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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 1993-94

Elected at the Annual General Meeting, 15th May 1993
President, Dr F. H Perring

Viceeere cnc Mr P. S. Green, Dr G. Halliday, Mr A. C. Jermy,
Mr A. O. Chater

Honorary General Secretary, Mrs M. Briggs

Honorary Treasurer, Mr M. Walpole

Editors of Watsonia

Papers and Notes, J. R. Edmondson, R. R. Mill, E. C. Nelson, B. S. Rushton*
Plant Records, C. D. Preston

Book Reviews, J. R. Edmondson

Obituaries, J. R. Akeroyd

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

Watsonia, 20, 1 (1994) |

Editorial

A NEW DIRECTION FOR WATSONIA

For many years, Watsonia has published Papers and Short Notes “concerning the taxonomy,
biosystematics and distribution of British and Irish vascular plants, as well as topics of a more
general or historical nature” and has rarely accepted papers on the conservation of plant species and
never accepted papers on their ecology. However, in recent years the Botanical Society of the
British Isles has found itself becoming more and more involved in the conservation and ecology of
the British flora through the Scarce Plants Scheme, the Monitoring Scheme, the Recovery
Programme, etc. Consequently, the Editorial Committee of the Journal and the Publications
Committee of the Society have agreed to increase the scope of the Journal to allow publication of a
wider range of articles in the areas of conservation and ecology, particularly the ecology of single
species. We do not anticipate including papers on community ecology nor the more physiological
aspects of ecology. Therefore, in future, the scope of Watsonia will be “‘the taxonomy, biosystema-
tics, ecology, distribution and conservation of British and Irish vascular plants, as well as topics of a
more general or historical nature’. We will welcome papers and notes related to topics such as
population monitoring (particularly of rare or endangered species but including other species as
well), conservation of individual species or habitats, habitat requirements of species, transplanta-
tion or relocation experiments, distribution of indictor species, the ‘performance’ of individual
species in particular habitats, germination requirements, etc., though this should not be seen as an
exhaustive list. In this way, we hope to provide a journal which is even more relevant to both the
members of the Society and the greater scientific community.

Prospective authors who are concerned whether their work falls within the now wider scope of the
Journal should consult the Honorary Receiving Editor before submission of any manuscripts.

BRIAN S. RUSHTON
Honorary Receiving Editor

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Watsonia, 20, 3-24 (1994) 3

Floristic change in English grazing marshes: the impact of 150
years of drainage and land-use change

J.O. MOUNTFORD

Institute of Terrestrial Ecology (Natural Environment Research Council), Monks Wood,
Abbots Ripton, PEI7 2LS

ABSTRACT

Grazing marshes have assumed a significant role in the conservation of British wetlands, but their biodiversity
has declined as a result of drainage and land use change. Using as wide a range as possible of published and
archive sources from 1840 to the present, changes in species abundance and distribution over 150 years were
reconstructed in three English grazing marshes: the Somerset Levels and Moors, the Romney and Walland
Marshes and the Idle/Misson Levels. Of 526 species studied, over half showed a decline, which was severe in 123
species. Only 34 species, often aliens or adventives, increased. Published accounts, land use and drainage data,
together with knowledge of the species’ ecology suggest that drainage has been the most important factor in
species decline. Raised bog and fen have experienced the greatest extinction of species. Species of wet grassland
and waiercourses survive, but often at much reduced abundance.

KeEyworbs: agriculture, drainage channel, grassland, mire, wetland, conservation.

INTRODUCTION

Much attention has been focused on wetland conservation and the relationship between land use
change and the survival of particular species and communities. This attention stems from the belief
that wetlands are particularly threatened by human activity intended to increase agricultural
production or to protect and extend urban areas. Evidence from species decline and extinction
supports this view (Ratcliffe 1984).

Concern for the survival of particular species of plants and animals has led ecologists and
conservationists to widen their research to include communities which in no sense are natural but
which hold populations of vulnerable species. The traditional grazing marshes of Britain represent a
Stage in the conversion of ‘virgin’ land into farmland and as such support vegetation that is neither
typical of primeval wetland nor of intensive cultivation (Moss 1907; Williams 1970).

