vegetative leaf width at base (mm) VEGBAS
Ligule length (mm) LIGL
Male spikelet length (mm) MSPIKE
Male spikelet peduncle length (mm) MPEDL
Beak length BEAKL
Perigynum length (mm) PERLE
Male glume length/width ratio MGLUM
Female glume length/width ratio FGLUM
No. bristles on one side of the perigynum BRISTLE
Nutlet length excluding beak (mm) NUTL

each variable would have a mean of 0 and a standard deviation of 1. PCA was carried out using the
Multivariate Statistical Package version 3.0 from Kovach Computing Services (Kovach 1998).
Data were further analysed using a stepwise DA (S.P.S.S. 1998).

Univariate statistical analyses (mean, range and standard deviation) were performed on all
characters using Microsoft Excel 97. Differences in the mean values for single characters between
all the taxa were tested using one way analysis of variance (ANOVA) with the data first being
tested for variance equivalence using the F-test.

RESULTS

Scatter plots of the principal components PC I vs PC II (Fig. 1) and for PC I vs PC III (Fig. 2) for
all the taxa and individuals depict two clusters. These can be identified as a C. flava/MT cluster
and C. viridula subsp. brachyrrhyncha/C. jemtlandica cluster. No sub-clusters were evident within
the C. flava/MT cluster. All individuals from the MT cluster with the C. flava individuals and are
not intermediate between C. flava and C. viridula subsp. brachyrrhyncha. From Figs 1 and 2, two
sub-clusters within the C. viridula subsp. brachyrrhyncha/C. jemtlandica cluster associated with
C. viridula subsp. brachyrrhyncha and C. jemtlandica may be identified. However, the separation
is not complete.

The first two principal components account for 46-6% of the variance within the data set, with
PC I accounting for 32-5% and PC II accounting for 14-1%. PC III accounts for a further 11-5% of
the variance within the data set. The variables with the highest component loadings (Table 3) on
PC I are, in descending order, variables PERL, BEAKL, BRISTLE, MPEDL and LIGL. The first
three variables are all associated with the perigynium. The highest component loading on PC II
and PC III are for characters MSPIKE, VEGBAS, LIGL and NUTL, FGLUM respectively.

Morphometric data from the 100 specimens of the C. flava agg. were further analysed using a
stepwise DA. The initial DA was constructed using the four a priori groupings listed above. The
program selected eight differentiating variables in the following descending order: MPEDL,
BRISTLE, NUTL, BEAKL, MGLUM, VEGLEAF, LIGL and FGLUM. DA succeeded in
correctly classifying 94% of the group cases (Fig. 3). The first canonical discriminant function
(CDF I) accounted for 76-1% of the variance, with 17-4% and 6-5% of the variance being
accounted for on CDF II and CDF III (not shown) respectively. The greatest contribution to variate
one is given, in descending order, by MPEDL, BEAKL, BRISTLE and FGLUM. Along variate
two NUTL, VEGLEAF and LIGL give the greatest contribution.

A second DA was then carried out on the same data set with the exception that the C. flava and
MT populations were condensed into one group. The discrimination efficiency improved to
classify 99% of the group cases correctly. However, condensing the groups gave a similar pattern
and spatial arrangement to that seen in Fig. 3. The first canonical discriminant function (CDF I)

PC Il

PC Ill

CAREX FLAVA AGG. AT MALHAM TARN

-0.4 -0.3 -0.2 -0.1 0 0.1 0.2 0.3 0.4
PC |

aC. flava AMTP @C. viridula subsp. brachyrrhyncha — © C. jemtlandica

FIGURE 1. Scatter plot of PC I vs PC II from PCA of the measurements of C. flava , MTP,
C. viridula subsp. brachyrrhyncha and C. jemtlandica.

-0.4 -0.3 -0.2 -0.1 0 0.1 0.2 0.3 0.4
PC |
4C. flava SMTP @C. viridula subsp. brachyrrhyncha — © C. jemtlandica

FIGURE 2. Scatter plot of PC I vs PC III from PCA of the measurements of C. flava , MTP,
C. viridula subsp. brachyrrhyncha and C. jemtlandica .

0.5

0.5

509

0.6

0.6

510 N. BLACKSTOCK AND P. A. ASHTON

TABLE 3. LOADINGS FOR THE FIRST THREE PRINCIPAL COMPONENTS
FROM PCA OF SPECIMENS OF C. FLAVA, THE MT POPULATION,
C. VIRIDULA SUBSP. BRACHYRRHYNCHA AND C. JEMTLANDICA.

Character PCI PC II PC Ill
VEGLEAF 0-160 0-294 0-230
VEGBAS 0-277 0-452 -0-084
LIGL 0-328 0-422 -0-126
MSPIKE -0-197 0-550 0-131
MPEDL -0-399 0-234 -0-071
BEAKL 0-415 -0-134 -0-016
PERLE 0-446 0-034 0-041
MGLUM 0-209 -0-203 0-288
FGLUM 0-075 -0-258 -0-633
BRISTLE 0-407 0-023 -0-118
NUTL 0-077 -0-217 0-634

Abbreviations follow Table 2.

accounted for 83-5% of the variance, with 16-5% of the variance being accounted for on CDF II.
The greatest contribution to variate one is given, in descending order, by MPEDL, BRISTLE ,
BEAKL and FGLUM. Along variate two NUTL, VEGLEAF and LIGL give the greatest
contribution.

Results for the univariate analyses of the mean, standard deviation and range for each character
are given in Table 4. A one-way ANOVA of each individual character showed no significant
differences in the mean values between the taxa included in the C. flava and MT cluster.
Significant differences were identified between the taxa in the C. flava/ MT cluster and the taxa
included in the C. viridula subsp. brachyrrhyncha/C. jemtlandica cluster. However no single
character could be identified that clearly delimited the taxa as all characters had some degree of
overlap in their range. This is consistent with previous studies on the C. flava agg. (e.g. Schmid
1980; Havlickova 1982; Crins 1985; Pykaéla & Toivonen 1994). The perigynium characters
BEAKL, PERLE and BRISTLE were found to be significantly different (p<0-05) between the
C. flaval MT cluster and the C. viridula subsp. brachyrrhyncha!/C. jemtlandica cluster.

Although the sample size of C. jemtlandica from Sweden is very small, PCA gives a clear

disjunction along PC I between these and the MT plants. DA gives a similar disjunction along >

CDF I. This is further supported by results of the ANOVA, which reveals statistically significant
differences in the mean values for characters MSPIKE, BEAKL, BRISTLE, PERLE and NUTL
(p<0-05) between the two groups. Within the C. viridula subsp. brachyrrhynchal/C. jemtlandica
cluster the mean length of characters MPEDL and NUTL gave the clearest differentiation
(p<0-001) between C. viridula subsp. brachyrrhyncha and C. jemtlandica with characters PERLE
and BEAKL also being significantly different (p<0-01).

DISCUSSION

Morphometric analysis does not support the classification of C. jemtlandica as a subordinate taxon
of C. flava. From PCA there is marked separation along PC I between C. jemtlandica and C. flava
and between C. jemtlandica and the MT plants. However, C. jemtlandica is not clearly
differentiated from C. viridula subsp. brachyrrhyncha although it does form a sub-cluster within
this group. Differentiation between C. jemtlandica and C. flava and between C. jemtlandica and
the MT plants is maintained in the results of the DA with marked separation along CDF I. Unlike
the results from PCA C. jemtlandica forms a distinct cluster when CDF I is plotted against CDF I.
However, this cluster is not clearly separated from the C. flava/MT or C. viridula subsp.
brachyrrhyncha clusters along either CDF I or CDF II individually. These results support the close
phenetic relationship that exists within the C. flava agg. C. viridula subsp. brachyrrhyncha and
C. jemtlandica have been recognised as two distinct species under the names C. lepidocarpa
(= C. viridula subsp. brachyrrhyncha) and C. jemtlandica respectively (Palmgren 1938;

CAREX FLAVA AGG. AT MALHAM TARN Sil

Canonical discriminant function II

6 -4 2 0 2 4 6
Canonical discriminant function |

4 C. flava 4 MTP @ C. viridula subsp. brachyrrhyncha

© C. jemtlandica x Centroid for Carex flava © Centroid for MTP

+ Centroid for Carex viridula subsp. brachyrrhyncha x Centroid for Carex jemtlandica
FIGURE 3. Scatter plot of CDF I vs CDF II for the measurements of C. flava , MTP, C. viridula
subsp. brachyrrhyncha and C. jemtlandica .

Halkka et al. 1992); however, they are generally united by Scandinavian botanists under the name
C. lepidocarpa and accorded subordinate rank as C. lepidocarpa subsp. lepidocarpa and
C. lepidocarpa subsp. jemtlandica (Palmgr.) Palmgr. (Palmgren 1926, 1959; Pykala & Toivonen
1994; Hedrén & Prentice 1996). C. lepidocarpa subsp. jemtlandica is described by Hedrén (1996)
as endemic to Fennoscandia, largely replacing C. lepidocarpa subsp. lepidocarpa north of the
limes norrlandicus. This northern distribution is reflected in the original name (nomen nudum)
given to this taxon in 1910 by Palmgren as C. lepidocarpa Tausch * septentrionalis Palmgr. forma
jemtlandica. Extensive studies on morphology (Hedrén 1994, 1996) and genetic differentiation
using allozymes (Hedrén & Prentice 1996) support the recognition of this taxon as a subordinate of
C. lepidocarpa (=C. viridula subsp. brachyrrhyncha).

On the basis of work on demography, ecology, karyology, hybridisation and morphology of
Swiss material Schmid (1983) reduced the status of C. lepidocarpa to that of a subspecies of
C. viridula Michx. He further considered that the name of brachyrrhyncha Celakovsky to have
priority over that of /epidocarpa. Former subspecies of C. lepidocarpa were thus recognised as
subordinate taxa of C. viridula subsp. brachyrrhyncha and given varietal status. The exception to
this was C. jemtlandica, which Schmid did not recognise at any level (cf. Schmid 1983). Based on
biometric analysis of herbarium specimens Crins (1985) similarly concluded that C. jemtlandica
did not merit recognition at any taxonomic level. However, the studies by Schmid (1983) and
Crins (1985) included specimens of C. jemtlandica collected from a greater geographical range,
extending as far south as Switzerland, than the range of the taxon recognised by Hedrén (1996).
Crins further renamed C. viridula subsp. brachyrrhyncha var. lepidocarpa as C. viridula subsp.
brachyrrhyncha var. elatior (Schltdl.) Crins.

Studies of allozyme variation by Bruederle & Jensen (1991) within the C. flava agg. concluded
that C. flava was enzymatically distinct from C. viridula, but that within C. viridula agg. clear
differentiation was not possible. However, they noted that C. viridula subsp. brachyrrhyncha
formed a relatively distinct group within the aggregate. They concluded that their data were
insufficient to justify a reclassification of this taxon and so they retained the classification of
Schmid. They also reported that C. jemtlandica was not separable from C. flava using allozymes.
This is in contrast to Hedrén & Prentice (1996) who found in their larger study of C. jemtlandica
that it was enzymatically distinct. Pykéla & Toivonen (1994) suggest that Bruederle & Jensen may
have included specimens of C. flava in their study rather than C. jemtlandica.

S12 N. BLACKSTOCK AND P. A. ASHTON
TABLE 4. MEAN, RANGE AND + 1 SD FOR MORPHOLOGICAL CHARACTERS
MEASURED FOR SPECIMENS OF C. FLAVA, THE MT POPULATION,
C. VIRIDULA SUBSP. BRACHYRRHYNCHA AND C. JEMTLANDICA.
Character C. flava MT C. viridula subsp. C. jemtlandica
(n=40) (n=10) brachyrrhyncha (n=40) (n=10)
VEGLEAF 53-41 (+18-48) 39-28 (45-97) 40-95 (+19-17) 55-82 (427-70)
21-41-105-05 31-77-52-79 11-22-107-76 21-27-109-83
VEGBAS 5-46 (+1-48) 5-10 (41-24) 4-23 (+1-28) 3-69 (+0-47)
3-02-11-78 3:50-7:29 2:30-8:79 2:89-4:54
LIGL 3-68 (+1-57) 2°97 (41-07) 2:04 (41-58) 1-21 (+0-37)
1-15-8-25 1-50-5-21 0-81—9-42 0-70-1-80
MSPIKE 11-48 (44-25) 9-72 (+1-80) 15-17 (45-18) 15-64 (+5-01)
3-81-22-01 6-79—13-12 5-32-24-89 9-63—25-51
MPEDL 1-61 (+1-85) 1-03 (41-21) 13-86 (+7-50) 6-41 (+3-06)
0:00-6:58 0-00-2-42 0-00-36-34 3-81-11-24
BEAKL 2-18 (+-038) 252 (20-27) 1-45 (40-31) 1-82 (+0-17)
1-17—3-00 1-94—2-82 0-73—2-38 1-61-2-01
PERLE 5-08 (+0-73) 5-14 (+0-46) 3-80 (40-27) 4-49 (+0-57)
4-02-6-63 4-51-5-74 3-12-4-48 3:72-5:28
MGLUM 3-19 (+0-44) 3-11 (+0-39) 2-73 (+0-38) 3-01 (40-51)
2-27-4-23 2-11-4-01 2-06-3-45 2-35—3-65
FGLUM 2-09 (40-29) 2-00 (40-31) 1-92 (+0-36) 1-69 (+0-21)
1-53-3-10 1:72-2:61 1-14—2-65 1-38—1-99
BRISTLE #33 (42-03) 8-40 (+1-71) 3-50 (+1-92) 4-10 (+0-74)
3-11 5-11 0-8 3-5
NUTL 1-60 (+0-13) 1-73 (40-11) 1-56 (+0-19) 1-96 (+0-17)
1-37-1-91 1-61-1-91 0-92-1-38 1-68-2:-21

Abbreviations follow Table 2.

Although the present study shows that morphological differences between C. viridula subsp.
brachyrrhyncha and C. jemtlandica are not fully developed there is a degree of separation
indicated by the PCA and DA analyses. The same chromosome number of 2n=68 and limited
morphological distinction supports the recognition of these taxa at a very close taxonomic level.
However, caricologists working in Fennoscandia, where both C. viridula subsp. brachyrrhyncha
and C. jemtlandica occur, consistently classify them as distinct taxa (e.g. Palmgren 1959; Pykala
& Toivonen 1994; Hedrén 1996). We support the recognition of C. jemtlandica at a subordinate
level to that of C. viridula subsp. brachyrrhyncha. In order to bring the current nomenclature for
C. jemtlandica into line with that used by Crins (1985) it is proposed that this taxon be renamed
C. viridula Michx subsp. brachyrrhyncha (Celak.) B. Schmid var. jemtlandica (Palmgr.)
Blackstock & P. A. Ashton comb. et stat. nov.

The hypothesis that the MT plants are of hybrid origin is not supported by this study. Jermy et
al. (1982) preferred to recognise hybrids as being both morphologically intermediate between the
parents and also being wholly sterile. However, they acknowledged that the MT plants are fertile.
It is implied that this fertility may be due to the population being a stabilised introgressed hybrid
between C. flava and C. viridula subsp. brachyrrhyncha, with the former species now being
extinct at this site. In Switzerland natural and cultivated hybrid populations between C. flava and
C. viridula subsp. brachyrrhyncha have been studied by Schmid (1983). Fl-hybrids were found to
be almost completely sterile with 0% seed set and pollen fertility of only 0-c.3% (Schmid 1983).
When directional backcrosses do arise, forms identical to the parent species in both morphology
and fertility may eventually result. In these populations it is C. viridula subsp. brachyrrhyncha that
may be out-competed and eventually ousted by C. flava, the Fl-hybrid or backcrosses with either
parent (Schmid 1980). This does not fit with the model cited by Jermy et al. (1982) who suggested
that C. flava was probably eliminated by introgression with C. viridula subsp. brachyrrhyncha,
and the putative hybrids at Malham Tarn, Greywell Moors and Lough Corrib being a product of

CAREX FLAVA AGG. AT MALHAM TARN 513

TABLE 5. PERIGYNIUM LENGTHS (MM) FOR C. FLAVA

Author and Year Area Perigynium length (mm)
Robertson (1984) North America 3-0-6-0
Hedrén, in itt. Sweden 3-5+
Havlickova (1982) Czechoslovakia 3-8-6:3
Crins (1985) North America 4-0-6:3
Mackenzie (1935) North America 4-5-6-0
Schmid (1985) Europe 4-5-6-0
Palmgren (1958) Scandinavia 4-5-6-7
Hultén (1968) North America 5-0-6-0
Chater (1980) Europe 5-0-7-0
Nelmes (1945) UK 5-0-7-0
Sell & Murrell (1996) UK 5-0-7-0
Davies (1953) UK 5-25-7-0
Stace (1997) UK 5-5-6-°5
Jermy et al. (1982) UK 6-6-5
Rose (1989) UK 6-6-5

this process. The hypothesis that the MT plants are intermediate in morphology is also
unsupported, as there is marked separation in the PCA along PC I between C. viridula subsp.
brachyrrhyncha and the MT plants, but no separation between C. flava and the MT plants. A
similar separation occurs along CDF I using DA. Based on the biometric data presented here the
MT plants are aligned with C. flava s. s.

A chromosome count of a specimen from the MT population by Davies (1953) gave a haploid
number of n=30. This is consistent with other chromosome counts for C. flava s. s. (Davies 1953;
Halkka et al. 1992; Schmid 1982) but inconsistent for chromosome counts for C. viridula subsp.
brachyrrhyncha (n=34) (Davies, 1953; Halkka et al. 1992; Schmid 1982) and C. x alsatica
(n=irregular) (Schmid 1982).

Difficulties over the correct classification of the MT plants may be due, in part, to its apparently
obvious morphological dissimilarity to the Roudsea Wood population of C. flava. It should be
noted that although both populations are situated at approximately the same latitude, the Roudsea
Wood population is situated in a semi-shaded wood near sea level whereas the MT population is
situated on an exposed site at approximately 380 m. It would appear that these two populations are
approaching the opposite extremes of morphological range within this species. When continental
European and North American material is taken into consideration it becomes apparent that both
populations fit within the phenotypic range of this species. Table 5 indicates that there may be an
over reliance upon large utricle size to separate C. flava from its closely related allies within the
British Isles.

In conclusion the population of yellow sedges discovered by G. A. Shaw in 1946 at the Tarn
Moss, Malham should be recognised as a second extant population of C. flava s. s. in the British
Isles. Furthermore, problematic populations whose identity is uncertain such as those already
identified as C. flava hybrids at Greywell Moors, Hants and by the River Corrib near Menlough
north-east of Galway City need to be reviewed as soon as material is available. In light of the
evidence presented here it is possible that C. flava is not as rare as currently thought in the British
Isles but has been overlooked or misidentified in the past.

TAXONOMIC TREATMENT

The following key should serve to discriminate between C. flava, C. viridula subsp.
brachyrrhyncha var. elatior and C. viridula subsp. brachyrrhyncha var. jemtlandica . However, it
should be noted that no single character can satisfactorily discriminate between the taxa and where
two or more of the taxa are sympatric the taxonomic affinities may be further confused. Where
possible several specimens from one population should be carefully compared and mean character
values used. Only perigynia from the middle of a pistillate spikelet should be used.

514 N. BLACKSTOCK AND P. A. ASHTON

KEY TO TAXA:

1 Utricles straight or curved, usually <5 mm long, abruptly contracting into a conspicuous beak
usually <2 mm long with usually <5 bristles on each side of the beak margin. Leaves of the
fertile tiller usually <4-5 mm wide, longer or shorter than the culm. Ligule usually <2-5 mm
long acute to rounded. Solitary staminate spikelet + sessile to conspicuously pedunculate,
PELMINAL ose.c 500k 55 vacantncs ivceenewssananagtevsicervecderiinectea Ueius enecesunebzoneccececnet heme auansnes Remnenee ees eae ee 2

1 Utricles curved usually >3-8 mm long, gradually tapering into a long 1-5-3 mm beak with a
conspicuously scabrous margin of usually 5—11 bristles on each side. Leaves of fertile tiller
usually >3 mm wide, usually + as long as the culm. Ligule usually >3 mm, acute. Solitary

Staiminate spikelet + sessile; terminal <.....:.0.1.:5:chccscienessscceeveeeese eer 1. C. flava
2 Utricles usually 3-5-5 mm long; beak 1-5-2 mm long. Leaves of fertile tiller usually <3 mm
WICC vis sciatic dcnancantccccess cada cedsiecedtanceaanebcdanddunetnudeeeciehlbebiaccespeaceeadeagecapeteceeees:aee eee eee ee 3
2 Utricles usually <4 mm long: beak usually <1-5 mm long. Leaves of fertile tiller 1-5—-4-5 mm
WYN sieora ices tice tynseisaauh tater tees ctipnceveas ee Renee aeem tenes 2. Other members of the C. viridula agg.

3 Utricles + dense; beak usually deflexed. Staminate spikelet usually conspicuously pedunculate.
Pistillate spikelets usually 2—3, + separate, ovate to cylindrical. Leaves of fertile tiller usually
shorter than half of the Culm scssssecececesesesenss 2aa. C. viridula subsp. brachyrrhyncha var. elatior

3 Utricles + lax; beak usually straight. Staminate spikelet usually sub-sessile to shortly
pedunculate. Pistillate spikelets usually 1-2 + contiguous to approximate, globose to ovate.
Leaves of fertile tiller usually longer than half of the culm

ee eee ere en er ere 2ab. C. viridula subsp. brachyrrhyncha var. jemtlandica

The following is a brief taxonomic treatment and includes only some of the most important
synonyms. For a fuller taxonomic treatment see Schmid (1983) and Crins (1985).

1. C. flava L., Sp. Pl. 2: 975 (1753). Type: LINN Savage Cat. No. 1100.40 (Holotype: LINN).

Synonyms:

C. patula Host., Icon. Descr. Gram. Austriac. 1: 48-49 (1801). Type: Austria (Holotype: W,
destroyed).

C. flava subsp. gaspensis Fernald, Rhodora 8: 200 (1906). Type: Canada, Quebec, gravelly banks
of Bonaventure River, 8 August 1904, J. F. Collins, M. L. Fernald & A. S. Pease s.n.
(Lectotype: GH, n.v. selected by Crins 1988).

2. C. viridula Michx. Fl. Bor-Amer. 2: 170 (1803). Type: Canada, between Montréal and Les Trois
Rivieres, Michaux in herb. Michaux (Holotype: P)

2a. C. viridula subsp. brachyrrhyncha (Celakovsky) B. Schmid, Watsonia 14: 317 (1983).

Basionym: ,

C. flava subsp. brachyrrhyncha Celakovsky, Prodr. Fl. Bom. 1: 71 (1876).

Synonyms:

C. lepidocarpa Tausch, Flora (Regensb.) 17: 179 (1834). Type: Czechoslovakia, Praha, (no
collector), no. 1636 (lectotype: PRC, isolectotype: PRC, selected by Crins 1985).

2aa. C. viridula subsp. brachyrrhyncha var. elatior (Schlectendal) Crins, Canad. J. Bot. 67(4):
1058 (1989).

Basionym:

C. flava var. elatior Schltdl., Fl. Berol. 1: 477 (1823). Type: Germany, Berolinensi
(Holotype: B n.v.).

Synonyms:

C. viridula subsp. brachyrrhyncha var. lepidocarpa (Tausch) B. Schmid Watsonia 14: 317 (1983).

C. lepidocarpa Tausch, Flora (Regensb.) 17: 179 (1834).

C. lepidocarpa var. lepidocarpa (Tausch) Petermann, Anal. Pflanzenschl. Leipzig 509 (1846), pro
autonym.

C. flava subsp. lepidocarpa (Tausch) Godr. Fl. Lorraine. 1: 118 (1843).

C. flava subsp. lepidocarpa (Tausch) Nyman, Consp. 771 (1882).

C. flava subsp. lepidocarpa (Tausch) Lange, Haandb. Danske. FI., 4th ed., 144 (1886).

CAREX FLAVA AGG. AT MALHAM TARN 515

2ab. C. viridula subsp. brachyrrhyncha var. jemtlandica (Palmgr.) Blackstock & P. A. Ashton
comb. et stat. nov.

Basionym:

C. lepidocarpa Tausch subsp. jemtlandica Palmgr. in Lindman. Sv. Fanerogamfl., ed. 2: 153
(1926). Type: Sweden, Jamtland, Ostersund, Frés6n, 13 August 1910, A. Palmgren, Caric.
Fulv. No. 53 (lectotype: H, selected by Crins 1985).

Synonyms:

C. lepidocarpa Tausch * septentrionalis Palmgr. forma jemtlandica Palmegr., Caric. Fulvellae Exs.
No. 52-54 (1910) (nom. nud.).

C. jemtlandica (Palmgr.) Palmgr., Mem. Soc. Fauna Flora Fenn. 13: 126 (1937).

C. jemtlandica (Palmgr.) Palmgr. var. gotlandiae Palmgr., Comment. Biol. 20(3): 12 (1958).
Type: Sweden, Gotland, par. Othem, 14 June 1910, A. Palmgren (Holotype: H, Isotype: H,
photo!).

C. jemtlandica (Palmgr.) Palmgr. var. kainuensis Palmgr., Comment. Biol. 20(3): 13 (1958).
Type: Finland, Ostrobottnia kajanensis, Suomussalmi, Ruhtinaansalmi, Rytisuo, 10 July 1937,
O. Kyyhkynen (Holotype: H, photo!).

C. flava L. subsp. jemtlandica (Palmgr.) P. D. Sell Fl. of Great Britain and Ireland 5: 110 (1996).

ACKNOWLEDGMENTS

This research has been carried out as part of a research studentship funded by Edge Hill College of
Higher Education. The authors would like to thank English Nature and The National Trust for
granting permission to collect samples from Roudsea Wood and Malham Tarn respectively. We
would also like to thank everyone who supplied seed and ramets for cultivation. Special thanks go
to Mikael Hedrén for suggesting localities for material from Sweden and his input on the
C. jemtlandica problem.

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(Accepted February 2001)

Watsonia 23: 517-523 (2001) DLT

Conservation of Britain’s biodiversity: Hieracium linguans
(Zahn) Roffey (Asteraceae), Tongue Hawkweed

T. C. G. RICH

Dept. Biodiversity and Systematic Biology, National Museum & Gallery, Cardiff CF 10 3NP

and
G. S. MOTLEY

Countryside Council for Wales, Cantref Court, Brecon Road, Abergavenny, Mons. NP7 7AX

ABSTRACT

Hieracium linguans, Tongue Hawkweed, is a very rare endemic Welsh species. It was first found in 1898 by
A. Ley in the Brecon Beacons (v.c. 42), Wales. It was initially named as a variety and later raised to species
level, its current status. It is a distinct member of Section Tridentata. The historical herbarium and literature
records were somewhat confusing, but indicate three localities in the Tawe Valley. In 1999 and 2000, only one
population of about 80 plants was found in Cwm Haffes (the type locality); it was not refound in the other two
localities. Records from Upper Nedd Glen, Llyn y Fan Fach and the Upper Tywi are probably errors. It is a
polycarpic perennial which flowers in July-August, and regenerates readily from wind-dispersed seeds.
Although it is very rare, it does not seem to be significantly at risk. Seed has been deposited in the Millennium
Seed Bank and it is being cultivated at the National Botanic Garden of Wales.

KEYWORDS: Compositae, Endemic, Lectotype, rare species, Wales.

INTRODUCTION

Hieracium linguans (Zahn) Roffey (Asteraceae), Tongue Hawkweed (from the Latin lingua,
meaning tongue), is a very rare, endemic species. Along with 79 other rare hawkweeds, it has been
included in the 3rd edition of the Vascular Plant Red Data Book (Wigginton 1999). In 1999 David
McCosh, who provided the original list of species for the Red Data Book, suggested that this
species would be worth re-surveying as there were no recent records. A joint project was therefore
set up between the National Museum & Gallery, Cardiff and the Countryside Council for Wales to
establish its current status and determine its need for conservation. In this paper the information
about H. linguans is summarised; full details can be found in Rich (1999), updated here with
additional field work in July 2000.

TAXONOMY

Plants that are now called Hieracium linguans were first found by Augustin Ley in the Brecon
Beacons on 2-3 August 1898. He subsequently distributed specimens through the Botanical
Exchange Club (B. E. C.) as ‘H. gothicum?’ (Ley 1900). When it became apparent that there was
no name for the plants, Ley described them as var. lingua of H. sparsifolium Lindeb. in Linton
(1905), and subsequently distributed many more cultivated specimens through the B. E. C. (Ley
1907, 1908). Zahn (1922) raised Ley’s variety to subspecies level, but transferred it to H.
laevigatum Willd. and provided it with the new subspecific name linguans Zahn in the process
(probably to avoid confusion with another taxon of uncertain status, subsp. lingua Dahlst.). Roffey
(1925) raised it to a species, its current status. The two synonyms of H. linguans (Zahn) Roffey are
thus H. laevigatum Willd. subsp. linguans Zahn (basionym) and H. sparsifolium Lindeb. var.
lingua Ley ex W. R. Linton.

518 T. C. G. RICH AND'G. S. MOTLEY

A type specimen (probably selected by H. W. Pugsley and designated here as the lectotype),
labelled ‘Haffes Glen, West Brecon, 2 August 1898, A. Ley’ is in the Natural History Museum
(BM).

Hieracium linguans belongs to sect. Tridentata (Fr.) Arv.-Touv., of which there are at least
twelve species in South Wales. Pugsley (1948) gave its distinguishing features as the oblong-
elliptic to linear-oblong, sinuate-dentate medium stem leaves, the racemose-corymbose
inflorescence with 4-8 large capitula on long, suberect branches, the greyish-green broad, densely
floccose phyllaries with numerous unequal, short, dark-glandular and fewer longer dark-based
pilose hairs, and dark styles. However, Pugsley’s description includes cultivated material, which is
more robust than the wild material. A comparison of plants in the field with wild-collected
specimens in herbaria suggests that the latter were also selected as the larger plants. Most plants
observed in the wild in 1999 (see below) were less than 20 cm tall, had only one stem (rarely two
stems) and up to four capitula (rarely up to 12 capitula).

DISTRIBUTION

HISTORICAL INFORMATION

Locality and habitat information was extracted from herbarium sheets at BIRM, BM, CGE, LIV,
OXF and NMW and from the literature (e.g. Perring 1968, Ellis 1983). Records from the
Hieracium database were provided by D. McCosh. No records are held in the Rare Plants Database
used to compile the Red Data Book (Wigginton 1999).

There has been significant confusion about the records, possibly partly due to a note from a B.E.
C. report probably referring to H. scoticum F. J. Hanb. being attached to a H. linguans specimen in
BM, and partly to some specimens which have subsequently been identified as different taxa. The
B.E.C. note (Ley 1900) indicated that ‘H. gothicum Backh?’ had been found in Cwm Haffes
(Capel Cellwen, now Callwen), Upper Nedd Glen and above Llyn y Fan Fach, but the style colour
was noted as pure yellow which is not a character of H. linguans. Ley in Linton (1905) noted H.
linguans from Cwm Haffes and the Upper Nedd Glen when he described it as a new variety and
later noted ‘this remarkable plant has not yet been detected except in West Brecon’ (Ley 1909).
Pugsley (1948) only cited material from Cwm Haffes.

Sell & West (1968) stated H. linguans was only recorded from rocky ledges in three localities in
Brecon, and records were plotted in SN81, SN82 and SN84 on the map. The Biological Records
Centre, Monks Wood holds details of the records in the Critical Supplement, but the cards for H.
linguans could not be traced in 1999 (J. Croft, pers. comm., 1999). The SN81 record appears to be
based on the well-known specimens from Cwm Haffes. The SN82 record may be based on 1906
material cultivated by A. Ley from Fan Fechan (CGE) originally determined as H. linguans in
1958 by P. D. Sell and C. West, but later redetermined as H. scoticum by P. D. Sell in 1983. The
SN84 record is probably based on a specimen collected in 1955 by J. E. Raven in Llanwrtyd
(CGE), later redetermined as ‘H. ?subcrocatum by P. D. Sell in 1983. Ellis (1983) cited a record
from the Upper Tywi based on a specimen collected by A. Ley in 1906 in CGE; the only such
specimen is from the Upper Tawe Glen, and thus probably a confusion with the two adjacent river
valleys with similar names. Records for the Nedd Glen (which would probably have been plotted
in either SN81 or SN91) do not appear to have been accepted by recent authors and no material
has been traced.

The historical records traced are summarised in Table 1. Some of the records may be duplicates
labelled slightly differently (e.g. ‘streamside near Capel Cellwen’ and ‘Haffes Glen’ on 2 August
1898; Ley was not consistent in his labelling of specimens), and some dates are suspiciously one
month or one year apart. There appear to be three distinct sites:

1. waterfall at the head of Cwm Haffes;
2. stony delta of the Haffes (“streamside near Capel Cellwen’ assumed to be this too);

3. riverside rocks, Upper Tawe Glen.

CONSERVATION OF HIERACIUM LINGUANS 519

TABLE 1. RECORDS OF HIERACIUM LINGUANS

Date Collector Site Source and notes
2/8/1898 A. Ley Capel Cellwen, streamside near BIRM, BM, LIV; det. P. D. Sell & C. West.
2/8/1898 A. Ley Haffes Glen, West Brecon BM; type of var. lingua Ley.
2/8/1898 A. Ley Haffys Glen, near Callwen BM; det. D. McCosh.
27/7/1899 A. Ley Haffys Glen CGE; det. P. D. Sell.
27/8/1899 A. Ley Waterfall in the Haffes Glen BM; det. P. D. Sell & C. West.
4/7/1906 - Haffes Glen, Callwen LIV; det. J. N. Mills.
5/7/1906 A. Ley riverside rocks, Upper Tawe Glen* CGE; det. P. D. Sell.
10/7/1906 A. Ley stony delta of the Haffes* CGE; det. P. D. Sell.
12/7/1906 H. J. Riddelsdell R. Haffes BM: det. D. McCosh
25/7/1955 J. E. Raven Cwm Haffes CGE; det. P. D. Sell & C. West.
25/7/1955 C.E. Andrews head of Cwm Haffes, rocks by water-BIRM; conf. T. Rich.

fall

28/7/1956 J. E. Raven (unlabelled, assumed Cwm Haffes) herb. D. McCosh.
26/7/1957 B.A. Miles side of large waterfall, Cwm Haffes CGE; det. P. D. Sell & C. West.

Cultivated material

23/6/1906 A. Ley West Brecon, cultivated at Sellack BM, CGE, LIV, NUW, OXF (Ley 1907).

muIso7 =A. Ley : West Brecon, cultivated BM (Ley 1908).

26/6/1908 A. Ley Brecon, cultivated BM (labelled incorrectly as syntype), CGE.

17/7/1909 E. F. Linton cultivated at Edmondsham NMW: det. D. McCosh.

Errors

2/8/1898 A. Ley Cliffs of Llyn-y-fan-fechan Ley (1900); confusion of B.E.C. labels.

3/8/1898 <A. Ley Nedd Glen, Breconshire Ley (1900); cited Upper Nedd Glen by Ley
in Linton (1905), no material traced.

1906 A. Ley Upper Tywi Ellis (1983); error for Upper Tawe.

14/6/1906 A. Ley Fan fechan, Carmarthen, cultivated CGE; redetermined as H. scoticum, P. D.

Sell 1983, LIV (lacks collector or
‘cultivated’ ).

24/7/1955 J. E. Raven Llanwrtyd, Brecon CGE; redetermined as H. ?subcrocatum by
PD Sell; 1983.

* = one specimen on a herbarium sheet representing two localities.

FIELD SURVEYS 1999-2000
The historical records were used to direct field work. Llyn y Fan Fach and the Nedd Glen were
also investigated as there were some possible records.

1. Cwm Haffes (also known as Glen Haffys, etc.)

Cwm Haffes is a large ravine cut by the River Haffes through the hillside above Callwen. It is the
type locality for H. linguans, and also contains at least two other species, variously named it
seems, one as H. diaphanum Fries, H. submutabile (Zahn) Pugsley and H. anglorum (Ley)
Pugsley, and the other as H. nidense (F. J. Hanbury) Roffey (incorrectly), H. substrigosum (Zahn)
Roffey and H. uiginskyense Pugsley s.1. (including H. trinitatis Pugsley).

The gorge was searched on 30 July 1999 by T.C.G.R. and G.S.M.. H. linguans was found on
east-facing ledges of Sgwd Ddu, the upper waterfall, at SN829179, c. 450 m altitude, none being
found elsewhere in the Cwm or its tributaries. The ledges occupied by H. linguans varied from dry
rocky ledges to wet moss carpets depending on proximity to the waterfall. Fifty-three flowering
and three vegetative mature plants were recorded. There were about 30 seedlings, mostly on moss
Carpets on the north side of the waterfall, indicating abundant potential for regeneration, though
most mature plants occurred in mineral soil at the back of the ledges.

520 Te@. G: RICEVAND’G: S: MOTLEY:

2. Stony delta of Cwm Haffes

The River Haffes delta below the Cwm Haffes gorge is about 300 m long and consists of many
huge boulders and shingle with patches of grassland and scrub. The upper part is heavily grazed
and no Hieracium plants were found on 30 July 1999. The lower part is partly included in a grazed
field, and partly in an ungrazed caravan site; H. cf. diaphanum was the only species found at the
latter on 19 July 2000.

3. Upper Tawe Glen
Ley appears to have recorded several Hieracium species from “Upper Tawe Glen’, but his meaning
is unclear. It might refer generally to the Tawe valley north of Callwen, or perhaps more strictly to
Nant y Llyn (SN8420) as he cited the locality of H. tavense using the same words (see Rich &
Houston 2000), or might even refer to Cwm Haffes. The unique combination of habitat and
locality (cf. Table 1) suggests it is a distinct site from the Cwm Haffes sites.

On 19 July 2000 all tributaries and ravines of the Tawe above Callwen were searched without
success, though at least seven other Hieracium species were found.

4. Llyn y Fan Fach

Whilst there is some potential for confusion, it seems very likely that the corrie above Llyn y Fan
Fach (c. SN8021) is the same site that A. Ley called ‘Fan-fechan, Carmarthen’ as there are many
of his other records which equate to this site (a Brecon “Fan Fechan’ at SN8319 has no suitable
habitat). Although the historical records are known to be in error a search was considered
worthwhile due to its proximity to Cwm Haffes and large areas of potential habitat. The cliffs
above the lake were searched from below on 5 August 1999 without success, although six other
Hieracium species were seen.

5. Upper Nedd Glen
The unconfirmed historical records for Nedd Glen and Upper Nedd Glen suggested a search would
be worthwhile. The area above Blaen-nedd Isaf (SN9114) is a shallow valley with some ravines
and waterfalls on the west side; it was searched twice unsuccessfully in 1999 (five other
Hieracium species seen, including H. scoticum). The area to the south, a wooded gorge over
limestone, was searched once but is unsuitable habitat (only one Hieracium species seen).

The fact that Ley specifically stated this was a site for his new variety is puzzling, but in the
absence of material no firm conclusion can be drawn as to whether it was there or not: it is
currently thought to be an error.

Thus H. linguans is now only known from one locality, with a total population of about 80 plants
in 1999, and it could not be refound at the other two localities. Given the general difficulty of
identifying Hieracium and the large areas over which it might occur, it would not be surprising if
other unknown populations existed. An updated distribution map is given in Figure 1.

ECOLOGY

LIFE CYCLE
Hieracium linguans is a perennial. It flowers in late July and early August and is probably an
obligate apomict like the other Hieracium species. The mean number of capitula per plant at Sgwd
Ddu in 1999 was 2-6 (ranging 1-12); over half of the plants had only one or two flowering heads.
Of the 53 flowering plants, five had aborted inflorescences. It produces abundant fruit, and the
seeds have a small pappus and are generally wind dispersed. Some seeds may have been washed
down Cwm Haffes, occurring on shingle far below the classic waterfall site. Seedlings were
observed in mid-summer before fruit was shed suggesting that germination mainly occurs in the
spring.

Ley (1908) noted it grew readily in the garden and increased by seed.
HABITAT

The rocks of the Sgwd Ddu waterfall are composed of the Plateau Beds of the Farlow Series of the
Devonian Upper Old Red Sandstone (Geological Survey of Great Britain 1979), which form a hard

CONSERVATION OF HIERACIUM LINGUANS 521

es 1. Distribution map of Hieracium linguans. ® 1999; x probable erroneous records. Plotted using
MAPW.

522 T. C. G. RICH AND G. S. MOTLEY

outcrop with ledges and exposed rocks. The H. linguans plants were rooted in immature rankers on
the ledges, directly into rock crevices with no apparent soil, or into moss carpets in the wetter areas
near the waterfall itself. Soil from the base of one plant was pH 5-7 (measured with a pHep2
Hanna pocket-sized pH meter in a 50:50 mixture with distilled water).

Vascular plants recorded on the ledges associated with H. linguans were Ajuga reptans L.,
Anemone nemorosa L., Arrhenatherum elatius (L.) P. Beauv. ex J. & C. Presl, Asplenium viride
Huds., C. pulicaris L., Carex viridula Michx., Cirsium palustre (L.) Scop., Festuca rubra L.,
Filipendula ulmaria (L.) Maxim., Galium palustre L., Holcus lanatus L., H. mollis L., Lychnis
flos-cuculi L., Molinia caerulea (L.) Moench, Phegopteris connectilis (Michx.) Watt, Pinguicula
vulgaris L., Rumex acetosa L., Taraxacum sp., Valeriana officinalis L., Vicia sepium L. and Viola
riviniana Rchb., with the bryophytes Aneura pinguis (L.) Dumort., Bryum pseudotriquetrum
(Hedw.) P. Gaertn., B. Mey. & Scherb., Ctenidium molluscum (Hedw.) Mitt., Dicranella palustris
(Dicks.) Crundw. ex E. F. Warb., Fissidens adianthoides Hedw., Palustriella commutata (Hedw.)
Ochyra var. commutata, Pellia endiviifolia (Dicks.) Dumort., Philonotis fontana (Hedw.) Brid. and
Preissia quadrata (Scop.) Nees. The vegetation on the ledges is somewhat heterogeneous and did
not fit any community of the National Vegetation Classification (Rodwell et al.1991 et seq.). Only
a few small shrubs were present and the waterfall was essentially unshaded.

CONSERVATION

Hieracium linguans is not protected under Schedule 8 of the Wildlife and Countryside Act 1981,
but is included in the Vascular Plant Red Data Book (Wigginton 1999), which should draw
attention to its rarity and be sufficient to ensure its survival. The Cwm Haffes waterfall is included
within the Mynydd Du (Black Mountain) S.S.S.I., and is also part of the Brecon Beacons National
Park. These designations give a significant degree of protection to the site, and thus to the species.
The land is common land managed under an agreement by the Brecon Beacons National Park
Authority. Re-surveys every five years are likely to be sufficient to monitor its survival.

Although the area around the waterfall is intensively grazed, the H. linguans cliff is inaccessible
to sheep, and thus essentially unmanaged. No management is currently required, and the only
potential management in the future might be control growth of scrub or trees.

The main threats to its survival in Cwm Haffes are potential rock falls (here considered
relatively unlikely) and hydroelectric schemes with consequent changes in water regime (a scheme
was proposed for this river in 1997). It is difficult to be certain of causes for loss elsewhere, but
grazing might have contributed to its disappearance from the delta below Cwm Haffes. Responses
to long-term climate change are also unknown, but as it grows readily in cultivation it is likely to
survive warmer temperatures. There are minor potential threats from collecting by botanists (most
of the plants are out of reach) or potential damage by rock climbers (most likely to be ice climbing
in cold winters).

Seeds from seven plants were collected on 30 July 1999 and sent to the Millennium Seed Bank
at the Royal Botanic Gardens, Wakehurst Place for long-term storage. It is also being cultivated at
the National Botanic Garden of Wales.

ACKNOWLEDGMENTS

This work was jointly funded by Countryside Council for Wales and National Museum and
Gallery, Cardiff as part of a collaborative project into conservation and ecology of critical species.

We would like to thank Andy Jones of C.C.W. for his help, Clive Stace and an anonymous
referee for nomenclatural advice, Tony Tipper for help with field work, David McCosh for records
from the B.S.B.I. Hieracium database, discussion and comments on the manuscript, and the
Keepers of the herbaria for access to specimens and libraries.

CONSERVATION OF HIERACIUM LINGUANS 523
REFERENCES

ELLIS, R. G. (1983). Flowering plants of Wales. National Museum of Wales, Cardiff.

GEOLOGICAL SURVEY OF GREAT BRITAIN (England and Wales) (1979). Sheet 231, Merthyr Tydfil. Solid and
drift edition. 1:50,000. Ordnance Survey, Southampton.

LEY, A. (1900). H. gothicum, Backh.?. Report of the Botanical Exchange Club of the British Isles 1: 580.

Ley, A. (1907). H. sparsifolium, Lindeb. var. lingua, Ley. Report of the Botanical Exchange Club of the
British Isles 2: 234.

Ley, A. (1908). H. sparsifolium, Lindeb. var. lingua, Ley. Report of the Botanical Exchange Club of the
British Isles 2: 300.

Ley, A. (1909). Brecon and West Yorkshire hawkweeds. Journal of botany 47: 8-16 and 47-55.

LINTON, W. R. (1905). An account of the British Hieracia. West, Newman & Co., London.

PERRING, F. H., ed. (1968). Critical supplement to the Atlas of the British flora. Botanical Society of the
British Isles, London.

PUGSLEY, H. W. (1948). A prodromus of the British Hieracia. Journal of the Linnean Society of London
(botany) 54: 1-356.

RICH. T. C. G. (1999). Distribution and conservation of Hieracium linguans, Tongue Hawkweed. Unpublished
report from National Museum of Wales to Countryside Council for Wales.

RicH, T. C. G. & Houston, L. (2000). Conservation of Britain’s biodiversity: Hieracium tavense
(Asteraceae), Black Mountain Hawkweed. Watsonia 23: 311-316.

RODWELL, J. S. et al., eds., (1991 et seg.). British plant communities. Cambridge University Press, Cambridge.

ROFFEY, J. (1925). Hieracium, in HANBURY, F. J., ed., The London catalogue of British plants. 1\th ed. G.
Bell & Sons, London.

SELL, P. D. & WEST, C. (1968). Hieracium L., in PERRING, F. H., ed., Critical supplement to the Atlas of the
British flora. Botanical Society of the British Isles, London.

WIGGINTON, M. J., ed., (1999). British Red Data Books. 1. Vascular Plants. 3rd ed. Joint Nature Conservation
Committee, Peterborough.

ZAHN, K. H. (1922). Compositae — Hieracium, in ENGLER, A., ed., Das Pflanzenreich IV. 280. Heft 79: 874.
Engelmann, Berlin.

(Accepted March 2001)

t

ria

Watsonia 23: 525-531 (2001) 525

A new subspecies of Bromus hordeaceus L. (Poaceae)
L. M. SPALTON

6 Marine Parade, Budleigh Salterton, Devon, EX9 6NS

ABSTRACT

A new subspecies of Bromus hordeaceus is described and named subsp. longipedicellatus. It has long pedicels
and branches and thus resembles B. commutatus, but the lemmas are papery which is a feature of the
B. hordeaceus group and its anthers are longer. The grass is fertile and comes true from seed. The reliability of
various taxonomic characters in Bromus is discussed.

KEYWORDS: B. commutatus, B. racemosus, diagnostic characters, taxonomy.

INTRODUCTION

In 1977 L. J. Margetts found an odd Bromus in Cornwall that had long branches and long pedicels
but also papery lemmas. He sent it to C. E. Hubbard at Kew who suggested that it might be ‘(?) B.
commutatus x B. mollis or B. commutatus var. pubens (pers. comm. Hubbard to L. J. Margetts
1978). Margetts later found it in Devon and further specimens all from one site in Devon were
determined as B. commutatus by P. J. O. Trist (pers. comm. to L. J. Margetts 1987), H. J. M.
Bowen (pers. comm.,1998) and T. A. Cope (pers. comm., 1998); as ‘(?) B. racemosus x B.
commutatus + B. hordeaceus’ by T. A. Cope (pers. comm. 1993); as ‘(?) B. commutatus var
pubens’ by R. M. Payne (pers. comm. to L. J. Margetts 1997) and A. Copping (pers. comm.,
1997); as ‘(?) B. hordeaceus x B. commutatus’ by C. Stace (pers. comm., 1998) and T. B. Ryves
(pers. comm.,1998); as ‘a new var. or ssp. of B. hordeaceus’ by M. Kerguélen (pers. comm.,
1998); as ‘a new taxon unnoticed until now with close affinities to B. hordeaceus by H. Scholz
(pers. comm., 1999); and as ‘an unusual variant of normal B. hordeaceus by T. A. Cope (pers.
comm., 1999).

Specimens of the grass were also sent to F. Llamas in Spain, F. Sales in Portugal, P. W.
StahIman in U.S.A., E. Pavlick in Canada, and E. Edgar in New Zealand. None of them reported
that they recognised the grass.

To determine its identity, in 1997 a comparative morphological study was commenced on this
puzzling grass and on the potentially related taxa B. hordeaceus subsp. hordeaceus, B. racemosus,
B commutatus Schrad. and B. commutatus var. pubens Wats. Botanists were requested to send
material for examination (Spalton pers. comm. to several botanists 1997, 1999, 2000), which many
of them did. At the same time herbarium specimens were examined and the grasses were
cultivated. The grass was given a provisional working name of ‘longipedicellatus .

METHODS

A total of 3021 specimens of Bromus were studied: 1232 were fresh or recently collected
specimens and 1789 were herbarium specimens from BDD, BM, BRISTM, BTN, DBN, E, GL,
GLAM, HAMU, HCCMS, K, LTR, MANCH, NMW, OLDM, OXF, RAMM, RNG, SLBI,
SUN, TCD, TOR, and WARMS. 2308 specimens were analysed morphometrically: 775 of
‘longipedicellatus’,1201 of B. hordeaceus subsp. hordeaceus, 183 of B. racemosus, 130 of
B. commutatus s. s. and 109 of B. commutatus var. pubens.

Before the main morphological study, there was a pilot investigation of arielet and floret
structure and development. Spikelets were collected at different times and at different stages of
development and all florets in each spikelet were opened and examined. It was found that anther
length became progressively smaller from the lowest floret towards the apex of the spikelet.

526 L. M. SPALTON

Chasmogamy usually only occurred in the lowest two or three florets and these rarely contained
fruit. In the other florets filament development was poor or absent and cleistogamy and fruit
formation was very rapid with the dehisced anthers being distorted and pushed out by the
developing fruit. In each panicle cleistogamous anthers varied widely in length but undehisced
chasmogamous anthers from the lowest florets did not. Smith (1972) and Clapham et al. (1962)
pointed out that cleistogamy changes anther size and according to Smith & Sales (1993) only
intact (undehisced) anthers should be measured. Pollen was also measured, but it varied widely in
size and too much to separate these taxa.

In the main study, culm height was first measured. In multi-culmed specimens the heights of
mature culms were averaged. Panicle length was measured from the node of the lowest branch or
pedicel to the apex of the terminal spikelet. Branches were measured from their node to the base of
their terminal spikelets. Pedicels were measured, and the pedicel and branch lengths were also
compared with the length of the spikelet that each of them bore. Nine spikelets from each panicle
were measured, three near the apex, three near the basal node and three from the middle region and
the measurements were averaged. Immature and atypical spikelets were ignored. Under lateral
lighting with a hand lens, it was noted whether a substantial majority of the lemmas had veins that
were protruding.

Some spikelets were removed and, whether fresh or dried, were soaked for two hours in water
containing a wetting agent. Soaking was to restore tissue, to facilitate dissection and produce a
more uniform condition for testing lemma texture. The lengths of the lowest lemmas and of the
lowest rhachilla-segments were measured. Mature undehisced anthers in the lowest florets were
measured and if the lowest florets were empty (indicating that chasmogamous anthers had already
been extruded and dropped) this was noted. Fruit development was always recorded and quantified
by the length of the developing fruit.

Lemma texture was assessed on a slide under a binocular microscope by holding the lemma at
the base or by the awn and probing it with a blunt needle. If necessary, previously soaked lemmas
of B. commutatus and B. hordeaceus were used as comparative controls. Papery lemmas were thin
and fragile and readily collapsed or bent under pressure. Leathery lemmas were thicker and stiffer
and therefore more resistant to pressure, and when they eventually give way it was over a wider
area and the lemma would resume its shape when the pressure was removed. In doubtful cases
additional lemmas were tested. If fruit was formed or was being formed it had to be removed
before assessing lemma texture.

Graticules in 30x binocular and 1000x monocular microscopes were used for all fine
measurements. Measurements in the text and in Tables 1 and 2 are range measurements obtained
with some extremes eliminated but the number of these extremes was never allowed to exceed 2%
from the upper and 2% from the lower limits. If there was any evidence of immaturity the data
were not recorded. If a spikelet was not fully developed, the upper glume appeared longer than
usual in comparison with the length of the spikelet. Immaturity was confirmed if under a high-
power microscope the anther cuticle was fragile and partially translucent and the pollen grains
lacked full content.

A character often used to pick out Bromus species in the field is the crowded or lax nature of the
panicle, and this was the first character which drew attention to ‘longipedicellatus’. In Stace
(1997), plants with a +dense panicle usually with all pedicels shorter than spikelets and with
papery lemmas are referred to B. hordeaceus, and plants with +lax panicles with at least some
pedicels longer than spikelets and with rather coriaceous lemmas are referred to B. racemosus and
B. commutatus. A somewhat similar wording was used by Smith (1980) and by Clapham ef al.
(1987). It is often difficult to decide whether a panicle is lax or dense and some plants of B.
hordeaceus often had one or two pedicels that were longer than the spikelets. Furthermore, lax
panicles frequently had long branches with or without long pedicels.

It was considered that the characters for determining lax panicles should be revised to recognise
the contribution of branching to laxness. After evaluating various possibilities including the
method used by Kerguélen (1995) in France, mentioned in Table 2, it was found that the panicle of
‘longipedicellatus’ could best be distinguished from that of subsp. hordeaceus by having at least
four pedicels and branches that are longer than the spikelets that they bear (branches measured to
the base of the terminal spikelet) When other diagnostic characters were also taken into
consideration there were few indeterminates. This formula also separated B. commutatus and most

BROMUS HORDEACEUS SUBSP. LONGIPEDICELLATUS 527

FIGURE 1. Panicle of Bromus hordeaceus L. subsp. longipedicellatus L. M. Spalton. L. M. Spalton 1926, v.c.
12, N. Hants, Hurstbourne Priors, SU445452, 20 May 2000. Scale bar = 1 cm.

specimens of B. racemosus from subsp. hordeaceus. It is suggested that this empirical formula,
while not perfect, is more satisfactory than the present descriptions of laxness and denseness that
the Floras have had to use (Table 2).

In 1999 and 2000, the grasses were cultivated in Cardiganshire, in Devon (two stations), in
Suffolk and, to a lesser extent, in Lancashire and Norfolk. Seed sources were as follows:
‘longipedicellatus’ from ten sites in Devon; subsp. hordeaceus from two sites in Devon; and
B. racemosus from one in Devon; B commutatus from two sites in Somerset; and B. commutatus
var pubens from Anglesey and Suffolk. Seed was sown in September (when these plants germinate
in the wild) in John Innes Compost in four or six inch pots, germinated initially under glass and
then placed outside for autumn growth and over-wintering. However, in one station the plants
remained under glass throughout. Some seedlings were also pricked out on to ordinary garden soil
and seed was also sown directly into the soil. From March 2000 one third of the plants of
‘longipedicellatus’ and of subsp. hordeaceus were enriched by doses of a fertiliser (Liquid
Growmore).

528 L. M. SPALTON
TABLE 1. MORPHOLOGICAL DATA ON THE FIVE BROMUS TAXA
Character B. hordeaceus B. hordeaceus B. racemosus B. commutatus B. commutatus s.s.

subsp. hordeaceus subsp. var. pubens
longipedicellatus

Indumentum of — usually villous variable and usually long variable from __ stiff thick patent
young, intermediate slender patent commutatus to hairs
lower leaf between subsp. hairs, often longipedicellatus
sheath hordeaceus and indeterminate types

racemosus

Culm height to 80 cm 80-75 cm 20-105 cm 60-120 cm 60-150 cm

Panicle usually dense, not lax; atleast4 narrow; usually at lax; atleast4 as for var. pubens
structure more than 3 pedicels and least 4 pedicels pedicels and but very spreading,

pedicels and branches and branches branches some pedicels or
branches exceeding the exceeding the exceeding the branches more
exceeding the length of their length of their length of their than 8 cm long
length of their spikelets spikelets spikelets
spikelets

Panicle length 1-10 cm 10-20 cm 4—20 cm 8-20 cm 10-25 cm

Spikelet length 11-20 mm 13-21 mm 11-18 mm 15-28 mm 15-30 mm

Lemma texture papery papery leathery + leathery leathery

Lemma length 7-10-5 mm 7-5—10-5 mm 7-9 mm 7-5—-11 mm 7-5-11 mm

Lemma pubescent or — pubescent (rarely glabrous pubescent glabrous
indumentum glabrous glabrous)

Shape of lemma __ indeterminate indeterminate usually smoothly often intermediate usually broadly
margin curved angled

Lower rachilla- 0-8-1-4 mm 0-8-1-5 mm 0-9-1-6 mm 1-1-7 mm 1-1-1-8 mm
segment
length

Lowest floret 1-2-6 mm 2-2—3-8 mm 1-5—3-5 mm 1-3—2:5 mm 1-3—2:5 mm
Undehisced
anther
length

RESULTS

The morphological data are summarised in Table 1. Lemma length is only of limited value to
separate these taxa but smaller lemmas do indicate B. lepidus. The lemmas of B. hordeaceus,
‘longipedicellatus’, B. x pseudothominei and B. lepidus were of papery texture and those of
B. racemosus, B. commutatus, B. secalinus, and B. pseudosecalinus were leathery. This difference
in texture is of fundamental importance to distinguish these two groups of grasses. In dried
material the veins of thin and papery lemmas tend to protrude but they do not in leathery lemmas.

Lemmas of subsp. hordeaceus are usually pubescent but in this study 6-1% were glabrous;
lemmas of ‘longipedicellatus’ were also pubescent but only 0-4% were glabrous. Lemmas of
B. commutatus were glabrous but those of B. commutatus var. pubens were pubescent and this
grass, which is more frequent than B. commutatus s. s., can be confused with ‘longipedicellatus’.
Pubescence was not found on the Iemmas of B racemosus and this was a good character for
distinguishing it. Glabrous lemmas in all the taxa were often scabrid. The margin of the lemma in
B. racemosus was smoothly curved and that of B. commutatus broadly angled; this was a
confirmatory character but often the difference was not clear. This character was of no diagnostic
value in B. commutatus var. pubens or in the B. hordeaceus group but a sharply angled margin was
indicative of B. lepidus.

The stiff, rather thick patent hairs on the lower leaf-sheath of young fresh B. commutatus were
very distinctive. In subsp. hordeaceus the indumentum was usually villous but in
‘longipedicellatus’, B. racemosus and in most specimens of B. commutatus var. pubens it was

BROMUS HORDEACEUS SUBSP. LONGIPEDICELLATUS 529

TABLE 2. DIAGNOSTIC CHARACTERS OF BROMUS HORDEACEUS SUBSP.
HORDEACEUS AND SUBSP. LONGIPEDICELLATUS

Taxon Source Culm Panicle Panicle structure Anther length
height length

Subsp. /ongipedicellatus L. M. Spaiton 80-175 10-20 at least 4 pedicels and 2-2—3-8 mm #

cm cm branches exceeding the
length of their spikelets
Subsp. hordeaceus L. M. Spalton up to 80 1-10 up to 3 pedicels and 1-2-6 mm
cm cm branches exceeding the
length of their spikelets
Clapham et al. 3-80 (3-)5-10 pedicels mostly shorter usually <1 mm
(1981) cm cm than spikelets
Hubbard (1968) 10-100 ~=—-1.5-16* pedicels 2-10 mm 0-2-2 mm
cm cm
Kerguélen (1995) 1-100 5-10 pedicels and branches 0-2—2 mm
cm cm < 2-5 cm
Smith (1980) 3-80 (3-)5-10 pedicels mostly shorter 0-2—2 mm,
cm cm than spikelets
Stace (1997) up to80 uptol0 — usually with all pedicels 0-5—1-5(—2) mm

cm (—16*) cm shorter than spikelets

* may include subsp. longipedicellatus. # Undehisced anthers from lowest florets only.

intermediate. The lengths of the lower rhachilla-segments were found to be unreliable for
distinguishing these taxa. These segments also elongated as fruit began to form.

In cultivated and in wild plants chasmogamy occurred in all taxa and was most frequent in
subsp. hordeaceus and ‘longipedicellatus’, but fertilisation was still mainly by cleistogamy.
Fertilisation and fruiting was most rapid in subsp. hordeaceus. Chasmogamous anthers were only
found in the lowest florets and were quickly extruded, dehisced and dropped. Within the other
florets, the anthers dehisced internally, cleistogamy occurred and these florets remained closed
until fruiting. In this study the anther measurements for subsp. hordeaceus differed from those
published elsewhere (Table 2). The anther lengths of B. commutatus (1-3—2:5 mm in Table 1) were
also different to the “1-1-5 mm’ in Stace (1997) and Clapham et al. (1987), ‘c. 1-5 mm’ in Smith
(1980) and ‘1-5-2 mm’ in Hubbard (1968). This may be because dehisced anthers had been
included.

The important characters distinguishing ‘longipedicellatus’ from subsp. hordeaceus are
summarised in Table 2.

In the cultivation experiments enrichment of ‘/ongipedicellatus’ and subsp. hordeaceus pots
with fertiliser did not increase culm height, nor the lengths of panicles, pedicels or branches. It is
likely that had these grasses been grown in nutrient-poor soil a different result would have been
obtained. However, the length of the spikelets of ‘longipedicellatus’ increased by 26% and the
enriched spikelets contained more florets The length of the lemmas only increased by 7%.
Enrichment of subsp. hordeaceus pots produced somewhat similar results but the sample was too
small to justify quantification. Smith (1972) had also found that spikelet size is susceptible to soil
nutrient status and that more florets are produced in rich soil. Stace (1997), Clapham et al. (1987)
and Smith (1980) did not use spikelet length in their keys to identify B. hordeaceus, but spikelet
length is employed to separate B. racemosus from B. commutatus (Table 1). Smith (1972) had also
found that lemma size was not much affected by richness of the substrate.

The cultivation experiments showed that ‘longipedicellatus’ was fertile and that it came true
from seed. At the main Devon station in May and June 2000 additional plants of
‘longipedicellatus’ were collected from the six marked sites where the seeds for cultivation had
been obtained in 1999. The cultivated plants of ‘longipedicellatus’ were very carefully compared
with these newly collected wild plants and with the herbarium specimens collected from these sites
in 1999. No significant differences were found (Table 3).

530 L. M. SPALTON

TABLE 3. COMPARISON OF WILD PARENT AND CULTIVATED PROGENY OF BROMUS
HORDEACEUS SUBSP. LONGIPEDICELLATUS

Locality Culm Panicle No. No. No. Spikelet Lemma Undehisced

height, length, pedicels > branches pedicels length, length, anther
cm mm their > their and mm mm _ length, mm
spikelets spikelets branches

Tiverton, wild (n=1) 15 14-1 8-0 3-0 11-0 19-0 9-1 3-4
Tiverton, cultivated (n=13) 141 14-7 7-1 2:3 9-4 21-0 9.2 3-6
Exeter, wild (n=1) 110 16-0 5-0 2-0 7-0 18-0 9-8 2:8
Exeter, cultivated (n=12) PF 15-8 6-2 1-1 73 19-3 10-2 2-9
Kennford, wild (n=1) 140 16-3 5-0 2-0 7-0 18-0 9-5 3-2
Kennford, cultivated (n=11) 147 15-9 4.4 3-1 7:5 19-9 9.2 2-9
Dartington, wild (n=1) 129 13-5 6-0 - 6-0 18-0 9-0 3-1
Dartington, cultivated (n=6) | 13-1 Dei - 5-7 17-6 9-3 2:9
Bicton, wild (n=1) 135 18-0 6-0 2-0 8-0 20-0 10-5 3-5
Bicton, cultivated (n=10) 122 16-4 4-] 2:3 6-4 19-1 11-0 3-3
Hams Barton, wild (n=1) 172 15-0 3-0 4-0 7-0 14-1 8-8 -
Hams Barton, cultivated (n=11) 161 14-3 3-8 4-] 7-9 14-3 9-0 3-5

Figures presented are the measurements from the single wild parents and mean of the cultivated progeny.
Parent plants were collected as follows: Tiverton, road lay-by $X996143, 5 May 1999. Exeter, bank above
road SX962916, 9 May 1999. Kennford, road verge, SX918875, 15 May 1999. Dartington, disturbed ground,
SX787621, 21 May 1999. Bicton, disturbed ground SX072868, 22 May 1999. Hams Barton, edge of wheat
field, SX882801, 26 May 1999.

DISCUSSION

The ‘longipedicellatus’ taxon was thus found to differ from the other taxa investigated. It is most
closely related to B. hordeaceus subsp. hordeaceus, but differs in its long culms, long panicles
with long pedicels and/or long branches and long chasmogamous undehisced anthers. It
superficially resembles B. commutatus var. pubens in being 80-175 cm tall with panicles 10-20
cm long but its lemmas are papery and like B. racemosus and B. arvensis it has long
chasmogamous anthers. As the grass is clearly a distinct taxon it is it is named Bromus hordeaceus
subsp. longipedicellatus.

Bromus hordeaceus L. subsp. longipedicellatus L. M. Spalton, subsp. nov.

Varietas robusta. Caules 80-175 cm. Paniculae 10-20 cm, quattuor minimum pedicellis aut ramis
longitudinem spicularum suarum excedentibus (ramis ad basem spiculae terminalis mensis).
Lemmae 7-5—10-5 mm, pubescentes, texturae papyraceae, venis in siccitate extantibus. Antherae
adhuc intactae infimorum florum 2-2—3-8 mm, ceterae breviores.

A robust grass with culms 80-175 cm long. Panicles 10-20 cm long with at least 4 pedicels or
branches exceeding the length of their spikelets (branches measured to the base of the terminal
spikelet). Lemmas 7-5—-10-5 mm long, of papery texture, with protruding veins when dried,
pubescent or rarely glabrous. Undehisced anthers from the lowest florets 2-2-3-8 mm long, other
anthers smaller. j

HOLOTYPUS: South Devon, v.c. 3, Aveton Gifford (SX691474), grassy bank on thin soil above
road, 23 May 1999, L. M. Spalton s.n. (BM).

Subsp. longipedicellatus is found on road verges and banks, on waysides, on the edges of arable
fields (especially wheat), on waste ground and less frequently in pasture. On road verges it is in
flower from the beginning of May and is very soon cut during roadside trimming. When in fruit
(and when dried) the spikelets lie close to the culm so the panicle no longer appears to be lax.

BROMUS HORDEACEUS SUBSP. LONGIPEDICELLATUS 531

Subsp. longipedicellatus has been confirmed in 46 vice-counties of England, eight of Wales, six
of Scotland and one of Ireland. Out of 1789 herbarium specimens that were examined only 28
were subsp. longipedicellatus, the earliest of which was collected by G. H. Douglas in Torbay,
Devon in 1926 (TOR). This suggests that, whatever the origin of subsp. longipedicellatus, it may
have spread rapidly in England and Wales. It is probable that the grass may have been
disseminated as a contaminant of Lolium perenne seed in agricultural and amenity grass seed
mixtures or of wheat seed.

Subsp. longipedicellatus might have resulted from a gene transfer from B. hordeaceus into
B. racemosus or B. arvensis with hordeaceus genes becoming dominant. B. commutatus var.
pubens has pubescent, less leathery (though still papery) lemmas, a variable indumentum of the
lower leaf-sheath and an earlier flowering time than B. commutatus and it might contain genes
from hordeaceus. On the rare occasions when B. racemosus and B. commutatus share the same
habitat, intermediates are produced. There are many problems in the genus and DNA or isoenzyme
studies might resolve at least some of them. A new key to the genus Bromus in Britain will be
produced when additional research has been completed.

An important consequence of the recognition of subsp. Jongipedicellatus is that the descriptions
of B. hordeaceus and subsp. hordeaceus now require revision because B. hordeaceus cannot be
reliably separated from B. racemosus and B. commutatus by panicles that are dense or lax. The
following descriptions are proposed:

Bromus hordeaceus L.: Culms usually erect 2-175 cm. Panicle dense or lax, 1-20 cm, sometimes
only 1 spikelet. Lemmas 6-5—10-5 mm, papery with protruding veins when dried, pubescent or
glabrous. Undehisced anthers from the lowest florets 1-3-8 mm.

Bromus hordeaceus subsp. hordeaceus: culms erect or ascending to 80 cm. Panicle 1-10 cm,
dense or somewhat lax with not more than 3 pedicels or branches exceeding the length of their
spikelets. Lemmas 7—10-5 mm, papery with protruding veins when dried, pubescent or glabrous.
Undehisced anthers from the lowest florets 1-2-6 mm.

ACKNOWLEDGMENTS

I thank C. J. Bruxner, S. P. Chambers, A. Copping, R. J. Hutchinson and L. J. Margetts for their
collaboration in cultivation experiments in their localities, the Principal of Bicton College of
Agriculture for providing me with experimental cultivation facilities, the grass specialists for
comments on the specimens sent to them, the Keepers of herbaria for the generous loan of
specimens and W. H. Tucker for the Latin diagnosis. Special thanks are due to L. J. Margetts for
first detecting this grass and for constructive comments on an earlier draft, to T. Rich for
additional comments and to the many botanists who have sent Bromus specimens to me. I hope
that they will continue to do so.

REFERENCES

CLAPHAM, A. R.,TUTIN, T. G. & WARBURG, E. F. (1962). Flora of the British Isles, 2nd ed. Cambridge
University Press, Cambridge.

CLAPHAM, A. R.,TUTIN, T. G. & Moore, D. M. (1987). Flora of the British Isles, 3rd ed. Cambridge
University Press, Cambridge.

HUBBARD, C. E. (1968). Grasses, 2nd ed. Penguin Books, London.

KERGUELEN, M. (1995). In PORTAL, R. Bromus de France. R. Portal, 43750, Vals-pres-Le Puy, France.

SMITH, P. M. (1972). Serology and species relationships in annual bromes. Annals of botany 36: 1-30.

SMITH, P. M. (1973). Observations on some critical Brome grasses. Watsonia 9: 319-332.

SMITH, P. M. (1980). Bromus L. in TUTIN, T. G. et al. eds. Flora Europeae 5. Cambridge University Press,
Cambridge.

SMITH, P. M. (1981). Ecotypes and subspecies in annual brome-grasses. Botanische Jahrbucher fur Systematik
102: 497-509.

SMITH, P. M. & SALES, F. (1993). Bromus L. Sect. Bromus: Taxonomy and relationships of some species with
small spikelets. Edinburgh journal of botany 50: 149-171.

STACE, C. A. (1997). New Flora of the British Isles, 2nd ed. Cambridge University Press, Cambridge.

(Accepted February 2001)

-t-

th

Watsonia 23: 533-538 (2001) 553

Festuca longifolia Thuill. (Poaceae) in Devon
R. E. N. SMITH
Institute of Grassland & Environmental Research, North Wyke, Okehampton, Devon EX20 2SP*
and
L. J. MARGETTS

16 Ashleigh Road, Honiton, Devon EX14 ITD

ABSTRACT

Festuca longifolia Thuill., Blue Fescue, was first recorded in Devon on limestone at Berry Head in 1936.
More recently it has been found on sea cliffs in South Devon between Start Point and Bolt Tail on both mica-
schist and hornblende-chlorite schist rocks and another site on limestone near Torquay. F. longifolia is not an
obligate calcifuge but occurs on dry soils in the absence of competition. It may be more widely distributed in
the south-west peninsula of England.

KEYWORDS: fescue, distribution, habitat, geology, soil type.

INTRODUCTION

There is some confusion over the name Festuca longifolia Thuill., which, especially in post-1945
Floras such as the popular Concise British Flora in Colour (Martin 1965), is used to describe an
introduced species, the Hard Fescue. However, recent taxonomic research (Wilkinson & Stace
1989), has shown that the correct name for Hard Fescue is F. brevipila Tracey. Wilkinson & Stace
(1991) have further shown that F. longifolia is not conspecific with the garden Blue Fescue, to
which the name F. glauca Vill. correctly belongs.

The British Red Data Book (Wigginton 1999) shows the distribution of Festuca longifolia
Thuill. in Great Britain and the Isle of Man to be restricted to a few sites in eastern England where
it occurs on acid soils in heathland communities in Suffolk, Lincolnshire and Nottinghamshire. In
fact, this species was recorded in Devon as long ago as 1936 when it was collected by G. T. Fraser
at Berry Head and determined by W. O. Howarth, the acknowledged Festuca expert at that time.
Two years later, Fraser returned to Berry Head to show the plants to three leading botanists - G. M.
Ash, J. F. G. Chapple and J. P. M. Brenan. These records were included in the Flora of Devon
(Martin & Fraser 1939) and vouchers deposited in TOR, RAMM and Herb. Druce (OXF).

In 1992 R. Arigho, a local botanist with a particular interest in grasses, collected a small fescue
from a rock outcrop near Prawle Point, the southernmost point of Devon. This was sent to T. A.
Cope who confirmed that it was F. huonii Aug. This new record for the mainland of Great Britain
was published locally (Margetts 1993) and nationally (Plant Records 1995). The following year, R.
Arigho and L.J.M. visited the South Hams coast in search for further colonies of F. huonii. Near
Gara Rock they found a small bluish fescue on rocks by the coastal footpath. They each collected
rooted offsets to grow on in pots. When these flowered, it was realised they were not F. huonii, but
almost certainly F. longifolia. The specimens were sent to T. A. Cope and P. J. O. Trist, both of
whom confirmed this identification. This record was also published (Margetts 1994; Plant Records
1995). Since then, further exploration of this part of the south Devon coast has led to the discovery
of F. longifolia at a number of sites between Start Point and Bolt Tail, and at a new station on the

“Address for correspondence: Email: roger.smith @bbsrc.ac.uk

534 R. E. N. SMITH AND L. J. MARGETTS

\ :
2 a i a

acre

Prawle Point

FIGURE 1. Distribution, by 1 km square, of Festuca longifolia (Thuill.) in Devon. The open circle represents
the 1936 record from Berry Head, the closed circles the 1993 and subsequent records.

Torbay limestone (Fig. 1; Appendix 1). It still exists on Berry Head (R. Arigho, pers. comm.,
1995). Festuca huonii has not been relocated or found elsewhere. It is possible the original
specimen may have been young F. longifolia.

GEOLOGY AND SOILS

Plants growing in eastern England and the Channel Islands have all been described as growing on
acid soils (Wilkinson & Stace 1991). In France it has generally been regarded as a calcifuge
(Portal 1999). However, the two populations found in the vicinity of Torquay occur on limestone
headlands. In addition, the majority of records come from the area between Start Point and Bolt
Tail where the rocks fall into two main groups, the mica-schists and hornblende-chlorite schists,
which were formed by the metamorphism of mafic lavas or sills (Durrance & Laming 1982). A
pulverised sample of the hornblende schist from near Prawle Point had a pH of 8-2. Consequently,
it was thought desirable to collect soil samples from below plants and to determine pH in the
laboratory. Testing was completed on fresh soil within 24 hours of collecting the samples from the
mica-schist and hornblende-chlorite schist sites and 48 hours for the limestone site (Table 1).
Standard laboratory procedures were followed.

HABITAT

On the south Devon coast F. longifolia occurs in sunny exposed situations, close to the edge of the
sea cliffs, mostly on flat or gently sloping surfaces of large outcrops of both rock types, usually
where there is some soil. It can also be found occasionally in vertical crevices. Soils on both the
mica and hornblende are immature, rich in raw humus but prone to dry out. There is little, if any,
root penetration of the rock surface. Where plants are established in very thin soil or growing in
fissured rock, they will be small and scattered but may be the dominant species in an open
community. Where the soil is a little deeper F. longifolia forms semi-continuous cover, with the
associated species growing amongst it. In these circumstances plants can be moderately robust.
and form tussocks 15 cm or more in diameter. However, in general, where the soil depth exceeds
about 10 cm F. longifolia is excluded by more competitive species such as Festuca rubra. In these
situations the occurrence of F. longifolia appears to be limited only by the size of the outcrop.
Between Start Point and Bolt Tail there are many such sites. The total population almost certainly
consists of thousands of individual plants.

FESTUCA LONGIFOLIA IN DEVON 535

TABLE 1. THE PH OF SOIL TAKEN FROM BELOW ESTABLISHED FESTUCA
LONGIFOLIA (THUILL.) PLANTS IN DEVON AT THE END OF APRIL 2000

Rock type Locality pH
Mica schist Bolberry Down 4.7
Cathole Cliff all
Hugh’s Hole 5-0
Whitechurch 5-3
Hornblende-chlorite schist Elanders Cove 6-1
Maceley’s Cove 5-9
Gammon Head 5:4
Hamstone Cove 5:3
Hamstone Cove 7),
Limestone Fishcombe Point 7:3
Fishcombe Point 7-6
Fishcombe Point 7:7
Fishcombe Point 8-5

In addition to the cliff edge populations reviewed here, F. /ongifolia has also been found a short
distance inland (R. Arigho, pers. comm., 1995) where rocky outcrops of both the mica and
hornblende schists occur a field or so behind the shore. It is known to occur here in somewhat
similar circumstances but the more maritime associates are, presumably, absent.

In the Torbay area F. longifolia grows at Fishcombe Point on sheltered south facing slopes in
thin soil amongst small limestone outcrops where small groups of plants occur in open
communities. The soils here are less organic and very freely draining. It probably occurs in similar
situations on Berry Head.

In Devon, 47 species were found growing in association with F. longifolia (Appendix 2). Only
Dactylis glomerata was more-or-less constant, as a dwarfed form, whilst Sedum anglicum was
found on all three rock types. Armeria maritima was constant on the schists but absent on the
limestone. The other associated species were either annuals, bulbs or dwarf forms of, presumably,
drought-tolerant species.

DISCUSSION

It is, perhaps, not very surprising that F. /ongifolia has been found on the coast of south Devon. In
the Channel Islands, it once occurred in Jersey and may still be found on Guernsey, Sark and Herm
where it occurs on coastal cliffs in shallow organic soils over granite or in cracks and crevices in
the rock (Trist 1996). Populations here are small, no more than 5-10 individuals at irregular
intervals, where the few constant associates include Armeria maritima, Dactylis glomerata, Sedum
anglicum, Trifolium campestre and Vulpia bromoides.

F. longifolia is also known from the western coast of Normandy, France (Portal 1999) where it
occurs between Avranches and Flamanville. These populations, described by Huon (1970) using
the name Festuca glauca Lam., occur on coastal schistose cliffs, fixed dunes and, in one locality,
on carboniferous limestone a short distance inland. The pH of these three types of substrate was
5-4, 8-0 and 7-6 respectively. Huon came to the conclusion, that in Normandy, F. longifolia was
indifferent to the nature of the substrate and simply occupied xerophyllous habitats.

For many years, the Berry Head record has been ignored and the plant assumed to be restricted
to acid soils. However, this record, together with that from Fishcombe Point, both on limestone,
indicate that F. longifolia is not an obligate calcifuge but occurs on soils with a very wide pH
range where taller growing species are excluded by environmental stress. This view is supported
by the evidence from Normandy. Given this tolerance of substrate, there is a great deal of
apparently suitable habitat on the coast of the south-west peninsula where this species could occur
and it seems likely that it may be more widely distributed.

536 R. E. N. SMITH AND L. J. MARGETTS

ACKNOWLEDGMENTS

We wish to thank R. Arigho for stimulating our interest in this species, Derek Wells for making
the manuscript and notes of the late P. J. O. Trist available to us, and Julia Tallowin for measuring
the pH of the soil samples. The distribution map was prepared using DMAP software produced by
Dr. Alan Morton.

REFERENCES

DURRANCE, E. M. & LAMING, D. J. C. (1982). The geology of Devon. University of Exeter.

Huon, A. (1970). Les Fetuques de l’ouest de la France - recherches de biosystematique et de biogeographie.
Thése Rennes.

MARGETTS, L. J. (1993). 85th report on botany, vascular plants. Report and transactions of the Devonshire
Association for the advancement of science, literature and the arts 125: 227-235.

MARGETTS, L. J. (1994). 86th report on botany, vascular plants. Report and transactions of the Devonshire
Association for the advancement of science, literature and the arts 126: 251-257.

MARTIN, W. K. (1965). The concise British Flora in colour. Ebury Press & Michael Joseph, London.

MARTIN, W. K. & FRASER, G. T. (1939). Flora of Devon, T. Buncle & Co., Arbroath.

PLANT RECORDS (1995). Watsonia 20: 419-437.

PORTAL, R. (1999). Festuca de France, R. Portal, Vals-prés-le Puy.

STACE, C. A. (1997). New Flora of the British Isles. 2nd ed. Cambridge University Press, Cambridge.

TRIST, P. J. O. (1973). Festuca glauca Lam. and its variety caesia (Sm.) Richt. Watsonia 9: 257-262.

TRIST, P. J. O. (1996). Unpublished manuscript on Festuca longifolia Thuill.

WIGGINTON, M. J., ed. (1999). British Red Data Books, 1 Vascular Plants, 3rd ed. Joint Nature Conservation
Committee, Peterborough.

WILKINSON, M. J. & STACE, C. A. (1989). The taxonomic relationship and typification of Festuca brevipila
Tracy and F. lemanii Bastard (Poaceae) Watsonia 17: 289-299.

WILKINSON, M. J. & STACE, C. A. (1991). A new taxonomic treatment of the Festuca ovina L. aggregate
(Poaceae) in the British Isles. Botanical journal of the Linnaean Society 106: 347-397.

(Accepted February 2001)

FESTUCA LONGIFOLIA IN DEVON 537
APPENDIX 1 THE DISTRIBUTION OF FESTUCA LONGIFOLIA THUILL. IN DEVON
10 km square Location Grid Year Recorder Det.
SX63 Bolberry Down SX687383 1994 LJM & LMS ci
Cathole Cliff SX693379 2000 RENS & CJS 2
Hugh’s Hole SX679388 2000 RENS & CJS £
nr Bolt Tail SX672393 2000 RENS & CJS ay
SX73 nr Sharp Tor, Salcombe SX729368 1994 LJM & LMS
nr Gara Rock SX742371 1993 LIM & RA ss
on Gara Rock SX751370 1994 LJM & LMS
Portlemouth Down SX747369 1994 LJM & LMS
Pigs Nose Valley SX762363 1994 DAB <
Gammon Head SX764357 1994 DAB 7
SX83 above Lannacombe Beach SX802372 1995 RA
Start Point SX826372 1993 RA
SX85 Fishcombe Point SX918570 1997 RENS & CJS £
SX95 Berry Head SX9456 1936 GTF Ist record |

GiE— G. T. Fraser

LJM = L. J. Margetts

LMS = L. M. Spalton

RENS é& CJS =R.E.N. & C. J. Smith

RA =R. Arigho

DAB = field meeting, Botany Section, Devonshire Association

o)
=}

feel:
Oni .J -O. Trist
onf. L. J. Margetts
et W. O. Howarth

Hh MW
Ho @ @

MH —-+ xX

In addition, a further seven sites in SX73, between Prawle Point and Evator Cove and based on
non-flowering material, were marked as provisional by RA. These are unlikely to be any other
species since the only other blue-leaved fescue in the F. ovina group is F. glauca Vill., the garden
plant, which is inconceivable here.

R. E.N. SMITH AND L. J. MARGETTS

538

APPENDIX 2. PLANT SPECIES ASSOCIATED WITH FESTUCA LONGIFOLIA IN DEVON

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Watsonia 23: 539-541 (2001) 539

Fallopia x conollyana The Railway-yard Knotweed
J. P. BAILEY

Biology Department, University of Leicester, United Kingdom, LEI 7RH

ABSTRACT

The hybrid between Fallopia japonica and F. baldschuanica has been known to exist in Britain as seed since
1983, and as an established plant since 1986. Over the last 15 years there have been a number of reports of this
hybrid from continental Europe, both as open-pollinated seed on F. japonica plants and as established plants.
In the light of the wide occurrence of the hybrid and the absence of a hybrid binomial, it is accordingly named
Fallopia x conollyana. The hybrid and its current distribution are described and its significance and potential
for further distribution are discussed briefly.

KEYWORDS: Fallopia japonica, Fallopia baldschuanica, wide-hybridisation, invasive plants, Japanese
Knotweed, Russian Vine.

INTRODUCTION

The hybrid between F. japonica (Houtt.) Ronse Decraene (Japanese Knotweed) and F.
baldschuanica (Regel) Holub (Russian Vine) was first discovered in Wales in 1983, as open
pollinated seed from F. japonica, and subsequently confirmed by artificial hybridisation the same
year (Bailey & Conolly 1984). Despite the almost ubiquitous occurrence of seed pollinated by F.
baldschuanica on F. japonica plants, it was not until 1987 (Bailey 1988) that a plant of some
considerable size and age was discovered by David Bevan in Haringey, Middlesex v.c. 21 (Bailey
1992). Since that date there have not been any confirmed sightings of further hybrids of this
constitution as established plants in the British Isles. However, increasing reports of this hybrid in
continental Europe convince me that this taxon should be given a hybrid binomial name.
Accordingly I am naming it Fallopia x conollyana in recognition of the decades of research into
alien Polygonaceae carried out by my close colleague Ann Conolly at Leicester.

Fallopia x conollyana J.P. Bailey, hybrida nova

Hybrida sterilis inter Fallopia japonica (Houtt.) Ronse Decr. et F. baldschuanica (Regel) Holub.
A Fallopia japonica caulibus basi lignosis minus quam 1 cm diametris tepalis per anthesin
validius carinatis differt. A F. baldschuanica planta rhizomatosa, caulibus cavis, foliis basi
subcordatis vel truncatis, stigmatibus fimbriatis differt.

HOLOTYPUS: Overgrown railway marshalling yard at Haringey (Railway Fields), Middlesex, v.c.
21, map reference TQ317882. 2 October 1987 J. P. Bailey & D. Bevan. (LTR)

Herbaceous perennial, younger plants weakly rhizomatous, but stout woody rhizomes in long-
established plants; stems erect but bowing over, hollow, up to 2 m long, slender with red blotches,
less than lcm diameter, becoming woody at the base; leaves glabrous with slender petioles 1-5—0-5
cm; lamina up to 13 x 6-5 cm, triangular ovate, acuminate to acuminate cuspidate at apex, sub-
cordate to truncate at base; inflorescence axillary and terminal panicles; flowers superficially
similar to, but larger (up to 3 mm diameter) and more conspicuous than those of F. japonica, the 3
keeled petals much more conspicuously winged than those of F. japonica; style trifid with 3 club
shaped stigmata with short fimbriae (resembling more closely the stigmas of the F. japonica
parent). Flowering occurs from September to early October. In spite of the woody lignified stem
with buds, the plant is herbaceous, overground growth dying back to ground level or near ground
level in winter, to be replaced by new canes in the spring. 2n=54

540 J; PALEY

Identification of this hybrid is problematic, particularly if only dried material is available.
Neither of its parents possesses distinctive epidermal trichomes and unfortunately the leaf shape
and overall growth-form are very similar to post-herbicide treatment regrowth of F. japonica. The
flowers are diagnostic, but even in well-established plants flowers are not often produced. It is
possible to distinguish young plants of the hybrid from weed-killed F. japonica, if the whole plant
complete with root system is collected, since young hybrid plants will not yet have developed
rhizomes. Chromosome counts are the most reliable means of identification, but are obviously of
limited utility.

F. x conollyana can be raised readily from seed, which germinates freely under greenhouse
conditions. However as the plants do not compete well with surrounding vegetation, and tend to
get blown down if not staked, flowering is very rare. Plants have occasionally flowered at
Leicester, but most of the information on flower structure comes from the plant at Haringey.
Meiotic behaviour is reported in Bailey & Stace (1992).

DISTRIBUTION

In 1994, Petr Horn informed me that he suspected that the plants that he had grown from seed
collected from open-pollinated F. japonica var. japonica plants in Ceske Budejovice (Czech
Republic) were hybrids with F. baldschuanica. Subsequent chromosome counts confirmed this as
the first occurrence of the hybrid outside Britain. Alberternst (1995) reported that seed collected
from a Japanese Knotweed plant growing near a plant of F. baldschuanica in Germany produced
plants as a result of pollination by F. baldschuanica. In 1995 I was sent a herbarium specimen of a
whole plant, including the root system, of F. x conollyana collected from an old railway yard in
Oldenberg, Germany by Bernd Hayen. He reported that he thought the plants were about six to
eight years old, certainly not much older since the railway yard had only been closed for 10 years
at that time. In 2000 Lajos Balogh sent me material from plants growing at the MTA Botanical
garden at Vacratét, Hungary. It is not known whether the plants originated spontaneously or were
deliberately planted, but they were quite well established. Fremstad & Elven (1997) reported the
hybrid growing in Stavanger, Norway.

TAXONOMY

Holub (1992) published the hybrid genus name x Reyllopia (Reynoutria x Fallopia) based on my
earlier report of the hybrid (Bailey 1988). Ironically, the existence of this hybrid was the final
piece of evidence needed to convince me that the genera Fallopia and Reynoutria should be
amalgamated; it follows that a hybrid generic name is no longer required.

DISCUSSION

F. x conollyana is a most unlikely hybrid for a number of reasons. Firstly, it is a cross between an
octoploid with a base number of 11 and a diploid with a base number of 10. Secondly its two
parents have contrasting growth-forms and over-wintering strategies, F. japonica is a herbaceous
perennial which stores reserves in underground stems or rhizomes, whilst F. baldschuanica is a
deciduous woody perennial that stores reserves in woody, overground stems. Hence F. x
conollyana may fall between two stools in terms of resource partitioning for over-wintering, and
this may contribute to its failure to establish more generally in the British Isles. Finally, the
parental species are not generally sympatric in their native distribution. Based on the somewhat
crude measure of herbarium specimen labels, there does appear to be some slight geographical
overlap of the two species in northern China.

Although vast amounts of seed are produced every year by F. japonica throughout Europe
bearing F. x conollyana embryos, only a minute proportion of them seem ever to have germinated
or become established in nature. I suspect that for this to occur, a long warm autumn (for
maximum seed ripening), followed by a cold winter (to stop the seed from rotting), followed by a
warm winter (for seedlings to survive the all-important first winter) are required - a rare but
admittedly not impossible combination!