Agricultural drainage has been a major factor in the reduced extent of wetlands (Sheail & Wells
1983). The environmental effects of drainage works are many (Hill 1976). They include:

a. Removal or modification of wetland plant communities (reduced species richness and altered
diversity).
. Reduction of breeding (and feeding) areas for animals.
Degradation and accelerated loss of organic matter.
. Higher mineralisation rate, leakage of nitrogen-nitrate to ground-water in the years following
drainage and run-off of other nutrients and chemical residues.
e. Change in the local water regime, with impacts on ground-water quantity and quality (channel
form, sediment load, water temperature and water chemistry).
f. Increased use of fertilisers and pesticides including aquatic herbicides (for drainage channel
maintenance).
g. Reduction of landscape and amenity values.
h. Removal of hedgerows, trees, scrub, small woodlands, ponds, field enlargements, etc. coinci-
dental with drainage works.
i. Drainage effects on land that has itself not been drained, but lies adjacent to such land.

aoc

In English grazing marshes, drainage for agriculture and associated land-use change have affected
the distribution and abundance of plants through: 1. changes in water-table; 2. peat cutting; 3.

+ J. 0. MOUNTFORD

channelisation; 4. changes in water-quality; 5. ditch elimination; and 6. management of the
watercourses.

In the present paper, the main objective has been to assess the relationship between drainage and
land use change since 1840 and the status of plant species in grazing marshes. The approach has been
to marshall the information provided by botanists over the period 1840-1990 and demonstrate
changes in species abundance and distribution, a method previously used in Broadland (Driscoll
1982), the Fenland (Sheail & Wells 1980) and Gwent Levels (Wade & Edwards 1980). An aim has
been to demonstrate the need for plant ecologists to pay attention to the historical context in which
the communities that they study developed (Salisbury 1927).

METHODS

THE STUDY AREAS

Different grazing marshes reflect different stages in the transition from primeval wetland to drained,
reclaimed and intensively farmed land. Three areas in England were selected that display the
increasing impact of modern agriculture (Fig. 1):

Idle/ Misson
Levels

Somerset Moors
and Levels ‘ ro
a
s

2

Romney and PBS
Walland tl @

AO

Ficure |. Location of three grazing marsh study areas in England. The sites are depicted at twice the base map
scale.

|
|

ee

ee

Oe

CHANGE IN ENGLISH GRAZING MARSHES 5

Portishead

Tickenham ordano ‘Bristol
~-— Portbury

iS ey Bourton
a Yatton 7
(361) Typical sand-pararendzinas
\Y
a

AN

(814) Pelo-calcareous alluvial gley

a (813) Pelo alluvial gley

] (1022) Earthy eu-fibrous peat
" ~ ;
Zs = <a ae |] ssa Humic~alluvial gley
Sy BRI XS iaale
a ae ac SROOLZ

Le KX
ye as en onbury Me (1021) Earthy oligo-fibrous peat

Bridgwater
VA

N Sear KA (532) Brown soil
Taunton NL / AL ei
al FT are “VY =,
7 ES a x ini a Shapwick
= AD Thorney \ Jay : [R] The Roughet
\ | 10 il Catcott Burtle
sy Kilometres

FiGureE 2. The Somerset Levels and Moors-— map of study area showing major soil types and localities mentioned
in the text. Upland (altitude >10 m) and sea areas left blank. Soil data derived from the Soil Survey of England

& Wales (1983).

1. THE SOMERSET LEVELS AND MOORS - chosen as a largely unimproved grazing marsh.
The study area takes in all the land on ground-water gley and peat soils between sea level and the 30’
(10m) contour and corresponds with the area defined by the 1:100,000 soil map (Soil Survey 1978)
(Fig. 2).

2. THE ROMNEY AND WALLAND MARSHES - chosen as an area where marked changes in
drainage and land use occurred after 1940. The study area includes the land south-east of the Royal
Military Canal, with Shirley Moor and the Rother Levels west to Potman’s Heath (Fig. 3).

3. THE LOWER IDLE VALLEY AND MISSON LEVELS - chosen to study the effect of a long
period of intense cultivation on the plants of pasture and drainage channels. The study area was
defined as that to benefit from the location of new pumps at West Stockwith, where the R. Idle joins
the R. Trent (i.e. between Epworth and Bawtry, and up the R. Idle to Mattersey) (Fig. 4).

SELECTION OF SPECIES

Over 500 species of vascular plant and Charophyte were included in an archive search. The species
chosen met one or more of the following habitat criteria, growing: 1. in standing or flowing fresh
water; 2. in mire vegetation at altitudes <300 m; 3. in mesotrophic grasslands below 300 m; 4. in
flood-plain or fen woodland; 5. on the banks of water-courses; 6. associated with occasionally
flooded sites, muddy ground, or where compaction of the soil leads to transient standing water; and

7. in or by brackish water or where saltmarsh gives way to freshwater communities. :

SOURCES OF FLORISTIC DATA
In order to trace changes in the distribution and abundance of species since 1840, as complete a set
of sources as possible was used to build up a picture of the history of individual species in each study

6 J. 0. MOUNTFORD

Bonnington
Hamstreet r

(814) Pelo-calcareous alluvial gley

y

Eel oS

yy (813) Pelo alluvial gley
Ve %
a ef Love)
é KO (532) Brown soil
ie > Ca, ’
P)

4, =
WY (361) Typical sand-pararendzinas

ISLA

V4 Denge Beach
Midrips ' ——— :
The Wicks Kilometres

FicurE 3. The Romney and Walland Marshes — map of study area showing major soil types and localities

mentioned in the text. Upland (altitude >10 m) and sea areas left blank. Soil data derived from the Soil Survey
of England & Wales (1983).

pees

(811) Typical

¢ Alluvial gley
yy, (813) Pelo
Epworth

(La

aes

LD (1022) Earthy eu-fibrous peat
Ls Z BES
OK > O59
Bes SaGA

IK (821) Sandy gley
SCS a 4

aaa 56

SOR

(861) Humic-sandy gley

NONE

xs
q
S

RS
RY
RS

eS

(532/551/ 572) Brown solls

Bawtry (711/712) Stagnogleys
Gainsborough

(851) Humic—alluvial gley

Mattersey

(372) Rendzina — like alluvial soils

Kilometres

Ficure 4. The Lower Idle valley and Misson Levels — map of study area showing major soil types and localities

mentioned in the text. Upland (altitude >10 m) and sea areas left blank. Soil data derived from the Soil Survey
of England & Wales (1983).

area. The sources used in the study areas are set out in Appendix |. There is great variation in the
detail and comprehensiveness of the earlier county or regional Floras. Recent plant atlases record
information systematically but often omit local detail useful in interpreting species distribution.

The journals of naturalists’ societies contain site-specific data, providing the detail required to
evaluate accounts in Floras and plant atlases. Manuscript data (card indices, etc.) were consulted
both in private hands and in herbaria, where accompanying specimens may be used to substantiate
the accuracy of records. When these sources were prepared by authors of (or contributors to)
Floras, they clarify and amplify the published accounts.

Many botanists have kept notebooks or annotated maps detailing their finds and the notes

CHANGE IN ENGLISH GRAZING MARSHES 7

referring to a single visit may be enough to reconstruct the contemporary vegetation. Such sources
were traced through the recorder network of the Botanical Society of the British Isles.

Both the national Biological Records Centre at Monks Wood and local centres provided species
distribution data and were consulted extensively in all three study areas. Local centres may also
store material on a site basis, giving an insight into floristic change at particular locations.

English Nature (formerly the Nature Conservancy Council) commissioned inventories of
watercourse vegetation for each of the study areas between 1978 and 1982. These accounts provided
detail on specific sites and a valuable baseline from which to measure change.

RECONSTRUCTION OF FLORISTIC CHANGE: THE EXAMPLE OF GOLDEN DOCK (RUMEX MARITIMUS L.)

The range and quality of material used can be demonstrated by a detailed example. Appendix 2
reproduces the information gathered about the Somerset distribution of Rumex maritimus. From
such material, trends in the distribution and frequency of R. maritimus can be discerned:

To 1900, it was common on the peat-moors, and also scattered by the R. Parrett and its estuary, with
outlying records near Wells and at Portishead.

Between 1901 and 1940, the dock may have spread further up the R. Parrett and near the R. Tone
also. It maintained its abundance on the peat-moors.

Between 1941 and 1975, R. maritimus was not seen in the Parrett valley (except on the coast and a
transient site at Thorney). It was present on the peat-moors but recorded only from specific sites
rather than described as generally ‘plentiful’.

After 1975, it was only found in a small area at the centre of the peat-moor between Catcott Burtle
and the R. Roughet.

There is a strong suggestion that R. maritimus declined on the Somerset Levels and Moors,
surviving only on the peat moors (the core of its former range) where peat cutting and ditch
clearance created the open, bare, wet conditions where the dock could germinate, flourish and seed.