FALLOPIA x CONOLLYANA 541
ACKNOWLEDGMENTS

I am very grateful to Clive Stace for preparing the Latin diagnosis and for reading through the
paper, and to Lajos Balogh and Bernd Hayen for sending me material of F. x conollyana.

REFERENCES

ALBERTERNST, B. (1995). Handbuch Wasser 2: Kontrolle des Japan-Knéterichs an Fliessgewdssern: II.
Untersuchungen zur Biologie und Okologie der neophytischen Knoteriche-Arten. Landesanstalt fiir
Umweltschutz Baden-Wiirttemberg.

BAILEY J. P. (1988). Putative Reynoutria japonica Houtt. x Fallopia baldschuanica (Regel) Holub hybrids
discovered in Britain. Watsonia 17: 163-164.

BAILEY, J. P. (1992). The Haringey Knotweed. Urban Nature Magazine 1 : 50-51.

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

BAILEY, J. P. & STACE, C. A. (1992). Chromosome number, morphology, pairing, and DNA values of species
and hybrids in the genus Fallopia (Polygonaceae). Plant Systematics and Evolution 180: 29-52.

FREMSTAD, E. & ELVEN, R. (1997). Fremmede planter 1 Norge. De store Fallopia-artene. Blyttia 55: 3-14.

HOLUvB, J. (1998). Reclassifications and new names in vascular plants 1. Preslia 70: 97-122.

(Accepted May 2001)

-

Watsonia 23: 543-551 (2001) 543

Notes

THE INTERSPECIFIC HYBRIDS OF THE RHIZOMATOUS COUCHES, ELYTRIGIA
DESV. (POACEAE)

Until about 1980 the rhizomatous couches were usually placed in the genus Agropyron Gaertn.
There are traditionally three species recognised in Britain:
A. repens (L.) P. Beauv.;
A. pycnanthum (Godr.) Godr. (A. pungens auct., non (Pers.) Roem. & Schult.);
A. junceum (L.) P. Beauv. (of which the British representative is ssp. boreoatlanticum Simonet
& Guin., sometimes recognised as a separate species, A. junceiforme (A. & D. Love) AD:
Love).

Following Melderis (1978) and Melderis & McClintock (1983) all these taxa were segregated into
the genus Elymus L. and the necessary new combinations were made:
E. repens (L.) Gould;
E. pycnanthus (Godr.) Melderis (FE. pungens auct., non (Pers.) Melderis) (later corrected to
E. athericus (Link) Kerguélen by Kerguélen (1983));
E. farctus (Viv.) Runemark ex Melderis ssp. boreoatlanticus (Simonet & Guin.) Melderis.

Plant breeders and other students of wheat and other Triticeae (e.g. Love 1984) have for long
further subdivided Elymus, particularly into the non-rhizomatous species (e.g. E. caninus (L.) L.)
and the rhizomatous species (e.g. the above three). This view is now the most usually accepted
one, and for this reason it was adopted by Stace (1991) and Kent (1992). The rhizomatous species
were separated into the genus Elytrigia Desv.:

E. repens (L.) Desv. ex Nevsk1;

E. atherica (Link) Kerguélen;

E. juncea (L.) Nevski ssp. boreoatlantica (Simonet & Guin.) Hyl.

Kerguélen (1987) actually cited the second species as E. atherica (Link) Kerguélen ex Carreras
Mart. The publication by Carreras Martinez (1986) quoted by Kerguélen was a thesis at the
University of Murcia, Spain. I have not seen a copy of this, and I have not traced one in this
country (not in BM or K), but I have been kindly informed by Valéry Malecot of Paris (who has
also not seen a copy) that his enquiries in Spain have shown that the thesis was not properly
published and was not available for sale or on request, so cannot have been a vehicle for the valid
publication of new names. It is quite likely that Kerguélen (deceased 1999) realised this, because
in his standard French checklist (Kerguélen 1993) he dropped the ““ex Carreras Mart.” part of the
author citation. Since Kerguélen (1987) quoted the full place of publication of the basionym, the
combination E. atherica (Link) Kerguélen can be dated from his 1987 publication. The same
applies to one other species (E. campestris (Godr. & Gren.) Kerguélen) and one subspecies (E.
elongata ssp. scirpea (C. Presl) Kerguélen), which Kerguélen (1987) had similarly ascribed to
Carreras Martinez. Kerguélen’s checklist is now available on-line at www.dijon.inra.fr/malherbo/
fdf/accueill.htm, providing a valuable resource (containing over 74000 names) which is being
maintained by Valéry Malecot. Strangely, although this was not made available until 1998 (fide
Malecot), it reinstates the Carreras Martinez ascriptions (possibly Kerguélen neglected to update
the electronic version of his work).

THE HYBRIDS

Hybrids between the three British species also occur in all three possible combinations. The
nomenclatural situation regarding these is best discussed separately.

Elytrigia juncea x E. repens

The epithet usually used for this hybrid is /axa, originally as Triticum laxum Fr., and the correct
combinations under the three genera are given below in the list of citations. Under Elytrigia
Kerguélen (1987) correctly gave the citation E. x laxa (Fr.) Kerguélen. There appear to be no

544 NOTES Watsonia 23 (2001)

nothosubspecific names to differentiate the hybrids involving the two subspecies of E. juncea. The
epithet /axa (type from Sweden) actually applies to our (the North Atlantic) subspecies
boreoatlantica, so a new nothosubspecific epithet would be needed for the hybrid involving the
Mediterranean subspecies juncea.

There is also another (later) epithet used for this species in the earlier British literature. Druce
(1907, p. 252) coined the name Agropyron x hackelii Druce; in that publication there is no
diagnosis, but the name was validated in Druce (1929b, p. 877) by reference to an even earlier note
(Druce 1905, p. 139-140), based on specimens labelled “A. junceum x repens” from Port Talbot
Docks, Glamorgan, collected by H. J. Riddelsdell in July 1904, and from Arbroath, Angus,
collected by W. A. Shoolbred in the same month. I choose the former as lectotype, represented by
a specimen in OXF, with the entry from the Report of the Distributor pasted on the sheet.
Riddelsdell commented: “I took the plant for this hybrid at first sight, and Mr Druce kindly
removed any doubts I had.” There follows a note to say that Hackel confirmed the determination. I
fully agree with this, despite the fact that in 1967 A. Melderis annotated the sheet “Agropyron
pungens x repens’. The lower spikelets are very remote and the leaves have many prickle-hairs on
the ribs on their upper surface, clearly showing the A. junceum influence. Despite the fact that
Melderis had identified this specimen (wrongly) as A. pungens x A. repens, Melderis &
McClintock (1983) listed A. x hackelii (wrongly) as a synonym of Elymus x obtusiusculus.
However, there is, strangely, another specimen in OXF collected from Port Talbot, Glamorgan,
again in July 1904, in this case by Druce. It is labelled on one of Druce’s own “Herbarium
Britannicum” labels: “Agropyron hackelti Druce Port Talbot Glamorgan July 1904 G. C. Druce”.
There is, however, not the slightest sign of any Elytrigia juncea characters in the specimen: the
spike bears closely overlapping spikelets and the leaves have almost smooth ribs on their upper
surface. The specimen was labelled “Agropyron pungens x repens” by A. Melderis in 1967, and I
fully agree: the anthers are indehiscent and there are a few small marginal cilia on the leaf-sheath
margins. Both hybrids were collected at Port Talbot in the same month, but the fact that the only
reference cited by Druce (1929b) that contained a diagnosis was the one relating to the Riddelsdell
plant fixes the name hackelii to the latter specimen, and places A. x hackelii within the synonymy
of E. x laxa. There are two other Druce specimens in OXF labelled A. x hackelii. One, from Sker,
Glamorgan (July 1904) is labelled A. pungens x A. repens by A. Melderis, and I agree with that.
The other, from Yarmouth, E. Norfolk (July 1904) is labelled A. pungens by A. Melderis; it has
dehisced anthers and again I agree with Melderis.

Elytrigia atherica x E. juncea

The relevant epithet is acuta, originally as Triticum acutum DC., and again the three correct
combinations are listed below. The type of Triticum acutum is from the French Mediterranean.
Kerguélen (1987) cited this taxon as E. x acuta (DC.) Kerguélen ex Carreras Mart, but this is
incorrect on two counts: the attribution to Carreras Mart. is erroneous as noted above; and the
combination E. x acuta had been made earlier by Tzvelev. In this case the nothosubspecies
involving E. juncea ssp. boreoatlantica has the epithet obtusiuscula, originally as Agropyron
obtusiusculum Lange, and as a subspecies under Elytrigia x acuta as nothossp. obtusiuscula
(Lange) Kerguélen.

Elytrigia atherica x E. repens

There has been extraordinary confusion regarding the name of this hybrid. The epithet used almost
universally over the past 25 years for this combination is oliveri, originally as Agropyron oliveri
Druce (Kerguélen 1975; Melderis 1975, 1980; Melderis & McClintock 1983; Stace 1991; Kent
1992). However, in his original diagnosis of “X Agropyron oliverii’, Druce (1912, p. 38) gave the
parentage of the plant (from Blakeney, E. Norfolk, August 1911) as A. pungens var. littorale x A.
junceum, and the diagnosis reiterated that parentage: “It differs from pungens by the more simple
[presumably meaning less congested] inflorescence and from junceum by the smaller spikelets and
less brittle stem”. But soon after this (Druce 1914, p. 514) he was using the name A. x oliveri for
the hybrid A. pungens x A. repens, and he remained of this opinion right up to the second edition
of his British Plant List (Druce 1919, p. 410; 1927, p. 143; 1928, p. 134; 1929a, p. 768; 1929b, p.
877) and presumably until his death in 1932. Kerguélen (1987), despite his earlier (Kerguélen
1975) view on the.parentage of A. x oliveri, stated “Melderis & McClintock (1983) rangent les
épithetes ‘laxa’ et ‘oliveri’ dans des taxons séparés, mais en leur donnant la méme formule hybride

NOTES Watsonia 23 (2001) 545
‘Elymus pycnanthus x E. repens’”’. This is not true! Melderis & McClintock clearly cited Elymus x
laxus as E. farctus x E. repens and Elymus x oliveri as E. pycnanthus x E. repens. Kerguélen
nevertheless cited Elytrigia x oliveri as a synonym of E. x laxa, under the parentage E. atherica x
E. repens. Hence Kerguélen presented a novel but erroneous interpretation of the parentage of E. x
laxa, while retaining Druce’s later interpretation of E. x oliveri. Kerguélen (1987) cited Druce’s
plant as Elytrigia x oliveri (Druce) Kerguélen ex Carreras Mart. In this case it is not possible to
correct the authority simply by dropping the “ex Carreras Mart.” part because Kerguélen cited the
name in synonymy and therefore it is invalid. It was most unusual for Kerguélen to make such an
error, and he corrected it in the electronic version of his Index Synonymique, where laxa is given as
the epithet for E. juncea x E. repens, and oliveri for E. atherica x E. repens. (The printed version
of his work (1993) did not cover these hybrids).

The true parentage of A. x oliveri can of course be cleared up only by examination of the type
specimen, and in Druce’s herbarium at OXF is a specimen which I deduce to be this. It is labelled
“Agropyron pungens x repens Blakeney Norfolk Aug. 1911 G.C. Druce Phyt.-Geog. Excurs.” on
one of Druce’s own “Herbarium Britannicum” labels. Also on the sheet is an undated label in the
handwriting of the eminent grass expert W. Hackel, to whom Druce sent much material for
naming: “Agropyron repens x junceum [signed] Hackel’. Thirdly, the sheet has attached an entry
cut from a Report of the Botanical Exchange Club and Society of the British Isles (the date and
exact origin of which I have been unable to trace, despite repeated searching through the printed
Reports, and which in any case might not be relevant to this specimen), reading: “X Agropyron
Hackelii Druce (= A. junceum x repens). This hybrid grass was found near Blakeney 28 in small
quantity, at Southport 59, but these much nearer A. junceum, and at North Bull, Co. Dublin. At
Blakeney a possible hybrid A. pungens x repens also occurred.” Taken together, these three labels
tell a confusing story. It seems that Druce collected his material from Blakeney in August 1911,
named it as A. pungens x A. repens, and so labelled it on his sheet. He also sent some (or the same)
material to Hackel, who named it A. repens x A. junceum, But why did Druce (1912) attribute a
third parentage (A. pungens x A. junceum) to his plant? He was clearly hedging his bets in the
undated Botanical Exchange Club Report quoted above, claiming that although the main material
at Blakeney was A. junceum x A. repens there was some A. pungens x A. repens present as well,
but there is no mention on that label of the original published parentage. Subsequently (1914
onwards) he used oliveri exclusively for A. pungens x A. repens. There seem to be no sheets at
OXF with the epithet ‘oliveri’ on the original label.

The type specimen of A. x oliveri is very clearly a hybrid of Elytrigia juncea. There can be little
doubt that, as Hackel stated, it is E. juncea x E. repens, and therefore a synonym of E. x laxa, as
was listed by Kerguélen (1987) albeit for the wrong parentage! Since it is merely a synonym I
have not bothered to validate the combination under Elytrigia here. As there seems to be no name
available for the hybrid E. atherica (pungens) x E. repens | have decided to name it E. x drucei,
based on Druce’s Port Talbot specimen, which he wrongly labelled Agropyron hackelii.

POSSIBLE FUTURE COMPLICATIONS

A further taxon in this group is Elytrigia repens ssp. arenosa (Spenn.) A. Léve, which was
carefully described by Trist (1995). Recently Scholz (1998) has pointed out that the epithet
arenosa does not apply to the Atlantic coastal plant, but to a Continental plant earlier typified by
Scholz (1993) on a specimen from Mainz, Germany. The epithet applicable to our coastal plant is
maritima Tzvelev. Moreover Scholz claimed that this taxon belongs better in E. campestris than in
E. repens, based mainly on the prominent, close veins on the leaf adaxial surface, and he made the
new combination E. campestris ssp. maritima (Tzvelev) H. Scholz. This effectively adds a new
British native species. Should it be found to form hybrids with the other three British species, new
hybrid names will be needed.

A further complication might loom. Many Triticeae specialists, e.g. Love (1984), further split
from Elytrigia the genus Thinopyrum A. Love for the species with a disarticulating rhachis, e.g. E.
juncea. as T. junceum (L.) A. Love. The evidence for doing this is not unequivocal, but, should it
become desirable, new combinations for the two hybrids concerned under a new nothogeneric
name for Elytrigia x Thinopyrum will be required.

546 NOTES Watsonia 23 (2001)

THE CORRECT HYBRID NAMES

Elytrigia x laxa (Fr.) Kerguélen, Lejeunia, n.s., 120: 88 (1987)

Basionym: Triticum x laxum Fr., Novit. Fl. Suec. Mant. 3: 13 (1842)

Agropyron x laxum (Fr.) Tutin in A.R. Clapham, Tutin & E.F. Warb., Fl. Brit. Isl. 1463 (1952)
Elymus x laxus (Fr.) Melderis & D.C. McClint., Watsonia 14: 394 (1983)

Agropyron x oliveri Druce, Bot. Exch. Club Soc. Brit. Isles 3: 38 (1912)

Elymus x oliveri (Druce) Melderis & D.C. McClint., Watsonia 14: 393 (1983)

Elytrigia x oliveri (Druce) Kerguélen, nom. inval., Lejeunia, n.s., 120: 88 (1987)

Agropyron x hackelii Druce, Bot. Soc. Exch. Club Brit. Isles 8: 877 (1929)

(E. juncea x E. repens)

Elytrigia x drucei Stace, hybr. nov.

Hybrida inter Elytrigiam repentem et E. athericam; ab E. repenti foliorum costis adaxialibus
approximatioribus et prominentioribus differt; ab E. atherica foliorum vaginis ciliis marginalibus
brevioribus et sparsioribus differt; antheris indehiscentibus, polline pro parte majore sterili.
Holotypus: Port Talbot, Glamorgan, v.c. 41, July 1904, G.C. Druce 28945 (OXF).

(E. atherica x E. repens)

Elytrigia x acuta (DC.) Tzvelev, Novosti Sist. Vyssh. Rast. 10: 32 (1973)
Basionym: Triticum x acutum DC., Cat. Pl. Horti Monsp. 153 (1813)
Agropyron x acutum (DC.) Roem. & Schult., Syst. Veg. 2: 751 (1817)

(E. atherica x E. juncea)

nothosubsp. obtusiuscula (Lange) Kerguélen, Lejeunia, n.s., 120: 86 (1987)
Basionym: Agropyron x obtusiusculum Lange, Haandb. Danske FI., ed. 2, 48 (1857)
Elytrigia x obtusiuscula (Lange) Hyl., Nordkdrlvaxt. Fl. 1: 369 (1953)

Elymus x obtusiusculus (Lange) Melderis & D.C. McClint., Watsonia 14: 394 (1982)
(E. atherica x E. juncea subsp. boreoatlantica)

ACKNOWLEDGMENTS

I am grateful to Dick Brummitt (K) for helping to clarify a point in the ICBN, to Valéry Malecot
(P) for investigating the thesis from Murcia and for alerting me to the web site for the Index
Synonymique, and to Stephen Harris (OXF) for arranging the loan of the Druce material.

REFERENCES

CARRERAS MARTINEZ, M. A. (1986). Estudio nomenclatural y corologico de las Gramineas del sureste de
Espana. Unpublished thesis, University of Murcia.

DRUCE, G. C. (1905). Report of the Distributor for 1904. Botanical Exchange Club and Society of the British

Isles 2: 108-140.

DRUCE, G. C. (1907). Report of the Distributor for 1906. Botanical Exchange Club and Society of the British

Isles 2: 206-252.

DRUCE, G. C. (1912). Plant Notes for 1911, etc. Botanical Exchange Club and Society of the British Isles 3:

7-6.

DRUCE, G. C. (1914). Report of the distributor for 1913. Botanical Exchange Club and Society of the British

Isles 3: 441-516. :

DRUCE, G. C. (1919). New county and other records [for 1918]. Botanical Society and Exchange Club of the

British Isles 5: 365-412.

DRUCE, G. C. (1927). New county and other records, 1926. Botanical Society and Exchange Club of the

British Isles 8: 102-144.

DRUCE, G. C. (1928). British Plant List, 2nd ed. T. Buncle & Co., Arbroath.

DRUCE, G. C. (1929a). New county and other records, 1928. Botanical Society and Exchange Club of the
British Isles 8: 722-769.

DRUCE, G. C. (1929b). Notes on the second edition of the “British Plant List’. Botanical Society and
Exchange Club of the British Isles 8: 867-877.

KENT, D. H. (1992). List of Vascular Plants of the British Isles. Botanical Society of the British Isles, London.

NOTES Watsonia 23 (2001) 547

KERGUELEN, M. (1975). Les Gramineae (Poaceae) de la flore francaise. Essai de mise au point taxonomique et
nomenclaturale. Lejeunia, n. s. 75.

KERGUELEN, M. (1983). Les Graminées de France au travers de “Flora Europaea” et de la “Flore” du C.N.R.S.
Lejeunia, n. s. 110.

KERGUELEN, M. (1987). Données taxonomiques, nomenclaturales et chorologiques pour une révision de la
flore de France. Lejeunia n. s. 120.

KERGUELEN, M. (1993). Index synonymique de la flore de France. Muséum Nationale d’ Histoire Naturelle,
Paris.

Love, A. (1984). Conspectus of the Triticeae. Feddes Repertorium 95: 425-521.

MELDERIS, A. (1975). Agropyron Gaertn., in STACE, C. A., ed. Hybridization and the flora of the British Isles,
pp. 568-570. Academic Press, London.

MELDERIS, A. (1978). Taxonomic notes on the tribe Triticeae (Gramineae), with special reference to the
genera Elymus L. sensu lato, and Agropyron Gaertner sensu lato. Botanical Journal of the Linnean
Society 76: 369-384.

MELDERIS, A. (1980). Elymus L., in TUTIN, T. G. et al., eds. Flora Europaea 5: 192-198. Cambridge
University Press, Cambridge.

MELDERIS, A. & MCCLINTOCK, D. (1983). The genera Elymus L. and Leymus Hochst. in Britain. Watsonia
14: 391-395.

SCHOLZ, H. (1993). Elytrigia arenosa (Gramineae) - ein mitteleuropdischer Relikt-Endemit. Botanische
Jahrbiicher fiir Systematik, Pflanzengeschichte und Pflanzengeographie 115: 351-366.

SCHOLZ, H. (1998). Taxonomy and nomenclature of “Elytrigia maritima” Tzvelev. Bulletin. Société pour
l’échange des plantes vasculaires de |’ Europe et du bassin Méditerranéen 27: 102-104.

STACE, C. A. (1991). New flora of the British Isles. Cambridge University Press, Cambridge.

TRIST, P. J. O. (1995). Elytrigia repens (L.) Desv. ex Nevski subsp. arenosa (Spenner) A. Love (Poaceae) in
north-western Europe. Watsonia 20: 385-390.

C. A. STACE
Department of Biology, University of Leicester, Leicester LEI 7RH

SALIX CAPREA L. x S. MYRSINIFOLIA SALISB. x S. PHYLICIFOLIA L.
IN PERTHSHIRE

During a programme involving field surveys and a general reassessment of Scottish montane
willow hybrids, a number of apparently new hybrid combinations have been discovered. One of
these which was collected on 7 August 1997 by the author near Milton Roro, Glen Lyon in mid-
Perths., v.c. 88, was suspected to be the triple hybrid, Salix caprea L. x S. myrsinifolia Salisb. x S.
phylicifolia L., due to its appearance and also because both S. caprea and S. myrsinifolia x S.
phylicifolia (S. x tetrapla Walker) were growing nearby. Further specimens were collected in July
2000 and sent to R. D. Meikle who confirmed the tentative determination. There appeared to be no
previous confirmed records for this hybrid in Britain according to Meikle (1975), although E. F.
Linton (1913) had earlier suggested that specimens collected by him near Clova, v.c. 90 (E.F.L.
reference numbers 36 and 78) might possibly be referred to this cross. The hybrid is not shown in
the British list (Kent 1992, 1997; Kent & Stace 2000). However, subsequent information has been
provided by R.D.M. which shows that he determined the same cross very recently from specimens
collected late in 2000 in Yorkshire, v.c. 64, by M. Wilcox (R.D.M. and M. Wilcox, pers. comm.).
As there were no certain records for this hybrid combination prior to these confirmations and the
fact that triple hybrids are always extremely difficult to determine, the alternative possibilities
were considered, by comparing the published descriptions in Linton (1913), and those of Meikle
(1975) with the specimens collected by the author in Perthshire, reference number D/T CR9/00,
although the latter lacked catkins, so a full comparison was not possible. There is very little doubt
that S. phylicifolia is one parent of the hybrid D/T CR9/00, therefore likely alternatives seemed to
be S. cinerea L. x S. phylicifolia (S. x laurina Sm.), or S. caprea x S. phylicifolia. Although Linton
(1913) suggested that some material determined as S. x /aurina might possibly refer to the triple
hybrid S. cinerea x S. myrsinifolia x S. phylicifolia (which is not only possible but very probable,
as S. phylicifolia may not occur in the pure state in Breadalbane) most if not all of its records have
been redetermined as S. x tetrapla Walker (R.D.M., pers. comm.), although it is possible that S. x
laurina could have been introduced in some areas. Similarly, it is also suggested that at least some
of the records, and accordingly the descriptions, of S. caprea x S. phylicifolia might refer to the

548 NOTES Watsonia 23 (2001)

triple hybrid S. caprea x S. myrsinifolia x S. phylicifolia, the same cross determined by R.D.M. for
the Perthshire hybrid, D/JT CR9/00. However, Linton (1913) suggested that S. caprea x S.
phylicifolia was extremely rare in Britain and had been much confused with the widely distributed
S. x laurina, so it cannot be assumed that the old records for the former refer to the triple cross
determined for the Perthshire hybrid D/T CR 9/00. Although a comparison of the descriptions in
Linton (1913) and by Meikle (1975) for hybrids involving S. phylicifolia which are recorded in
Perthshire shows that the descriptions which fit the hybrid D/T CR 9/00 best are those which were
published as S. caprea x S. phylicifolia, and therefore suggest that these do not refer to S. x laurina
or to the similar triple cross S. cinerea x S. myrsinifolia x S. phylicifolia, but for the reasons given
above most probably refer to S. caprea x S. myrsinifolia x S. phylicifolia, and therefore provide
additional evidence that the hybrid DJT CR 9/00 has been correctly determined as the latter.
A description of the main characters of the hybrid DJT CR 9/00 is as follows:

A bush approximately 2-5 m high. Mature twigs dark reddish-brown, glossy and becoming
glabrous; shoots moderately pubescent. Leaves coriaceous, dark, bright green and glossy on the
upper surface, and rather thinly pubescent with the hairs in the majority confined to the midrib and
near to the apex, or subglabrous, the lower surface pale green, glaucous and glabrous with
prominent nervation, the majority with the lamina obovate or broadly obovate to subrotund, the
largest typically 5 cm long, 3:5 cm wide, shortly mucronate to cuspidate at apex, the margins
narrowly recurved, shallowly serrate to remotely serrulate, the smaller leaves narrowly rounded,
the larger broadly rounded at base; petioles moderately pubescent. Foliage not turning black when
dried.

CONCLUSIONS

Based on the published evidence the description of the Perthshire hybrid DJT CR 9/00 best fits the
descriptions given for S. caprea x S. phylicifolia, other than a slight variance in the leaf
indumentum, which may not be significant. However, the records of S. phylicifolia in the area in
question have been shown to refer to S. myrsinifolia x S. phylicifolia, so the descriptions of S.
caprea x S. phylicifolia very probably refer to the triple hybrid S. caprea x S. myrsinifolia x S.
phylicifolia, and provide additional support for the determination of the Perthshire hybrid D/JT CR
9/00 as this cross and to confirm its presence in Britain. The recent record from Yorkshire gives
further confidence in its existence. It is, however, possible that this hybrid has been overlooked in
the past. In the absence of S. phylicifolia, at least some of the older records for S. caprea x S.
phylicifolia might have more correctly referred to the same triple cross. Ideally, it would have been
advisable to locate the herbarium material which represents the old records on which the
descriptions were based in order to confirm the status of the Perthshire record, however, in the
absence of a prolonged and perhaps unsuccessful search it is necessary to rely on the present
specimens and the original descriptions, and assume that they refer to the hybrid combinations as
stated. The Perthshire hybrid has now been propagated from cuttings so further information may
be obtained if catkins are eventually produced.

ACKNOWLEDGMENTS

I should like to thank Desmond Meikle for his confirmation of identity and other helpful
comments, David Mardon for local field information, Mr. M. Wilcox for additional details of the
Yorkshire record, and finally, the Science and Research Committee of the B.S.B.I. for their
support and a grant towards the costs of part of this project.

REFERENCES

KENT, D. H. (1992). List of vascular plants of the British Isles. Botanical Society of the British Isles, London.
KENT, D. H. (1997). List of vascular plants of the British Isles, Suppl. 1. Botanical Society of the British Isles,
London.

KENT, D. H. & STAGE, C. A. (2000). List of vascular plants of the British Isles, Suppl. 2. Botanical Society of
the British Isles, London.

LINTON, E. F. (1913). A monograph of the British Willows. Journal of Botany, London 51 (Suppl.): 1-92.

NOTES Watsonia 23 (2001) 549

MEIKLE, R. D. (1975). Salix L., in STACE, C. A., ed. Hybridization and the flora of the British Isles, pp. 304—
338. Academic Press, London.

D. J. TENNANT

Low Missise Farm, Laverton, Ripon, North Yorkshire, HG4 3SY

A NEW SPECIES OF RUBUS SECT. MICANTES (ROSACEAE) FORMERLY KNOWN
AS A VARIANT OF R. CRINIGER (E. F. LINTON) ROGERS

A bramble which is locally abundant in the Colchester area of Essex (v.c. 19) and along the Stour
valley on the Suffolk side (v.c.c. 25 and 26) and is scattered in North West Suffolk (v.c. 26), West
Norfolk (v.c. 28) and South Essex (v.c. 18) with one known station in East Kent (v.c. 15) has been
dismissed as “var. trifolius” of Rubus criniger in recent years. Previously to this, E. S. Edees had
considered it to be a form of Rubus conspersus W. C. R. Watson, though Watson himself had
appended the name Rubus egregius var. effeminatus Focke, to a sheet sent him in 1923 through the
Botanical Exchange Club by G. C. Brown, and collected from Stanway near Colchester. This sheet
is now in BM. (D. E. Allen, pers. comm.)

The present author has had ample opportunity to study the plant, with annual visits to Essex for
nearly twenty years, as well as its stations in the other counties mentioned, and has long held the
view that it is distinctly and constantly different from its supposed parent. During the summer of
2000, the plant was collected as widely as possible in Essex, West Suffolk and West Norfolk and
comparisons made with R. criniger, also from as many sites as possible, including some from
Herts. v.c. 20 and Cambs. v.c. 29, where the present plant has not been recorded, and it was
discovered that the differences between the two plants, even when they are growing in close
proximity, as they do at Waterhouse Plantation, Tottington, v.c. 28, were even greater than was at
first thought. Neither have any plants been found which could be regarded as intermediate between
the two. It would be true to say that the only similarities between the two are the overall greyish
green coloration, the hairiness of the primocanes and the structure and dimensions of the stem
prickles. It is therefore proposed to name the plant with three leaflets as:

Rubus trinovantium A. L. Bull sp. nov.

A R. crinigero his notis differt. Turiones glandibus brevistipitatis numerosis vel copiosis, in
partibus turionis ejusdem diversis quoad numerum variabilibus, vestiti. Aculei aliquot in turionum
paginis sicut in angulis interdum reperiantur. Folia ternata vel rarissime quatuor vel quinque
foliola ferentia; foliola terminalia elliptica vel obovatocuspidata, cuspide 1-5-2 cm longa praedita.
Paniculae laxae, relative pauciflorae, parte superiore subracemosa brevi truncata; pedicelli 2-4 cm
longi. Adsunt 2-4 rami inferiores ascendentes ad 15 cm longi. plerumque modo 3-7 floribus
instructi. Flores magni, 3-3-5 cm diametro, stellati; petala elliptica, 1-5-1-75 cm _ longa,
emarginata; antherae glabrae.

Rubus trinovantium differs from R. criniger in the following characters. The stems are clothed
with numerous to abundant short-stalked glands, variable in number on different parts of the same
stem. A few prickles may occasionally be found on the faces of the stems as well as on their
angles. The leaves are ternate or very rarely bear four or five leaflets; the terminal leaflets are
elliptic to obovate-cuspidate, with the cusp 1-5 to 2 cm long. The panicles are lax and relatively
few-flowered, with a short truncate subracemose upper part; the pedicels are 2-4 cm long. There
are 2-4 ascending lower branches up to 15 cm long and usually only 3—7 flowered. The flowers
are large, 3-3-5 cm in diameter and starry the petals are elliptic, 1-5—1-75 cm long and notched,
and the anthers are glabrous.

Rubus criniger belongs to the Series Vestiti, but bearing in mind that the armature of R.
trinovantium is somewhat variable, to the extent that shade plants occasionally have consistently
longer stalked glands reminiscent of the group Radulae as at Lexden Gathering Ground near
Colchester, and some prickles may be found which are not on the angles of the stem. it is felt that
the new species should be placed in the Series Micantes. It has been found in 16 hectads to date.

HOLOTYPE: Tiptree Heath, Essex. v.c. 19., TL883148 July 12th 2000. BM.
Isotypes are in Herb. A. Newton and Herb. A. L. Bull.

550 NOTES Watsonia 23 (2001)

Representative Exsiccatae from High Woods, Colchester, TL9926, July 12th 2000; Lexden
Gathering Ground, TL9725, July 12th 2000; Layer Breton Heath, TL9118 July 16th 1981; East
Donyland, Colchester, TM02 July 23rd 1972, all in Essex; Denstead Wood, East Kent, TROS July
19th 1990; Arger Fen, West Suffolk, TL9335 July 10th 2000; Assington Thicks, West Suffolk
TL9337, July 10th 2000; Letch Moor, Icklingham, West Suffolk, TL7971, Aug. 20th 1971 and
July 17th 2000; South Runcton, West Norfolk TF6407, July 23rd 1977 and Waterhouse Plantation,
Tottington, West Norfolk, TL9094, July 15th 2000 are all in Herb. A. L. Bull.

R. trinovantium is a bramble of acid sands and gravels which characterise the formerly extensive
heathland that stretched from Tiptree to both north and south of Colchester. The West Suffolk site
at Arger Fen is on the edge of the former Leaven Heath, whilst nearby Assington Thicks is ancient
woodland with some patches of acid soils. Letch Moor at Icklingham and Waterhouse Plantation,
Tottington, are both areas of overgrown wet acid heathland and South Runcton lies on the Norfolk
Greensand.
The name trinovantium derives from Trinovantes, the ancient British tribe whose territory
centred on the Colchester area at the time of the Roman invasion.
My thanks are due to Mr Philip Oswald for writing the Latin description, to Dr D. E. Allen for
tracing specimens in BM and to Mr A. Newton for advice on the name trinovantium.
A. L. BULL
“Hillcrest”, East Tuddenham, Dereham, Norfolk, NR20 3JJ

DATES OF PUBLICATION OF COUNTY FLORAS

It seems to have been generally overlooked that, for various reasons, county Floras are liable to
bear an erroneous date on the title page. This is a matter for concern not merely bibliographically,
for in some of these publications new taxa have been described or valid new combinations
perpetrated unwittingly. The increasing attention being given to establishing the history of rare
species at individual sites also makes precision in published dates of records a matter of
importance.

The usual reason for misdating would seem to have been the tendency for small printers, in the
days before computerised typesetting, to treat such major and often typographically complex jobs
as ‘fillers’. to be worked on whenever business was slack and to be laid aside for extended periods
when more urgent or more profitable commitments intervened. In a publication process so
leisurely and protracted, including the dispatch of bound copies maybe some considerable time
after the receipt of the final corrected proofs, it could easily happen that an obsolete scheduled date
was left unamended. A particularly glaring instance where this is presumed to have been the cause
is the first edition of F. Townsend’s Flora of Hampshire, including the Isle of Wight. Though
allegedly published in 1883, an addendum was inserted (just before the index) at a clearly very late
stage and the dates of some of the records in that show that it cannot have appeared before 1884 -
assuming that all the bound copies initially issued included it. There is also reason to suspect that
the second edition of that work came out in 1905 instead of, as stated, 1904 (Allen 1986). Two
more recent cases have been pointed out by Mitchell (2000). J. P. Brunker’s Flora of the county
Wicklow, though bearing the date *1950’, actually appeared in the year following, as mentioned at
the time by Praeger (1951) and since confirmed by the publisher’s records. J. Harron’s Flora of
Lough Neagh was similarly published a year later that the indicated one. Printing delays are not
invariably responsible for this phenomenon, though. The Flora of the Isle of Man (Allen 1986),
though ready for issue by the date on the title page, was held back by the publisher for two years in
order for its appearance to coincide with ‘Manx Heritage Year’.

Publication can also be earlier than the year stated. T. Whilde’s The natural history of
Connemara, in which vascular plants are included in a list in the appendix, appeared in reality in
1993, not ‘1994’ (Mitchell 2000). Copies of the section on the botany contributed by T. H. Cooper
to the second volume (1835) of T. W. Horsfield’s The history, antiquities and topography of the
county of Sussex were distributed by Cooper as a separate pre-print a year before the publication of
the book itself, as shown by one that has survived in the W. J. Hooker Letters in the archives of the
Royal Botanic Gardens, Kew, accompanied by a dated covering letter.

NOTES Watsonia 23 (2001) 551
REFERENCES

ALLEN, D. E. (1986)(°1984’). Flora of the Isle of Man. Manx Museum & National Trust, Douglas.
ALLEN, D. E. (1996). Some earlier workers on the Hampshire flora, in BREWIS, A., BOWMAN, P. & ROSE, F.,
eds., The flora of Hampshire. Harley Books, Colchester.
MITCHELL, M. E. (2000). The Irish floras: a checklist of non-serial publications. Glasra 4: 47-57.
PRAEGER, R. L. (1951). Review of J. P. Brunker, Flora of the county Wicklow. Irish Naturalist’s Journal 10:
192-195.
D. E. ALLEN
Lesney Cottage, Middle Road, Winchester, Hampshire, SO22 5EJ

Watsonia 23: 553-592 (2001) 553

Plant Records

Records for publication must be submitted to the appropriate Vice-county Recorder (see BSBI Year Book
2001), and not to the Editors. The records must normally be of species, hybrids or subspecies of native or
naturalised plants (listed in Kent (1992) or Stace (1997)), belonging to one or more of the following
categories: Ist or 2nd v.c. record; Ist or 2nd post-1930 v.c. record; only extant v.c. locality, or 2nd such
locality; a record of an extension of range by more than 100 km. Such records will also be accepted for the
major islands in v.cc. S, 102-104 and 110. Only Ist records can normally be accepted for Rubus, Hieracium
Taraxacum 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 for Taraxacum must normally be additional to
those in Dandelions of Great Britain and Ireland by A. A. Dudman & A. J. Richards (1997).

Records are arranged in the order given in the List of vascular plants of the British Isles and its supplements
by D. H. Kent (1992), from which the species’ numbers, taxonomy and nomenclature are taken. Taxa not in
that book follow New Flora of the British Isles Ed. 2 by C. A. Stace (1997). The Ordnance Survey national
grid reference follows the habitat and locality. With the exception of collectors’ initials, herbarium
abbreviations are those used in British and Irish herbaria by D. H. Kent & D. E. Allen (1984). Records are
field records if no other source is stated.

Records from the following vice-counties are included in the text below (superscript figures indicate
pmnbemniaac) ta 2 34. 576.8 9°, 10’, 11°, 127.13", 15°, 17 21, 23°; 25°, 26, 28", 28°, 33°,
Beeman), 39°, 40°41, 42”, 437, 44° 45°, 461%, 47°, 48°, 49°, 50, 51”, 53’, 54’, 58)°, 59°, 60), 61°,
fremG 687,69), 70,717, 77°, 78, 79, 80 , 81, 83°, 89, 91°, 93°, 94", 95, 97’, 99", 100’, 1017, 104°,
Mise, 109, 110’, H10°, H16', H21', H27°, H30", H31°, H35°.

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.

1/3.1. LYCOPODIUM CLAVATUM _ 28, W. Norfolk: One small plant on floor of extensive gravel
pit, formerly heath, Leziate RSPB Reserve, TF682195, S. Fisher, 1997, conf. G. Beckett. 1st post-
1930 record.

1/4.1. DIPHASIASTRUM ALPINUM 80, Roxburghs.: NE facing bank, old forestry quarry,
Ramsaygrain West Forest, Teviothead, NT332022, R.W.M. Corner, 1996. 2nd extant site.

1/4.2a. DIPHASIASTRUM COMPLANATUM subsp. ISSLERI *108, W. Sutherland: Stony ground,
Canisp, NC2216, A.G. Kenneth, 1985, conf. A.C. Jermy.

72/1.2. SELAGINELLA KRAUSSIANA *50, Denbs.: Naturalised in mown grass by stream,
Bodnant, SH799721, J.A. Green, 1997.

3/1.1. ISOETES LACUSTRIS 94, Banffs.: Sandy bay, Loch Avon, NJO25032, J. Edelsten, 1989.
Not seen at this site since 1867. Pool by Loch Avon, NJO18026, D. Law, 1989. Not seen at this site
since 1866. Both BM, det. A.C. Jermy. Ist and 2nd recent records.

3/1.1x2. ISOETES ECHINOSPORA x I. LACUSTRIS (I. x HICKEYI) *46, Cards.: NE corner of Llyn
Pendam, SN709839, A.O. Chater & A.C. Jermy, 1997, BM, conf. D. Britton. Ist British record.

4/1.3. EQUISETUM VARIEGATUM 64, Mid-W. Yorks.: Riverside marsh, Dunsop Bridge,
SD6551, P. Jepson, 1997. 2nd record.

4/1.4x8. EQUISETUM FLUVIATILE x E. PALUSTRE (E. x DYCEI) *46, Cards.: With both parents
on peaty mud of summer-dry leadmine reservoir, Bog Pond, 2km NW of Ponterwyd, SN732824,
A.O. Chater, 1994, NMW, conf. A.C. Jermy. Ist record for Wales.

4/1.5x8. EQUISETUM ARVENSE x E. PALUSTRE (E. x ROTHMALERI) *73, Kirkcudbrights.:
Shoulder high rushy area, Southness, Gillfoot side, NX977546, O.M. Stewart, 1994, E, det. C.N.
Page. Presumed Ist record. *80, Roxburghs.: Flushed bank, Crookedsike Head, Cheviot Burn,
Cocklawfoot, NT871183, C. Dixon, 1997, herb C.N.P., det. C.N. Page.

4/1.6x7. EQUISETUM PRATENSE x E. SYLVATICUM (E. x MILDEANUM) *89, E. Perth: Road
verge, N side of minor road 1km E of Loch of Lowes, NO064439, C.I. Pogson, 1996, det. C.N.
Page.

5/1.1. OPHIOGLOSSUM VULGATUM 81, Berwicks.: Parkland, Newton Don, NT711374, M.E.
Braithwaite, 1995. 2nd extant record.

554 PLANT RECORDS

5/2.1. BOTRYCHIUM LUNARIA 28, W. Norfolk: Dune slack, Wells-next-the-Sea, TF909454, E.
Banham, 1996. 2nd record since 1973, a new area.

7/1.1. CRYPTOGRAMMA CRISPA = *95, Moray: Only four plants on rocky outcrops on mountain,
N facing, Creag an Tarmachain, Grantown-on-Spey, NJ154312, I.P. Green, 1997.

8/1.2. HYMENOPHYLLUM WILSONII *89, E. Perth: North facing mica schist cliff, Kindrogan
Crag, NO049628, H.S. McHaffie, 1985.

9/1.1. PILULARIA GLOBULIFERA 47, Monts.: Three populations on lake-edge, shallow
(c.30cm) and deep (1-6m) water, Llyn Gwyddior, SH9307, R.A. Jones, E.J. Lomas & T. Teearu,
1997. Ist recent records. 51, Flints.: Mud, edge of lake, Llyn Helyg, SJ115775, W. Hay, 1993.
Ist record for 20 years.

10/2.1. TRICHOMANES SPECIOSUM (gametophyte) *44, Carms.: Abundant in dry rock cavity,
under overhang on NW facing slope in wood, by track along bottom of Allt Penrhyn wood, Forge-
Cych, SN254413, A.O. Chater, 1994.

11/1.1x2. POLYPODIUM VULGARE x P. INTERJECTUM (P. x MANTONIAE) —_*37, Worcs.: Mossy
bank, West Malvern, SO7645; North Quarry, Malvern, SO7646; both P.G. Garner, 1989, det. R.H.
Roberts. Ist and 2nd records. *58, Cheshire: Wooded bank, Barnston, Wirral, SJ279837, E.F.
Greenwood, 1995, LIV, det. R.H. Roberts. Mixed with P. vulgare.

15/2.5a. ASPLENIUM TRICHOMANES subsp. TRICHOMANES 50, Denbs.: Roadside wall, Capel
Garmon, SH8157, J.A. Green, 1996. 2nd record.

15/2.5c. ASPLENIUM TRICHOMANES subsp. PACHYRACHIS *42, Brecs.: Limestone rocks near
Pontneddfechan, SN9108, M. Porter, 1995, det. J.C. Vogel & A.C. Jermy. *58, Cheshire:
Sandstone cliffs, Maiden Castle, Bickerton, SJ4952, J.P. Guest. Sandstone cliffs, Bickerton Hill,
SJ5053, G.M. Kay; both 1997, det. J. Vogel. Ist and 2nd records.

15/2.5x8. ASPLENIUM TRICHOMANES x A. SEPTENTRIONALE (A. x ALTERNIFOLIUM), , 89, E.
Perth: Stenton, Dunkeld, NO04, J.H. Penson, 1966, herb. J.H.P. Volcanic rocks, Craig Tronach,
NO053402, J. Castle-Smith & N.W. Taylor, 1991, E. 1st and 2nd recent records.

15/2.6. ASPLENIUM VIRIDE *47, Monts.: In crevices on vertical rock face in rocky stream
gorge, Lake Vyrnwy, SH996205, P. Parker, 1997, NMW, det. P.M. Benoit.

+16/1.1. MATTEUCCIA STRUTHIOPTERIS 6, N. Somerset: Over 50 plants under trees in marshy
area of wood on stream edge, Vobster, ST710489, P.R. Green, 1997.

17/1.1. POLYSTICHUM SETIFERUM 67, S. Northumb.: Wooded slope above stream bank,
Dovecrag Burn, NT945024, G.A. & M. Swan, 1997, herb. G.A.S. 2nd confirmed record.

17/1.1x2. POLYSTICHUM SETIFERUM x P. ACULEATUM (P. x BICKNELLII) +7 Comvwall:
With both parents in woodland by the River Tamar, Lamerhooe Ford, W of Latchley, $X398732,
M. Atkinson, 1996, conf. A.M. Paul. *37, Worcs.: Park Wood, Malvern Hills, SO7644, R.
Maskew, 1991, conf. A.M. Paul. *50, Denbs.: Bank of old railway, Pwllglas, SJ125551, M.
Keene, 1995, det. A. Paul.

+17/1.1xmun. POLYSTICHUM SETIFERUM x P. MUNITUM *17, Surrey: One plant of several
fronds with both parents on bank of B.P., Winkworth Hill, SU996403, A.C. Leslie, 1995, herb. A.
C.L.

17/3.2x3. DRYOPTERIS FILIX-MAS x D. AFFINIS (D. x COMPLEXA) *37, Worcs.: Streamside
wooded bank, Southstone Rock, Teme Valley, SO7063, P.G. Garner & R. Maskew, 1989, det. A.
C. Jermy,.

17/3.3a. DRYOPTERIS AFFINIS subsp. AFFINIS *S, Channel Islands: Frequent in wooded valley
near stream, Dixcart Valley, Sark, WV467754, R.M. Veall, 1996, det. M. Gibby.

17/3.3b. DRYOPTERIS AFFINIS subsp. CAMBRENSIS *2, E. Cornwall: Two plants only in
unshaded roadside hedge on granite, Kit Hill, near Callington, SX383714, R.J. Murphy & M.G.C.
Atkinson, 1999, det. A.C. Jermy. Ist confirmed record. *11, S. Hants.: Beside main ride in
wood, Crowdhill Copse, Fair Oak, SU487198, P.J. Selby, 1995, herb. R.P. Bowman, det. H.
Corley.

719/1.1. AZOLLA FILICULOIDES 41, Glam.: Settling pond, Aberthaw Cement Works, ST0367,
M. Hampton, 1996. Only extant locality.

PLANT RECORDS 555

720/1.1. ABIES ALBA *$1, Berwicks.: Maturing saplings established at plantation edge,
Woodhead, Lauder, NT531468; Plantation edge, Blinkbonny Hill, NT566395; both M.E.
Braithwaite, 1996. Ist & 2nd records as self-seeded.

+20/1.nor. ABIES NORDMANNIANA *46, Cards.: About 15 self-sown trees 2-4m tall in estate
woodland, Llanerchaeron, SN479601, S.P. Chambers, 1996.

+20/2.1. PSEUDOTSUGA MENZIESII *46, Cards.: Shingle by Afon Ystwyth 2km W of
Pontrhyd-y-groes, SN720721, A.O. Chater, 1990. Estate woodland, Coed Ffynnon Caradog,
Llangorwen, SN617831, A.O. Chater, 1991. Ist and 2nd records. *81, Berwicks.: Bank, A68
near Chapel-on-Leader, NT564412, M.E. Braithwaite, 1996. Ist record as self-seeded.

+20/3.1. TSUGA HETEROPHYLLA *47, Monts.: Cwm wood, Aberhafesp, SO0595, IC.
Trueman & M. Oliver; Hafren Forest, SN8685, A.J. Morton, both 1988. Ist and 2nd records.
Regeneration observed in both sites.

+20/4.1. PICEA SITCHENSIS *S, Channel Islands: One small tree (planted) about Im tall in
wooded valley, Dixcart Valley, Sark, WV466754, R.M. Veall, 1997, JSY, det. R.A. Waterman.
*46, Cards.: Roadside slopes by plantations, Cwm Einion, SN695942, A.O. Chater, 1983. Edge of
reservoir by plantations, Llyn Pendam, SN702842, A.O. Chater, 1985. Ist and 2nd records. Widely
naturalised in Cards.

+20/4.2. PICEA ABIES *46, Cards.: Roadside banks and plantation, 300m NW of Panthirion,
Brynafan, SN701729, A.O. Chater, 1993. Roadside slope by plantation, W of Blaenmyherin,
SN785795, A.O. Chater, 1993. 1st and 2nd records. Rarely naturalised.

+20/5.1x2. LARIX DECIDUA x L. KAEMPFERI (L. x MARSCHLINSID) *12, N. Hants.: Self-sown,
Alice Holt Forest, Glenbervie Enclosure, SU8142, A. Brewis, 1996. *81, Berwicks.: Plantation
edge, Mellerstain, NT644401, M.E. Braithwaite, 1996. 1st record as self-seeded.

720/5.2. LARIX KAEMPFERI *46, Cards.: Roadside verge and slopes by plantations by A44, W
of Nantyrarian, SN709811, A.O. Chater, 1988. Rocky slopes by plantations, Coed Bwlchgwallter,
Hafod, SN770717, A.O. Chater, 1993. Ist and 2nd records. Widely naturalised. *47, Monts.:
Plentiful regeneration on forestry tracksides, Uwch-y-garreg, SN7793, A.J. Morton & W.A.
Thompson, 1990.

720/7.2. PINUS NIGRA *46, Cards.: Edges of mixed plantation, Coed Ty’n-y-garth, Cwm
Einion, SN689946, A.O. Chater, 1991. Roadside verges by plantations, 800m ENE of Bryn
Llychese, SN832810, A.O. Chater, 1993. 1st and 2nd records. Widely naturalised.

720/7.5. PINUS RADIATA *35, Mons.: A line of 10 trees at field edge on B4293 for at least the
last 40 years, Devauden, ST4898(Z), T.G. Evans, 1995, herb. T.G.E.

+20/7.6. PINUS STROBUS *46, Cards.: In middle of pasture, below the Rookery Wood NR,
Nanteos, SN616781, A.O. Chater, 1991, NMW. *50, Denbs.: Probably planted near farm,
Nantglyn, SJO061, P. Williams, 1996.

720/7.mug. PINUS MUGO-_ *46, Cards.: S facing sheepwalk by side of old estate lodge, Angler’s
Retreat (Plas y Mynydd), SN746923, A.O. Chater, 1992, NMW. = *81, Berwicks.: Heather bank,
A68 at Soutra, NT476574, M.E. Braithwaite, 1997. lst record as self-seeded.

+20/7.mur. PINUS MURICATA _ S, Channel Islands: A mature tree, edge of maritime scrub, near
hotel, near Jaspellarie, overlooking Dixicart Bay, Sark, WV467252, R.M. Veall, 1983, det. R.A.
Waterman. Confirmed R.A. Waterman (1997) as obviously planted but no longer present. Noted
by M.H.Marsden in 1994.

*20/7.wal. PINUS WALLICHIANA *35, Mons.: One old tree producing cones at woodland edge
at top of cliffs, Piercefield, ST5295, T.G. Evans, 1995.

t20A/CRY jap. CRYPTOMERIA JAPONICA *46, Cards.: Open mixed plantation, Cockshead
Wood, 3km NE of Llangybi, SN630554, A.O. Chater, 1996. *49, Caerns.: Probably planted on
roadside, Bettws-y-Coed, SH7853, J.A. Green, 1997.

+20A/SEQ.sem. SEQUOIA SEMPERVIRENS *S, Channel Islands: In wooded valley bordering
hotel garden, Dixcart Valley, Sark, WV465754, R.M. Veall, 1984, det. R.A. Waterman. Two trees
obviously planted; one on edge of hotel garden subsequently destroyed by gale, survivor is inside
hotel garden. *47, Monts.: A single seedling, about 3 yrs old, under the largest trees in the
Redwood Grove, Leighton near Welshpool, $J2403, A. Markwick, 1987. The first seedling to have
occurred naturally here and still flourishing in 1997.

556 PLANT RECORDS

421/1.1x2.1. CUPRESSUS MACROCARPA x CHAMAECYPARIS NOOTKATENSIS
(x CUPRESSOCYPARIS LEYLANDII) *S, Channel Islands: Established plantation of mixed, mature
trees, near L’ Ecluse, Sark, WV461766, R.M. Veall, 1997, JSY, det. R.A. Waterman, Mature trees
(also at other sites on Sark), all apparently planted, up to 9m or more.

+021/2.1. CHAMAECYPARIS LAWSONIANA ~— *81, Berwicks.: Plantation edge, Blinkbonny Hill,
NT566395, M.E. Braithwaite, 1996. Ist record as self-seeded.

*21/3.1. THUJA PLICATA *46, Cards.: Estate woodland by stream, Tyglyn, Ciliau Aeron,
SN498598, A.O. Chater, 1992. Slope above railway and in adjacent woodland, Glandyfi,
SN694968, A.O. Chater, 1995. Ist and 2nd records. Widely naturalised.

+022/1.1. ARAUCARIA ARAUCANA *17, Surrey: Margin of raised area, sandy soil, Dorking,
TQ174494, J.F. Leslie, K. Page & J.E. Smith, 1997.

23/1.1. TAXUS BACCATA _+*81, Berwicks.: Cliffs, Newton Don, NT714374, M.E. Braithwaite,
1995. Ist record as established.

+26/2.3. NUPHAR ADVENA _ *81, Berwicks.: Established in artificial loch, Spottiswoode Loch,
NT6049, A.G. & D.G. Long, 1964, HAMU. Originally det. as N. /utea. Confirmed by D.G. Long,
1996, E. Ist record.

28/12.1. CLEMATIS VITALBA 73, Kirkcudbrights.: Growing up 12m around a 2m diameter
dead tree, near Grove Hospital, NX9278, O.M. Stewart, 1997. 2nd extant record.

28/13/7. RANUNCULUS PARVIFLORUS 12, N. Hants.: Large patch near Range Gate No. 2,
Woolmer Forest, SU788327, J.S. Denton, 1994. Ist recent record, not seen since.

+28/13.16. RANUNCULUS ACONITIFOLIUS flore pleno *68, Cheviot: Naturalised in wet grass
by stream, Shawdon, NZ083151, G.A. Swan, 1997, herb. G.A.S.

28/13.24. RANUNCULUS PELTATUS 94, Banffs. : Ditch, Nether Dallachy, NJ352644, J.
Edelsten, 1994, E, det N.T.H. Holmes. 2nd record.

28/13.25a. RANUNCULUS PENICILLATUS subsp. PENICILLATUS 50, Denbs.: River Conwy,
Llanrwst, SH7962, J.A. Green, 1996. 2nd record.

28/17.4. THALICTRUM MINUS _+*46, Cards.: Roadside verge and scrub 250m W of Post Office,
Penbontrhydybeddau, SN676836, A.O. Chater, 1992. Roadside verge on river bank, 50m E of
Pont Tanycastell, Rhydyfelin, SN589788, A.O. Chater, 1992. 1st and 2nd records.

28/17.5. THALICTRUM ALPINUM *77, Lanarks.: Rocky outcrop and bank above Gana Burn,
Gana Hill, NS9501, K. Watson, 1997.

+28/4.1. ERANTHIS HYEMALIS *12, N. Hants.: Hedgebank, South Warnborough, SU720453,
C.R. Hall, 1989. Planted in woodland, dominant, spreading to adjoining fields, Wyck Place, near
Alton, SU759392, A. Brewis, 1996. Ist and 2nd records.

+28/5.1. NIGELLA DAMASCENA —_*49, Caerns.: Waste ground near railway station, Llandudno,
SH7881, W. McCarthy, 1997.

+28/6.1xvar. ACONITUM NAPELLUS x A. VARIEGATUM (A. x CAMMARUM) *12,.N. Hants::
One slowly spreading clump with violet flowers, Odiham Common, SU750524, J. Moon, 1996.

+28/7.1. CONSOLIDA AJACIS *46, Cards.: Disturbed slopes of grassed-over rubbish tip,
Pendinas, Aberystwyth, SN584798, A.O. Chater, 1991, NMW.
+28/9.bla. ANEMONE BLANDA *49 Caerns.: Naturalised on roadside, Bodafon Lane,

Llandudno, SH8081, W. McCarthy, 1997.

*29/1.2. BERBERIS THUNBERGII *73, Kirkcudbrights.: Wood, Loch Ken side of Kenmuir
Castle, NX635764, O.M. Stewart, 1995.

729/1.5. BERBERIS AGGREGATA *95, Moray: Several plants, self sown on wall an edge of
wood, W of Elgin, NJ170630, P.R. Green, 1997.

+29/1.9. BERBERIS DARWINII *11, S. Hants.: Scrub on heath in brambles, East Boldre,
SZ366991, A.E. Bolton, 1997, det. Everton Nurseries.

+30/1.1. PAPAVER PSEUDOORIENTALE *81, Berwicks.: Established on road verge, A6105
-Longformacus road end, NT772532. Known here for many years; Established on and under Town
walls, Coldstream; NT842395; both M.E. & P.F. Braithwaite, 1997. Ist and 2nd records. *99,
Dunbarton: Waste ground, bank near Hillfoot Station, Bearsden, NS5572, A.McG. Stirling & A.
Rutherford, 1983.

PLANT RECORDS 557

30/1.5b. PAPAVER DUBIUM subsp. LECOQHI *81, Berwicks.: Soil heap, housing estate, Oxton,
NT498534, M.E. & P.F. Braithwaite, 1997. Well established around village; recorded here as a
casual in 1960.

+30/6.1. ESCHSCHOLZIA CALIFORNICA 46, Cards.: Established for several years and abundant
on sandy slope above shore, Penyrergyd, Gwbert, SN164485, A.O. Chater, 1996. 2nd record.

731/1.1. DICENTRA FORMOSA *44, Carms.: Escaping along stream from garden, near
Blaenwaun, SN219280, G. Hutchinson & J. Bevan, 1995, conf. A.O. Chater.

731/3.2. PSEUDOFUMARIA ALBA *44, Carms.: Gelli Aur Country Park, SN5920, D. Smith,
1988, NMW, det. D. McClintock. Garden escape.

31/5.1b. FUMARIA CAPREOLATA subsp. BABINGTONII *37, Worcs.: Old wall, Witley Court,
SO7764, R. Maskew & P. Whitehead, 1991, conf. M.G. Daker. 39, Staffs.: One plant in flower
and fruit in hedgebank, Workhouse Lane, Keele, SJ805452, IJ. Hopkins, 1995. Ist record since
1954.

31/5.3. FUMARIA BASTARDIT —108, W. Sutherland: Garden weed, Clachtoll, NC039280, I.M.
Evans, 1997. 2nd record.

31/5.6. FUMARIA PURPUREA 51, Flints.: Hedgebank along roadside between Bodfari and
Tremeirchion, SJ085717, G. Wynne, 1992, det. M. Daker. Ist record since 1942.

31/5.10. FUMARIA VAILLANTII 28, W. Norfolk: Garden weed, Fakenham, TF934298, M.
Keene, 1997, conf. K. & G. Beckett. Ist record since 1958.

+32/1.occxori. PLATANUS OCCIDENTALIS x P. ORIENTALIS (P. x HISPANICA) *35, Mons:.:
Field/road edge, Piercefield Park, Chepstow racecourse, ST5294, T.G. Evans, 1995.

33/1.3. ULMUS MINOR = *43, Rads.: In corner of field, near Shaky Bridge, Llandrindod Wells,
SO0081614, A. Orange, 1984, NWVW.

+33/1.lae. ULMUS LAEVIS _ S, Channel Islands: Clump of trees near well in valley meadow, S
of Le Manoir, Sark, WV465757, R.M. Veall, 1975, det. T.R.G. Rountree. One mature tree
presumably planted, storm damaged 1987, dead 1990. *46, Cards.: Well established by suckers
from two mature trees in mixed woodland, Y Glog, Rhydyfelin, SN593793, A.O. Chater, 1995,
NMW.

nos-2-L. HICUS CARICA 58, Cheshire: Canal bank, Winsford, SJ703663, J.H. Clarke, 1997.
Only extant locality.

739/2.ant. NOTHOFAGUS ANTARCTICA *S, Channel Islands: Planted tree about 2-5m tall,
roadside verge, Le Manoir, Sark, WV466758, R.M. Veall, 1997, JSY, det. R.A. Waterman.

39/4.2xsub. QUERCUS CERRIS x Q. SUBER (Q.x CRENATA) *35, Mons.: 2 trees in churchyard
at Trevethin, SO284020, C. Titcombe, 1997.

139/4.3.+ QUERCUS ILEX 47, Monts.: One large tree in council car park, Newtown, SO1091,
C.A. Small, 1980. Still there in 1997. 2nd record.

740/1.pap. BETULA PAPYRIFERA _*S, Channel Islands: Roadside verge, Aval du Creux, Sark,
WV471759, R.M. Veall, 1997, JSY, det. R.A. Waterman. Planted.

40/2.1x2. ALNUS GLUTINOSA x A. INCANA (A. x HYBRIDA) *77, Lanarks.: Self sown on to a
coal bing, Hamilton, NS7253, P. Macpherson, 1996, herb. P.M., det. C.S. Crook.

740/2.3. ALNUS CORDATA *46, Cards.: Marshy waste ground NW of Pen-y-bont Bridge,
Llanbadarn Fawr, SN594804, A.O. Chater, 1997.

40/3.1. CARPINUS BETULUS *%+81, Berwicks.: Established on bank, Carolside, NT565397, M.
E. & P.F. Braithwaite,1995. 1st record as an established alien.

43/1.10. CHENOPODIUM HYBRIDUM 3, S. Devon: Pottles Farm, Exminster, $X924873, L.M.
Spalton, 1996. 2nd record.

43/1.6. CHENOPODIUM RUBRUM _ *43, Rads.: On heaps of farm manure on roadside tip, S of
Upper Pits Farm, Knighton, SO315710, R.G. Woods, 1990, NMW.

43/3.2x4. ATRIPLEX PROSTRATA x A. LONGIPES (A. x GUSTAFSSONIANA) *35, Mons.: Many
plants mixed with A. procumbens on upper shore and rocks of sea wall, Uskmouth, $T3382, T.G.
Evans, 1996, NMW. Upper saltmarsh, Blackrock, ST5188, T.G. Evans, 1996, NMW. Ist and 2nd
records.

558 PLANT RECORDS

+45/2.1. CLAYTONIA PERFOLIATA 46, Cards.: Among gorse on S facing scree W of Cliff
Railway, Constitution Hill, Aberystwyth, SN583826, A.D. Hale, 1996. 2nd record.

745/2.2. CLAYTONIA SIBIRICA — 41, Glam.: North bank of River Rhymney, N of Llanrumney
school, ST209803, 1995, J.P. Woodman. Only extant locality.

46/5.3. STELLARIA PALLIDA *8, S. Wilts.: Windmill Hill, East Knoyle, ST873310, D. Green,
1997. Not seen subsequently. *39, Staffs.: Locally frequent on thin soil over rock ledge, Ilam,
Hall Dale, SK133538, R.E. Groom, 1997, herb. B.R. Fowler.

46/5.4. STELLARIA NEGLECTA *81, Berwicks.: Wood, Coldingham Bay, NT9166, E.P. Beatie,
1956. Shoreline, Coldingham West Loch, NT893679, M.E. Braithwaite, 1996. Ist and 2nd records.

46/7.2x3. CERASTIUM ARVENSE x C. TOMENTOSUM = *37, Worcs.: With both parents in sandy
heathland, Hartlebury Common, SO8170, W.A. Thompson, 1993, conf. J.J. Day & R. Maskew.

46/7.7b. CERASTIUM FONTANUM subsp. HOLOSTEOIDES *108, W. Sutherland: Shady seacliff,
Fanagmore. NC177502, I.M. Evans, 1997.

46/7.11. CERASTIUM PUMILUM *31, Flints.: Open calcareous grassland on rocky, W facing
slopes, Prestatyn, SJ067810, D. Guest, 1996.

46/8.1. MYOSOTON AQUATICUM *$1, Berwicks.: Sandy bank, River Tweed between
Fireburnmill and the Lees, NT832389, M.E. Braithwaite, 1997, herb. M.E.B.

46/9.1. MOENCHIA ERECTA *28, W. Norfolk: Three colonies of c.12 plants in short grass on
shingle, Snettisham, TF650304, S. Fisher, 1996, conf. G. Beckett. Ist record since 1959. = *41,
Glam.: By paths in Agrostis heath, on S facing slopes of hill, Penmaen, Gower, $S537885, Q.O.N.
Kay, 1992, NMW.

46/10.7a. SAGINA APETALA subsp. APETALA *67, S. Northumb.: Forest roadside, Black Rigg,
NU753747, G.A. & M. Swan, 1967, herb. G.A.S., det. F.N. Hepper.

46/10.8. SAGINA MARITIMA _ 81, Berwicks.: Road verge, A68 at Renshawrig, NT477571, M.E.
Braithwaite, 1996. Ist inland record.

46/15.1. POLYCARPON TETRAPHYLLUM *6, N. Somerset: Sparse open grassland (school
playing field), Weston-super-Mare, ST320593, R. Higgins, 1997, herb. I.P. Green.

46/17.3. SPERGULARIA MARINA — +*77, Lanarks.: Verge of A721, Melbourne, NT0944; herb.
M.E.B. Verge of A721, near Harpenhall, NT0443; both M.E. Braithwaite, 1996. lst and 2nd
records. Abundantly established.

46/17.5. SPERGULARIA BOCCONEI 1, W. Cornwall: In sandy patch by inlet off the Camel
Estuary, near Dennis Farm, S of Dinas and Padstow, SW919744, H.M. Meredith & M. Tempest,
1996, conf. R.J. Murphy. A completely new locality.

+46/18.1. LYCHNIS CORONARIA *49, Caerns.: Naturalised by footpath, Wellington near
Llanbedrog, SH3030, BSBI meeting, 1997.

46/20.9x10. SILENE LATIFOLIA x S. DIOICA (S. x HAMPEANA) *47, Monts.: One plant with
S. latifolia, in waste ground on old lead-mine site, Fan Pool, SN9487, W.A. Thompson, 1986.

46/20.11. SILENE GALLICA *43, Rads.: One plant on recently constructed road verge, N of
entrance to sewage works, Rhayader, SO978673, R.G. Woods, 1987.

46/25.5. DIANTHUS DELTOIDES +39, Staffs.: Eight patches in short sunny grassland, landfill
site, Dudley Wood, near Mushroom Green, $0939863, T. Beynon, 1996, BM, det. F. Rumsey &
A.M. Paul. Known here for 2 -3 years. *+73, Kirkcudbrights.: Well naturalised by ruined
cottage, Craigaploch, Dundrennan, NX724444, O.M. Stewart, 1995.

46/25.7. DIANTHUS ARMERIA 41, Glam.: 98 flowering and 140 non-flowering plants in open
stoney waste in old, disused, sandstone quarries, Cwm Clydach, Rhondda, $S961926, J.P.
Woodman; 74 plants counted on dry bank by Baglan Reservoir, SS741930, C.R. Hipkin. Known in
this area for several years. Both 1997. 2nd extant localities.

t47/1.1. PERSICARIA ALPINA *77, Lanarks.: Grassy slope above burn, Cambuslang Golf
Course, NS6460, K. Watson, 1987, GL. Seen well established in 1996 by P. Macpherson.

+47/1.2. PERSICARIA CAMPANULATA *50, Denbs.: Wet wood near garden, Capel Garmon,
SH8157, BSBI group, 1996.

PLANT RECORDS 559

+47/1.5. PERSICARIA MOLLIS *11,S. Hants.: Steep E-facing grass bank overlooking W side of
roadside pond, St Leonards Grange, St Leonards, Beaulieu, SZ406983, R.A. Barrett & E.J.
Clement, 1995, herb. R.P. Bowman.

747/1.7. PERSICARIA AMPLEXICAULIS *46, Cards.: Grassy roadside verge, Upper Borth,
SN607889, J.E. Halfhide, det. A.J. Silverside, 1977. Hedgebank, Capel Salem graveyard,
Brongest, SN324450, A.O. Chater, 1997. lst and 2nd records.

47/1.16. PERSICARIA MINOR 50, Denbs.: Oxbow, Llanrwst, SH7962, R. Lewis. 1996. 2nd
post-1930 record.

+47/1.cap. PERSICARIA CAPITATA *49, Caerns.: Pavement cracks, Abersoch, SH3128 &
Criccieth, SH501380, J. Hawksford, 1997. Ist records.

47/4.6. POLYGONUM RURIVAGUM *46, Cards.: Bean field, 150m SW of Pen, SN216521, A.O.
Chater, 1989, NMW. = _+*81, Berwicks.: Established in stubble, Fireburnmill, NT8139, A. Espir,
1992, det. D.R. McKean. Established in arable field, Coldstream Hospital, NT832393, M.E.
Braithwaite, 1997, herb. M.E.B., det. J.R. Akeroyd. 1st and 2nd records.

47/5.1x2. FALLOPIA JAPONICA x F. SACHALINENSIS (F. x BOHEMICA) *35, Mons.: Scattered
along ditch / roadside, E bank of River Rhymney, NE of Michaelstone Bridge, ST2486, T.G.
Evans, 1996, NMW, conf. A.P. Conolly. Some F. japonica present. *41, Glam.: Along about
30m stretch of roadside, N of Michaelstone Bridge, ST2485, T.G. Evans, 1996, NMW. *64,
Mid-W. Yorks.: Large patch by old mill-race, Kirkstall Abbey, Leeds, SE261361, K. Pyne, 1996,
LTR, det. J.P. Bailey & A.P. Conolly. *95, Moray: Two large stands on verge of B9010,
growing with F. sachalinensis at roadside, Kellas, near Dallas, NJ163538, IP. Green, 1996, conf.
A.P. Conolly. Now recorded from several sites.

747/5.3. FALLOPIA BALDSCHUANICA = *50, Denbs.: River bank, Bryn Rhyd-yr-arian, SH9568,
J.A. Green, 1996.

747/7 rhaxpal.t RHEUM RHAPONTICUM x R. PALMATUM (R. x HYBRIDUM) *12, N. Hants.:
Lay-by, roadside, near Wootton St Lawrence, SU598534, J. & P. Rollinson, 1997, conf. A.
Brewis.

47/8.8. RUMEX LONGIFOLIUS *108, W. Sutherland: Marshy bank of Ledmore River,
NC253116, I.M. Evans, 1997, conf. J.R. Akeroyd.

47/8.10x 14. RUMEX HYDROLAPATHUM x R. CONGLOMERATUS (R. X DIGENEUS) *47, Monts.:
Rare in Montgomery Canal near Arddlin, SJ265165, R. Lansdown, 1997, RNG, conf. J.R.
Akeroyd.

47/8.10x19. RUMEX HYDROLAPATHUM x R. OBTUSIFOLIUS (R. x LINGULATUS) *37, Worcs.:
Old brick-pit, Grimley, SO8460, C.B. Westall, 1989, conf. J.R. Akeroyd. *47, Monts.: Rare in
Guilsfield Arm of Montgomery Canal SSSI, $J239133, R. Lansdown, 1997.

47/8.13b. RUMEX CRISPUS subsp. LITTOREUS *35, Mons.: Sea wall near River Severn, Lamby,
ST228777, T.G.Evans, 1995, NMW. *49, Caerns.: Sea-shore, Morfa Nefyn, SH2940, J.
Hawksford, 1997. Sea-shore, Abersoch, SH3128; Waste ground by sea-shore, Pwllheli, SH3734;
Beach, Criccieth, SH4937, J. Hawksford, 1997. Ist and 2nd records. *61, S.E. Yorks.: Welwick
Saltmarsh, 1-6km S of Welwick, TA3318, P.J. Cook, 1997, det. J.R. Akeroyd.

47/8.13c. RUMEX CRISPUS subsp. ULIGINOSUS *46, Cards.: Edge of tidal ditch, Teifi estuary,
Cardigan, SN14, A. Ley, 1885, BM, det. J.R. Akeroyd. Flood terrace, Leri estuary E of Borth
church, SN616899, J.R. Akeroyd & A.O. Chater, 1988. 1st and 2nd records.

47/8.13x19. RUMEX CRISPUS x R. OBTUSIFOLIUS (R. x PRATENSIS) 50, Denbs.: Waste ground,
Pontcysyllte, SJ266419, J.A. Green, 1997. 2nd record. *73, Kirkcudbrights.: Waste ground,
Newton, Dumfries, NX9477, O.M. Stewart, 1996.

47/8.14x21. RUMEX CONGLOMERATUS x R. MARITIMUS (R. x KNAFII) *58, Cheshire: Muddy
shore, Rostherne Mere, SJ740840, G.M. Kay, 1997, herb. G.M.K., det. J.R. Akeroyd.

47/8.15x9. RUMEX SANGUINEUS x R. OBTUSIFOLIUS (R. x DUFFTII) *58, Cheshire: Ditch,
Adlington, SJ905796, J.H. Clarke, 1997, det. J.R. Akeroyd.

47/8.16. RUMEX RUPESTRIS 41, Glam.: 21 fruiting plants on largely inaccessible tufaceous
cliff-ledge near Dunraven, SS8872, Q.O.N. Kay, 1996. Ist record since 1934.

560 PLANT RECORDS

47/8.19x21. RUMEX OBTUSIFOLIUS x R. MARITIMUS (R. x CALLIANTHEMUS) Sie WV OICS.:
Damp laneside verge, Heightington, SO7671, W.A. Thompson & B. Westwood, 1996, det. J.R.
Akeroyd.

47/8.20. RUMEX PALUSTRIS 15, E. Kent: On sandy banks and edges of newly dug pits,
Dungeness, TRO618, E.G. Philp, 1997. 1st record since 1632; c.100 plants still present in 1997.

48/1.11. LIMONIUM PARVUM 45, Pembs.: Several hundred plants on blow-hole rim in
limestone sea cliff, E of Mewsford Point, Castlemartin, SR9493, S.J. Leach, 1993, det. M.
Ingrouille. Splash zone in gulley near top of limestone sea cliff, Stackpole NNR, SR9894, S.J.
Leach, 1993, det. M. Ingrouille. 2nd British records.

+49/1.off. PAEONIA OFFICINALIS *69, Westmorland: S side of fell road far from houses, Fell
Yeat, Casterton, SD636795, C.E. Wild, 1990, LANC. *77, Lanarks.: Grassy bank, Shiels,
Glasgow, NS5366, P. Macpherson, 1985, herb. P.M. Still there and flowering in 1989.

*51/1.1. HYPERICUM CALYCINUM — 50, Denbs.: Edge of car park, Mold, SJ199627, J. Philips,
1997. Ist recent record.

+51/1.3x4.¢ HYPERICUM ANDROSAEMUM x H. HIRCINUM (H. x INODORUM) *49, Caerns.: Old
quarry, Llangystennin near Colwyn Bay, SH8279, G. Battershall & W. McCarthy, 1997.

+51/1.4., HYPERICUM HIRCINUM *50, Denbs.: Cracks in stone bridge, Tervor, $J262415, J.A.
Green, 1997.

51/1.6x7. HYPERICUM PERFORATUM x H. MACULATUM (H. x DESETANGSII) *35, Mons.: Side
of forest track, Thicket Wood, Rogiet, ST4488, T.G. Evans, 1991; small wood, Cwmcarvan,
SO4807, J. Harper, 1991. Ist and 2nd records.

*51/1.for. HYPERICUM FORRESTIL *46, Cards.: Mixed woodland on N bank of Afon Ystwyth,
Chwarel Goch, Pontrhyd-y-groes, SN725721, A.O. Chater, 1994, det. N.K.B. Robson.

52/1.2. TILIA CORDATA = 29, Cambs.: Regrowth from large old coppice stool c.300 years old,
Ditton Park Wood, TL662570, S. Leatherdale, 1995, CGE, det. C.D. Pigott. Ist record in E of
county of native tree. *73, Kirkcudbrights.: Wood, by Loch Roan, NX745693, O.M. Stewart,
1995. +*30, Roxburghs.: A single large roadside tree planted W of Fairnington, Roxburgh,
NT643278, R.W.M. Corner, 1995, herb. R.W.M.C., det. C.D. Pigott.

+52/l.euc. TILIA x EUCHLORA *29, Cambs.: In Park, by SW corner of Cherry Hill, Ely,
TL540798, G. Crompton, 1993, det. C.D. Pigott. Widely planted in Cambridge; it is sterile and
probably all one genetic clone.

752/1.tom. TILIA TOMENTOSA ~~ *S, Channel Islands: Two mature trees (planted), taller about
8m, roadside verge, La Vaurocque, Sark, WV462758, R.M. Veall, 1997, JSY, det. R.A.
Waterman..

753/6.1. ABUTILON THEOPHRASTI *42, Brecs.: 3 flowering plants in swede field, Ty Mawr,
Llysdinam, SN9957, K. Hughson & R.G. Woods, 1996.

57/1.2. VIOLA HIRTA =.28, W. Norfolk: Chalk downland, Ringstead Downs, TF691402, T.C.E.
Wells, 1997. 2nd extant locality in W. Norfolk.

57/1.4x5. VIOLA RIVINIANA x V. REICHENBACHIANA (V. x BAVARICA) *50, Denbs:.:
Deciduous woodland, Ruthin, $J125554, S. Chambers, 1996, det. D. Moore. Ist record.

57/1.7. VIOLA LACTEA = 10, Wight: Clifftop heath, Bouldor Copse, $Z379903, T.D. Dines & C.
D. Preston, 1996. 2nd extant locality.

57/1.9b. VIOLA PALUSTRIS subsp. JURESSI 46, Cards.: Damp rocks by stream, 900m W of
Glasbwll, Llyfnant, SN729974, A.O. Chater & J.P. Woodman, 1995, NMW. 2nd record.

+57/1.11x12xalt. VIOLA ALTAICA x V. LUTEA x V. TRICOLOR (V. xX WITTROCKIANA) *50,
Denbs.: Weedy roadside verge, Llyn Brenig, SH983592, BSBI group, 1997.

61/1.1x2. POPULUS ALBA x P. TREMULA (P. x CANESCENS) _ S, Channel Islands: On wall W of
Seineusie grounds, road to La Moinerie, Sark, WV463764, R.M. Veall, 1997, JSY, det. R.A.
Waterman. Ist record as naturalised. Trees now about 3m tall and suckered from tree about 30m
distant.

61/1.3a. POPULUS NIGRA subsp. BETULIFOLIA *44, Carms.: tMorfa, Llanelli, SS59, J.A.Webb,
1945, NMW, det. G. Hutchinson & I.K. Morgan; tree now probably destroyed. Four trees on edge
of footpath adjacent to reed-swamp, Dyfatty Marsh, Burry Port, SN454008, G. Hutchinson, 1993,
NMW, conf. R.D. Meikle. Ist and 2nd records.

PLANT RECORDS 561

¥61/1.3xdel. POPULUS NIGRA ‘PLANTIERENSIS’ x P. DELTOIDES ‘CORDATA’ (P. x CANADENSIS
‘“ROBUSTA’) *S, Channel Islands: One tree (planted) about 4m tall, roadside, foot of hedgebank,
Rue de la Coupée between La Vaurocque & Dixcart Lane, Sark, WV461755, R.M. Veall, 1997,
JSY, det. R.A. Waterman.

+61/1.3xdel. POPULUS x CANADENSIS “‘MARILANDICA’ x P. x CANADENSIS ‘SEROTINA’ (P. x
CANADENSIS ‘REGENERATA’ ) *S, Channel Islands: Three mature trees (planted), tallest now
18m, near stream in small, partly wooded, valley, Harbour Hill, Sark, WV473758, R.M. Veall,
1997, JSY, det. R.A. Waterman. Known to R.A.W. for many years.

*+61/1.delx4. POPULUS DELTOIDES x P. TRICHOCARPA (P. xX GENEROSA) *S, Channel Islands:
Two trees (planted), taller about 6m, roadside, foot of hedgebank, Rue de la Coupée between La
Vaurocque & Dixcart Lane, Sark, WV461755, R.M. Veall, 1997, JSY, det. R.A. Waterman.

61/2.5lam. SALIX PURPUREA subsp. LAMBERTIANA _*77, Lanarks.: Bank of Carmichael Burn,
Carmichael, NS9238, P. Macpherson, 1997, herb. P.M., det. R.D. Meikle.

61/2.10x11. SALIX CAPREA x S. CINEREA (S. x REICHARDTII) *47, Monts.: Canal tow-path,
Red Bridge, Arddlin, S$J2514, F.H. Perring, 1975. Beside River Rhiw, NE of Manafon, SJ122028,
E. Roberts & M. Wainwright, 1991, det. C.A. Sinker. Ist and 2nd records.

61/2.10x12. SALIX CAPREA x S. AURITA (S. x CAPREOLA) *44, Carms.: Edge of scrub, acid
moorland valley, Glanrhyd Farm, Maudsland, SN332321, R.D. Pryce, 1992, NMW, det. R.D.
Meikle. 1st confirmed record.

61/2.10x15. SALIX CAPREA x S. PHYLLICIFOLIA *73, Kirkcudbrights.: Wood, Hardgate
Quarry, NX817672, O.M. Stewart, 1996, E, det. R.D. Meikle.

61/2.11bx16. SALIX CINEREA subsp. OLEIFOLIA x S. REPENS (S. x SUBSERICEA) *46, Cards.:
Damp meadow, Peris Brook, Llanon, SN56, J.H. Salter, 1930, NMW, det. R.D. Meikle.

61/2.12x15. SALIX AURITA x S. PHYLICIFOLIA (S. x LUDIFICANS) *108, W. Sutherland: Edge
of burn, Strath Feith an Leothaid, NC200212, P.A. Evans, 1997, conf. R.D. Meikle.

61/2.12x16. SALIX AURITA x S. REPENS (S. x AMBIGUA) 77, Lanarks.: Boggy ground, W of
Longriggend, NS8070, P. Macpherson, 1997, herb. P.M., det. R.D. Meikle. 1st record since 1923.

61/2.14. SALIX MYRSINIFOLIA 81, Berwicks.: Scrub, Winding Burn, NT815665, M.E.
Braithwaite, 1996. Only extant record.

61/2.14x20. SALIX MYRSINIFOLIA x S. MYRSINITES (S. x PUNCTATA) *94, Banffs.: Scree,
Shelterstone Crag, NH999015, J. Edelsten, 1994, E, det. R.D. Meikle.

61/2.16. SALIX REPENS 47, Monts.: Damp lake margin with Calluna, Molinia and Sphagnum,
Llyn Gwyddior, SH938074, T. Teearu, 1997, conf. R.D. Meikle. 1st recent record.

61/2.21. SALIX HERBACEA = 95, Moray: Barish areas on top of mountain, Creag an Tarmachain,
Grantown-on-Spey, NJ154312, IP. Green, 1996, conf. P.R. Green. 2nd record.

61/2.9x10x11. SALIX VIMINALIS x S. CAPREA x S. CINEREA (S. x CALODENDRON) 137,
Worcs.: Waste ground, Bury Hill, Oldbury, SO9789, W.A. Thompson, 1990, det. R.D. Meikle.

61/2.9x11. SALIX VIMINALIS x S. CINEREA (S. x SMITHIANA) *50, Denbs.: Streamside by
road, Llangwm, SH9844, A.O. Chater, 1996. Riverbank, Llanrwst, SH7961, J.A. Green, 1996. Ist
and 2nd records.

762/7.1. ERYSIMUM CHEIRANTHOIDES 81, Berwicks.: Established in turnip field, by
Craighouse Quarry, NT605356, M.E. Braithwaite, 1995, herb. M.E.B. Ist record since 1931 and
only extant locality.

762/10.lon. MATTHIOLA LONGIPETALA *49, Caerns.: Several plants on waste ground, Port
Dinorwic, SH5267, G.& I. Battershall, 1996.

762/11.4. BARBAREA VERNA *89, E. Perth: New verge near Information Centre, Killicrankie,
NN96, M. McCallum Webster, 1976, E. Shingle, Tay Marshes, Perth, NO129217, O.M. Stewart,
1982, E. Ist and 2nd records.

62/12.3. RORIPPA ISLANDICA 41, Glam.: Small population of about 5 plants in muddy
disturbed ground in marshy grassland, Cwm Risca Meadow SSSI, SS8784, J.P. Woodman, 1996,
conf. A.O. Chater. 2nd record.

62/12.4. RORIPPA PALUSTRIS 49, Caerns.: Shore of Coedty Reservoir above Dolgarrog,
SH7566, R. Lewis, 1996. 2nd record.

562 PLANT RECORDS

62/12.5, RORIPPA SYLVESTRIS 93, N. Aberdeen: Garden of old farm, Old Meldrum,
NJ823279, D. Elston, 1996, conf. D. Welch. 2nd record.

62/14.6. CARDAMINE IMPATIENS 50, Denbs.: Shady deciduous wood, Pandy, SJ1935, J.A.
Green, 1996. 2nd post-1930 record.

762/16.1. AUBRIETA DELTOIDEA *50, Denbs.: Limestone rocks, Mold, SJ199627, J. Phillips,
1997. *81, Berwicks.: Established on town walls, Lauder, NT530476, M.E. Braithwaite, 1997.

762/2.1. DESCURAINIA SOPHIA *46, Cards.: Weed in garden of Tawelfan, Penrhyn-coch,
SN641840, A.G. Bates, 1983, NMW, det. A.O. Chater.

62/21.4. DRABA MURALIS 8, S. Wilts.: In chippings at base of stone wall, Stratford Tony,
SU091266, E.J.B. & P.M. Rollinson, 1994. 1st record for 60 years. +81, Berwicks.: Established
on dry bank, Grantshouse Quarry, NT/810652, M.E. Braithwaite, 1996. 2nd record.

62/22.1. EROPHILA MAJUSCULA *3,S. Devon: A garden weed, Newton Ferrers, SX552480, R.
E.N. Smith, 1997, det. T.T. Elkingion. *37, Worcs.: Little Malvern Quarry, SO7641, P.G.
Garner, 1991, conf. T.T. Elkington. *42, Brecs.: Limestone outcrop, Ystradfellte, SN915134, M.
Porter, 1997.

62/22.3. EROPHILA GLABRESCENS 3, S. Devon: Forword Point, SX9049, R.E.N. Smith, 1993,
det. T.T. Elkington, 2nd record. *37, Worcs.: West Malvern, SO7644, P.G. Garner, 1990, conf.
T.T. Elkington.

62/23.3. COCHLEARIA OFFICINALIS *+42, Brecs.: Roadside about 2:‘5km NW of Erwood,
S0075444, R.G. Woods, 1994. Ist record. This record replaces that for Cochlearia danica in
Watsonia 20: 423 (1995)

+62/24.1. CAMELINA SATIVA 51, Flints.: “Y Fedwen Arian’, Cilcain, SJ1865, G. Harvey,
1996, det. T.C.G. Rich. Ist post-1930 record

62/28.4. THLASPI CAERULESCENS *89, E. Perth: Four plants only on steep, scree-strewn,
craggy S-facing slope, across Lochsie Burn from Dalmunzie Hotel, Glen Taitneach, NO094714,
D. Lloyd-Thomas & M. Sutton, 1992.

62/30.5. LEPIDIUM RUDERALE 46, Cards.: Minute pavement flowerbed, Llanrhystud,
SN538697, A.O. Chater, 1991, herb. A.O.C. Ist record since 1941.

62/33.1. DIPLOTAXIS TENUIFOLIA *46, Cards.: Waste ground 500m SE of Aberystwyth
Station, SN588811, A.O. Chater, 1997. *91, Kincardines.: Disturbed ground at abandoned
railway yard, Crathes, NO7496, D. Welch, 1996, ABD.

+62/34.4. BRASSICA JUNCEA *51, Flints.: Garden weed, “Gwylfa’, Licswm, SJ1771, G.
Wynne, 1996, det. T.C.G. Rich. Possibly from bird-seed.

+62/38.1. HIRSCHFELDIA INCANA 42, Brecs.: Road verge, Talgarth, SO153337, M. Porter,
1997. 2nd record.

62/41.1. CRAMBE MARITIMA *91, Kincardines.: Coastal cliffs, Stonehaven, NO887874, V.
McAdam & U. Urquhart, 1988.

62/42.1b. RAPHANUS RAPHANISTRUM subsp. MARITIMUS 46, Cards.: Rank vegetation on edge
of Ystwyth estuary, Aberystwyth, SN579806, S.P. Chambers & P. Amis, 1996, NMW, conf. A.O.
Chater. 2nd record.

+62/42.sat. RAPHANUS SATIVUS —_*S, Channel Islands: In field near derelict vegetable garden,
La Rondellerie, Sark, WV460763, M.H. Marsden, 1983, JSY.

762/ERU.ves. ERUCA VESICARIA subsp. SATIVA *44, Carms.: Garden, below bird table,
Llwynhendy, SS537993, I.K. Morgan, 1993. 51, Flints.: Disturbed ground, New Brighton, near
Mold, $J2564, J. Phillips, 1996. 2nd record.

63/1.3. RESEDA LUTEA *91, Kincardines.: Roadside, Crathes, NO7496, D Welch, 1996.

+65/12.arb. ERICA ARBOREA *17, Surrey: One fair sized shrub in hedgerow by footpath,
Brook, SU931380, J.F. Leslie, K. Page & J.E. Smith, 1992. Parent visible in garden behind high
wall opposite.

765/2.2. RHODODENDRON LUTEUM *47, Monts.: Steep side of Afon Hirddu and roadside,
Lake Vyrnwy, SH974213, J. Clarke, 1997. | *50, Denbs.: Wet areas around ponds in woodland,
Coed Cerrig-y-Wyallt, near Tal y Cafn, SH786702, R. Lewis, 1988, NMW.

PLANT RECORDS 563

+65/8.1. GAULTHERIA SHALLON *49, Caerns.: Naturalised on verge, Llanystumdwy, SH4738,
BSBI meeting, 1997. *91, Kincardines.: Policies of mansion, Glen Dye, NO649862, D. Welch,
1997.

*65/8.2. GAULTHERIA PROCUMBENS *46, Cards.: Under oaks in estate grounds S of
Trawsgoed mansion, SN671730, A.O. Chater & R.G. Woods, 1989.

66/1.3. PYROLA ROTUNDIFOLIA *50, Denbs.: Several patches over 50m in hazel-birch-ash
wood on waste from limestone quarry, Minera, near Wrexham, SJ264519, M. Stapley, 1989.

769/1.1xpru.; PRIMULA VULGARIS x P. x PRUHONICENSIS *41, Glam.: Hybrid swarms in
chapel graveyard with both parents, Rhyd y Fron, near Pontardawe, SN713059, R.G. & J.D.
Woods, 1997.

69/2.1. HOTTONIA PALUSTRIS *71, Man: Boundary ditch on maritime heath, Ayres,
NX432026, M. Devereau, 1988, L.S. Garrard, 1994. Means of arrival unknown but conceivably
bird carried. Size of colony fluctuates.

+69/3.1. CYCLAMEN HEDERIFOLIUM 46, Cards.: Shaded bank of Afon Einon by Ynys-hir
bridge, Eglwys-fach, SN684957, E. Greenwood, & P. & W.M. Condry, 1995. 2nd record. *47,
Monts.: Bank of Bachan Brook adjoining roadside, Aberbechan, SO1393, C.A. Small, 1989. Ist
localised record.

69/8.1. SAMOLUS VALERANDI “*12, N. Hants.: Shallow pool, Longmoor Airstrip, SU8031, N.
A. Sanderson, 1994.

+71/DEU/sca. DEUTZIA SCABRA *49. Caerns.: Edge of woodland W side of B5106, Pont
Dolgarrog, SH7766, R. Lewis, 1996, NMW. Ist Welsh record.

7+71/HYD.mac. HYDRANGEA MACROPHYLLA = _*17, Surrey: One shrub, pale blue, in woodland,
Hindhead, SU881360, J.B.S. Hodge, 1993. Not obviously planted — possibly self-sown.

+72/2.4. RIBES SANGUINEUM *50, Denbs.: Canal bank, Trevor, $J256413, J.A. Green, 1997.

773/1.1. CRASSULA TILLAEA *45, Pembs.: Abundant in compact soil amongst car park blocks,
Manorbier car park, SS0697, J.W. Donovan, 1996.

+73/1.3. CRASSULA HELMSI *44, Carms.: Abundant in pond, Wern Fendigaid, SN638455, I.
K. Morgan, 1987. *45, Pembs.: Small patch with abundant Lemna trisulca in small pond on
slope in woodland, Pwll Dyfrig, Glyn-y-mel, Lower Fishguard, SM967371, S.B. Evans et al.,
1997. *67,S. Northumb.: Blackworth, NZ37, S. Lowe, 1990. East Cramlington, NZ292759, M.
Teffner, 1992. Ist and 2nd records. *73, Kircudbrights.: Newly dug pools in unimproved
meadow, Craig, NX681755, O.M. Stewart, 1995. *77, Lanarks.: The Moat, Golf Course, Cadder
in Glasgow, NS6072, P. Macpherson, 1996, herb. P.M.

+73/3/1. SEMPERVIVUM TECTORUM *50, Denbs.: Roadside wall, Llangwm, SH9745, A.O.
Chater, 1996. Chapel wall, Llangwm, SH9645, S.P. Chambers, 1996. Ist and 2nd records.

73/5.18. SEDUM VILLOSUM 73, Kirkcudbrights.: By a small burn, Cairnsmore of Carsphairn,
NX584980, D. Hawkes, 1997. Ist record since 1854.

773/5.4. SEDUM SPECTABILE *11, S. Hants.: Under bramble near car park along track,
Keyhaven, SZ308917, P.A. Budd, 1997.

+74/4.1. DARMERA PELTATA %*58, Cheshire: Stream bank, Adlington, $J912807, J.H. Clarke,
1996, conf. G.M. Kay. Wooded streambank, Disley, SJ969844, G.M. Kay, 1997. Ist and 2nd
records.

774/5.2. SAXIFRAGA CYMBALARIA *46, Cards.: Wall, Alltyrodyn, Capel Dewi, SN450443, M.
L. Lewis, c.1921, ABS, det. A.O. Chater & A.D.Q. Agnew.

+74/5.9. SAXIFRAGA SPATHULARIS *46, Cards.: Mixed woodland, Plas Einion, Furnace,
SN684948, A.O. Chater, 1991.
74/5.9x10. SAXIFRAGA SPATHULARIS x S. HIRSUTA (S. x POLITA) *93, N. Aberdeen:

Woodland by lochan, Cortes, NK003593, D. Welch, 1997, herb. D.W.

74/5.19. SAXIFRAGA TRIDACTYLITES *94, Banffs.: Disused airfield, Boyndie, NJ615646, J.
Edelsten, 1993, E, ABD, det. D. Parker.

+74/7.1. TOLMIEA MENZIESII *67, S. Northumb.: Cross roads, Oakerland near Hexham,
NY942624, R.M. Burton, 1994, herb. G.A. Swan. *68, Cheviot: On bank of streamlet in part
shade, Ingram, NU018162, G.A. & M. Swan, 1995, herb. G.A.S.

564 PLANT RECORDS

+74/8.1.+ TELLIMA GRANDIFLORA *31, Flints.: Probable garden escape in woodland, near
houses, Wepre Country Park, Connah’s Quay, SJ291683, J. Phillips, 1997. | *67, S. Northumb.:
Partly shaded on N bank of River Wansbeck, near Morpeth, NZ215865, H.E. Ellis, 1997, herb. G.
A. Swan. *89, E. Perth: Kinnaird Burn Den, NN952585, R.E. Youngman, 1996.

775/2.1. PHYSOCARPUS OPULIFOLIUS *95, Moray: Several bushes in former garden, rough
area on edge of forest, Darnaway Forest, near Forres, NJO06568, I.P. Green, 1996, conf. E.J.
Clement.

+75/3.1. SPIRAEA SALICIFOLIA *35, Mons.: Railside bank, Cadicot Pill, ST48, T.G. Evans,
1985, NMW.

+75/3/1x2. SPIRAEA SALICIFOLIA x S. ALBA (S. x ROSALBA) *49, Caerns.: Hedgerow, near
Ysbyty Ifan, SH8348, G. & I. Battershall, 1996. *77, Lanarks.: Near Abington, NS92, P.
Macpherson, 1997, herb. P.M., det. A.J. Silverside.

175/3/2x3. SPIRAEA ALBA x S. DOUGLASII (S.x BILLARDII) *46, Cards.: Hedges by road at N
end of Ponthirwaun, SN262452, A.O. Chater, 1980, det. A.J. Silverside. Widely planted and
naturalised in hedges.

+75/3.3a. SPIRAEA DOUGLASII subsp. DOUGLASII *44. Carms.: Railway cutting, Burry Port,
SN444008, G. Hutchinson, 1988, NMW. *46, Cards.: Scrub, Ty n-y-garth, Cwm Einion,
SN691945, A.O. Chater, 1991, NMW. Roadside hedges, 100m S of Pwll-y-whil, Pontgarreg,
SN352540, A.O. Chater, 1992, NMW. Ist and 2nd records. *51, Flints.: Waste ground on site
of former garden, Sealand, $J3568, G. Wynne, 1996, det. G. Hutchinson. *81, Berwicks.: Two
large colonies established on roadside, Blyth road end, NT592489, M.E. Braithwaite, 1997, herb.
M.E.B.

+75/3.canxtri. SPIRAEA CANTONIENSIS x S. TRILOBATA (S. x VANHOUTTEI) *46, Cards.:
Roadside hedges SW of Neuadd-lwyd chapel, 3-5km SSE of Aberaeron, SN473595, A.O. Chater,
1997, NMW.

+75/4.1. ARUNCUS DIOICUS *42, Brecs.: Probably a garden outcast established on river bank,
Sennybridge, SN9228, R.G. Woods, 1997.

75/6.1. FILIPENDULA VULGARIS *41, Glam.: Limestone grassland on cliff-top at Oxwich
Point, far from roads and houses and apparently native, SS507848, B. Pawson, 1997.
+75/8.2. RUBUS TRICOLOR *11, S. Hants.: Footpath by streambank, Beckton Bunny,

SZ243935, V. Scott, 1997, herb. R.P. Bowman.

75/8.6. RUBUS BRIGGSIANUS *46, Cards.: One bush only in roadside hedgebank 250m ESE of
Penwaun, St Dogmaels, SN157442, D.E. Allen & A.O. Chater,1995, BM, conf. A. Newton.

75/8.7x321. RUBUS IDAEUS x R. CAESIUS *42, Brecs.: Hedge, Tretower, SO188211, M.
Porter, 1997, herb. M.P., conf. A. Newton.

175/8.9. RUBUS SPECTABILIS *73, Kirkcudbrights.: Wood edge and roadside, Damhead,
Barnbarroch, NX8456, O.M. Stewart & A. White, 1996.

+75/8.10. RUBUS LOGANOBACCUS *12, N. Hants.: Waste ground, Woolmer Forest, near
Borden, SU798367, A. Brewis, 1997, herb. A.B.

+75/8.11. RUBUS COCKBURNIANUS _ 11, S. Hants.: Grassy roadside by gate into wood, nearly
opposite cottage, A35 Southampton road, Mallard Wood, Lyndhurst, SU319093, R.P. Bowman,
1997, herb. R.P.B. 2nd record.

75/8.20. RUBUS FISSUS *8, S. Wilts.: Wardour [Castle], near Tisbury, ST9226, F.A. Rogers,
1895, BM, det. W.M. Rogers, conf. D.E. Allen. [12, N. Hants.: the BM vouchers for the record
in Journal of Botany 45: 72 (1907) are R. nessensis, det. D.E. Allen.] *27, E. Norfolk: North
Walsham to Westwick, TG2727, E.F. Linton, 1893, BM, conf. D.E. Allen.

75/8.23 RUBUS NESSENSIS *73, Kirkcudbrights.: Roadside through Dalbeattie Forest, NX85,
J.R.I. Wood, 1972, BM, comm. D.E. Allen.

75/8.27. RUBUS PLICATUS *37, Worcs.: Wooded heath, Devil’s Spittleful, Kidderminster,
SO8075, A. Newton et at., 1989. | *H27, W. Mayo: Between Keel and Doogort, Achill Island,
Fc.6507, J. Roffey, 1911, BM, conf. D.E. Allen.

75/8.32. RUBUS VIGOROSUS *37, Worcs.: Hartlebury Common, Stourport-on-Severn,
SO8270, A. Newton, 1987.

PLANT RECORDS 565

75/8.34. RUBUS ALBIONIS *35, Mons.: Wooded path edge, Mescoed Mawr, ST2789, T.G.
Evans, M. Porter & R.D. Randall, 1996.

+75/8.51. RUBUS ERRABUNDUS' _*54, N. Lincs.: Lane near Walesby railway crossing, TF1091,
F.A. Lees, 1877, BM, det. D.E. Allen.

75/8.52. RUBUS GRATUS *64, Mid-W. Yorks.: Bank of Claude’s Clough, Brogden,
SD848419, D.P. Earl, 1997.

75/8.53. RUBUS HESPERIUS *H10, N. Tipperary: Wooded bogland, near Slevoir Bay, Lough
Derg, M8904, D.E. Allen, 1988, BM, conf. A. Newton.

775/8.55. RUBUS LACINIATUS *78, Peebless.: Wallace Hill Wood plantation, NT3136, A.
Buckham, 1987, det. D.J. McCosh.

75/8.58. RUBUS LEPTOTHYRSOS *78, Peebless.: Quarry, near Innerleithen, NT3336, R.
Learmouth, 1984, det. A. Newton.

75/8.61. RUBUS LUDENSIS *46, Cards.: Roadside hedgebank 100m S of Waunwhiod, St
Dogmaels, SN145450, D.E. Allen & A.O. Chater, 1995, BM, conf. A. Newton.

75/8.62. RUBUS MACROPHYLLUS *37, Worcs.: Edge of deciduous plantation in Ockeridge
Wood, SO7962; By woodland ride in Monk Wood, SO8060; both R. Maskew, A. Newton & M.
Porter, 1995. 1st and 2nd records.

75/8.76. RUBUS PYRAMIDALIS *37, Worcs.: Trackside in coniferous woodland, Kyre near
Tenbury Wells, SO06264, R. Maskew & A. Newton, 1992.

75/8.92. RUBUS BOUDICCAE *5, S. Somerset: Simonsbath, SS7739, E.S. Marshall, 1918 (as R.
pulcherrimus), E, det. D.E. Allen. *40, Salop: Diddlebury Common, SO5085, A. Ley, 1909,
BM, det. D.E. Allen. *46, Cards.: Hedgebank on sandy soil 200m N of Warren Farm, Penparc,
SN201477, D.E. Allen & A.O. Chater, 1996, BM, conf. A. Newton. *48, Merioneth: Dolgellau,
SH71, W.C. Barton & H.J. Riddelsdell, 1923, BM. det. D.E. Allen. *100, Clyde Is.: Lochranza,
Isle of Arran, NR9350, M.L. Wedgwood, 1926, BM, E, det D.E. Allen. *101, Kintyre: Skipness
Castle, NR908577, A.R. Church, 1997; sides of B842 in Saddell area, NR786314, G.H.
Ballantyne, 1997, BM, conf. D.E. Allen. *H16, W. Galway: Near Aughnanure Castle,
Oughterard, M1543, J. Roffey, 1911 (as R. pyramidalis?), det. D.E. Allen. *H27, W. Mayo:
Beside Lough Beltra, M0898, D.E. Allen & G. Sharkey, 1992, BM. *H35, W. Donegal: Wood
by roadside, Ardnamona, G963857, M. Dowlen & P. Hodson, 1993, BM, det. D.E. Allen.
*H38, Co. Down: Meenan, Aghaderg parish, J0939, W.M. Rogers, 1901, BM, det. D.E. Allen.
Field next to Rowallane Gardens, Saintfield, J414579, A. Newton & P. Hackney, 1985, BEL, det.
D.E. Allen. Ist and 2nd records. *H40, Co. Londonderry: Hedge, station yard, Limavady
Junction, C637255, P. Hackney, 1978, BEL, det. D.E. Allen.

75/8.93. RUBUS CARDIOPHYLLUS *100, Clyde Is.: One bush on rocky ground by shore,
Kingscross Point, Arran, NS056282, D.E. Allen & A.R. Church 1997, BM.

75/8.102. RUBUS ELEGANTISPINOSUS *78, Peebless.: Roadside, A701 S of Broughton, NT13,
G.H. Ballantyne, 1986. Possibly planted.

+75/8.102. RUBUS ELEGANTISPINOSUS *9, Dorset: Wessex Ridgeway through woods, Iwerne
Minster, ST8813, D.E. Allen, 1996, BM, conf. A. Newton.

75/8.112. RUBUS NEMORALIS *78, Peebless.: Roadside, by junction of A703 and A72,
Peebles, NT2540, G.H. Ballantyne, 1982.

75/8.113. RUBUS PAMPINOSUS *1, W. Cornwall: Among bracken below railway bridge,
Ponsanooth, SW7737, W.C. Barton & F. Rilston, 1930, BM, det. D.E. Allen, conf. A. Newton.
*46, Cards.: Colony at edge of larch plantation 100m W of Ty’n-y-cewm, Cwm Einion, SN699943,
D.E. Allen & A.O. Chater, 1995, BM, conf. A. Newton. *47, Monts.: Llyfnant Valley, SN79,
W.C. Barton, 1923, BM, det. D.E. Allen, conf. A. Newton,

75/8.118. RUBUS PROLONGATUS *37, Worcs.: Trackside in Wissett’s Wood near Bayton,
S06772, R. Maskew & A. Newton, 1993.

75/8.122. RUBUS RUBRITINCTUS *8, S. Wilts.: East Copse, between Redlynch and
Whiteparish, SU226219, Wiltshire Botanical Society, 1996, det. D.E. Allen. *64, Mid-W.
Yorks.: Roadside, Gisburn Forest, SD7356, A. Newton, 1997.

566 PLANT RECORDS

75/8.125. RUBUS SUBINERMOIDES *8, S. Wilts.: East Copse, between Redlynch and
Whiteparish, SU226219, Wiltshire Botanical Society, 1996, det. D.E. Allen. *37, Worcs.: Scrub
on limestone grassland, Tinker’s Coppice, Horsham, $O7357, J.J. Day, R. Maskew & A. Newton,
1992.

75/8.132. RUBUS SPRENGELI] *37, Worcs.: Trench Wood, Oddingley, SO9258, A. Newton et
al., 1988.

175/8.134. RUBUS ARMENIACUS “HIMALAYAN GIANT’ = *46, Cards.: Slope above road 200m S
of Cardigan Bridge, SN178456, D.E. Allen & A.O. Chater, 1996, NMW. *H31, Co. Louth:
Clogher Head, 01785, M. Campbell, 1991, BM, det. D.E. Allen.

75/8.140. RUBUS ROSSENSIS *8, S. Wilts.: Roadside between Redlynch and Whiteparish,
SU221218, Wiltshire Botanical Society, 1996, det. D.E. Allen.

75/8.140x 142. RUBUS ROSSENSIS x R. ULMIFOLIUS *37, Worcs.: Scrub in woodland clearing,
Papermill Coppice, near Alfrick, SO7451, P.G. Garner, R. Maskew & A. Newton, 1993.

75/8.141. RUBUS STENOPETALUS *9, Dorset: Two bushes on Wessex Ridgeway through
woods, Iwerne Minster, ST8813, D.E. Allen, 1996, BM, conf. A. Newton.

75/8.142x321. RUBUS ULMIFOLIUS x R. CAESIUS *37, Worcs.: Hedge, New Mill Bridge,
Shelsley Beauchamp, SO7262, J.J. Day, R. Maskew & A. Newton, 1992.

75/8.143. RUBUS WINTERI *33, E. Gloucs.: Hailes to Farmcote, SP0529, M.A. Rogers, 1913,
BM, det. D.E. Allen, conf. A. Newton. *35, Mons.: Alder carr, Henllys Fen, ST2692, T.G.
Evans, 1996, det. A. Newton.

75/8.144. RUBUS ADSCITUS *17, Surrey: Queen’s Cottage grounds, Royal Botanic Gardens,
Kew, SU1776, H.J. Riddelsdell, 1929, BM, conf. D.E. Allen. *37, Worcs.: Hagley Wood,
Hayley Green, SO9381, A. Newton & M. Porter, 1993.

75/8.150. RUBUS CRINIGER *9, Dorset: Wessex Ridgeway near Iwerne Minster, ST8813, D.E.
Allen, 1996, BM, conf. A. Newton.

75/8.158. RUBUS ORBUS *9, Dorset: Conifer plantation, France Firs, Blandford Forum,
ST880095, D.E. Allen, 1996, BM, conf. A. Newton.

75/8.161. RUBUS SURREJANUS *35, Mons.: Woodland edge, N of Gray Hill, ST4293, R.D.
Randall, 1993. *100, Clyde Is.: Bushy waste ground, Glen Cloy, Brodick, Arran, NSO11358, D.
E. Allen, 1997, BM.

75/8.163. RUBUS VESTITUS *78, Peebless.: Roadside, E of A72, Peebles, NT2540, G.H.
Ballantyne, 1988.

75/8.167. RUBUS EGREGIUS = *17, Surrey: Tooting Common, TQ2972, J. Roffey, 1910, BM,
det. R.D. Randall, conf. A. Newton.

75/8.169. RUBUS FUSCICORTEX *46, Cards.: Llyfnant Valley, SN79 or SN69, W.C. Barton,
1923, BM, conf. D.E. Allen.

75/8.171. RUBUS MUCRONATIFORMIS *46, Cards.: Roadside hedgebank and field margin,
Warren Farm, Penparc, SN201477, D.E. Allen & A.O. Chater, 1997, BM, conf. A. Newton.

75/8.174. RUBUS WIRRALENSIS *37, Worcs.: Scrub on site of disused army camp, Burlish
Top, Kidderminster, SO8073, A. Newton et al., 1989.

75/8.176. RUBUS AEQUALIDENS *33, E. Gloucs.: Whiteshill, near Stroud, SO8407, E.M. Day,
1914, BM, det. D.E. Allen, conf. A. Newton. *34, W. Gloucs.: Berry Hill, SO5612, E.M. Day,
1913, BM, det. D.E. Allen. *46, Cards.: Track verge in conifers, Long Wood, Llangybi,
SN610515, D.E. Allen & A.O. Chater, 1996, conf. A. Newton.

75/8.180. RUBUS DIVERSUS *8, S. Wilts.: Coniferised ancient woodland, Barnell Copse
portion of Whiteparish Common, SU250218, Wiltshire Botanical Society, 1996, BM, det. D.E.
Allen, conf. A. Newton.

75/8.184. RUBUS GLAREOSUS *35, Mons.: Wooded path edge, Mescoed Mawr, ST2789, T.G.
Evans, M. Porter & R.D. Randall, 1996.

75/8.185. RUBUS GRIFFITHIANUS *H30, Co. Cavan: Lane on E side of Slieve Glah, H460002,
D.E. Allen & P. Reilly, 1987, BM, conf. A. Newton.

PLANT RECORDS 567

75/8.188. RUBUS HETEROBELUS *4, N. Devon: Track in forestry, Powler’s Piece near
Melbary, $S365190, R.W. Gould, 1997, herb. R.W.G, det. A. Newton.

75/8.195. RUBUS MOYLEI *8, S. Wilts.: West Dean, SU257273. W.C. Barton, 1941, BM, det.
D.E. Allen.

75/8.197. RUBUS NORVICENSIS *21, Middx.: Wooded enclosure, Holland park, London,
TQ248800, D.E. Allen, 1997, BM, conf. A. Newton. *46, Cards.: Woodland by bridge 300m
SSW of Capel Dewi church, SN451421, D.E. Allen & A.O. Chater, 1996, BM, conf. A. Newton.
lst Welsh record.

75/8.200. RUBUS TRICHODES *11, S. Hants.: Gorse hedge by Bedhampton hypermarket,
Havant, SU679077, D.E. Allen, 1992, BM, det. A. Newton. *23, Oxon: Chinnor Hill, SU7699,
W.O. Focke & W.M. Rogers, 1894, BM, det. D.E. Allen & R.D. Randall, conf. A. Newton.

75/8.203. RUBUS ADAMSII *37, Worcs.: Trackside in woodland, Pitcher Oak Wood, Redditch,
SP0267, R. Maskew, A. Newton & W.A. Thompson, 1996.

75/8.206. RUBUS ANISACANTHOS *97, Westerness: Streamside, Resipol, Sunart, NM76, S.M.
Macvicar, 1897-8, BM, det. A. Newton. *H21, Co. Dublin: Sides of Bohernabreena larger
reservoir, 00923, D.E. Allen, 1987, BM, conf. A. Newton.

75/8.207. RUBUS BILOENSIS *6, N. Somerset: Rough pasture near Stourton, ST7633, R.P.
Murray, 1894, BM, det. A. Newton & M. Porter.

75/8.209. RUBUS DENTATIFOLIUS *37, Worcs.: Clent, SO9279, A. Newton & M. Porter, 1993.

75/8.211. RUBUS DREJERI *100, Clyde Is.: Roadside 2km N of Dougarie, Arran, NR871389,
A.R. Church, 1997, BM, conf. G.H. Ballantyne.

75/8.214. RUBUS FORMIDABILIS *12, N. Hants.: Heathy wood margin, Wheatsheaf Common,
Liphook, SU835305, D.E. Allen, 1996, BM, conf. A. Newton.

75/8.216. RUBUS HIBERNICUS *9,. Dorset: Beech-—fir plantation, Okeford Hill, ST8109, D.E.
Allen, 1996, BM, conf. A. Newton.

75/8.217. RUBUS INFESTUS *78, Peebless.: Lay-by on A72, Woodend, NT3038, G.H.
Ballantyne, 1981.

75/8.218. RUBUS LEYANUS *37, Worcs.: Scrub in farmyard, Stoke Bliss near Tenbury Wells,
S06463, R. Maskew & A. Newton, 1992.

75/8.224. RUBUS BLOXAMII *37, Worcs.: Trackside in woodland, Pepper Wood near
Broadheath, SO9374, J.J. Day, R. Maskew & A. Newton, 1991.

75/8.225. RUBUS BOTRYEROS *37, Worcs.: Trackside in woodland clearing, Ramscombe
Coppice, Great Witley, SO7666, J.J. Day, R. Maskew & A. Newton, 1990.

75/8.227. RUBUS CANTIANUS *17, Surrey: Bridleway through woodland, Croham Hurst,
Croydon, TQ343629, D.E. Allen, 1997, BM, conf. A. Newton.

75/8.231. RUBUS ECHINATOIDES *37, Worcs.: Edge of woodland, Black Meadow, Chaddesley
Woods, SO9173, J.J. Day, R. Maskew & A. Newton, 1991.

75/8.239. RUBUS INSECTIFOLIUS *37, Worcs.: Clent, SO9279, A. Newton & M. Porter, 1993.

75/8.243. RUBUS MALVERNICUS *46, Cards.: Replanted conifer plantation, Allt Maestir,
3-5km NW of Lampeter, SN546503, D.E. Allen & A.O. Chater, 1997, BM, conf. A. Newton.

75/8.245. RUBUS PALLIDUS *37, Worcs.: Edge of woodland, Black Meadow, Chaddesley
Woods, SO9173; Trackside in woodland, Pepper Wood near Bournheath, $09374; both J.J. Day,
R. Maskew & A. Newton, 1991. Ist and 2nd records.

75/8.249. RUBUS RADULA __ *78, Peebless.: W side of Walkerburn, NT3537, G.H. Ballantyne,
1981. *100, Clyde Is.: Trackside, Kilmory, Arran, NR959210, A.R. Church & G.H. Ballantyne,
1996; A.R. Church & D.E. Allen, 1997.

75/8.253. RUBUS RUDIS *37, Worcs.: Scrub in woodland clearing, Papermill Coppice near
Alfrick, SO7451, P.G. Garner, R. Maskew & A. Newton, 1993. *64, Mid-W. Yorks.: Edge of
woodland, Gisburn, SD821496, D.P. Earl & J. Buckley-Earl, 1997. New northern limit?

75/8.257. RUBUS SECTIRAMUS *11, S. Hants.: Upper Grenville Copse, Hambledon,
S$U635172, D.E. Allen, 1996. *12, N. Hants.: Wood margin, Hartley Wood Common, Hartley
Wespall, SU692581, D.E. Allen, 1994. *13, W. Sussex: Oakwood by sea, Chalkdock Point, near
West Itchenor, SU790013, D.E. Allen, 1996. All BM, conf. A. Newton.

568 PLANT RECORDS

75/8.264. RUBUS ATREBATUM *11, S. Hants.: Margin of chestnut plantation, Hall Copse, near
Romsey, SU329209, D.E. Allen, 1981, 1989, BM, conf. A. Newton.

75/8.266. RUBUS BERCHERIENSIS *46, Cards.: Bank below road, Cwm Degwell, 200m S of St
Dogmaels Abbey, SN163454, D.E. Allen & A.O. Chater, 1995, BM, conf. A. Newton.

75/8.268. RUBUS DASYPHYLLUS *78, Peebless.: Old railway, Peebles, NT253401, G.H.
Ballantyne, 1988.

75/8.270. RUBUS HYLOCHARIS *50, Denbs.: Roadside, Padog, SH8351, G. Battershall, 1996.
[100, Clyde Is.: 1965 Arran specimen det. E.S. Edees in NMW is R. pictorum, det. D.E.Allen.]
*101, Kintyre: Car park, Skipness, NR903579, A.R. Church, 1997, BM, det. D.E. Allen, conf. A.
Newton.

75/8.273. RUBUS MARSHALLII *9, Dorset: Heath and conifer plantation, Black Down, ST6087
& 6187, D.E. Allen, 1997, BM, conf. A. Newton.

75/8.275. RUBUS ASPERIDENS *37, Worcs.: Scrub in coniferous plantation, Island Coppice,
Kyre near Tenbury Wells, S06464, R. Maskew & A. Newton, 1992.

75/8.276. RUBUS MURRAYI = _*37, Worcs.: Trackside in deciduous plantation, Ellbatch Wood
near Abberley, SO7366, J.J. Day, R. Maskew & A. Newton, 1990.

75/8.278. RUBUS NEWBRIDGENSIS [12, N. Hants.: Delete sole record in Fl. Hants., 158
(1996); further study has shown population is of a different but unnamed species, D.E. Allen & A.
Newton. |

75/8.284. RUBUS RILSTONEI *46, Cards.: Roadside hedgebanks 250m ESE of Penwaun, St
Dogmaels, SN157442, D.E. Allen & A.O. Chater, 1995, BM, conf. A. Newton.

075/8.286. RUBUS SCABRIPES [17, Surrey: Delete record published in Watsonia 20: 291
(1995).]

75/8.299. RUBUS PEDEMONTANUS *37, Worcs.: Trackside in coniferous woodland,
Ribbesford Wood near Bewdley, SO7872, R. Maskew, A. Newton & W.A. Thompson, 1997.

75/8.307. RUBUS CONJUNGENS _*S, Channel Is.: Bouley Bay, Jersey, WV6654, W.M. Rogers,
1897, BM, conf. A. Newton.

75/8.308. RUBUS EBORACENSIS *78, Peebless.: Roadside, Nether Harsburgh, NT304394, D.J.
McCosh, 1983, det. D.E. Allen.

75/8.310. RUBUS HEBRIDENSIS *94, Banffs.: Road junction, Dufftown, NJ323392, D. Welch,
1992, ABD, det. A. Newton.

75/8.312. RUBUS LATIFOLIUS *71, Man: Open hillside, The Creggans, Knockaloe, SC238818,
D.E. Allen, 1975, BM, conf. A. Newton.

75/8.314. RUBUS PICTORUM *35, Mons.: Hedge, NW of White House, SO4222, M. Porter &
R.D. Randall, 1993

75/8.315. RUBUS PRUINOSUS *46, Cards.: Disused railway embankment, Trawsgoed,
SN663733, D.E. Allen & A.O. Chater, 1997.
75/8.317. RUBUS TENUIARMATUS 42, Brecs.: Hedge and lane verge, Digedi valley near

Llanigon, SO2138, M. Porter, 1997, herb. M.P. 2nd record.

75/8.319. RUBUS TUBERCULATUS *100, Clyde Is.: Edge of drive, Whiting Bay, Arran,
NS046254, A.R. Church & A. Smith, 1997, BM, conf. D.E. Allen.

75/8.arl. RUBUS ARICONIENSIS *35, Mons.: Woodland, Kingswood, SO4612, M. Porter & R.
D. Randall, 1993.

75/8.isc. RUBUS ISCANUS *35, Mons.: Woodland, Kingswood, SO4612, M. Porter & R.D.
Randall, 1993.

75/8.per. RUBUS PERCRISPUS *46, Cards.: Laneside hedge 300m WSW of Bryneithyn,
Llanfarian, SN579780, D.E. Allen & A.O. Chater, 1997, BM.

75/8.vag. RUBUS VAGENSIS *35, Mons.: Woodland, Kingswood, $O4612, M. Porter & R.D.
Randall, 1993. *37, Worcs.: Hay Slad, West Malvern, SO7644, P.G. Garner, R. Maskew & A.
Newton, 1993. *42, Brecs.: Woodland, Park Wood, Talgarth, SO168338, M. Porter, 1991, herb.
M.P.

PLANT RECORDS 569

75/8.ven. RUBUS VENETORUM *104, N. Ebudes: Laig Bay, Isle of Eigg, NM4802, S.M.
Macvicar, 1895, BM, det. D.E. Allen.

75/9.5. POTENTILLA ARGENTEA 95, Moray: One plant on waste ground, Rafford, NJ053551,
P.R. Green, 1996. Ist record for well over 100 years.

75/9.13x14. POTENTILLA ERECTA x P. ANGLICA (P. x SUBERECTA) *37, Worcs.: Trackside
verge, Lickey Hills, SO9875, J.J. Day & R. Maskew, 1992, conf. B. Harold. +*77, Lanarks.:
Junction between grassy bank and gravelly shore, Hillend Reservoir, NS8467, P. Macpherson,
1996, herb. P.M., det. B. Harold.

75/9.14. POTENTILLA ANGLICA *95, Moray: Woodland ride, Quarry Wood, W of Elgin,
NJ174631, P.R. Green, 1997. Ist confirmed record; possibly brought in with forestry vehicles.

75/9.14x15. POTENTILLA ANGLICA x P. REPTANS (P. x MIXTA) — *37, Worcs.: Rough grassland,
Clows Top, SO7171, R. Maskew, 1994, det. B. Harold.

775/11.3. FRAGARIA x ANANASSA *49, Caerns.: Side of disused railway, Bontnewydd,
SH4757, G. Battershall & W. McCarthy, 1996. Waste ground, Trevor, SH3847, G. Battershall &
R. Lewis, 1996. Ist and 2nd records.

+75/11.chi. FRAGARIA CHILOENSIS *S, Channel Islands: Roadside bank, Rue La Rade outside
Beauvoir, Sark, WV465764, R.M. Veall, 1933, JSY, det. A. Chevalier. Established on bank until
site cleared about 1980. Portions of plants planted in garden and still present (1997)

75/15.2. AGRIMONIA PROCERA 61, S.E. Yorks.: Woodland, Hagg Wood, 0-8km E of
Dunnington, SE6852, M. Hammond, 1996. Only extant locality.

75/17.3. SANGUISORBA MINOR *91, Kincardines.: Disturbed ground at abandoned railway
yard, Crathes, NO7496, D. Welch, 1996, ABD.

+75/17.3b. SANGUISORBA MINOR subsp. MURICATA *50, Denbs.: Roadside near café,
Redbrook, SJ508412, K. Watson, 1991, NMW.

+75/18.4. ACAENA INERMIS *44, Carms.: Disturbed river shingle, Llangadog, SN706277, I.K.
Morgan & R.D. Pryce, 1987, NMW.

75/19.10b. ALCHEMILLA FILICAULIS subsp. VESTITA *46, Cards.: Flushed SW facing slope,
sheep-grazed pasture with boulders, Cwm Berwyn 800m WNW of Diffwys, SN726780, A.O.
Chater, 1988, NMW.

775/19.15. ALCHEMILLA MOLLIS 49, Caerns.: Grassy bank W side of B5106, between Lodge
Hotel and Bedal Inn, Talybont, SH7669, R. Lewis, 1996. Grassland near Elephant’s Cove, Great
Orme, SH7783, W. McCarthy, 1996. 2nd v.c. records. 50, Denbs.: Edge of car park, Llanrhaiadr
ym Mochnant, SJ1226, J.A. Green, 1996. Probable garden escape, 2nd record.

775/21.1. ROSA MULTIFLORA *11, S. Hants.: Thicket with R. canina, Linford Bottom,
Linford, SU179070, G.D. Field, 1997, det. A.L. Primavesi.

175/21.3. ROSA LUCIAE *41, Glam.: Scrub at edge of old colliery tip below Kenwood
Kennells, adjoining salt-marsh, W of Llanmorlais, Gower, SS526945, Q.O.N. Kay, 1995, herb. Q.
O.N.K. Ist Welsh record. Presumably introduced or a garden throw-out but appears established.

75/21.4x11. ROSA ARVENSIS x R. STYLOSA (R. x PSEUDORUSTICANA) *4, N. Devon: Lane
hedge, Sourton Quarry, SX521897, W.H. Tucker, 1997, conf. L.J. Margetts. Ist record under this
name.

75/21.4x12. ROSA ARVENSIS x R. CANINA (R. X VERTICILLACANTHA) 4, N. Devon: Climbing
through bushes by River Taw, North Wyke, SX6598, R.E.N. Smith, 1997, conf. A.L. Primavesi.
2nd record.

75/21.4x13b. ROSA ARVENSIS x R. CAESIA subsp. VOSAGIACA —_*37, Worcs.: Scrub on shore of
reservoir, Upper Bittell Reservoir, Cofton Hackett, SP0275, R. Maskew, 1995, conf. A.L.
Primavesi as hybrid with R. arvensis as female parent.

75/21.4x14. ROSA ARVENSIS x R. OBTUSIFOLIA (R. x ROUYANA) *37, Worcs.: Scrub at edge
of woodland, Brazier’s Coppice, Stoke Bliss near Tenbury Wells, S06464, R. Maskew, 1991, det.
as hybrid with R. obtusifolia as female parent.

75/21.4x16. ROSA ARVENSIS x R. SHERARDII *37, Worcs.: Hedge near Harpley, SO6860, R.
Maskew, 1995. Hedge near Tanner’s Brook, SW of Wyre Forest, SO7273, R. Maskew & W.A.
Thompson, 1995; both det. R.M. as hybrids with R. sherardii as female parent. Ist and 2nd
records.

570 PLANT RECORDS

75/21.4x19. ROSA ARVENSIS x R. MICRANTHA (R. x INELEGANS) *37, Worcs.: Woodland,
Coalpits Wood near Knightwick, $O7254, R. Maskew & W.A. Thompson, 1995, conf. A.L.
Primavesi as hybrid with R. micrantha as female parent.

75/21.5x17. ROSA PIMPINELLIFOLIA x R. MOLLIS (R. x SABINIT) *78, Peebless.: Innerleithen,
NT33, A. Craig-Christie, 1873, E, det. A. L. Primavesi.

75/21.5x18. ROSA PIMPINELLIFOLIA x R. RUBIGINOSA (R. x BITURIGENSIS) 93, N. Aberdeen:
Roadside, Wells of Ythan, NJ623368, D. Welch, 1996, herb. D.W., det. G.G. Graham. Ist post-
1930 record.

+75/21.8. ROSA FERRUGINEA *95, Moray: One bush self-sown on old tombstone, Alves, near
Elgin, NJ134629, I.P. Green, 1996.

75/21.11x20. ROSA STYLOSA x R. AGRESTIS *11, S. Hants.: Heathland roadside, B3078
Godshill Ridge, SU186156, P.D. Stanley, 1997, det. A.L. Primavesi. 2nd British record.

75/21.12x13a ROSA CANINA x R. CAESIA subsp. CAESIA (R. x DUMALIS) _*37, Worcs.: Scrub
on side of Tanners Brook, SW of Wyre Forest, SO7273, R. Maskew & W.A. Thompson, 1995.

75/21.12x15. ROSA CANINA x R. TOMENTOSA (R. x SCABRIUSCULA) 17, Surrey: Two bushes
in hedgerow, Byfleet, TQ071606, B.W. Phillips & K. Page, 1997, conf. A.C. Leslie & P. Stanley.
2nd record.

75/21.12x17. ROSA CANINA x R. MOLLIS (R. x MOLLETORUM) *94, Banffs.: Hedge,
Broomhills farm road, NJ582646, J. Edelsten, 1997, det. R. Maskew.
75/21.12x18. ROSA CANINA x R. RUBIGINOSA (R. x NITIDULA) *37, Wores:: Scrub-on

limestone grassland, Walsgrove Hill, Great Witley, SO7465, R. Maskew, 1995, det. as hybrid with
R. canina as female parent.

75/21.12x19. ROSA CANINA x R. MICRANTHA (R. x TODDIAE) *37, Worcs.: Calcareous
grassland, Gadbury Bank near Eldersfield, SO7931, R. Maskew, 1989, det. A.L. Primavesi as
hybrid with R. micrantha as female parent. *47, Monts.: Llanymynech Rocks, SJ/26.21, R.
Maskew, 1995.

75/21.12x20. ROSA CANINA x R. AGRESTIS (R. x BELNENSIS) *11, S. Hants.: Heathland
roadside, B3078 Godshill Ridge, SU186156, P.D. Stanley, 1997, det. A.L. Primavesi. *49,
Caerns.: Wood, near Keeper’s Cottage, Penrhynside, Llandudno, SH8081, W. McCarthy, 1996. Ist
Welsh record.

75/21.13x16. ROSA CAESIA x R. SHERARDII *94, Banffs.: Hedge, Mains of Durn farm road,
NJ586651, J. Edelsten, 1997, det. R. Maskew.

75/21.13bx14. ROSA CAESIA subsp. VOSAGIACA x R. OBTUSIFOLIA *37, Worcs.: Scrub on
limestone grassland, Walsgrove Hill, Great Witley, SO7465; Scrub on limestone grassland,
Bredon Hill, S09439; both R. Maskew, 1995. 1st and 2nd records.

75/21.11x14. ROSA STYLOSA x R. OBTUSIFOLIA *37, Worcs.: Roadside bank by A44,
Broadwas, SO7555, R. Maskew, 1997, conf. A.L. Primavesi as hybrid with R. obtusifolia as
female parent.

75/21.14x15. ROSA OBTUSIFOLIA x R. TOMENTOSA *37, Worcs.: Scrub in deciduous
plantation in Trench Wood, Oddingley, SO9259, R. Maskew, 1996, det. as hybrid with R.
tomentosa as female parent.

75/21.14x19. ROSA OBTUSIFOLIA x R. MICRANTHA *37, Worcs.: Scrub on calcareous
grassland near Broadwas, SO7755, R. Maskew, 1997, conf. A.L. Primavesi as hybrid with R.
micrantha as female parent. :

75/21.15x16. ROSA TOMENTOSA x R. SHERARDII (R. x SUBERECTIFORMIS) *37, Worcs.: Scrub
on roadside at Broad Heath, SO6765, R. Maskew, 1994, conf. A.L. Primavesi as hybrid with R.
sherardii as female parent.

75/21.17. ROSA MOLLIS 50, Denbs.: Cliffs in Country Park, Mold, SJ1962, W. McCarthy,
1996. Limestone grassland, Maeshafn, SJ2061, BSBI group, 1996, det. A.L. Primavesi. 2nd v.c.
records.

75/21.18. ROSA RUBIGINOSA 50, Denbs.: Limestone hillside, Rhyd y Foel, SH912763, W.
McCarthy, 1997. Canal bank, Trevor, $J258414, J.A. Green, 1997. Ist & 2nd recent records.

PLANT RECORDS Sal

75/21.19x20. ROSA MICRANTHA x R. AGRESTIS (R. x BISHOPII) *8, S. Wilts.: Hedge, Toyd
Farm, 5km S of Coombe Bissett, SU089206, P.D. Stanley, 1995, det. A. L. Primavesi.

75/21.20. ROSA AGRESTIS 8, S. Wilts.: Hedge, Toyd Farm, 5km S of Coombe Bissett,
SU089206, P.D. Stanley, 1995, det. A.L. Primavesi. Ist record since before 1957. #12, N.
Hants.: Old chalk pit, Noar Hill, SU7431, A. Brewis, 1978, herb. A.B., det. R. Melville. "37,
Worcs.: Scrub on Malvern Common, SO7744, P.G. Garner & R. Maskew, 1989, det. G.G. Graham
& A.L. Primavesi. 49, Caerns.: Limestone quarry, Bodafon, Llandudno, SH807820,
W. McCarthy, 1995, det. A.L. Primavesi. Limestone hillside, Great Orme, SH7683, W. McCarthy,
1996. Ist & 2nd records this century. *50, Denbs.: Limestone hillside, Colwyn Bay, SH829799,
W. McCarthy, 1996, det. A.L. Primavesi.

+75/22.13. PRUNUS LUSITANICA *46, Cards.: Estate woodland, Llanerchaeron, SN480602, A.
O. Chater, 1982. Mixed woodland, Coed Penglanowen, Nanteos, SN610786, A.O. Chater, 1992.
Ist & 2nd v.c. record; widely naturalised throughout Cards. *49, Caerns.: Steeply sloping copse
near roadside, between Tyn-y-groes and Talycafn Bridge, SH785719, R. Lewis, 1993, NMW.
Originally planted but now spreading from bird-sown plants. [The record for this species from
Llanbedr-y-cennin in Watsonia 20(3): 292 (1995) should be deleted and replaced by the above].
Edge of woodland above B5106, Gwydir Castel, Llanrwst, SH/79.60, R. Lewis, 1976. Ist and 2nd
records.

75/22.4x5. PRUNUS SPINOSA x P. DOMESTICA (P. x FRUTICANS) *37, Worcs.: Hedgerow,
Eckington, SO9141, A.W. Reid, 1997, conf. R. Maskew & W.A. Thompson.

+75/28.18. SORBUS DECIPIENS *47, Monts.: Berriew, SJ1600, P.J. Nethercott, 1990.

+75/28.23. SORBUS LATIFOLIA *47, Monts.: Several plants with S. aria cut back into roadside
hedge, Brooks, SO1499, M. Wainwright & V. Morgan, 1989, det. P.J. Nethercott. One medium-
sized tree in hedgebank beside road, Berriew, SJ1600, P.J. Nethercott, 1989. Ist and 2nd records.

+75/28.7. SORBUS INTERMEDIA 42, Brecs.: One young plant c.15cm tall in waste ground,
Brynmawr, SO167119, M. & C. Porter, 1997. 2nd record. *44, Carms.: On waste ground,
Llanelli, SS500999, I.K. Morgan, 1992, NMW, det. G. Hutchinson. *47, Monts.: One small tree
on roadside bank, Berriew, SJ1600, P.J. Nethercott, 1989. Roadside hedge and woodland, Brooks,
SO1499, P.J. Nethercott, 1990. 1st and 2nd records.

175/31.1. PHOTINIA DAVIDIANA *11, S. Hants.: Amongst gorse at edge of wood along
heathland path, Fields Heath, Fawley, SU456023, R.P. Bowman, 1988, herb. R.P.B., det. R.P.
Bowman & E.J. Clement.

175/32.34. COTONEASTER BULLATUS *4, N. Devon: Hedge near limestone quarry, Bampton,
SS$969221, M. Greenwood, 1995, det. J.R. Palmer. Plantation, Meddon, SS281189, W.H. Tucker,
1997, det. J. Fryer. lst and 2nd record.

175/32.35. COTONEASTER REHDERI *4. N. Devon: In plantation, Windcutter Hill, Lee,
SS487461, E. Stenger & W.H. Tucker; Old railway cutting, Landcross, SS459237, W.H. Tucker;
both 1997, det. J. Fryer. 1st and 2nd records.

175/32.37. COTONEASTER DIELSIANUS *35, Mons.: Bird sown on old wall with
C. horizontalis, Abergavenny, SO2914, I.K. Morgan, 1993, NMW, det. G. Hutchinson. *46,
Cards.: Rocky scrub 150m SSW of Gwynfa, Tresaith, SN283511, A.O. Chater & P. Culyer, 1995,
NMW.

+75/32.39. COTONEASTER FRANCHETII *49, Caerns.: Bank in old quarry, Llangystenium near
Colwyn Bay, SH8279, G. Battershall & W. McCarthy, 1997. Roadside, Morfa Bychan,
Porthmadog, SH5337, BSBI meeting, 1997. Ist & 2nd v.c. record.

*75/32.40. COTONEASTER STERNIANUS __*46, Cards.: Self-sown bush in rocky scrub N of road
at W end of Cwm Rheidol Reservoir, SN695796, A.O. Chater, 1996, NMW. *49 Caerns:.:
Limestone grassland bank, Great Orme, Llandudno, SH7782, W. McCarthy, 1997.

+75/32.mai. COTONEASTER MAIREI *11, S. Hants.: Roadside in thick shade, Somerley area,
SU121075, G.D. Field, 1998, det. J. Fryer.

775/32.mar. COTONEASTER MARGINATUS *50, Denbs.: Quarry floor, Llandulas, SH9177, W.
McCarthy, 1994, det. J.Fryer.

+75/33.1. PYRACANTHA COCCINEA *46, Cards.: Several plants in rocky scrub above road at W
end of Cwm Rheidol Reservoir, SN695796, A.O. Chater, 1996, NMW.

ay 2 PLANT RECORDS

75/35.7x8. CRATAEGUS MONOGYNA x C. LAEVIGATA (C. x MEDIA) 28, W. Norfolk: Roadside
hedge, Rougham, TF836206, K. Beckett, 1996. 2nd record, c.600m from C. laevigata. *46,
Cards.: One bush in hedgebank 130m SW of Tanllan, Llanbadarn Odwyn, SN633609, A.O.
Chater, 1996.

75/35.8. CRATAEGUS LAEVIGATA 35, Mons.: Two trees on wet heath, S of Tredegar,
SO151068, T.G. Evans, 1995, NMW. Ist record since 1946.

+77/7.1b. ANTHYLLIS VULNERARIA subsp. POLYPHYLLA *49, Caerns.: Side of disused railway
track, Bontnewydd, SH4757, G. Battershall & W. McCarthy, 1996.

77/7.1e. ANTHYLLIS VULNERARIA cf. subsp. LAPPONICA *45, Pembs.: Seacliff vegetation, E
side of Porthclais Harbour, St David’s, SM742239, E. Norman, 1997, NMW,, det. J.R. Akeroyd.

+77/12.1. HIPPOCREPIS EMERUS “*12, N. Hants.: Hedgerow, Monk Sherborne, SU607556, M.
Taylor, 1997, herb. A. Brewis, det. E.J. Clement.

77/14.1. VICIA OROBUS 109, Caithness: Valley side, Sandside, by Reay, NC950660, R.E.C.
Ferreira, 1996. Ist localised record.

+77/14.3. VICIA TENUIFOLIA 28, W. Norfolk: Pine forest ride in long grass, Roudham,
TL931884, K.A. & G. Beckett, 1997, conf. E.J. Clement. 2nd record. c.3 plants far from habitation
or disturbance.

177/14.5. VICIA VILLOSA *35, Mons.: Newport Docks, ST38, T.G. Evans, 1985, NMW.

77/14.11b. VICIA SATIVA subsp. SEGETALIS *49, Caerns.: Road bank, Uwchmynydd near
Aberdaron, SH1526, M. & A. Atkinson, 1997. Hedge bottom, Machroes near Abersoch, SH3226,
M. & A. Atkinson, 1997. Ist & 2nd localised records of this subspecies.

+77/14.13. VICIA LUTEA *35, Mons.: Less than ten plants on railway embankment near bridge
over Afon Honddu, Llanvihangel Crucorney, SO3220, S.A. Rippin, 1987.

+77/15.10. LATHYRUS HETEROPHYLLUS *35, Mons.: Extensive area on old railway ballast,
Pye Corner, Bassaleg, ST2787, T.G. Evans, 1996. Ist Welsh record.

+77/15.11. LATHYRUS HIRSUTUS *58, Cheshire: Looking established on rough bank, Wharton,
Winsford, $J655667, J.P. Guest, 1997, herb. G.M.K., det. G.M. Kay.

177/15.12. LATHYRUS NISSOLIA *49, Caerns.: Grassy area at side of track leading to Llyn
Parc, about 1.75km SW Llanrwst, SH7860, W. McCarthy, 1996. 51, Flints.: Roadside verge
near crossroads, | km NW of Nercwys church, $J230613, J. Phillips, 1997. Frequent component
of new amenity grassland on site of old rubbish tip, Etna Country Park, Buckley, $J285650, P.
Hodges, 1997. Ist confirmed records post-1930.

+77/15.6. LATHYRUS TUBEROSUS _ *42, Brecs.: Edge of disused railway, Cradoc, SO014304,
E. Tompkins, 1997, NMW. At this site for at least 10 years.

*+77/15.7. LATHYRUS GRANDIFLORUS 44, Carms.: Established in roadside hedgerow, N side of
A476 at Gate Square, near Penygroes, SN578148, R.D.Pryce, 1991, conf. G. Hutchinson. 2nd
record, still present in 1996. *95, Moray: Rough area of former garden on edge of forest,
Darnaway Forest, near Forres, NJO06568, I.P. Green, 1996.

+77/15.9. LATHYRUS LATIFOLIUS *50, Denbs.: Grassy hillside, Rhyd y Foel, SH914774, J.V.
Robertson, 1997.

77/16.4b. ONONIS REPENS subsp. MARITIMA *46, Cards.: Dune at S end of Tan-y-bwlch
Beach, Aberystwyth, SN579798, A.O. Chater, 1996, CGE, det. P.D. Sell. Sandy slope above sea,
Traeth y Mwnt, SN194519, A.O. Chater, 1996, NMW. Ist and 2nd records.

+77/16.mit. ONONIS MITISSIMA * *41, Glam.: Garden lawn, Dinas Powis, ST154709, J.P.
Curtis, 1992, NMW, det. R.M. Burton.

177/17.3. MELILOTUS OFFICINALIS *10, Wight, Disturbed ground, Bembridge Harbour,
SZ632885, S.A. Campbell, 1997, herb. C.R. Pope, det. R.M. Burton. Ist confirmed record this
century, despite intensive searches.

77/18.5. MEDICAGO ARABICA +81, Berwicks.: Sandy bank, The Lees, NT840390, M.E. & P.F.
Braithwaite, 1997. 2nd extant record.

77/19.3. TRIFOLIUM OCCIDENTALE *45, Pembs.: Several flowering patches on coastal cliff
grassland, Stack Rocks, Castelmartin, SR/92.5944 and Short Point, Dale, SM798044, R.S.
Cropper, 1995. 1st and 2nd records.

PLANT RECORDS 29/5)

+77/19.4b. TRIFOLIUM HYBRIDUM subsp. ELEGANS *77, Lanarks.: Coal bing slope, S of
Garnkirk, NS6767, P. Macpherson, 1997, herb. P.M.

77/19.6. TRIFOLIUM SUFFOCATUM ~—_ 10, Wight: Frequent on sandy coastal bank, Cande Lake,
Ryde, SZ602926, C.R. Pope, 1996, herb. C.R.P. 2nd extant locality.

+77/19.10. TRIFOLIUM AUREUM _ 5, S. Somerset: Rough grassland at S end of Otterhead Lakes,
Royston Water, ST226130, C.J. Cornell, 1997, conf. P.R. Green. 1st record for 73 years. Most
likely sown with grass seed several years ago when work was done on lakes.

*77/22.1x2. LABURNUM ANAGYROIDES x L. ALPINUM (L. x WATERERI) *44, Carms.: Pure
stand in hedge, Banc Blodeuyn, Mynydd Llanllwni, SN4835, P.A. Smith & A.O. Chater, 1996.
Mainly a pure stand in hedgerow, Ffos-y-Gaseg, about 4km SW of Brechfa, P.A. Smith & A.O.
Chater, 1996. Ist and 2nd records.

177/23.2. CYTISUS MULTIFLORUS *49, Caerns.: Side of disused railway, Port Dinorwic,
SH5267, G. Battershall et al., 1996. 1 plant on grass verge, near Bangor, SH5971, G. & I.
Battershall, 1997. 1st and 2nd records.

+77/24.1. SPARTIUM JUNCEUM *44, Carms.: One plant (self-sown?), roadside, Felinfoel,
SN5202, I.K. Morgan, 1996. Amenity plantings, Machynys, SS5097, I.K. Morgan, 1996. Ist and
2nd records.

77/26.1x2. ULEX EUROPAEUS x U. GALLI *46, Cards.: Gorsey slope, W side of Pendinas,
Aberystwyth, SN582804, A.O. Chater, 1997, NMW, conf. P.M. Benoit.

+80/1.man. GUNNERA MANICATA *49, Caerns.: Riverside, Pont Eidda near Ysbyty Ifan,
SH8350, G. & I. Battershall, 1997.

84/1.1x8. EPILOBIUM HIRSUTUM x E. CILIATUM (E. x NOVAE-CIVITATIS) *3, S. Devon: Near
River Axe, Whitford Bridge, SY2695, L.J. Margetts, 1996, conf. T.D. Pennington. *37, Worcs.:
Interfield, SO7749, P.G. Garner, 1993, det. T.D. Pennington.

084/1.8x12. EPILOBIUM CILIATUM x E. BRUNNESCENS _*44, Carms.: Acid quarry spoil heap,
Pen-y-Dinas Quarry, Llansawel, SN629355, A.O. Chater, 1995, NMW, det. T.D. Pennington. Ist
Welsh record.

784/1.13. EPILOBIUM PEDUNCULARE 45, Pembs.: Woodland garden, Colby Lodge, SN156078,
G. Kitchener, 1995.

84/1.2x3. EPILOBIUM PARVIFLORUM x E. MONTANUM (E. x LIMOSUM) *46, Cards.:
Hedgebank, Clawdd-moel, Temple Bar, SN531531, A.O. Chater, 1996, NMW, conf. G.
Kitchener. *77, Lanarks.: Sloping grassland in light wood, Burnside, near Glasgow, NS6259, P.
Macpherson, 1995, herb. P.M., det. J.R. Akeroyd.

84/1.2x8. EPILOBIUM PARVIFLORUM x E. CILIATUM *37, Worcs.: Garden weed, Wollaston,
Stourbridge, SO8884, M.E. Smith, 1989, det. G.D. Kitchener. *49, Caerns.: Streamside gravels,
Aberdaron, SH1726, M. & A. Atkinson, 1997. *58, Cheshire: By pool, Sandbach, SJ762610, J.
H. Clarke, 1997, det. G. Kitchener. *77, Lanarks.: Base of wall by River Clyde, Blantyre,
NS6958, A. McG. Stirling & P. Macpherson, 1996, herb. P.M., det. G.D. Kitchener.

84/1.3x6. EPILOBIUM MONTANUM x E. OBSCURUM (E. x AGGREGATUM) *47, Monts.:
Laneside near v.c. 50 border, Ffridd Llanerch, SJ1422, P.M. Benoit, 1988. Ist localised record.

84/1.3x8. EPILOBIUM MONTANUM x E. CILIATUM *77, Lanarks.: Roadside, Plains, Airdrie,
NS7966, P. Macpherson, 1995, herb. P.M., det. J.R. Akeroyd. *81, Berwicks.: One plant with
both parents on waste ground, Lauder, NT527477, M.E. Braithwaite, 1997, herb. M.E.B., det. G.
Kitchener.

84/1.4x6. EPILOBIUM LANCEOLATUM x E. OBSCURUM (E. x LAMOTTEANUM) *49 Caerns:.:
Quarry near Aberdaron, SH1929, A.P. Conolly, 1959, det. C.A. Stace.

84/1.6x8. EPILOBIUM OBSCURUM x E. CILIATUM 44, Carms.: Swampy ditch between road and
Afon Teifi, 300 m SE of Pont Stephan, Cwmann, SN583474, A.O. Chater, 1997. 2nd confirmed
record. *46, Cards.: Shaded verge of A475, 700m E of Llandyfriog church, SN340409, C.D.
Preston & A.O. Chater, 1996. Felled conifer plantation, Lodge Park, Tre’r-ddol, SN662931, A.O.
Chater, 1997, NMW. Ist and 2nd records.

84/1.7. EPILOBIUM ROSEUM 89, E. Perth: By road along river, Blairgowrie, NO178457, O.M.
Stewart, 1981. 2nd record.

574 PLANT RECORDS

84/1.9x12. EPILOBIUM PALUSTRE x E. BRUNNESCENS *46, Cards.: With both parents on damp
shaley ground by Forestry road 2km S of Hafdre, Llyn Brianne, SN804512, A.O. Chater, 1995. E,
NMW., conf. T.D. Pennington, G. Kitchener & D. McKean. Ist British record.

84/4.1x3. OENOTHERA GLAZIOVIANA x O. BIENNIS (O. x FALLAX) *28. W. Norfolk:
Roadside, Garboldisham, TL990816, J. Hawksworth, 1994, det. J.C. Bowra. *51, Flints.:
Between dunes and small marshy area near old lighthouse, Point of Ayr, SJ18, R.G. Ellis. 1980.
NMW., det. K. Rostansk1.

84/4.3x4. OENOTHERA BIENNIS x O. CAMBRICA *95, Moray: Many plants on road bank.
Blackhills, W of Fochabers, NJ276596, IP. Green, 1996, det. J.C. Bowra. Scattered along road
bank. W of Forres, NJO06585, I.P. Green, 1996, conf. J.C. Bowra. 1st and 2nd records.

84/4.4. OENOTHERA CAMBRICA *44. Carms.: Churchyard, St Paul’s Church. Llanelli,
SS508997, A.M. Pell, 1985, NMW, det. R.D. Pryce.
84/6.2. CIRCAEA ALPINA *46, Cards.: Wooded ravine of Llyfnant stream 200m W of

Cwmrhaiadr, SN752962, A.O. Chater & T.D. Dines, 1997, NMW, conf. P.M. Benoit.

784/CLA.amo. CLARKIA AMOENA —_*35, Mons.: Rubbish tip, Newport, ST3085, T.G. Evans.
1985.

785/2.1. AUCUBA JAPONICA *50, Denbs.: Old hedge, Llangollen, $J184439, J. Clarke & A.
Franks, 1997.

88/1.1. EUONYMOUS EUROPAEUS 81, Berwicks.: Riverside woodland, Birkwood Heaugh, M.
E. & P.F. Braithwaite, 1995. 2nd extant record.

788/1.2. EUONYMUS LATIFOLIUS “*11, S. Hants.: Edge of wooded strip along track to beach,
Cadland, Stanswood Bay, SZ470998, R.P. Bowman, 1997, herb. R.P.B., det. E.J. Clement.

+89/1.1xper. ILEX AQUIFOLIUM x I. PERADO (I. x ALTACLERENSIS) S. Channel Islands: One
bushy tree (planted), now about 8m tall, verge of footpath, opposite Flagpole Cottage
(Beauregard), Sark, WV459756, R.M. Veall, 1997, JSY, det. R.A. Waterman. *50, Denbs.:
Farm lane, Llangollen, SJ183434, J. Clarke & A. Franks, 1997. *73, Kirkcudbrights.: Wood
edge, Netherlaw, NX740449, O.M. Stewart, 1996. *95, Moray: Self sown in roadside hedgerow,
Forres, NJO58577, P.R. Green, 1996.

791/2.7. EUPHORBIA SERRULATA *4, N. Devon: Waste ground, Colleton Mills, SS659147, E
Stenger, 1997, conf. L.J. Margetts. | *41, Glam.: Thirty plants on calcareous waste ground, The
Orchards, Llanishen, Cardiff, ST1781, D.Green, 1995.

91/2.15. EUPHORBIA CYPARISSIAS +81, Berwicks.: Established 2m up wall, Lauder.
NT529477, M.E. Braithwaite, 1997. 2nd record.

791/2.16b. EUPHORBIA AMYGDALOIDES subsp. ROBBIAE *46, Cards.: Roadside hedgebank S
of Upper Bridge, Aberaeron, SN458623, A.O. Chater, 1996. Woodland just W of Cymerau Hall.
Glandyfi, SN696962, S.P. Chambers, 1996. Ist and 2nd records.

+91/2.0bl1. EUPHORBIA OBLONGATA _*44, Carms.: Established on garden rubbish tip, Llanelli,
SN4901, LK. Morgan, 1995, NMW, det. E.J. Clement. Ist Welsh record. *46. Cards.: Scrub
slope above church hall, Llanbadarn Fawr, SN598810, S.P. Chambers, 1991, conf. B. Wurzell &
E.J. Clement.

7+93/2.2. PARTHENOCISSUS INSERTA *95. Moray: Creeping over gorse in waste area,
Cummingstown, NJ135692, P.R. Green, 1996.

799/1.5. ACER SACCHARINUM *77, Lanarks.: Surviving seedling on pathside in wood,
Corehouse, NS8841, P. Macpherson, .1996, herb. P.M., det. C.S. Crook.

7100/1.1. RHUS TYPHINA *46, Cards.: Well naturalised thicket of suckers in paddock,
Penrhyncoch, SN646842, A.O. Chater, 1997. *49, Caerns.: Side of path, Glynllifon, SH4655, G.
& I. Battershall, 1997. *77, Lanarks.: Bank of River Clyde, Linthouse, NS5466, P. Macpherson.
1992, herb. P.M., conf. J.R. Akeroyd. One plant surviving in 1998.

7102/1.3. OXALIS CORNICULATA *50, Denbs.: Wall in village, Nantglyn, SJO062, BSBI
group, 1996. 51, Flints.: Abundant around disused glasshouses, Thornleigh Park, Sealand.
SJ3666, G. Kay & BSBI group, 1996, conf. M.F. Watson. Ist post-1930 record.

7102/1.4. OXALIS EXILIS 73. Kirkcudbrights.: Grass by road, Mavis Grove road S of
Maxwelltown, O.M. Stewart, 1996. 2nd record.

PLANT RECORDS 575

*102/1.8. OXALIS ARTICULATA 46, Cards.: Roadside hedgebank N of Afon Cledan, 150m E of
A487, Llanon, SN515668, A.O. Chater, 1996. 2nd record.

*102/1.10. OXALIS DEBILIS 46, Cards.: Several plants in rough grass on floor of old sandpit
just N of A487(T), Banc y Warren, Penparc, SN203477, A.O. Chater, 1996. 2nd record.

7102/1.14. OXALIS INCARNATA *46, Cards.: Roadside 100m W of Llangranog church,
SN315540, A.O. Chater, 1996.

+103/1.1. GERANIUM ENDRESSII 81, Berwicks.: An established clump in damp grassland, near
Lauder, NT521481, M.E. Braithwaite, 1997. Ist localised record.

+103/1.1x2. GERANIUM ENDRESSII x G. VERSICOLOR (G. x OXONIANUM) *46, Cards.:
Streambank by churchyard, Llanilar, SN623751, A.O. Chater & S.P. Chambers, 1994, NMW.

103/1.4. Geranium ROTUNDIFOLIUM 51, Flints.: Spontaneous garden weed, Gronant, SJ0983,
J. Phillips, 1996. 2nd v.c. record.

+103/1.13. GERANIUM IBERICUM *50, Denbs.: Edge of stream, Llanrwst, SH799623, R.
Lewis, 1997.

7103/1.13xpla. GERANIUM IBERICUM x G. PLATYPETALUM (G. x MAGNIFICUM) *49, Caerns.:
Naturalised on a bank, Port Dinorwic, SH5267, G. & I. Battershall, 1996.

103/1.15. GERANIUM PUSILLUM 46, Cards.: Disturbed flowerbed, Art Dept. grounds, Buarth
Mawr, Aberystwyth, SN588815, S.P. Chambers, 1996. Sandy swede field, Nantyferwig,
SN168482, A.O. Chater, 1997. lst and 2nd extant localities.

+103/1.17. GERANIUM MACRORRHIZUM __—- *46, Cards.: Footpath verge, SE of National Library,
Aberystwyth, SN595814, A.O. Chater, 1992. Salix cinerea scrub, 200m NW of Cross Inn,
SN542642, A.O. Chater, 1997. Ist and 2nd records. *73, Kircudbrights.: Woodland edge, Port
Ling, NX883540, O.M. Stewart, 1996.

7103/1.bru. GERANIUM BRUTIUM *29, Cambs.: Melbourn by-pass, Meldreth, TL384461, P.D.
sell’ 1997, CGE.

103/2.2. ERODIUM MOSCHATUM 46, Cards.: Shaley track margin and in adjacent field, 700m
ESE of Capel Tygwydd, SN277432, A.O. Chater, 1996. 2nd record.

7+103A/LIM.dou. LIMNANTHES DOUGLASII *S, Channel Islands: Cliff path, Little Sark near
Les Fontaines, Sark, WV451740, M.W. Angel, 1991, JSY, det. D. McClintock from photograph.
*58, Cheshire: Newly-seeded roadbank, Stockport, $J904902, G.M. Kay & E. Kearns, 1997.

+104/1.maj. TROPAEOLUM MAJUS = *44, Carms.: Behind rubble at edge of sea shore, St Johns
Hill, Laugharne, SN302104, G. Hutchinson, 1991, NMW. *49, Caerns.: Waste ground,
Porthmadog, SH5739, G. & I. Battershall, 1996, and J. Clarke & A. Franks, 1997.

105/1.1. IMPATIENS NOLI-TANGERE *46, Cards.: In a gully at Capel Bangor, SN67, C. Morley,
1939, in MS notebook, Ipswich Museum, per A.P. Fowles.

106/1.1., HEDERA COLCHICA *46, Cards.: Scrub and wood, Caerhedyn, Llyfnant, SN709974,
few). Chater, 1997. *47, Monts.: Scrambling up lakeside bank and trees, Lake Vyrnwy,
SH962222, J. Clarke, 1997, det. A. Rutherford (as var. dentata).

106/1.2b. HEDERA HELIX subsp. HIBERNICA *50, Denbs.: Ruin, Llangwm, SH9844, A.O.
Chater, 1996. Roadside wall, Llanrhaiadr ym Mochnant, SJ1226, J.A. Green, 1996. Ist and 2nd
records.

7 107/1.ran. HYDROCOTYLE RANUNCULOIDES *11, S. Hants.: Stream flowing through grounds,
Southampton University, Highfield, Southampton, SU424151, C.M. Cockerill, 1997, herb. R.P.
B., det. R.P. Bowman. Browndown, Gosport, SZ583996, M.J. Southam, 1997. Ist and 2nd records.
*35, Mons.: 4 large patches, Broadway Reen, Marshfield, ST2681, T.G. Evans, 1996.

+107/3.1. ASTRANTIA MAJOR *46, Cards.: Well established on shaley slope below Forestry
road in conifer forest 2-3km SE of Eisteddfa Gurig, SN811820, A.O. Chater, 1996, NMW. With
Fuchsia magellanica and Cotoneaster horizontalis and presumably deriving from throw-outs,
although 2km from nearest garden.

107/16.1. BERULA ERECTA 93, N. Aberdeen: Marsh, Logie Buchan, NK0029, D. Welch,
1996, ABD. Only extant locality.

576 PLANT RECORDS

107/19.3. OENANTHE PIMPINELLOIDES *61, S.E. Yorks.: Seventy plants in neutral grassland,
on edge of playing field, Ennerdale Sports Centre, Hull, TA0933, R. Middleton, 1997, HLU, det.
M.J. Southam, Ist record N of Ipswich.

+107/30.1. SISON AMOMUM *44, Carms.: About fifty plants, presumably introduced with
material from Slimbridge, in marshy area at edge of path, S of main wildfowl lake, Penclacwydd
Wildfowl and Wetland Trust Centre, Llanelli, SS530986, B. Stewart, 1991, det. I.K. Morgan. Ist
post-1930 record.

107/31.1. CICUTA VIROSA —_*37, Worcs.: Marginal reed-swamp, Podmore Pool, Kidderminster,
SO8477, C.B. Westall, 1989, conf. J.J. Day & R. Maskew.

+107/38.1. LEVISTICUM OFFICINALE *35, Mons.: One plant in gap in pavement, Chepstow,
ST5393(G), C.A. Shirley, 1994, NMW. *95, Moray: Several plants self sown, waste ground on
dunes, Finehorn, NJ043643, I.P. Green, 1997.

+ 107/39.3. PEUCEDANUM OSTRUTHIUM *67, S. Northumb.: At W end of enclosure around
long-deserted farm house, Grindon Green, NY729736, G. Simpson, 1992, herb. G.A. Swan. Only
extant locality. *91, Kincardines.: Den by old graveyard, Garvock, NO743705, D. Welch, 1997.

107/40.1. PASTINACA SATIVA +42, Brecs.: Edge of railway track, Garth Station, SN953494, L.
J. Smith, 1996. 2nd record. +*73, Kirkcudbrights.: 6-7 plants on roadside, A75 just E of road off
to Girton, NX6154, O.M. Stewart & A. White, 1996.

+107/41.2. HERACLEUM MANTEGAZZIANUM *46, Cards.: Naturalised for c. 27 years,
farmyard, Ynys Edwin, Eglwys-fach, SN678962, A.O. Chater & W.M. Condry, 1997. =O1"
Kincardines.: By River North Esk, Morphie, NO708636, D. Welch, 1994. Well established so
present here for some years.

108/4.1. BLACKSTONIA PERFOLIATA _*77, Lanarks.: Very widespread in abandoned industrial
estate, Netherton, NS7855, W. Brackenridge, 1997, herb. P. Macpherson.

108/5.2. GENTIANELLA CAMPESTRIS 83, Midlothian: About twenty plants in rich grassland,
Fullarton Water, only 2m away from lay-by after bridge, NT283564, C. Dixon, 1997, det. D.R.
McKean. Ist record post-1934.

108/5.5. GENTIANELLA ANGLICA subsp. ANGLICA *37, Worcs.: Limestone grassland, Tunnel
Hill, SP0247, J.W. Meiklejohn, 1992, conf. A.W. Reid. Limestone grassland, Highclere, SP0147
& SP0148, D.T. Holyoak, 1992. Ist and 2nd records.

7110/1.1. NICANDRA PHYSALODES 44, Carms.: Occasional garden weed on disturbed soil,
Maesquare, W of Crymlyn Manor, Bethlehem, SN653233, M. Williams, 1995, NMW, det. G.
Hutchinson. 2nd record.

110/8.1x3. SOLANUM NIGRUM x S. PHYSALIFOLIUM (S. x PROCURRENS) *11, S. Hants.: Potato
field, North Hayling, SU728030, P.D. Stanley & E.J. Clement, 1996, herb. R.P. Bowman, det. E.
J.C.

+110/8.3. SOLANUM PHYSALIFOLIUM *49, Caerns.: Flower beds, town centre, Llandudno,
SH7782, W. McCarthy, 1996.

+110/8.4. SOLANUM SARACHOIDES *95, Moray: One plant on waste ground, rubbish tip,
Elgin, NJ236631, P.R. Green, 1997.

+110/8.7. SOLANUM TUBEROSUM _ *46, Cards.: Naturalised for at least 5 years in gully on sea
cliffs, 300m ENE of Mwnt church, SN198521, A.O. Chater, 1997.

+110/8.ros. SOLANUM ROSTRATUM *11, S. Hants.: Edge of refuse tip in gravel extraction area,
along footpath, Efford, Keyhaven, $Z310922, M.E. Young, 1997, herb. R.P. Bowman.

+110/8.vil. SOLANUM VILLOSUM *35, Mons.: Farmyard, Rogiet, ST48, T.G. Evans, 1990s,
NMW.

111/3.2b. CALYSTEGIA SEPIUM subsp. ROSEATA *46, Cards.: Grassy railway embankment,
500m W of Ynys Edwin, Eglwys Fach, SN673962, A.O. Chater & W.M. Condry, 1991, NMW.

111/3.2x4. CALYSTEGIA SEPIUM x C. SILVATICA (C. x LUCANA) _*37, Worcs.: Waste ground,
Billesley Common, SP0880, J.W. Partridge, 1995, det. R.K. Brummitt.

111/3.3x4. CALYSTEGIA PULCHRA x C. SILVATICA (C. x HOWITTIORUM) *¥1, S. Hants.: In

roadside hedge opposite house and garden, twining over Symphoricarpos, A3057 Lower Brook,
Kings Somborne, SU339278, R.P. Bowman, 1997, herb. R.P.B., det. R.K. Brummitt.

PLANT RECORDS a7

113/2.1. NYMPHOIDES PELTATA 751, Flints.: Established from planted stock in shallow
muddy water, Greenfield Country Park, SJ193773, G. Wynne, 1997. 2nd record. +*60, W.
Lancs.: Old mill pond, Fleetwood, SD327465, D. & J. Steeden, 1985.

7114/1.1. POLEMONIUM CAERULEUM *50, Denbs.: Naturalised on rough banks, Carrog,
SJ1046, S.P. Chambers, 1996.

115/1.1., PHACELIA TANACETIFOLIA *35, Mons.: Horsington Yard, Abergavenny, SO21, R.
Fraser, 1988, NMW. About 300 plants in strip across rape field, E of Gwern-eiddig, SO4106, J.D.
R. Vernon, 1996; planted to keep away aphids. 1st and 2nd records. *41, Glam.: Established
garden throw out, Whitchurch, Cardiff, ST152795, G. Hutchinson, 1991, NMW. *42, Brecs:.:
Weed on garden path, Cwm Gwdi, 3km SW of Brecon, SO0226, C.H.G. Allum, 1996. Introduced
with bird seed? *43, Rads.: Garden path, Presteigne, SO3164, S. Voelcker, 1996, det. D.R.
Humphreys.

*+116/2.4. ECHTUM PININANA *3, S. Devon: Self seeding in a lane, Ringmore, SX6545, M.
Catt, 1997. :

¥116/12.2. AMSINCKIA MICRANTHA *58, Cheshire: New earth bank, Moreton, S$J263901, P.G.
Gutteridge, 1997, det. G.M. Kay. 77, Lanarks.: Well established in recently planted woodland,
Blantyre, NS6829, K. Watson, 1997, GLAM, det. E.J. Clement. Ist record since 1923, presumably
an accidental introduction.

116/15.3. MYOSOTIS STOLONIFERA *77, Lanarks.: Flushes by the Carsehope Burn, SW of
Daer Reservoir, NS9505, K. Watson, 1997, herb. P. Macpherson.

7116/4.2. SYMPHYTUM ASPERUM *830, Roxburghs.: One large established clump, roadside
verge, Pinnacle, Ancrum, NT589254, H. Noltie, 1992, herb. R.W.M. Corner.

+116/4.4. SYMPHYTUM GRANDIFLORUM *49, Caerns.: Edge of wood, Edern near Nefyn,
SH2839, G. Battersall et al., 1996. Grassy roadside, Bethesda, SH6067, J. Hawksford, 1997. Ist
and 2nd records.

7¥116/9.2. BORAGO PYGMAEA ~—_*35, Mons.: Narrow band of plants among foxgloves, bracken
and brambles in strip of tall plants in middle of steep field, Yew Tree Cottage, Long Lane, Penallt,
$0527092, S.J. Tyler, 1995, herb. T.G.E., det. T.G. Evans, conf. J.R. Akeroyd.

+118/4.1c. LAMIASTRUM GALEOBDOLON subsp. ARGENTATUM *50, Denbs.: Road verge,
Bryneglwys, SJ1446, S.P. Chambers, 1996.

118/5.2. LAMIUM MACULATUM 50, Denbs.: Established on bank, Maerdy, SJ0144, S.P.
Chambers, 1996. 2nd record.

7118/17.1. SATUREJA MONTANA = *44, Carms.: About thirty-five plants well naturalised, street-
side brick wall, Bryn Avenue, Barry Port, SN453009, I.K. Morgan, 1993.

118/18.5. CLINOPODIUM ACINOS *10, Wight: Fifty-six plants counted in old chalk pit,
Nunwell Down, Brading, S$Z592872, A. Campbell, 1997, herb. C.R.P., det. C.R. Pope. Only
extant locality.

*+118/23.3x4. MENTHA SUAVEOLENS x M. SPICATA (M. x VILLOSA) *46, Cards.: Scrub above
coast path, W end of Tresaith beach, SN277514, A.O. Chater, 1994, NMW.

118/23.4. MENTHA SUAVEOLENS +*81, Berwicks.: Established on riverside, River Tweed
below Clintmains, NT606324, M.E. Braithwaite, 1995, herb. M.E.B., det. R.M. Harley.

+118/24.1. ROSMARINUS OFFICINALIS 49, Caerns.: Sand dunes, West shore, Llandudno,
SH7781, W. McCarthy, 1996. 2nd record.

118/25.4. SALVIA VERBENACA 50, Denbs.: Limestone rock outcrops, Henllan, SJO27697, C.
Welsh, 1997, det. J.A. Green. Grassland, Rhyd y Foel, SH914762, J.V. Robertson, 1997. Ist recent
records

120/1.2. CALLITRICHE TRUNCATA 15, E. Kent: Marsh dyke, Denge Beach, TRO517, E.G.
Philp, 1996, MNE. Ist record for about fifty years.

120/1.6. CALLITRICHE BRUTIA *110, Outer Hebrides: Beinn na h-Aire, S Uist, NF8435, P.
Smith, 1994.

121/1.1. PLANTAGO CORONOPUS 47, Monts.: Sandy saltmarsh, near Dovey Junction Station,
SN694987, Flora project field party, 1989. 2nd record.

578 PLANT RECORDS

121/1.3b. PLANTAGO MAJOR subsp. INTERMEDIA *50, Denbs.: Sandy bank, Sutton Green,
SJ417489, K. Watson, 1993, NMW.

*122/1.2. BUDDLEJA DAVIDII *91, Kincardines.: Waste ground by petrol station, Banchory,
NO709960, D. Welch, 1997.

+123/1.susxvir. FORSYTHIA SUSPENSA x F. VIRIDISSIMA (F. x INTERMEDIA) ONE Hants:
Lane, Kingsley, SU787380, A. Brewis, 1997. *35, Mons.: Bank of River Ebbw, Risca, ST29, J.
Harper, 1992. The River Ebbw bank has been reinforced and raised and many strange, planted?
species have appeared there; this may be one of them.

7123/2.ang. FRAXINUS ANGUSTIFOLIA *29, Cambs.: Roadsides, Cambridge, TL45, C.D.
Preston, G. Crompton & G.M.S. Easy, 1986 & 1987. ?Frequently planted now.

124/1.2. VERBASCUM VIRGATUM *79, Selkirks.: Pathside (now part of Southern Upland
Way), near Galafoot, Galashiels, NT511351, J. Murray, 1995, herb. R.W.M. Corner, conf. D.R.
McKean.

124/1.2x7. VERBASCUM VIRGATUM x V. THAPSUS (V. x LEMAITREI) *83, Midlothian: With
both parents in waste ground, opposite Duddington caravan site, NT285721, S. Maxwell, 1996, E,
det. D.R. McKean. New Scottish record.

+124/1.6. VERBASCUM DENSIFLORUM *35, Mons.: One plant with three main branched stems,
on trackside in Council road maintenance site, Dixton, Monmouth, SO5113, T.G. & U.T. Evans,
1997, herb. T.G.E.

124/8.2. CHAENORHINUM MINUS *91, Kincardines.: Railway cess, Newtonhill, NO910935, D.
Welch, 1997.

124/1.11. VERBASCUM PULVERULENTUM “*44, Carms.: Waste ground, E of Llanelli Railway
Station, SS508993, T. Davies, 1991.

+124/10.1. ASARINA PROCUMBENS _*58, Cheshire: Old wall, Alderley Edge, SJ851781, A.R.
Franks, 1997. Present for at least 10 years.

*124/11.2. CYMBALARIA PALLIDA *81, Berwicks.: Established on lower town walls, Lauder,
NT529476, M.E. Braithwaite, 1997.

+124/11.3. CYMBALARIA HEPATICIFOLIA *40, Salop: Garden escape, growing on a wall,
Grinshill, SJ515238, T. Preece, 1994.

124/12.2. KICKXIA SPURIA *43, Rads.: On disturbed ground, S of Penlan Wood, Clyro,
S0208440, R.G. Woods, 1988, NMW.

124/13.1x4. LINARIA VULGARIS x L. REPENS (L. x SEPIUM) *37, Worcs.: Railway track near
Langley Green Station, SO9988, B.R.W. Fowler, 1987.

124/13.3x4. LINARIA PURPUREA x L. REPENS (L. x DOMINII) *11, S. Hants.: Edge of building
site on bank between pavement and fence, Milford on Sea, SZ289914, V. Scott, 1997, herb. R.P.
Bowman, det. R.P. Bowman & C.A. Stace. *37, Worcs.: Waste ground, Moseley, SP0882, J.J.
Day, P.G. Garner & B. Westwood, 1993.

+124/15.1. ERINUS ALPINUS *95, Moray: Well naturalised in masonry on on Dulicht railway
bridge, Grantown-on-Spey, NJ024282, J.R. Edelsten, 1996.

124/16.19. VERONICA AGRESTIS 10, Wight: Cracks in paving stones, Yarmouth, $Z357897, P.
& G. Stanley, 1997, herb. P.S. 2nd recent record.

7124/16.25. VERONICA LONGIFOLIA *35, Mons.: Waste ground outside walled garden,
Llanfoist Church, SO2813, T.G. Evans, 1990, NMW, det. E.J. Clement. *46, Cards.: Rough
grass by long-overgrown tip, Blaendolau, Llanbadarn Fawr, SN600804, A.O. Chater, 1990.

124/20.1x5. EUPHRASIA ROSTKOVIANA x E. ARCTICA *42, Brecs.: Heathy field, Coelbren,
SN8612, M. Porter, 1980, herb. M.P., det. A.J. Silverside. Ist confirmed Welsh record.

124/20.3x7. EUPHRASIA ANGLICA x E. NEMOROSA *42, Brecs.: Rough grassy bank, Cwm
Fforch, SO0937, M. Porter, 1982, det. A.J. Silverside.

124/20.7x19. EUPHRASIA NEMOROSA x E. SCOTTICA __*42, Brecs.: Flushes along forestry ride,
Garwnant, SN9913, M. Porter, 1989, det. A.J. Silverside.

124/20.8. EUPHRASIA PSEUDOKERNERI *35, Mons.: Turf over Carboniferous Limestone, S of
Little Dinham Wood, T.G. Evans, 1994, det. A.J. Silverside.

PLANT RECORDS 572

124/20.9x19. EUPHRASIA CONFUSA x E. SCOTTICA *46, Cards.: Flush by drainage adit,
Cwmystwyth leadmine, SN810751, S.P. Chambers, 1992, det. A.J. Silverside. *49, Caerns.:
Damp montane grassland, Cwm Idwal, SH6458, A.J. Silverside, 1994.

124/20.13x14. EUPHRASIA CAMBRICA x E. OSTENFELDII *49, Caerns.: Rocky ledge, Cwm
Idwal, SH6458, A.J. Silverside, 1994. 1st Welsh record.

124/20.14. EUPHRASIA OSTENFELDII *42, Brecs.: Flush by rocky outcrop, Nant Irfon,
SN8453, M. Porter, 1995, det. A.J. Silverside. Ist record south of Snowdonia.

124/23.1. PARENTUCELLIA VISCOSA 10, Wight: Over 250 plants in donkey pasture, Wootton
Common, $Z533918, C.R. Pope, 1997. New and only extant site. +42, Brecs.: Single plant
probably introduced in grass-seed mixture on road verge, near Llyswen, SO141375, M. & C.
Porter, 1997. 2nd record.

124/24.1. RHINANTHUS ANGUSTIFOLIUS *11, S. Hants.: Species-rich acidic grassland over
gravel/sand surface of former tip area, W of Fort Cumberland, Eastney, Portsmouth, $Z679991, D.
P.J. Smith, 1997, det. R.P. Bowman (from colour transparencies). *35, Mons.: On bank
damming stream to create a duck pond, Green Meadow Community Farm, $T2994, C. Titcombe,
1992, NMW, det. T.G. Evans, 2nd Welsh record.

124/24.2f. RHINANTHUS MINOR subsp. BOREALIS *49, Caerns.: Moel Siabod, SH75, E.
Roberts, 1950s, det. D.J. Hambler.

124/25.2b. PEDICULARIS SYLVATICA subsp. HIBERNICA *41, Glam.: Abundant on Molinia
bog, especially in areas burnt two years previously, Broad Pool, Gower, SS5191, E. Lughadha, P.
A. Smith, R. Warren & T.C.G. Rich, 1993, NMW.

*125/1.2. LATHRAEA CLANDESTINA — _*48, Merioneth: Shoreline of estuary, Dwyryd Estuary,
SH589357, R.G. Hughes, 1993, NMW. *5(0, Denbs.: Under rhododendron, Bodnant Garden,
SH7982, A. Parry Jones, 1995. 80, Roxburghs.: Under riverside willows, River Tweed, N side
below Allan Water, NT525354, J. Murray, 1997. 2nd record.

125/2.10. OROBANCHE MINOR ~~ *50, Denbs.: Bare gravelly ground, Llay, SJ3354, K. Davies,
1996.

*+127/1.1. ACANTHUS MOLLIS *49, Caerns.: Large, well-naturalised clump on bridleway,
Rhiw, near Aberdaron, SH2328, G. & I. Battershall, 1996. *73, Kirkcudbrights.: Waste ground,
Castle Douglas, NX7562, O.M. Stewart & A. White, 1995.

128/1.4. PINGUICULA GRANDIFLORA +*48, Merioneth: Basic boggy rill with P. vulgaris,
Brithdir, near Dolgellau, SH71, P.M. Benoit, 1987, NMW. Ist record for Wales. Two plants only
in 1983; increased to 11 in 1987. In a completely natural habitat but probably originally planted.

128/2.1. UTRICULARIA VULGARIS 73, Kirkcudbrights.: Abundant in loch by shore, Loch Ken,
NX6970, O.M. Stewart, 1996, E. Ist recent record.

129/1.10.; CAMPANULA POSCHARSKYANA *49 Caerns.: Boulders on bank of River Roe,
Rowen, Conwy, SH7670, W. McCarthy, 1996. Wall base, Llandudno, SH7882, W. McCarthy,
1996. Ist and 2nd records. *50, Denbs.: Old wall at edge of waste ground,Llanrwst, SH7961, R.
Lewis, 1996.

129/1.12. CAMPANULA TRACHELIUM *46, Cards.: Naturalised in thick grass and nettles,
Tyglyn Aeron, SN5059, M.L. Lewis, 1908, ABS, conf. A.O. Chater.

129/1.7. CAMPANULA GLOMERATA 51, Flints.: Semi-improved calcareous grassland, Pentre-
cwm, SE side of Moel Hiraddug, Dyserth, SJ0678, CCW ‘Phase 1’ survey, 1994. 2nd record.

7129/7.2. LOBELIA ERINUS 42, Brecs.: Disturbed roadside verge beside B4358, Beulah,
SN9251, R.G. Woods, 1996. 2nd record. *44, Carms.: River shingle, River Teifi, N of
Pentrecwrt, SN392395, G. Hutchinson, 1991. *47, Monts.: Roadside parking place, NW of
Talywern, SH820009, C.A.Small, 1989. Stoney beach beside Afon Rhiw, NW of Felindre,
SJ160022, E. Roberts & M. Wainwright, 1989. Ist and 2nd records. *49, Caerns.: Waste
ground, Llanberis, SH5759, G. Battershall et al., 1996. Pavement crack, Abersoch, SH3128, J.
Hawksford, 1997. Ist and 2nd records.

129/7.3. LOBELIA DORTMANNA *94, Banffs.: Pool near Loch Avon, NJO18026, J. Edelsten,
1994, E, det. D. McKean.

580 PLANT RECORDS

7130/5.arv. ASPERULA ARVENSIS *S, Channel Islands: Under bird-table in garden, The
Kennel, Rue du Fort, Sark, WV466764, B. Gurden, 1979, JSY, det. D. McClintock.

130/6.3. Galium ULIGINOSUM 46, Cards.: Flushed slope W of Llyn Gwngu, SN/837.728, J.P.
Woodman & A.O. Chater, 1997. 2nd extant locality, and the only upland one.

*+130/8.tin. RUBIA TINCTORUM *53, S. Lincs.: Growing over a Jurassic limestone wall,
Boothby Graffoe, SK9859, P. Porter, 1996, det. I. Weston, conf. E.J. Clement.
131/1.4. SAMBUCUS EBULUS +*46, Cards.: Waste ground, Picton Terrace, New Quay,

SN388600, R.E.N. Smith, 1997. Only extant locality. 59, S. Lancs.: Well-established clump,
fruiting well on disturbed mesotrophic grass/herb between industrial premises, Knowsley
Industrial Park, Kirkby, SJ428972, P. Gateley, 1996. Ist record since 1960.

+131/2.3. VIBURNUM TINUS *49, Caerns.: One shrub in hedgerow between Trefriw and
Gower Bridge, near Llanrwst, SH7862, R. Lewis, 1997. One shrub in hedgerow of lane near Afon
Crafnant, W of Trefriw, SH7763, R. Lewis, 1997. 1st and 2nd records.

+131/3.orbxmic. SYMPHORICARPOS ORBICULATUS x S. MICROPHYLLUS (S.x CHENAULTII)
*11, S. Hants.: Scrub, in hawthorn, gorse and brambles, Pennington Common, SZ307952, M.E.
Young, 1997, det. Everton Nurseries. *39, Staffs.: One prostrate plant on old dumped spoil, W
side of Bilston Cosely Road, SO944957, B.R. Fowler & IC. Trueman, 1996, det. D. Gardner.
*70, Cumberland: Plantation ride, Green Rigg, Catlowdy, NY485784, R.E. Groom, 1990, LANC.
73, Kirkcudbrights.: Disused quarry, near Hardgate, NX812668, O.M. Stewart, 1995. 2nd & only
extant record.

131/4.1. LINNAEA BOREALIS 93, N. Aberdeen: Sitka spruce plantation, previously Scots pine
woodland, The Bin, NJ4943, E. Shand, 1989. Ist post-1930 record, confirming 1906 record for
this pinewood.

7131/6.3. LONICERA INVOLUCRATA *43, Rads.: Streamside, Davids Well, SO0678, D.R.
Humphreys & E.R. Dean, 1996, det. R.G. Woods.

+131/6.6. LONICERA JAPONICA *46, Cards.: Scrub by old allotments, Felin-y-mor Road,
Aberystwyth, SN581805, A.O. Chater, 1994, NMW. Scrub on riverbank W of Llansanntffraed
church, SN511674, A.O. Chater & J.P. Woodman, 1997. Ist and 2nd records.

7131/6.tat. LONICERA TATARICA *17, Surrey: Bush 2m high in woodland strip, Send,
TQ026548, J.F. Leshe & K. Page, 1993. Established garden throw-out.
133/1.2. WALERIANELLA CARINATA *108, W. Sutherland: Stony drive to house, Nedd,

NC137319, ILM. Evans, 1997.

133/1.3. VALERIANELLA RIMOSA 12, N. Hants.: Over 100 plants in arable field, Bramdown
Copse, Overton, SU528471, J.R. Moon, 1994, det. F. Rose. Only extant locality.

4133/2.2. VALERIANA PYRENAICA *91, Kincardines.: Policies of mansion, Glenbervie,
NO768805, D. Welch, 1996.

7133/2.phu. VALERIANA PHU 46, Cards.: Roadside hedgebank 100m SW of Neuadd-lwyd
chapel, 3-5km SSE of Aberaeron, SN473595, A.O. Chater, 1997, NMW. Roadside bank 850m
NNE of Dihewyd, SN491566, A.O. Chater & J.P. Woodman, 1997. Only extant localities in
Britain.

+134/1.2. DIPSACUS SATIVUS = *51, Flints.: Probable garden escape in waste ground, near car
park, Bridge Inn, Pontblyddyn, $J278604, S.P. Chambers, 1997.

+134/1.lac. DIPSACUS LACINIATUS *17, Surrey: Over 20 plants in roadside verge, S of
Hascombe, TQ001382, J.F. Leslie, K. Page & J.E. Smith, 1993. *44, Carms.: About twelve
plants on cinders and ballast by main railway line, Pwll, Llanelli, SN463007, I.K. Morgan, 1997.

+135/1.3. ECHINOPS BANNATICUS *73, Kirkcudbrights.: Several clumps in open wood, near
Kenmure Castle, NX635764, O.M. Stewart & A. White, 1995.

135/3.2c. ARCTIUM MINUS 73, Kirkcudbrights.: Disused railway depot, E of Lochanhead,
NX9171, O.M. Stewart, 1997. 2nd recent record.

135/4.1. SAUSSUREA ALPINA —_*77, Lanarks.: Rocky outcrop, Gana Hill, NS9501, K. Watson,
1997.

135/5.3x4. CARDUUS CRISPUS x C. NUTANS (C. x STANGII) *37, Worcs.: With both parents in
horse pasture, Cookley, SO8480, B. Westwood, 1996, conf. R. Maskew & W.A. Thompson.

PLANT RECORDS 581

135/11.1. CENTAUREA SCABIOSA = *91,, Kincardines.: Disturbed ground at abandoned railway
yard, Crathes, NO7496, D. Welch, 1996.

7135/11.2. CENTAUREA MONTANA *81, Berwicks.: Strong colony established in railway
cutting, Lauder, NT498531, M.E. Braithwaite, 1997.

135/11.3xjac. CENTAUREA NIGRA x C. JACEA (C. x MONCKTONII) *61, S.E. Yorks.: Neutral
grassland, verge of bypass, near Beverley, TA03, J. Dews, 1997, det. C.A. Stace.

135/16.1b. LEONTODON AUTUMNALIS subsp. PRATENSIS [58, Cheshire: Goyts Moss,
SD001718. Delete record in Watsonia 21: 396 (1997). ]

135/16.3. LEONTODON SAXATILIS 95, Moray: Grassy road verge, N verge of A96, W of Elgin,
NJ166631, P.R. Green, 1996. 2nd record.

135/18.1. SCORZONERA HUMILIS *41, Glam.: Flushed marshy grassland, Cefn Cribwr
Meadows SSSI, SS8583, J.P. Woodman, 1996, conf. P.D. Sell & A.O. Chater. Single patch in
species-rich meadow at Little Hills Farm, $S539928, J.P. Woodman, 1997. 1st and 2nd records.

135/19.1b. TRAGOPOGON PRATENSIS subsp. MINOR *79, Selkirks.: Edge of path, near Gala
Water & College of Textiles, Galashiels, NT507353, M. Little, 1994. Ist localised record.

135/21.4gla. SONCHUS ASPER subsp. GLAUCESCENS *46, Cards.: Roadside by Teifi estuary,
Nantyferwig, Gwbert, SN166483, A.O. Chater, 1995. Disturbed ground by supermarket
development, Parc-y-llyn, Llanbadarn Fawr, SN593806, A.O. Chater, 1995. 1st and 2nd records.

+135/23.3. CICERBITA PLUMIERI *95, Moray: Many plants on road verge and open woodland,
Blackhills near Fochabers, NJ276585, I.P. Green, 1996.

135/28.62. TARAXACUM FULGIDUM *42, Brecs.: Meadow, Llangasty Tal-y-Llyn, SO1225, A.
J. Richards et al., 1996.

135/25.65. TARAXACUM HAEMATICUM *35, Mons.: Short turf over coal waste, Cum Du,
SO2502, T.G. Evans, 1997, NMW, det. A.J. Richards.

135/25.93. TARAXACUM LANCIDENS *42, Brecs.: Meadow, Llangasty Tal-y-Llyn, SO1225, A.
J. Richards et al., 1996.

135/25.96. TARAXACUM PRIONUM *42, Brecs.: Lane verge, grassy bank, 3km N of Ponsticill,
SO0514, M. Porter, 1995, det. A.J. Richards.

135/25.124. TARAXACUM CURTIFRONS *42, Brecs.: Bank of old lane near Llanbedr,
S$O0225193, M. Porter, 1996, herb. M. P., det. A.J. Richards.

135/25.125. TARAXACUM CYANOLEPIS *35, Mons.: River Wye meadow covered by Spring
tides, N of Chepstow Castle, ST531951, T.G. Evans, 1997, NMW, det. A.J. Richards.

135/25.134. TARAXACUM EXSERTIFORME *35, Mons.: Forest road, N of Howick, ST5095,
Hans @llg. et al., 1996.

135/25.140. TARAXACUM HORRIDIFRONS *35, Mons.: Roadside, Cleddon, SO5103,
International Taraxacum Group, 1996, det. A.J. Richards, Hans @llg., et al.

135/25.159. TARAXACUM LUCIDUM _ *35, Mons.: Shady bank, Caerllan, SO4982, International
Taraxacum Group, 1996, det. A.J. Richards, Hans Ollg., et al. | *42, Brecs.: Pasture at edge of
Llangorse Lake, Llangasty Tal-y-Llyn, SO1326, M. Porter, 1995, det. A.J. Richards.

135/25.170. TARAXACUM NITIDUM ~~ *35, Mons.: Shady bank, Caerllan, SO4908, International
Taraxacum Group, 1996, det. A.J. Richards, Hans @llg., et al.

135/25.173. TARAXACUM OBTUSIFRONS *42, Brecs.: River bank, Llangynidr, SO1420, A.J.
Richards et al., 1996.

7135/25.189. TARAXACUM PSEUDORETROFLEXUM *35, Mons.: Cleddon Lane verge, Cleddon,
$O5103, T.G. Evans, 1996.

135/25.199. TARAXACUM SINUATUM *61, S.E. Yorks.: Neutral grassland, Hollym Cross,
TA3325, P.J. Cook, 1997, det. A.J. Richards.

135/25.215. TARAXACUM TUMENTILOBUM *35, Mons.: Roadside bank, New House
Roundabout, M48, ST5391, T.G. Evans, 1996, det. A.J. Richards.

135/25.bre. TARAXACUM BRECONENSE *35, Mons.: Short turf over coal waste, Cwm Du,
$O02502, T.G. Evans, 1997, NMW, det. A.J. Richards.

582 PLANT RECORDS

135/25.edm. TARAXACUM EDMONSONIANUM *42, Brecs.: Bank of old lane, Llangynidr,
SO1419, M. Porter, 1996, det. A.J. Richards.

135/25.ope. TARAXACUM OPERTUM H.@llg. ined. *35, Mons.: Forest roadside, N of Howick,
ST5095, Hans Ollg. et al., 1996.

135/26.3. CREPIS BIENNIS *46, Cards.: Abundant on roadside and field banks 500m SE of
Glynarthen, SN313482, A.O. Chater, 1997, NMW. 47, Monts.: Semi-improved neutral
grassland, Gweunydd Ty-Brith SSSI and Nature Reserve, Deuddwr, SJ243178, S.L. Smith & D.
Guest, 1997. 2nd record. 80, Roxburghs.: One plant on road verge, Threepwood Bridge,
NT519441, L. Gaskell, 1997, herb. M.E. Braithwaite. Ist record since 1878. +*81, Berwicks::
Perhaps established on road verge near old quarry, Threepwood Bridge, NT519441, L. Gaskell.
1997, herb. M.E.B.

+135/26.set. CREPIS SETOSA *35, Mons.: Near meadow boundary fence, near Llanfoist,
$0300119, T.G. Evans, 1991, NMW, det. R. Fraser.
7135/27.2x6. PILOSELLA OFFICINARUM x P. AURANTIACA (P. x STOLONIFLORA) *4 oN.

Devon: Grass, Parracombe churchyard, $S669448, W.H. Tucker, 1997, herb. L.J.M.. conf. L.J.
Margetts. *46, Cards.: E part of chapel graveyard, Tan-y-groes, SN284493, A.O. Chater, 1997,
CGE, det. P.D. Sell. This population agrees with subsp. schuriana Naegeli & Peter and appears to
be the first record for Britain for this nothomorph.

+135/27.3a. PILOSELLA FLAGELLARIS subsp. FLAGELLARIS *80, Roxburghs.: Old road, now
gated off, near A7 N of Galashiels, NT476378, J. Murray, 1995, herb. R.W.M. Corner, det. D.R.
McKean. *$1, Berwicks.: Established on road verge, Al near Grantshouse, NT814655, M.E.
Braithwaite, 1996.

135/28.3. HIERACIUM RIGENS *37, Worcs.: Occasional in sandy heathland, Devil’s Spittleful,
SO08074, SO8174, SO8175, J. Bevan & W.A. Thompson, 1991.

135/28.6a. HIERACIUM UMBELLATUM subsp. UMBELLATUM *49, Caerns.: Side of disused
railway, Caernarfon, SH4863, G. & L Battershall, 1996.
135/28.21. HIERACIUM TRICHOCAULON *37, Worcs.: Small colony in sandy heathland,

Devil’s Spittleful, SO8174, J. Bevan & W.A. Thompson, 1993.

135/28.34. HIERACIUM SPARSIFOLIUM *47, Monts.: Rocky banks of Afon Trannon, E of
Staylittle, SN9193, A.J. Morton, 1987, conf. J. Bevan.

135/28.67. HIERACIUM SUBAMPLIFOLIUM *37, Worcs.: Stone walls, Croft Bank, West
Malvern, SO7646, J. Bevan, P.G. Garner & W.A. Thompson, 1990.

135/28.78. HIERACIUM ORCADENSE *91, Kincardines.: Outcrop of Old Red Sandstone,
Dunnottar Castle, NO878838, D. Welch, 1995, herb. D.W., det. D.J. McCosh.

135/28.115. HIERACIUM SCOTOSTICTUM *37, Worcs.: Sandy laneside bank, Upper Rochford,
SO06367, R. Maskew & W.A. Thompson, 1994, det. J. Bevan.

135/28.123. HIERACIUM STENSTROEMII *37, Worcs.: Small colony in woodland, Armley
Bank, Broadway Hill, SP1136, J. Bevan & W.A. Thompson, 1994.

135/28.133. HIERACIUM SUBCRASSUM _*37, Worcs.: Broken wooded slope, The Dingle, West
Malvern, SO7645, J. Bevan, P.G. Garner & W.A. Thompson, 1996, det. P.D. Sell. Fourth British
record.

135/28.140. HIERACIUM EXOTERICUM *47, Monts.: Large population with other members of
this section, railway cutting, S of Talerddig, SN9399, A.J. Morton, 1989.

135/28.147. HIERACIUM CALEDONICUM *94, Banffs.: Serpentine outcrop, Craigs of Succoth,
NJ435363, D. Welch, 1992, ABD, det. D. McCosh.

135/28.165. HIERACIUM LASIOPHYLLUM *44, Carms.: One plant only on acid crag, Craig Allt-
y-berau, SN7746, J. Bevan, R.D. Pryce & G. Hutchinson, 1995.

135/30.1. FILAGO VULGARIS 46, Cards.: Dry bank by road, RAE Site, Aber-porth, SN251516,
A.O. Chater, 1997. 2nd extant locality.

135/30.4. FILAGO MINIMA _ 45, Pembs.: Hundreds to low 1,000s of plants scattered in short
open turf on small stones, Trecwn Burning Ground, Llanychaer, SM9934, S.B. Evans et al., 1996.
2nd record. 64, Mid-W. Yorks.: Hundreds of plants on floor of old sandstone quarry, Ireland
Woods, Leeds, SE254382, M. Wilcox, 1997. Ist record this century.

—————

PLANT RECORDS 583

135/33.6. GNAPHALIUM LUTEOALBUM *15, E. Kent: Sandy area at edge of newly dug pits,
Dungeness, TRO618, E.G. Philp, 1996, MNE. In 1997 several hundred plants & two further

colonies were present in the area. +*95, Moray: One plant on waste ground on Gallow Hill,
Hopeman, NJ149686, P.R. Green, 1997, herb. I.P. Green, conf. E.J. Clement.
7135/40.4. SOLIDAGO GIGANTEA *49, Caerns.: Village green (jettisoned garden relic?),

Bryncroes, SH2231, J.P. Bailey & A.P. Conolly, 1985, NMW.

+135/40.5. SOLIDAGO GRAMINIFOLIA *42, Brecs.: Naturalised on waste ground, Brynmawr,
S$O185116, M. Porter, 1997.

+135/41.3x4. ASTER LAEVIS x A. NOVI-BELGII (A. x VERSICOLOR) *46, Cards.: Saltmarsh by
Afon Rheidol, Aberystwyth, SN584811, A.O. Chater, 1993, CGE, conf. P.D. Sell & P.F. Yeo.
*49, Caerns.: Edge of neglected car park, Pwllheli, SH3734, J. Hawksford, 1997. *77, Lanarks.:
Well established in scrubby wood, Gartsherrie, NS7266, P. Macpherson, 1996, herb. P.M., det. P.
Feo:

+135/41.4. ASTER NOVI-BELGI 81, Berwicks.: Established in ditch by road, A1107, near Old
Cambus Wood, NT8368, M.E. Braithwaite, 1996. Large colony on road verge, A68 near Lauder
Barns, NT547463, M.E. Braithwaite, 1997, herb. M.E.B. Ist and 2nd post-1930 records and only
extant localities.

*4135/41.4x5. ASTER NOVI-BELGI x A. LANCEOLATUS (A. x SALIGNUS) *44, Carms.:
Roadside bank, Railway station approach, Kidwelly, SN401065, G. Hutchinson, 1995, NMW, det.
P.F. Yeo. Ist authenticated record.

+135/41.5. ASTER LANCEOLATUS *49, Caerns.: Scrub, edge of river, Pwllheli, SH3734, J.
Hawksford, 1997.

+135/41.con. ASTER CONCINNUS *46, Cards.: NE bank of Afon Cletwr, Tre’r-ddol,
SN654929; N bank of Afon Rheidol, Aberystwyth, SN587812; both A.O. Chater, 1990, CGE &
NMW,, det. P.F. Yeo & P.D. Sell. Ist and 2nd records.

+ 135/43.1. ERIGERON GLAUCUS *49, Caerns.: Naturalised at side of track near Aberdaron,
SH1626, G. & I. Battershall, 1996. Roadside verge, near Penmaenmawr, SH7176, G. & I.
Battershall, 1997.

7+135/43.4. ERIGERON KARVINSKIANUS 50, Denbs.: Lowest W facing crags above woodland,
W side of Bryn Euryn, Colwyn Bay, SH8379, R. Lewis, 1993, det. Mrs R. Lever. [The determiner
was eroneously given as Mrs R. Lewis in Watsonia 20(3): 298. ]

7135/44.1. CONYZA CANADENSIS *95, Moray: Two plants in set-aside field, Roseisle near
Duffus, NJ156668, I.P. Green, 1996. Many plants on waste ground on new housing estate, Forres,
NJ033588, P.R. Green, 1997. 1st and 2nd records.

7135/44.2. CONYZA SUMATRENSIS *12, N. Hants.: Rough grass near A331, Spring Lakes,
Aldershot, SU886516, C.R. Hall, 1997, herb. A. Brewis.

7135/45.2. OLEARIA MACRODONTA _*49, Caerns.: Probably planted originally on side of track,
Machroes near Abersoch, SH3226, G. & I. Battershall, 1997.

7135/48.2. TANACETUM MACROPHYLLUM _*77, Lanarks.: Well established between railings of
Botanic Gardens & River Kelvin, Kelvinside, Glasgow, NS5667, K. Watson, 1996, GLAM, det.
E @lement.

135/54.1. CHAMAEMELUM NOBILE 10, Wight: Non-flowering but frequent in mown amenity
grassland, Puckpool Park, Ryde, SZ613922, C.R. Pope, 1996, herb. C.R.P. 2nd extant site.

7135/57.1. LEUCANTHEMELLA SEROTINA 12, N. Hants.: Bridleway called Alresford Lane,
Cheriton, SU600292, R.M. Veall, 1996. 2nd record.

7135/58.lacxmax. LEUCANTHEMUM LACUSTRE x L. MAXIMUM (L. x SUPERBUM) *81,
Berwicks.: Large colony established on road verge, A697 E of Fireburnmill, NT824393, M.E.
Braithwaite, 1997.

135/60.1. TRIPLEUROSPERMUM MARITIMUM _ 47, Monts.: Saltmarsh, Morben-isaf, SN7098, T.
Kohler, T. Teearu & E. Lomas, 1988. 2nd record.

135/60.1x2. TRIPLEUROSPERMUM MARITIMUM x T. INODORUM *51, Flints.: Behind coastal
dunes, Talacre Warren, SJ1184, BSBI group, 1996.

584 PLANT RECORDS

+135/61.1. COTULA CORONOPIFOLIA 11, S. Hants.: Sandy shingle bank S side of Hermitage
Stream, Bedhampton Mill, SU704062, J.R.W. Hollins, 1997, herb. R.P.B., conf. R.P. Bowman.
2nd record.

+135/62.1. SENECIO CINERARIA *81, Berwicks.: Established at field edge, Burnmouth,
NT955611, J. Muscott, 1997.

135/62.10x11. SENECIO JACOBAEA x S. AQUATICUS (S. x OSTENFELDII) 50, Denbs.: Rough
field, Cerrig-y-druidion, SH952484, J.A. Geen, 1997. 2nd record.

135/62.12. SENECIO ERUCIFOLIUS *48, Merioneth: One plant with S. jacobaea, roadside bank,
near Trawsfynydd, SH7037, P.M. Benoit et al., 1994, NMW. Ist certain record.

135/62.13x15. SENECIO SQUALIDUS x S. VULGARIS (S. x BAXTERI) *46, Cards.: Garden,
Llanbadarn Fawr, SN598810, J.H. Salter, 1935, (Fl. Pl. Ferns Cardiganshire (1935)).

+135/62.3. SENECIO INAEQUIDENS _*77, Lanarks.: Unused cul de sac, King George V Dock,
Glasgow, NS5366, P. Macpherson, 1995, herb. P.M., det. R.M. Burton. One plant 1995, three
plants 1996 and 1997.

+135/66.comxlax. BRACHYGLOTTIS COMPACTA x B. LAXIFOLIA (B. ‘SUNSHINE’) *46, Cards.:
Gully on sea cliffs below Grogal, W of New Quay, SN373593, A.O. Chater & A.P. Fowles, 1989,
BM, conf. D.H. Kent. *77, Lanarks.: Grassy, open area near river, Meadowside, Glasgow,
NS5566, P. Macpherson, 1988, herb. P.M., det. D. McClintock.

+135/69.1x2x3. DORONICUM PARDALIANCHES x D. PLANTAGINEUM x D. COLUMNAE (D. x
EXCELSUM) *67, S. Northumb.: Roadside verge, Acumb Fell, NY955675, G.A. Swan, 1996,
herb. G.A.S., det. P S. Green.

7135/71.2. PETASITES JAPONICUS *S, Channel Islands: Small shaded valley beside stream,
Harbour Hill, Sark, WV473758, R.M. Veall, 1988, det. D. McClintock. Apparently planted — seen
in 1988 by Botanical Group from Guernsey — still present and spreading.

*135/73.1. CALENDULA OFFICINALIS 31, Flints.: Waste ground, W end of Marsh Road, Rhyl,
SJ002803, J. Phillips, 1997. 2nd record and Ist since 1942.

*135/73.1. CALENDULA OFFICINALIS *43, Rads.: Spoilheap in lay-by at roadside, between
Newbridge & Disserth, SO030589, R.G. Woods, 1980s, NMW.

*135/78.1. RUDBECKIA HIRTA *41, Glam.: Coal dump, Cardiff Docks, ST197749, G.
Hutchinson, 1988, NMW.

+135/79.1. HELIANTHUS ANNUUS *49. Caerns.: Demolition site, Llandudno, SH784821, W.
McCarthy, 1995.

+135/79.2xrig. HELIANTHUS TUBEROSUS x H. RIGIDUS (H. x LAETIFLORUS) *81, Berwicks::
Over 2m up wall, Lauder, NT528477, M.E. Braithwaite, 1997.

+135/81.3. BIDENS CONNATA var. ANOMALA _*58, Cheshire: Canal bank, Sandbach, $J732505,
G.M. Kay, 1997, herb. G.M.K., conf. E.J. Clement. Canal bank, Hassall Green, SJ778584, G.M.
Kay, 1997. 1st and 2nd records.

135/SCO.his. SCOLYMUS HISPANICUS *46, Cards.: Two plants by waste ground, N side of
Afon Rheidol, Aberystwyth, SN586811, R. Birch, 1997.

138/2.1. STRATIOTES ALOIDES *5, S. Somerset: Many plants in pond in Wayford Woods,
Wayford, ST399064, I.P. Green, 1997. +*35, Mons.: Planted in artificial pond in Water Park
Pond, near Ahtophon Factory, St Mellons, ST249815, G. Hutchinson, 1994. Pond in Industrial
Park, Pwll-mawr, ST223787, T.G. Evans, 1996. 1st records. A series of ponds and lakes have been
created between Newport and Cardiff with some strange plantings.

+138/4.2. ELODEA NUTTALLII *49, Caerns.: Margin of Llyn Padarn, Llanberis, SH5662, A.
Jones, 1997. *95, Moray: Very plentiful in pond, Sunbank Park, Lossiemouth, NJ231694, I.P.
Green, 1996.

+138/6.1. LAGAROSIPHON MAJOR *46, Cards.: In two ponds in grounds of Ffynnon Berw,
lkm NNW of Ffynnonddewi, SN381538. A.O. Chater, 1994. Upland ponds 2km NNE of Ffair-
thos, SN750697, M.D. Sutton, 1997. 1st and 2nd records.

141/1.1. TRIGLOCHIN MARITIMA 47, Monts.: Dovey Junction, SN7098, M. Hogan & C.
Fieldhouse, 1986. 2nd record.

PLANT RECORDS 585

142/1.3. POTAMOGETON COLORATUS *44, Carms.: Pond in sand dunes created about 5 years
previously, Pembrey Country Park, SN3900, N.F. Stewart, 1996.

142/1.5x6. POTAMOGETON LUCENS x P. GRAMINEUS (P. x ZIZII) *41, Glam.: SW part of
Kenfig Pool, SS796812, N.F. Stewart, 1996.

142/1.6x9. POTAMOGETON GRAMINEUS x P. PERFOLIATUS (P. x NITENS) *41, Glam.: Pool on
Margam Dunes, SS782838, N.F. Stewart, 1996.

142/1.7. POTAMOGETON ALPINUS 81, Berwicks.: Pond, Eden water at Nenthorn House,
NT673371, M.E. Braithwaite, 1995, herb. M.E.B., det. N.T.H. Homes. Large colony in slow
moving burn in fen, Lauder Burn, NT516452, M.E. & P.F. Braithwaite, 1997, herb. M.E.B. Ist
and 2nd recent records and only extant localities.

142/1.16. POTAMOGETON TRICHOIDES *4, N. Devon: Portledge Pool, SS392245, W.H.
Tucker, 1996, conf. L.J. Margetts.

142/1.19. POTAMOGETON CRISPUS +*46, Cards.: Pond in pasture 200m N of Clogfryn,
Aberaeron, SN449624, A.O. Chater, 1996, NMW, conf. C.D. Preston.

142/1.20x21. POTAMOGETON FILIFORMIS x P. PECTINATUS (P. x SUECICUS) 93, N. Aberdeen:
Shallow water, Loch of Strathbeg, NK0858, C.D. Preston & P.M. Hollingsworth, 1994. Ist post-
1930 record.

143/1.1. RUPPIA MARITIMA *47, Monts.: Many plants found in four different brackish pools in
salt-marsh, Dovey Junction, SN695985, F. Evans, E. Gwynn & A. Law, 1993.

+147/1.1. ACORUS CALAMUS 4, N. Devon: Pond edge, Simpson, Holsworthy, SS364038, W.
H. Tucker, 1997. 2nd post-1930 record.

+147/2.1. LYSICHITON AMERICANUS =*12, N. Hants.: Riverside Railway Walk, River Rother,
Liss, SU780283, J. Ockenden, 1997, det. F. Rose. *91, Kincardines.: Roadside ditch, Durris,
NO785968, D. Welch, 1997. Probably escaped from Durris House, about 1km away. Shingle
terrace on island in River Dee, Maryculter, NO815983, D. Welch, 1997. Ist and 2nd records.

+147/5.2b. ARUM ITALICUM subsp. ITALICUM _*41, Glam.: Caswell, SS5897, A.S. Lewis 1997.

148/1.1. SPIRODELA POLYRHIZA 41, Glam.: Abandoned Carboniferous Limestone sheep-dip,
Tythegston, SS864781, P.S. Jones & S. Thomas, 1997. Abundant in a concrete drinking-pond in
the SW corner of a field near Tresilian, SS953680, Q.O.N. Kay, 1997. Ist recent records.

148/2.3. LEMNA TRISULCA *45, Pembs.: Abundant with small patch of Crassula helmsii in
shady shallow pond on slope in woodland, Pwll Dyfrig, Glyn-y-mel, Lower Fishguard,
SM967371, S.B. Evans et al., 1997.

+148/2.4. LEMNA MINUTA —_ *41, Glam.: Locally abundant, with L. minor, in shallow water in
moat & fish pond in Botanic Garden, University College of Swansea, SS6291, Q.O.N. Kay, 1996.
46, Cards.:. Garden pond, Old Castle Farm, Cardigan, SN165463, A.O. Chater, 1996. 2nd record

151/1.13x14. JUNCUS ARTICULATUS x J. ACUTIFLORUS (J. x SURREJANUS) *71, Man: Isolated
pond, Ballavarran Dub, SC365972, B.A. Tregale, 1999, DGS.

151/1.25x26. JUNCUS EFFUSUS x J. INFLEXUS (J. x DIFFUSUS) *50, Denbs.: Wet ground near
lake, Painters Green, SJ502499, J.A. Green, 1997, NMW.

151/1.6. JUNCUS FOLIOSUS *35, Mons.: Wet meadow, Ty’r Sais Farm, near Pen-y-fan Pond,
SO1800, T.C.G. Rich, P.A. Smith & S.E. Erskine, 1996. *108, W. Sutherland: Wet trackside,
Aultanrynie, Loch More, NC349356, C. Dixon, 1997.

151/1.8. JUNCUS AMBIGUUS 47, Monts.: Frequent in upper saltmarsh, Dovey Junction, with
J. bufonius, SN695983, P.M. Benoit, 1987. 2nd record. *51, Flints.: Damp depression in
disturbed dunes, Point of Ayr, SW of lighthouse, SJ1185, A.O. Chater, 1996, det. T. Cope.

*151/2.4. LUZULA LUZULOIDES 81, Berwicks.: Open woodland, Castle hill, Thirlstane Castle,
NT534477, M.E. & P.F. Braithwaite, 1997. Only extant locality.

152/11.1. CYPERUS LONGUS +*44, Carms.: SW corner of lake margin, Sandy Water Park,
Llanelli, SN493004, I.K. Morgan, 1995. Source unknown, lake constructed c.5 years ago.

152/16.1x15. CAREX PANICULATA x C. REMOTA (C. x BOENNINGHAUSIANA) *80,
Roxburghs.: Side of ditch, Linton Loch, Morebattle, NT793254, O.M. Stewart, 1997, E.

586 PLANT RECORDS

152/16.5. CAREX OTRUBAE 109, Caithness: Coastal turf, near Rattar, ND2574, R.E.C.
Ferreira, 1995. 1st localised record.

152/16.5x15. CAREX OTRUBAE x C. REMOTA (C. x PSEUDOAXILLARIS) *45, Pembs.: Grassy
verge of track (re-surfaced for motors) with C. remota, Cardigan Wildlife Park (entrance road
from Cilgerran), SN185436, R.W. David, 1978, NMW, conf. A.O. Chater. The second parent (C.
otrubae) is frequent on the banks of the River Teifi nearby.

152/16.9b. CAREX DIVULSA subsp. LEERSII *73, Kirkcudbrights.: Few plants on bank of waste
ground and rough grassland, Castle Douglas, NX750624, O.M. Stewart, 1996, E, det. D.A.
Pearman & A.O. Chater. 81, Berwicks.: One large clump in flower under beech, The Lees,
NT/840390, M.E. & P.F. Braithwaite, 1997, herb. M.E.B. 2nd extant record. Status unknown.

152/16.21. CAREX LACHENALII *94, Banffs.: Cairngorms, NH994010, J. Edelsten, 1996, E,
det. A.C. Jermy.

152/16.25x26 CAREX ACUTIFORMIS x C. RIPARIA (C. x Sool) *25, E. Suffolk: Large stand,
Framlingham Mere, TM285637, A.C. Jermy, 1990. Ist record for Suffolk.

152/16.27. CAREX PSEUDOCYPERUS *49, Caerns.: Fen, Cors Llyferin S of Abersoch, SH3126,
D. Jones, 1995, NMW.

152/16.28x29. CAREX ROSTRATA x C. VESICARIA (C. x INVOLUTA) *80, Roxburghs.: With
both parents in fen, Tilery Moss, SW of Muirhouse Law, NT623274, R.W.M. Corner, 1997, herb.
R.W.M.C., det. A.O. Chater.

152/16.31. CAREX PENDULA 81, Berwicks.: Spring in former woodland, Eye Water above
Grantshouse, NT810650, M.E. Braithwaite, 1996. 2nd extant record. 91, Kincardines.: Marsh in
woodland, Maryculter, NJ860999, D. Welch, 1997. Ist definite post-1930 record.

152/16.39. CAREX LAEVIGATA *89, E. Perth: Bridge of Cally, NO15, J.H. Penson, 1960,
herb. J.H.P., conf. A.McG. Stirling.

152/16.44x46. CAREX HOSTIANA x C. VIRIDULA (C.x FULVA) 83, Midlothian: Marsh, S of
Gladhouse Reservoir, NT296502, C. Dixon, 1997, E, det. D.R. McKean. 2nd record.

152/16.44x46a. CAREX HOSTIANA x C. VIRIDULA subsp. BRACHYRRHYNCHA (C. x FULVA)
*79, Selkirks.: With both parents in basic flush, Roughhope Burn, Philhope, Borthwick Water,
NT383078, R.W.M. Corner, 1997, herb. R.W.M.C., det. A.O. Chater. *80, Roxburghs.: With
both parents in basic flush, Grindstone Burn, Upper Hindhope, Kale Water, NT768086, R.W.M.
Corner, 1997, herb. R.W.M.C., det. A.O. Chater.

152/16.59a. CAREX MAGELLANICA subsp. IRRIGUA *49, Caerns.: Few plants in swamp with
Carex rostata, Mignient near Ysbyty Ifan, SH7744, G. Battershall, 1996.

152/2.2b. TRICHOPHORUM CESPITOSUM subsp. CESPITOSUM *108, W. Sutherland: Schoenus
nigricans mire, Ceathramh Garbh, Rhiconich, NC228515, R.W.M. Corner, 1997, conf. G.A.
Swan.

152/2.2axb. TRICHOPHORUM CESPITOSUM subsp. CESPITOSUM x subsp. GERMANICUM
(proliferous form) *46, Cards.: Small flush with Carex dioica, Rhynchospora alba, etc., Bryn
Mawr, SE of Hafod, SN771717, A.O. Chater, 1963, NMW, det. G.A. Swan.

153/12.9. FESTUCA VIVIPARA *93, N. Aberdeen: Trackside in heather moorland, Hill of
Millmedden, NJ5323, D. Welch, 1997, ABD, conf. C.A. Stace.

153/12.10. FESTUCA FILIFORMIS 47, Monts.: Hummocks in boggy field, NE of Llanfihangel-
yng- ngwynfa, SJO915, P.M. Benoit, 1989. 2nd record.

153/12.4. FESTUCA ALTISSIMA .*43, Rads.: In quantity on rock outcrops & decaying wood,
Bach Howey Gorge, $SO119435, R.G. Woods & M. Porter, 1980s. (81, Berwicks.: Cromwells.
Brunta Burn, NT596505, M.E. Braithwaite, 1992, det. D.R. McKean. Delete record in Watsonia
19: 293 (1993). The plant has flowered and is Elymus caninus, M.E. Braithwaite, 1995, det. D.R.
McKean. ]

*153/12.5. FESTUCA HETEROPHYLLA *28, W. Norfolk: Edge of woodland ride, West Briggs,
TF656105, S. Fisher, 1997, conf. R.M. Payne. *95, Moray: Well naturalised in grassy area
under trees, in grounds of small chapel at Gordonstown School, Duffus, NJ192689, I.P. Green,
1996, conf. R.M. Burton.

PLANT RECORDS 587

153/12.6. FESTUCA ARENARIA *44, Carms.: Sparse population on seaward Ammophila
arenaria ridge, Pendine Ranges, Laugharne Burrows, SN311071, A.O. Chater, 1997, NMW.

153/12.7b. FESTUCA RUBRA subsp. JUNCEA *46, Cards.: Storm beach W of road, Tan-y-bwlch
Beach, Aberystwyth, SN579805, S.P. Chambers, 1994, herb. S.P.C., det. C.A. Stace. Rocky sea
cliff, NNW side of Foel y Mwnt, SN192521, A.O. Chater, 1994, NMW. Ist and 2nd records.
*49_. Caerns.: Cliff-top turf, Porth Ceriad, near Abersoch, SH3124, M. & A. Atkinson, 1997.

153/12.7c. FESTUCA RUBRA subsp. LITORALIS *46, Cards.: Saltmarsh E of Afon Leri, Ynys-
las, SN616930, A.O. Chater, 1993. Saltmarsh, Teifi estuary, 1km S of Nantyferwig, SN168480, A.
O. Chater, 1997, NMW. Ist and 2nd records.

153/12.7d. FESTUCA RUBRA subsp. COMMUTATA *44, Carms.: Llansteffan Dunes, Llansteffan,
SN31, H. Sealy-Lewis, 1987.

153/12.7g. FESTUCA RUBRA subsp. MEGASTACHYS *44, Carms.: Roadside verge, near
Edwinsford, SN635345, A.O. Chater, 1995. Roadside near Maesteilo, Capel Isaac, SN592265, J.
A. Green, D.M. George & E. Stephenson, det. A.O. Chater, 1995, NMW. Ist and 2nd records.
*46, Cards.: Grassy slope in village, Ponterwyd, SN751807, A.O. Chater, 1988, NMW, det. A.K.
Al-Bermani. Laneside verge just N of Pont Pant-mawr, 2km WNW of Llanilar, SN607757, A.O.
Chater, NMW, 1992. Ist and 2nd records.

153/12.8c. FESTUCA OVINA subsp. OPHIOLITICOLA *44, Carms.: Rocky calcareous grassland,
Pal-y-Cwrt, SN678182, A.O. Chater et al., 1997.

153/12.9. FESTUCA ‘VIVIPARA *93, N. Aberdeen: Trackside in heather moorland, Hill of
Millmedden, NJ530230, D. Welch, 1997, ABD, conf. C.A. Stace.

7153/12.gau. FESTUCA GAUTIERI *3, S. Devon: Road embankment, Exeter, SX926927, R.
Takagi-Arigho, 1990, herb. R.T.-A., det. T.A. Cope.

153/12.1x13.1. FESTUCA PRATENSIS x LOLIUM PERENNE (x FESTULOLIUM LOLIACEUM) *50,
Denbs.: Wet meadows, near Overton, SJ396425, D. Tinston, 1987, NMW, det. P.M. Benoit.

153/12.2x13.1. FESTUCA ARUNDINACEA x LOLIUM PERENNE (x FESTULOLIUM HOLMBERGII)
*28, W. Norfolk: Wiggenhall St Mary, TF575142, R.M. Payne, 1992, conf. R.M. Payne & A.
Copping. 2nd record for Norfolk.

153/12.2x13.2. FESTUCA ARUNDINACEA x LOLIUM MULTIFLORUM *11, S. Hants.: Bishop’s
Palace, Bishop’s Waltham, SU553175, P.D. & C.A. Stanley, 1997, herb. R.P. Bowman, det. E.J.
Clement.

153/12.7x14.3. FESTUCA RUBRA x VULPIA MYUROS 35, Mons.: By railway line, Mitchell
Troy, SO4910, E. Nelmes, 1951, K, det. C.A. Stace. Precedes TGE’s record of 21/7/1988 as Ist
record. C.A. Stace writes ‘This .... is the earliest from any part of the world.’

153/14.1. VULPIA FASCICULATA *51, Flints.: Fixed sand dunes, Point of Ayr, $J123851, J.A.
Green, 1985, NMW, det. G. Hutchinson.

153/14.3. VULPIA MYUROS *89, E. Perth: Waste ground, Invergowrie, NO349300, A.A.
Slack, 1980. Virtually on boundary between v.cc 89 & 90. Apparently Ist localised record.

153/16.2. PUCCINELLIA DISTANS +*77, Lanarks.: Verge of A73, N of Newmains, NS8256,
herb. M.E.B. Road verge of A74, Melbourne, NT0844; both M.E. Braithwaite, 1996. 1st and 2nd
records.

153/16.2b. PUCCINELLIA DISTANS subsp. BOREALIS *95, Moray: Road verges, Speybridge,
NJ040261, P.R. Green, 1997, conf. H. Bowen. Plentiful along many of the roads and also on
saltmarshes; greatly on the increase.

153/18.4. POA HUMILIS *61, S.E. Yorks.: On old brick wall, Sledmere, SE9364, M.E.
Braithwaite, 1997. On old brick wall, Patrington, TA3122, P.J. Cook, 1997. 1st and 2nd records.
Subsequently found on other brick walls in Holderness.

7153/18.7. POA CHAIxIE *43, Rads.: Among trees, Rock Park, Llandrindod Wells, SO055609,
R.G. Woods, 1986. 47, Monts.: Park woodland under beech, Berriew, SJ1900, E. Roberts &
M. Wainwright, 1989. 2nd record; possibly planted for pheasant cover.

153/20.1. CATABROSA AQUATICA 91, Kincardines.: Muddy pool close to HWM, Nigg Bay,
NJ967044, D. Welch, 1997, ABD. Ist post-1930 record.

588 PLANT RECORDS

153/21.2. CATAPODIUM MARINUM *35, Mons.: On exposed former shelly beach on River
Severn bank, Rumney Great Wharf, ST248786, P. Millman, et al., 1997. Though looking native in
a suitable habitat, much material has been imported to build up nearby sea wall and status is
uncertain.

153/22.1. SESLERIA CAERULEA *35, Mons.: Chepstow, ST59, T.G. Evans, 1988, NMW.

153/24.1. GLYCERIA MAXIMA 46, Cards.: Dominant in pond in fields 300m SSW of Coed-y-
cwm, Cwm Cou, SN275428, A.O. Chater, 1996. 2nd record.

+153/28.2. AVENA FATUA *46, Cards.: Several plants on disturbed ground, Ffair-Rhos,
SN743679, A.O. Chater, 1991, NMW.

+153/29.1. GAUDINIA FRAGILIS 11, S. Hants.: Horse pasture in closed sward of fairly rich area
of grassland, on clay, Hounsdown, Eling, SU353117, J. Rowe, 1997, herb. R.P.B., conf. R.P.
Bowman. 2nd record.

+153/30.1pur. TRISETUM FLAVESCENS subsp. PURPURASCENS *46, Cards.: Grassy slope by
pond, Gwern-medd, Llangoedmor, SN213454, A.O. Chater & L. Gander, 1997, NMW.

+153/37.2. ANTHOXANTHUM ARISTATUM *12, N. Hants.: Roadside B2070, Petersfield,
SU752240, P. Stanley, 1994, herb. E.J.C., det. E.J. Clement. Also present in 1995.

153/39.1x4. AGROSTIS CAPILLARIS x A. STOLONIFERA (A. x MURBECKII) 46, Cards.: River
shingle by Afon Ystwyth downstream of road bridge, Llanfarian, SN589777, S.P. Chambers,
1997. 2nd record.

153/40.2. CALAMAGROSTIS CANESCENS 80, Roxburghs.: Edge of fen, Linton Loch,
Morebattle, NT794255, R.W.M. Corner, 1997, herb. R.W.M.C., 2nd extant locality.

+153/43.1. LAGURUS OVATUS *49, Caerns.: Waste ground, Pwllheli, SH364341, G.
Battershall & W. McCarthy, 1996.

153/45.1. MIBORA MINIMA *59, S. Lancs.: In low fore-dune, NE end of Marine Lake,
Southport, SD337185, D.P. Earl & J. Buckley-Earl, 1996, conf. T.A. Cope.

153/46.1. POLYPOGON MONSPELIENSIS *58, Cheshire: Old sludge bed and tracks, Wharton,
Winsford, SJ656677, J.P. Guest, 1997, herb. G.M. Kay, det. E.J. Clement. +*95, Moray:
Several plants on waste ground, Hopeman, NJ149686, I.P. Green, 1997.

+153/48.1. BECKMANNIA SYZIGACHNE *59, S. Lancs.: Ditch bank between Marsh Farm and
River Douglas, Hesketh Bank, SD452240, D.P. Earl, 1996, conf. T.A. Cope.

153/50.3. BROMUS RACEMOSUS 51, Flints.: Hay meadow, Coed Duon, Tremeirchion,
SJ073715, A.O. Chater, 1997. 1st post-1930 record.

+153/50.5. BROMUS LEPIDUS 12, N. Hants.: Grassy part of garden, Warren Road, Liss Forest,
SU7828, det. F. Rose. Sandy tank tracks, Hogmoor Enclosure, SU7834; both A. Brewis, 1997,
herb. A.B. 1st and 2nd recent records.

*+153/50.7. BROMUS SECALINUS “*11, S. Hants.: Edge of bean field, S of Twyford, SU485232,
P.J. Selby, 1997, herb. R.P.B., conf. R.P. Bowman & E.J. Clement.

*153/52.2. ANISANTHA RIGIDA *29, Cambs.: Large patches by fence between sown area and
old shelter belt, SE side Magog Down, TL490531, S.M. Walters, 1997, CGE, det. D.A. Wells.

¥153/52.4. ANISANTHA TECTORUM ~~ *58, Cheshire: New earth bank, Moreton, $J263901, P.G.
Gutteridge, 1997, herb. G.M.K., conf. G.M. Kay.

7153/52.5. ANISANTHA MADRITENSIS *12, N. Hants.: Among rubbish, Woolmer Trading
Estate, SU795350, A. Brewis, 1997, herb. A.B.

+153/53.1. CERATOCHLOA CARINATA — *95, Moray: Very well naturalised on road verge and
edge of arable field, Roseisle, NJ138670, P.R. Green, 1997. Well naturalised on verge of lane,
Duffus, NJ174687, P.R. Green, 1997. Ist and 2nd records.

153/53.3. CERATOCHLOA CATHARTICA 51, Flints.: College grounds, Welsh College of
Horticulture, Northop, SJ2369, J.A. Green & BSBI group, 1996. Ist post-1930 record.

153/54.1. BRACHYPODIUM PINNATUM _ 89, E. Perth: Ungrazed grassland on sloping calcareous
gravels, banks of River Tay near Boat of Murthly, NO0O57396, N.W. Taylor, 1992, PTH, det. T.A.
Cope.

PLANT RECORDS 589

153/56.1x3. ELYTRIGIA REPENS x E. JUNCEA (E. x LAXA) 3, S. Devon: At head of creek,
Salcombe, Lower Batson, SX735397, L.J. Margetts & L.M. Spalton, 1997, det. T.A. Cope. At
head of saltmarsh creek, River Erme, Holbeton, SX619491, L.M. Spalton, 1997, det. T.A. Cope.
Ist and 2nd post-1930 records.

153/56.2. ELYTRIGIA ATHERICA *12, N. Hants.: Sown on sandy waste ground, Bordon,
SU7936, A. Brewis, 1997, herb. A.B. 73, Kirkcudbrights.: Shore of marsh, Kirkdale,
NX518527, O.M. Stewart, A. White & C. Jermy, 1996. Ist recent record.

7153/59.1. HORDEUM DISTICHON *47, Monts.: Farm manure heap, SE of Llanwyddyn,
S10213> CJ: Tyas, 1986.

7153/59.3. HORDEUM JUBATUM *42, Brecs.: Well established on verges of A465, between
Brynmawr and Rhymney, SO1812, T.G. Evans, 1997. 81, Berwicks.: Established on road verge,
A698 below Lochton, NT777385, M.E. & P.F. Braithwaite, 1995, herb. M.E.B. 2nd record.

+153/60.1. TRITICUM AESTIVUM *S, Channel Islands: Roadside, La Vaurocque, Sark,
WV461757, M. Marsden, 1982, JSY. *46, Cards.: Field crop, 300m W of Tegfan, near
Llechryd, SN226451, A.O. Chater, 1992, NMW.

7153/62.1. CORTADERIA SELLOANA *49, Caerns.: Edge of pool, Spinnies Nature Reserve,
near Bangor, SH613721, G. Battershall, 1994.

7153/62.ric. CORTADERIA RICHARDII *44, Carms.: Waste ground, below Cwmbach Road,
Llanelli, SN498012, G. Hutchinson & I.K. Morgan, 1994, det. T.B. Ryves. *46, Cards.: Gorse
slope above tip, Pendinas, Aberystwyth, SN584799, A.O. Chater, 1993, NMW. Streambank 100m
S of Ynys-hir bridge, Eglwys-fach, SN684956, P. Condry, 1995, det. A.O. Chater. 1st and 2nd
records.

7 153/65.cil. ERAGROSTIS CILIANENSIS *29, Cambs.: In scattered bird seed, near Kidman’s
Plantation, Whittlesford, TL447488, A. Arbon, 1994, conf. R.M. Payne.

7153/68.1. ECHINOCHLOA CRUS-GALLI *49, Caerns.: Pavement at base of wall, Llandudno,
SH794819, W. McCarthy, 1995. Roadside, Bethesda, SH6066, J. Hawksford, 1997. Ist and 2nd
records.

7153/68.col. ECHINOCHLOA COLONA *4, N. Devon: Garden paving, Instow, SS473303, M.
Tulloh, 1989, det. T.A. Cope. Probably introduced in bird-seed.

7 153/68.esc. ECHINOCHLOA ESCULENTA *39, Staffs.: Waste ground by RSPB centre, by River
Tame, 3km E of West Bromwich, SP035927, J.P. Martin, 1989. *44, Carms.: Shingle of river,
Afon Lliedi, near Buckley’s Brewery, Llanelli, SN509006, I.K. Morgan, 1991, NMW, det. G.
Hutchinson. 2nd Welsh record.

7153/70.pum. SETARIA PUMILA *49, Caerns.: Base of wall, Jubilee Street, Llandudno,
SH7831, W. McCarthy, 1997.

7153/71.san. DIGITARIA SANGUINALIS 39, Staffs.: Two plants on roadside, Railway Drive,
Madeley, SJ772441, I.J. Hopkins, 1995. 1st record since 1932.

7153/FAR.spa. FARGESIA SPATHACEA *S, Channel Islands: Roadside verge, near fence,
Dixcart Lane, near Dixcart Hotel, Sark, WV465754, R.M. Veall, 1997, JSY, det. D. McClintock.
Planted; one clump now about 1-5m tall. *46, Cards.: Several large thickets in damp estate

woodland, Highmead, Llanybydder, SN499431, A.O. Chater, 1985, NMW, det. C. Stapleton.
[previously erroneously published as 153/1.1. Semiarundinaria anceps (under Arundinaria
jansaurensis), Watsonia 16:196 (1986) and BSBI Welsh Bull. 44: 29(1986)]. Colony in wood near
gardens, Y Glog, Rhydyfelin, SN593793, A.O. Chater, 1985, NMW. Ist and 2nd records.

7153/PAN.sch. PANICUM SCHINZII *29, Cambs.: Two plants by grain silo, Newmarket,
TL646629, G.MLS. Easy, 1984.

154/1.1b. SPARGANIUM ERECTUM subsp. MICROCARPUM _*44, Carms.: Ditch on boundary of
two farms, Carway, SN463058, R.D. Pryce, 1979, NMW.

154/1.1d. SPARGANIUM ERECTUM subsp. OOCARPUM *3, S. Devon: Stream, Thorverton
Village, SS927021, L.J. Margetts, 1997.

590 PLANT RECORDS

155/1.1x2. TYPHA LATIFOLIA x T. ANGUSTIFOLIA (T. x GLAUCA) *12, N. Hants.: On the
marshy edge of the promontory that extends into the pond from the W bank (probably acidic),
Shortheath Pond, SU775368, B.A. Gale, 1985, herb. B.A.G., det. A.C. Leslie. Possibly introduced
by an angling club. *37, Worcs.: With both parents in small reed-swamp, Broadwaters,
Kidderminster, SO8478, W.A. Thompson & B. Westwood, 1994, conf. M. Briggs.

1156/5.2. HEMEROCALLIS LILIOASPHODELUS *77, Lanarks.: Rough grass at base of scrubby
trees, East Flatt, East Kilbride, NS6551, P. Macpherson, 1995, herb. P.M., det. E.J. Clement.
Presumably a result of fly-tipping; obviously surviving (still present in 1997).

+158/6.1. KNIPHOFIA UVARIA __*46, Cards.: One non-flowering plant among gorse on heathy
ground among leadmine spoil heaps, Cwmsymlog, SN698837, A.O. Chater, 1996. *81,
Berwicks.: Established where cast out in rough ground near railway, Burnmouth, NT955610, V.
Muscott, 1997.

7158/6.2. KNIPHOFIA PRAECOX *46, Cards.: Scrub on streambank W of bridge by
Llansantffraed church, SN512674, S.P. Chambers, 1997.

158/9.1. GAGEA LUTEA = +95, Moray: Hundreds of plants on W bank of River Spey, Orton, S
of Fochabers, NJ317523, P.R. Green, 1996. 2nd record, Ist native site.

+158/12.bulxmac. LILIUM BULBIFERUM x L. MACULATUM (L. x HOLLANDICUM) *58,
Cheshire: Sand dunes, Hoylake, $J207877, G.M. Kay, 1997. A small colony has developed from
original introduction.

158/20.2x3. HYACINTHOIDES NON-SCRIPTA x H. HISPANICA *46, Cards.: Scrub slope W of
church, Llanbadarn Fawr, SN598810, S.P. Chambers & A.O. Chater, 1991.

*158/22.1. CHIONODOXA FORBESII *26, W. Suffolk: Old garden, now plantation of beech
trees, Ousden, TL735595, R. Addington, 1988, det. by a local gardener. *49, Caerns.:
Naturalised on roadside, Bodafon Lane, Llandudno, SH8081, W. McCarthey, 1997. *Q9.
Dunbarton: Well established in wood with rich loamy soil, behind Cardross Park, NS3477, A.
McG. Stirling & A. Rutherford, 1985.

7158/23.1. MUSCARI NEGLECTUM *46, Cards.: Grassy roadside slope, Fron-deg, Llanbadarn
Fawr, SN600811, A.O. Chater, 1993. Scrub slope W of church, Llanbadarn Fawr, SN598810, S.P.
Chambers, 1997. Ist and 2nd records.

7158/23.2. MUSCARI ARMENIACUM *46, Cards.: Grassy roadside verge, road to Upper Forest,
Lampeter, SN576490, A.O. Chater, 1992. Copse by Primrose Hill, Llanbadarn Fawr, SN600811,
A.O. Chater, 1993. 1st and 2nd records.

7158/24.12. ALLIUM CARINATUM *$1, Berwicks.: Established on limestone rocks, Birgham »

Haugh Wood, NT798389, M.E. Braithwaite, 1995, herb. M.E.B.

7158/24.2. ALLIUM UNIFOLIUM *99, Dunbarton: Spreading well in damp, basic soil in
deciduous woods, behind Cardross Park, NS3477, A.McG. Stirling & A. Rutherford, 1985, det C.
D. Brickell. Considered first British escape.

158/31.1. LEUCOJUM AESTIVUM ~~ +*89, E. Perth: Among Phragmites in Tay Marshes, Perth,
NO129217, O.M. Stewart, 1982, E.

158/31.2. LEUCOJUM VERNUM 8, S. Wilts.: Boggy ground near pond, Britford Green near
Salisbury, SU1627, M. Marks, 1991, det. D.P.E. Stevens. 1st record for 90 years. +*49, Caerns:.:
7 clumps by footpath, Glynllifon Park near Caernarfon, SH4555, D. Jones, 1991.

+158/32.1x3. GALANTHUS NIVALIS x G. PLICATUS *46, Cards.: Damp grassland, Aberstringell
limekilns, Llanrhystud, SN519684, A.O. Chater, 1995, NMW. *95, Moray: Kincorth Wood
with both parents, Kintessack, NJO11617, P.R. Green, 1997.

7 158/32.4a. GALANTHUS PLICATUS subsp. PLICATUS *95, Moray: Well naturalised in Kincorth
Wood, Kintessack, NJO11617, P.R. Green, 1997.

7158/32.1ka. GALANTHUS IKARIAE *95, Moray: Naturalised in waste ground, N of Forres,
NJ026608, P.R. Green, 1996.

+158/33.3. NARCISSUS POETICUS *46, Cards.: Sea cliff at W side of Aber-porth, SN257515,
A.O. Chater, 1992. Roadside hedgebank, Clogfryn, Aberaeron, SN449623, A.O. Chater, 1994. Ist
and 2nd records. °

PLANT RECORDS 591

+158/33.3a. NARCISSUS POETICUS subsp. POETICUS *46, Cards.: Copse NW of Ffosrydygaled,
Chancery, SN578765, A.O. Chater, 1994. Roadside bank at NE end of Llwyncelyn, SN443598, A.
O. Chater, 1995. 1st and 2nd records.

+158/33.3b. NARCISSUS POETICUS subsp. RADIIFLORUS *46, Cards.: Abundant under trees
100m E of Lovesgrove mansion, SN629816, A.O. Chater, 1995. Hedgebank of road by
Tanycastell, Llanychaiarn, SN588788, A.O. Chater, 1994. 1st and 2nd records.

*+158/33.3x5. NARCISSUS POETICUS x N. PSEUDONARCISSUS (N. x INCOMPARABILIS) *46,
Cards.: Wooded streambank 200m W of Coed Wileirog, Wallog, SN601854, A.O. Chater, 1992.

7158/33.3x5c. NARCISSUS POETICUS x N. PSEUDONARCISSUS subsp. MAJOR (N. x
INCOMPARABILIS [N. x BERNARDII]) *46, Cards.: Roadside hedgebank, Tanycastell, Rhydyfelin,
SN588788, A.O. Chater, 1994.

7158/33.5c. NARCISSUS PSEUDONARCISSUS subsp. MAJOR *46, Cards.: Sand dunes by N end
of road, Ynys-las Dunes NNR, SN611940, A.O. Chater, 1993. Marshy scrub 200m S of Felin Geri,
Cwm Cou, SN300421, A.O. Chater, 1994. Ist and 2nd records.

+158/33.5x6. NARCISSUS PSEUDONARCISSUS x N. CYCLAMINEUS (N. x MONOCHROMUS) *46,
Cards.: Scrub by old cottage site 100m E of Cwmtudu, SN357575, A.O. Chater, 1994.

¥158/33.bic. NARCISSUS BICOLOR *46, Cards.: Ash wood SW of Pen-y-wern, New Cross,
SN636765, A.O. Chater, 1994. Mixed woodland on ridge E of Crugiau, Rhydyfelin, SN592794, A.
O. Chater, 1996. Ist and 2nd records.

7158/35.2. RUSCUS HYPOGLOSSUM *12, N. Hants.: Well-naturalised in deciduous woodland,
Ashford Chace, Steep, near Petersfield, SU741261, J.D. Fryer, 1998, herb. J.D.F., det. J.M.
Mullin, Probably bird-sown.

+158/36.1. ALSTROEMERIA AUREA “*41, Glam.: Eastcliff, Pennard, SS5567, A.S. Lewis, 1997.

7158/ERY.den. ERYTHRONIUM DENS-CANIS _*17, Surrey: Many seedlings of varying ages in
cemetery, Brookwood, SU9656, J.F. Leslie & K. Page, 1988. Seeded from grave.

7159/2.5. SISYRINCHIUM STRIATUM *35, Mons.: Two plants on earth mound in road
development, Cleppa Park, Newport, ST278847, G. Hutchinson, 1994.

7159/8.1. CROCUS VERNUS *95, Moray: Well naturalised on dunes, Hopeman, NJ146698, P.
R. Green, 1996.

7159/8.6. CROCUS CHRYSANTHUS *46, Cards.: Rough grass in SE corner of Llanafan
churchyard, SN685721, A.O. Chater, 1997.

7159/8.7xang. CROCUS FLAVUS x C. ANGUSTIFOLIUS (C. x STELLARIS) *46, Cards.: Roadside
verge 400m NNW of Moelifor, Llanrhystud, SN548710, A.O. Chater, 1997.

7 159/13.2. CROCOSMIA MASONIORUM ~— _*95, Moray: One patch on waste ground, Miltonduff,
NJ201587, IP. Green, 1996, conf. P.S. Green.

162/3.7. EPIPACTIS PHYLLANTHES *37, Worcs.: Old quarry, Fish Hill, Broadway, SP1136, T.
D. Knight & R. Maskew; Wooded bank on edge of Hartlebury Common, $O8270, J.J. Day; both
1987, det. R. Maskew, conf. A.J. Richards. 1st and 2nd records.

162/5.1. NEOTTIA NIDUS-AVIS_ 47, Monts.: Old valley woodland, Abermule Dingle, SO19, J.
Clarke & A. Franks, 1992. 2nd record and only extant locality. *108, W. Sutherland: Deciduous
woodland, Loch Dubh, Ardroe, NC072238, R.E.C. Ferreira, 1981.

162/8.1. GOODYERA REPENS *109, Caithness: Peat moor, Aukengill Loch, ND353652, J.K. &
S.I. Butler, 1996.

162/11.1. CORALLORHIZA TRIFIDA %*91, Kincardines.: In Sphagnum in willow carr, Banchory,
N0687946, D. Welch, 1992.

162/15.1. PSEUDORCHIS ALBIDA 50, Denbs.: Eryrys, SJ2057, M. Prosser, 1986. Ist record
since 1950.

162/16.1b. GYMNADENIA CONOPSEA subsp. DENSIFLORA *78, Peebless.: West Linton, NT15,
A. Craig-Christie, 1873, E, det. F. Rose.

162/16.1c. GYMNADENIA CONOPSEA subsp. BOREALIS 46, Cards.: Heathy pasture, Cae’r-
meirch, Pontrhyd-y-groes, SN7573, A.O. Chater, 1996. 2nd record

592 PLANT RECORDS

162/16.1x18.1. GYMNADENIA CONOPSEA x DACTYLORHIZA FUCHSI (x DACTYLODENIA ST-
QUINTINI) *39, Staffs.: One plant with parents on N facing roadside bank on limestone, Ecton,
SK098586, B.R. Fowler, 1987, det. R.H. Roberts.

162/17.1. COELOGLOSSUM VIRIDE 83, Midlothian: Thirty-one, very small plants seen in
heavily grazed rich grassland, Fullarton Water, NT282562, C. Dixon, 1997, det. D.R. McKean. 1st
record for over 100 years.

162/18.8. DACTYLORHIZA LAPPONICA *108, W. Sutherland: Schoenus mire, Ardbhair,
NC155333; Kinloch Lodge, NC566510; both R.E.C. Ferreira, post 1985. Ist and 2nd records.

162/23.3. OPHRYS APIFERA 67, S. Northumb.: Reclaimed pit-heap, near Rising Sun Farm,
NZ296688, M. Jones, 1996. 2nd record.

7+162/SER.par. SERAPIAS PARVIFLORA *2, E. Cornwall: Natural, coastal grassland, Penlee
Point, Watchhouse Field, SX439490, P. Cobbing, 1989, det. P. Cribb. New to the British Isles;
probably deliberately introduced.

Watsonia 23: 593—597 (2001) 593

Book Reviews

Plant life of the Quaternary cold stages. R. G. West. Pp. xi + 320. Cambridge University Press,
Cambridge, 2000. Hbk £65.00. ISBN 0-521-59397-2.

Every once in a while it is important to take stock and assess the mass of palaeobotanical
information accumulated. This book does that and gives the reader an excellent, much needed
review of plant life in the British Isles during the cold periods that characterised a large proportion
of the last two million years (the Quaternary Period). Both palynological (pollen and spores) and
plant macrofossil (seeds, fruits, etc.) evidence is examined and integrated into the discussion of
vegetation history of the British Quaternary cold stages, an approach that should be noted and
copied by other palaeobotanists.

After setting out the proposed aims for the analysis of cold stage floras there are a number of
introductory chapters. One of these is an illuminating and important chapter on the type of
sedimentary environments encountered in the fossil record and how investigations of taphonomic
processes acting upon plant parts before they are deposited in modern analogue environments can
allow more precise interpretations of cold stage fossil assemblages.

A compact disc accompanies the book which contains the database which is used as the
foundation for the analysis of the cold stage fossil assemblages. Chapter 4 describes the structure
of this database which contains information on the fossil sites, their chronology, the botanical
remains recovered from each sample, the age of those samples, certain information on the biology,
habitat and present distribution of the fossil taxa recorded, and a bibliography.

The latter part of the book analyses the data collected. Querying the database shows which taxa
are found where, when and in what quantity. Discussions are presented about how to translate the
quantity of remains into actual representation in the past vegetation. It appears that many taxa
represented in cold stage deposits are present in certain parts of Britain today. Another interesting
revelation is that many taxa are recorded in both temperate and cold stages, indicating how
Linnean species can survive climatic fluctuations. A knowledge and understanding of the biology
of the flora (life forms, life spans and variability of species) is essential in explaining such results
and these are discussed. Vegetation, habitat, and climate is reconstructed for the cold periods, with
the contribution of coleopteran, non-marine molluscan and vertebrate faunas to a wider view of
cold stage environments also assessed. Climatic changes in a cold stage can be difficult to
determine from the fossil assemblages, because extensive grassland, which is often represented in
the cold stage fossil record, can exist under a range of climatic regimes. This is a difficult problem
which is addressed.

The reviewer recommends this book to any member of the scientific community or general
public who is interested in an account of the flora that once existed in Britain during the
Quaternary cold stages, and its implications for understanding present flora and environment
(including climate). The price tag of £65 is not too expensive for a hard-back book (plus CD
database) of this type.

M. H. FIELD

British Plant Communities. Volume 5. Maritime Communities and Vegetation of Open Habitats.
Edited by J. R. Rodwell. Pp. xiv + 512. Cambridge University Press, Cambridge, 2000. Hbk.
£70.00. ISBN 0-521—39167-9. Pbk £32.50. ISBN 0-521-64476-3.

Work on the ‘National Vegetation Classification’ project began in 1975 and now with the
appearance of the fifth Volume of ‘British Plant Communities’ it reaches a triumphant conclusion.
Its completion is a tribute to those who initiated and fostered it, to its dedicated and indefatigable
editor John Rodwell, and to his army of co-workers and helpers. The five volumes, taken together,
display not only the size and complexity of the task they undertook but also the uniformly high
quality of the final product. The preceding volumes have already amply proved its practical value
and are widely accepted as essential tools in the hands of vegetation surveyors, managers and
conservationists. Now that coastal vegetation and communities of agricultural weeds and open
habitats are covered, this comprehensive and unique phytosociological treatment of British
vegetation is complete.

594 BOOK REVIEWS

A Preamble (as in earlier volumes) explains the background to the work and the style of
presentation. The main part of the book consists of Community descriptions, arranged in four
sections: Salt-marsh communities (28 in number); Shingle, strandline and sand-dune communities
(19); Maritime cliff communities (12) and Vegetation of open habitats (42). Each section includes
a key to the communities (described as ‘a crude guide to identifying the types of vegetation’ —
helpful though not infallible!), and begins with an Introduction. These Introductions are a
particularly useful feature and will appeal to the many botanists who have an interest in the flora
and ecology of coastal vegetation, or in weeds and ruderals. At the close of this volume is a 14-
page ‘Phytosociological Conspectus of British Plant Communities’ which users of the series have
already looked for but of necessity had to await completion of the whole. This will facilitate cross-
reference between different parts of the work, and will also enable comparisons with Continental
schemes of classification.

The dynamic nature of much coastal vegetation, notably that of salt-marshes and dunes, creates
certain problems for community classification, and its application in the field. A country-wide
scheme such as this may not, in all cases, fit comfortably with the floristic variation and patterning
of local sites. However, with experience of working in a range of sites and with a number of
examples it will normally provide the user with a convenient and instructive framework, bearing in
mind that it is based solely on floristic groupings and does not necessarily reflect ecological
processes very closely. Special problems also confront the task of sampling and classifying
communities of open habitats and arable fields, which in any case constitute a more heterogenous
grouping than most of the other vegetation types. However, they do have features in common, and
in view of the scant attention paid in Britain to their phytosociology this is a most welcome
component of the whole.

This volume, with its predecessors, will become for the foreseeable future the standard and
indispensable handbooks for survey, classification and mapping of vegetation in Britain. Their
intrinsic interest also extends to all students of the British flora, for whom they contain a wealth of
relevant and valuable information. It is of the utmost importance, however, to heed the Editor’s
warning that this should not be treated as a ‘static edifice’ but as a working tool, which with use
and in the light of further sampling and new data, will require up-dating in the future.

C. H. GIMINGHAM

The Flora of Dorset. Humphry Bowen. Pp viii + 373. Pisces Publications, Newbury, 2000. Hbk
£45.00. ISBN 1—87—4357-16-1.

Dorset may be seen as a nursery of modern botanical thought. For the first half of the twentieth
century Ronald Good developed an understanding of the correlation of plant distribution with
external factors such as climate, drainage and soil type. Good’s 1948 Geographical Handbook, and
its subsequent revisions, have been the principal published sources of botanical information in
Dorset until the recent production of Humphry Bowen’s Flora.

In common with other recent floras the principal author is joined in the preparation of the
introductory chapters. David Allen’s account of some earlier workers on the Dorset Flora is of
interest for its perception of field botany as a social phenomenon. The section on archaeobotany of
the county, by Pat Hinton, is intriguing. Post-glacial evidence of vascular plants would be a
valuable addition to any other County flora in preparation.

The Flora of Dorset is a flora of the tetrad recording tradition. The accounts and associated maps
concentrate on post-1984 records. For each species the number of tetrads for post-1984 records are
given together with historic records. The maps and the accounts, however, do not always tally. For
example, the maps of Ophrys insectifera and Platanthera bifolia show eight and ten post-1984
records respectively. The text however only refers to four post-1984 sites for Ophrys and five for
Platanthera. Those extracting data for use elsewhere should check the full records which are held
in the Dorset Environmental Records Centre.

Most species accounts only give general locations such as a parish name within a tetrad. The
more sought after groups such as the localities of orchids and sedges are more precise. The
absence of a general map of the vice-county, combined with not using four figure grid references
to identify tetrads, adds to the difficulty in identifying localities.

Judged against the two aims set out in its introduction the Flora may be judged as successful by

BOOK REVIEWS 595

those searching for attractive plants. The leisure botanist will undoubtedly gain great pleasure from
the Flora. Similarly keen gardeners and arboriculturalists will appreciate the many accounts of
selected localities of some ornamental plants in cultivation.

It is unfortunate that our vice-county floras are now drawn upon by more people than just those
who love wild flowers. With the advent of legal protection for wild plants and their habitats, and
with the integration of biodiversity planning into the work of government, biological data are
becoming very important. Both data, and any interpretation, is now subject to detailed scrutiny by
those whose interests may not be primarily botanical.

It was therefore a disappointment that the excellent data that are contained within the Flora are
prone to misinterpretation and challenge. At the heart of this vulnerability is the apparent lack of
consistency of approach. For example, whether or not a species is extinct, or nearing extinction, in
a county is of great importance. The Flora does record some species as extinct, 1.e. Pulicaria
vulgaris and Oenanthe aquatica. Other species with equally long periods since being last recorded
and with similar histories of habitat change are in contrast not recorded as extinct, 1.e. Eleocharis
parvula and Eriophorum gracile. A clear and consistent use of criteria is not apparent.

The emphasis in the Flora on aliens, garden specimens and casual occurrences similarly
complicates interpretation. Complications arise where accounts reproduce all records for a species
but do not differentiate between persistent native populations, casuals and possible introductions.
For example, the status of Mentha pulegium described in the Flora is difficult to reconcile with that
in the 1994 Scarce Plant accounts reproduced in the 1999 Red Data Book. The discovery of the
important population of Mentha pulegium in Purbeck in 1996 is masked by the other parts of the
account. As the presence of this protected species has considerable significance not only to
botanists, but also to the landowners, a proper understanding of its status is essential.

The Flora of Dorset will bring pleasure to many. It also provides a valuable point of reference
upon which others may build. The interpretation of the data gathered for the Flora will need to be
progressed elsewhere. The Flora of Dorset has raised many fundamental issues as to what a flora is
for. If a flora intends to inform scientists or civil servants whose decisions can help or hinder the
wellbeing of our wildlife and their habitats then a different form to that adopted by the Flora of
Dorset is required. If we wish our county floras to serve a community wider than just botanists, we
need to find clearer ways to communicate with those whose decisions determine the fate of the
natural world.

C. CHATTERS

Wild Flowers of the Peak District. Text by Patrick Harding. Art Editor Valerie Oxley. Pp. 144.
Hallamshire Press, Sheffield, 2000. Price £ 25.00. ISBN 1—874718-—53-9.

Offering an interesting marriage between science and art, this book has a curious pedigree. It is the
very professional outcome of a selection of work from students on a Botanical Illustration
Certificate course for adults at Sheffield University. A well crafted appearance and articulate text
brings credit to both the tutors and the students who produced the illustrations. Clearly the painters
have worked from a basis of sound botanical knowledge. However, if you know and love the Peak
District as a meeting point for highland and lowland Britain, you might be disappointed to find that
it is represented by only sixty specimens of its flora, and few of the area’s rarities find a place. But,
among the criteria for a plant’s inclusion have been the twin aims for a chronological treatment of
the flowering year and the need to select paintable plants that can be depicted life-size. These are
understandable restrictions. This reviewer misses the convention, usually found in botanical
illustration, that adds enlarged details for a plant whose fruit, flower, leaf or root characteristics are
particularly hard to define. Photographs are included in each species double spread and they
sometimes clarify the situation; they also set the plant usefully within its environment. Regrettably,
there is considerable variation between the printing of a painting, usually excellent, and its
companion photograph where the colour is harsher.

The standard of the painting is consistent and very good, with each page beautifully laid out.
The delicate drawing of some of the root systems, done, we are assured, from archive specimens,
is to be applauded. A minor carp would be that there is considerable similarity of style in the
painting of each plant. A plus is the absence of any text upon the actual illustration page, which, in
some cases, becomes a work of art. I particularly liked the very painterly depiction of the
ubiquitous Rosebay Willowherb, a notoriously difficult subject.

596 BOOK REVIEWS

The text which is scholarly in detail but relatively light-hearted in delivery, is very fine. Where
Latin nomenclature has been changed in recent years both old and new names are included. There
is a lot of useful information in the introductory paragraph for each plant. For those who enjoy
their Grigson, the inclusion of folk-names, folk-lore, herbal, medical and domestic uses associated
with the plant, make good reading. An early chapter briefly discusses the area’s climate, geology
and biotic factors.

This is not a volume for the pocket or rucksack, but would be a happy addition to a flagging
botanist’s bookshelf, or a coffee table. Furthermore, if your fieldwork is increasingly done more by
wistful memory from an armchair than by being part of the muddy boots brigade, this book could
be a delight. You would not need another mortgage in order to buy it!

M. M. MARTIN

Zander. Handworterbuch der Pflanzennamen. Dictionary of Plant Names. Dictionnaire des noms
de Plantes. 16th edition. W. Erhardt, E. Gotz, N. Bédeker & S. Seybold. Pp. 990. Eugen Ulmer
GmbH & Co., Stuttgart, 2000. Hardback. ISBN 1—903257-01—8. Marketed by The Plant Press,
Lewes, East Sussex.

Professor Robert Zander (1892-1969) produced the first edition of his Handwéorterbuch in 1927,
and went on to revise it up to the ninth edition in 1964. Since then ‘Zander’, as it is generally
known, has been revised by two or more other authors; of the current four only Seybold was
involved before the 16th edition, which is the first to be trilingual and which includes about 3,640
genera, 20,000 species and 10,000 synonyms. The alphabetical listing of these genera and species
occupies the bulk of the book (626 pp.), but comprises only one (no. 6) of 11 chapters. It claims to
cover the most important cultivated plants and nearly all the native plants of central Europe, and
hopes in future editions to take in plants of western and northern Europe as well.

Zander is a much used reference work, and in turn follows the most generally accepted standard
publications such as the International Code of Botanical Nomenclature (1994), Brummitt &
Powell’s Authors of Plant Names (1992), Kubitzki’s The Families and Genera of Vascular Plants
(1990 onwards), Cronquist’s The Evolution and Classification of Flowering Plants (1988),
Dahlgren, Clifford & Yeo’s The Families of Monocotyledons (1985) and Mabberley’s The Plant
Book (1997). It is unique in scope. Perhaps closest to it is Griffiths’ Index of Garden Plants
(1994), which covers about three times as many taxa and includes potted descriptions of them, but
omits many purely wild plants, has fewer supplementary chapters (see below), is less wieldy and
costs £40. Hence Zander has 43 and Griffiths 15 species of Trifolium; but the figures for Solidago
are 18 and 38 respectively. To give an idea of the coverage of a purely cultivated genus, Zander
has 75 and Griffiths 120 species of Opuntia.

For each species Zander provides the Latin name (plus the gender of each genus); authority:
German, English and French name(s); a series of symbols that denote growth form, flowering
period, garden usage (e.g. border plant, hanging plant, cut-flower plant, hothouse plant, decorative-
fruit plant), whether scented, officinal (its designation ‘Off is omitted in error from the list of
abbreviations), poisonous or otherwise useful, and conservation/protection status; and a detailed
list of its native territories. The English names were apparently compiled by Charles Quest-Ritson
(Salisbury); in the main they are the familiar ones, but there are also many much less used names
and many well-known ones are omitted. For example, Montia fontana is given no English name,
while M. perfoliata is called Miner’s Lettuce and Winter Purslane; Spring Beauty is (?mis)applied,
along with Siberian Purslane, to M. sibirica. Hence the standard English name is omitted from all
three! The French names seem to me even more patchily quoted; for example Tussilago farfara is
not provided with one even though its called Pas d’ane in all my French Floras. It is not difficult
to find other minor errors, such as spelling mistakes, in the book, but on the whole it is well
produced and authoritative.

The other ten chapters provide a wealth of additional and often surprising information: 1 —
biography of the authors of all 16 editions; 2 — explanation of botanical nomenclature and of the
pronunciation of Latin names; 3 — systematic list of families of vascular plants (following
Cronquist and Dahlgren ef al.); 4 — alphabetical list of families with the genera listed
alphabetically under each; 5 — an elaborate (17 pp.) hierarchical listing of the world’s
phytogeographical-regions; 7, 8 & 9 — indices of German, English and French plant names (the

BOOK REVIEWS 597

above comments on these are reinforced by the 96, 71 and 19 pages respectively devoted to each);
10 — list of abbreviations and full names of authorities of plant names; and 11 — a short
bibliography. All the free text in these chapters is triplicated in the three languages.

For all people seriously interested in plants, Zander is a most valuable handbook of information.
First and foremost it provides a list of virtually all the plants that one is likely to encounter, with a
wealth of information about each. Many of the other chapters are most likely to be of use to those
without access to a botanical library, enabling them to refer to, inter alia, the most commonly used
systems of classification, the rules of botanical nomenclature and the standard abbreviations of
authorities. Probably no other book provides quite this range of information, certainly none in such
a handy compact format. This first trilingual version of Zander deserves a much wider circulation
among English speakers than previous editions have enjoyed. If the next edition succeeds in
improving the list of English names and in adding or emending the entries of the wild and
cultivated plants that grow in the Atlantic fringes but not in central Europe, it will be
indispensable.

C. A. STACE

Causes of Change in British Vegetation. Ecofact Volume 3. L. G. Firbank, S. M. Smart, H. M. van
de Poll, R. G. H. Bunce, M. O. Hill, D. C. Howard, J. W. Watkins and G. J. Stark. Pp. 98. Dept. of
Environment, Transport and the Regions, London, 2000. Price £8.00. ISBN 1—870393-51-1.

This is the third volume in this series. It has as its stated aims to: 1. Identify the causes of observed
changes in botanical diversity, 2. Assess the relative importance of land management and other
factors, such as pollution, 3. Recommend land management practices for the maintenance and
enhancement of diversity, 4. Develop predictive techniques for determining ecological impacts.
With aims like these, this could well be described as being the botanical conservation holy grail —
not only does it confirm in hard scientific terms what we felt we knew all along; that nutrients are
up, diversity is down, with an increase in competitors, but also it offers reasons, solutions and a
method for stopping this happening again (or at least for letting us know if it’s happening again).
So, does it deliver?

The authors identify four major drivers of vegetation change, being |. Changes in agriculture, 2.
Changes in the management of roadside verges, 3. Conifer planting and 4. Inputs of nitrogen and
other nutrients. These are further broken down into individual factors such as waterside
management, and the impact of leisure. There is an informative table on how these drivers shifted
between the surveys and their effects on various environmental factors. They then go on to define
twelve Indicators of Botanical Diversity (or I.B.D.s), such as Ellenberg scores per plot, frequency
of scarce species, and food plants for animal groups. These drivers and the indicators are then
linked by chapters on the causes of change and policy implications. Hidden in the latter chapter is
a table which is perhaps the most informative part of the document, listing policy initiatives such
as S.S.S.I. designations, Nitrate Sensitive Areas or Biodiversity Action Plans against eighteen
drivers such as drainage and conifer planting, then assessing the impact that these policies have
had on the drivers. The remaining forty five pages consist of twenty explanatory ‘boxes’ and
annexes, making this a highly readable and digestible document. Is there a down side? My only
grumble was that there is much emphasis on Ellenberg’s continentality scores, which were ignored
in the technical annexe, while salt tolerance was covered but not in the plots analysed; but this is
pretty much a minor point.

So, in answer to my question, yes, it does deliver. These are the hard data, the ‘smoking gun’
that our campaigning conservation cousins should use as evidence to influence policy, although it
would be interesting to know if we will ever get the chance to see original data. This series of
Ecofact publications should continue to contribute in an objective, dispassionate way to the
conservation lobby’s armoury and together with the forthcoming Atlas, form a formidable data
bank on which to base their case.

S. J. WHILD

Watsonia 23: 599-600 (2001) 599

Obituary

JOHN MICHAEL MULLIN
(1947-1999)

Michael Mullin, or Mike as he was known to his botanical colleagues, family and friends, was
born in 1947 in Tyneside into a large family of three brothers and two sisters. A few years later his
family moved to Stevenage New Town, and his natural history interests were shown early; when
he was five years old Michael went missing for most of a day, until found happily and totally
absorbed in a study of the local frogs. As a teenager Michael’s main interest was firmly focused on
botany, and with a school friend he organised an exchange scheme of insectivorous plants. At
fourteen he was seriously recording wild plants, helping John Dony on his Flora of Hertfordshire
project at weekends and in school holidays, and later he worked with the Upper Teesdale Group
from Durham University. Schooling was at Alleyne Grammar School from which he left with
eight ‘0’ level passes, but always with a tendency against authority, he left with a rude message to
the headmaster watered in weedkiller onto the cricket square. Later from South Shields Grammar
Technical School for Boys, he left with three ‘A’ level passes - one with distinction in Biology.

Mike started work as a Biology lecturer in various schools, soon adding field trips for Extra-
mural Associations, and slide presentations. As a keen walker, cyclist and camper with a
passionate interest to see every British plant, by the time that he joined the staff at The Natural
History Museum (then the British Museum (Natural History)) Mike had an exceptional knowledge
of field localities all around the British isles. Mike had met his wife Anne on field work at Malham
Tarn in 1966, and together they had continued camping, walking and hitch-hiking to field
meetings or to search for plants, collecting for herbarium specimens where appropriate.

From 1973-1990 Mike worked at the BM (Natural History), first as a student then as a Scientific
Officer when he worked mainly in the British Herbarium curating collections. He participated in
the British Museum Outer Hebrides project and was co-author with R. J. Pankhurst of the Flora of
the Outer Hebrides (1991). Working in the Museum Mike acquired a phenomenal knowledge of
plants from around the world, and was always prepared to spend much time researching plants
brought to him for identification, and other botanical queries that came to him in the Department.
As a member of the B.S.B.I. Meetings Committee, Michael had special responsibility for the
organisation of the Annual Exhibition Meeting, which for many years was held in the Department
of Botany each November, attended by several hundred members and guests - in the General
Herbarium. Mike cycled to work and around London and his bicycle became his logo, his letters
and memoranda notes being signed ‘Mike the Bike’ with a small cycle drawing below. He was
also regrettably accident prone, and managed to get knocked off his bike on what surely must be a
record number of times? In 1990 Michael was Assistant leader on a botanical tour to the
Peloponnisos, but returned to find that staffing at the Museum had been re-organised and his post
no longer existed.

For a number of years Michael then worked on short-term contracts, including with London
Wildlife Trust as Project Manager/Warden of Gunnersbury Triangle Local Nature Reserve, as an
Ecologist with a consultancy firm, as a Lecture Course Organiser at St Mary’s College, University
of Surrey, and as an Ecologist with the London Ecology Unit. In 1997 he was appointed as a Tour
Leader with Naturetrek Tours, taking groups to the Himalayas, Madeira, Portugal etc. Here, Mike
had really found his niche and he could have continued as a travelling lecturer for a long time. As
a brilliant teacher in the field, he had a talent for making plants and natural history interesting for a
wide range of people. I recall meeting him unexpectedly on Wengen Station in the Bernese
Oberland, surrounded by his group and recently returned from Sikkim; he was dressed in white
with a topee, a cane and looking every inch the genial traditional botanical explorer.
Unfortunately, after Sikkim the first signs of the illness which was to take his life became
apparent. In spite of a courageous fight against the cancer, Michael died on 2 August 1999.

Colleagues from the Department of Botany attended his funeral with representatives of his other
interests. Michael described himself as having ‘eclectic tastes in music ranging from classical and
folk to New Age etc.’ He had a deep interest in dancing with the Cotswold Morris, and with his

600 OBITUARY

family and friends travelled to dance in Britain and also to Spain and France. Michael Mullin died
sadly young leaving much promise and unfinished projects. We extend sympathy to his wife Anne
and their daughter Fiona, and to his many friends — he will be sadly missed. On April 9 2000, a
memorial bench to Mike at Gunnersbury Triangle L.N.R. was celebrated with ‘a toast and some
tales’. Friends commented ‘it was a grand party and Mike would have loved it’ but Anne said ‘I
am not so sure, he would have been gnashing his teeth at not being able to get at the beer.’

MARY BRIGGS

INSTRUCTIONS TO CONTRIBUTORS

Scope. Authors are invited to submit Papers and Notes concerning British and Irish vascular
plants, their taxonomy, biosystematics, ecology, distribution and conservation, as well as topics of
a more general or historical nature. Authors should consult the Hon. Receiving Editor for advice
on suitability or any other matter relating to submission of manuscripts.

Papers and Notes must be submitted in duplicate, typewritten on one side of the paper, with wide
margins and double-spaced throughout. Pages should be numbered. Submission of final edited
copy on computer disc will be requested, but two hard copies of the text are acceptable if computer
facilities are not available.

Format should follow that used in recent issues of Watsonia. Underline where italics are required.
Names of periodicals should be given in full, and herbaria abbreviated as in British and Irish
herbaria (Kent & Allen 1984). The Latin names and English names of plants should follow the
New Flora of the British Isles (Stace 1997). Further details on format can be obtained from the
Hon. Receiving Editor or by viewing the website at:
http://www.rbge.org.uk/data/BSBI/Activities/watsonia.htm

Tables, figure legends & appendices should be typed on separate sheets and attached at the end
of the typescript.

Figures should be drawn in black ink or be laser-printed and identified in pencil on the back with
their number and the author’s name. They should be no more than three times final size, bearing in
mind they will normally be reduced to occupy the full width of a page. Scale-bars are essential on
plant illustrations and maps. Lettering should be of high quality and may be done in pencil and left
to the printer. Black and white photographs can be accepted if they assist in the understanding of
the article. If you are able to submit figures on disc please contact the Receiving Editor to check
they are in a suitable format.

Contributors must sign a copyright declaration prior to publication which assigns the copyright of
their material to the Botanical Society of the British Isles. Twenty-five offprints are given free to
authors of Papers and Notes; further copies may be purchased in multiples of 25 at the current
price. The Society takes no responsibility for the views expressed by authors of Papers, Notes,
Book Reviews or Obituaries.

Submission of manuscripts

Papers and Notes: Mr M. N. Sanford, c/o The Museum, High Street, Ipswich, Suffolk, IP] 3QH.

Books for Review: Mr D. A. Pearman, The Old Rectory, Frome St Quintin, Dorchester, Dorset
DT2 OHF.

Plant Records: the appropriate vice-county recorder, who should then send them to Dr C. D
Preston, Biological Records Centre, Monks Wood, Abbots Ripton, Huntingdon, PEI7 21S.

Obituaries: Mrs M. Briggs, 9 Arun Prospect, Pulborough, West Sussex, RH20 IAL.

Back issues of Watsonia are available from the official agents for BSBI Publications,
Summerfield Books (John & Sue Atkins), Summerfield House, High Street, Brough, Kirkby
Stephen, Cumbria CA17 4BxX. Tel: 017683 41577. Fax: 017683 41687. E-mail:
bsbipubs @beeb.net to whom orders for all issues prior to Volume 22 part 1 should be sent.

Recent issues (Vol 22 part 1 onwards) are available from Mr M. Walpole, B.S.B.L.,
68 Outwoods Road, Loughborough, Leicestershire, LE11 3LY.

Watsonia

August 2001 Volume twenty three Part four

Contents

RICH, T. C. G. What is fie vulneraria L. ae corbierei aes & a
Cullen (Fabaceae)?

KHALAF, M. K. & STACE, C. A. ‘The Aeincnen eiasente Cerastium tomentosum L.
and C. biebersteinii DC. (on bene and their occurrence in the wild in
Britain ,

HENDERSON, S. A. ‘The Sercatan acetal aah ne ee
Cham. (Orchidaceae), Irish Lady’ s-tresses, on the Isle of Coll, Inner Hebrides

BLACKSTOCK, N. & ASHTON, P. A. A re-assessment of the putative Carex flava
agg. (Cyperaceae) hybrids at Malham Tarn (v.c. 64): A morphometric analysis

RICH, T. C. G. & MOTLEY, G. S. Conservation of Britain’s biodiversity: Hieracium
linguans (Zahn) Roffey (Asteraceae), Tongue Hawkweed “ie ar

SPALTON, L. M. A new subspecies of Bromus hordeaceus L. (Poaceae)

SMITH, R. E. N. & MARGETTS, L. J. Festuca longifolia Thuill. (Poaceae) in Devon
BAILEY, J. P. Fallopia x conollyana The Railway-yard Knotweed

NOTES

Stace, C. A. The interspecific hae ae of the rhizomatous couches, ies Desv.
(Poaceae) be

Tennant, D. J. Salix caprea L. x S. “myrsinifoia Salisb. x S. phylicifoia L. in
Perthshire

Bull, A. L. A new species ea Rubus Sect. Micantes fRosaceue) Sonne es as

a variant of R. criniger (E. F. Linton) Rogers
Allen, D. E. Dates of publication of county floras
PLANT RECORDS
Book REVIEWS
OBITUARY

Published by the Botanical Society of the British Isles
ISSN 0043-1532

Typeset by D. K. & M. N. SANFORD
Printed in Great Britain by PALLADIAN PRESS, UNIT E, CHANDLERS ROW, PORT LANE, COLCHESTER, ESSEX CO1 2HG

469-480

481-491

493-503

505-516

517-523
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539-541

543-547

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549-550
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593-597
599-600

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