A similar marshalling and assessment was undertaken for each of the 500 species covered by the
archival search, though for the commonest taxa there may only be summary data. Judgement had to
be exercised in the interpretation of a wide range of disparate sources. Any trends identified in the
abundance of species might be 1. marked, 2. moderate but unambiguous or 3. slight and based upon
data of doubtful quality (Mountford 1994). Species with similar environmental needs which show
consistent population or distribution changes were grouped together. Where the abundance or
distribution of a species changed between 1840 and 1990, the extent of its preferred habitat can often
be shown to have changed also. When species from one habitat show similar changes, there is
circumstantial evidence that the cause of the change is some alteration to that habitat. Land-use and
drainage data were examined to test whether any changes in the habitat had occurred. Changes in
the study areas are described in detail elsewhere (Sheail & Mountford 1984; Mountford & Sheail
1989).

RESULTS

MAJOR TRENDS IN PLANT DISTRIBUTION AND ABUNDANCE

Trends in distribution are summarised in Tables 1—5, which list all taxa for which the sources used
provide some evidence of change 1840-1990 (References and Appendix 1). Over half the taxa
showed a population change in one or more area.

SPECIES SHOWING NO CHANGE IN POPULATION OR DISTRIBUTION
There were 206 species for which no convincing evidence of population change could be found. This
total included:

— Species where the published sources differed in precision (giving an impression of change), e.g.
Carex pilulifera L.

8 J. 0. MOUNTFORD

— Taxonomically difficult plants, identified by very few botanists, producing insufficient data from
which to infer trends, e.g. Rorippa X anceps (Wahlenb.) Reichb.

— Species where opposing pressures (some favourable, others unfavourable) balanced, resulting in
no perceptible change, e.g. Phragmites australis (Cav.) Trin. ex Steudel.

— Species whose habitat requirements limited them to a few sites from 1840-1990, e.g. Potamogeton
coloratus Hornem.

— Ubiquitous species occupying habitats which were as widespread in 1990 as 1840, e.g. Agrostis
stolonifera L.

SPECIES SHOWING AN INCREASE IN ABUNDANCE AND DISTRIBUTION

50 species showed evidence of spread in at least one study area, although 16 of these declined in
another of the three areas (Tables 1 & 2). Of the remaining 34, there is some doubt attached to the
quality of the data for 18 (due to taxonomic problems or unevenness of coverage). Only seven
spread markedly and unambiguously — Azolla filiculoides Lam., Chamerion angustifolium (L.)
Holub, Elodea canadensis Michaux, E. nuttallii (Planchon) H. St John, /mpatiens glandulifera
Royle, Nymphoides peltata Kuntze and Puccinellia distans (Jacq.) Parl. Table 1 includes many
aliens and colonists (species introduced into grazing marshes from other habitats or parts of
Britain). Increasing abundance might be presumed for some species whose preferred habitat
became more widespread between 1840 and 1990, but cannot be demonstrated from the sources.
These species were classified as unchanged in abundance.

SPECIES INCREASING IN SOME AREAS AND DECREASING IN OTHERS

In most cases, presumed causes of population change and the resulting trends in species were similar
in all study areas (Table 6). However, 16 species showed conflicting trends in distribution when the
study areas are compared (Table 2). Wade & Edwards (1980) note several species in the Gwent

TABLE 1. SPECIES SHOWN TO HAVE INCREASED IN POPULATION SIZE OR DISTRIBUTION
FROM 1840 TO 1990

(a) In all three study areas

Elodea canadensis Potamogeton berchtoldii?
Elodea nuttallii Ranunculus trichophyllus?
Populus alba? Rorippa microphylla?
(b) In two of the study areas (absent/unchanged in third)
Azolla filiculoides Mimulus guttatus
Callitriche platycarpa? Myriophyllum spicatum?
Carex acuta? Nymphoides peltata
Chamerion angustifolium Petroselinum segetum
Festuca arundinacea? Potamogeton pusillus?
Glyceria declinata? Salix triandra?
Impatiens glandulifera Veronica catenata?
Lemna gibba
(c) In one study area (absent/unchanged in other two)
Alopecurus bulbosus Leucojum aestivum
Barbarea stricta Myosurus minimus?
Chara globularis Nitella mucronata?

var. annulata? Potamogeton pectinatus

var. virgata? Puccinellia distans
Eleocharis uniglumis? Rorippa X sterilis?
Elodea callitrichoides Salix viminalis?

Leersia oryzoides

Notes: i. Where a question mark follows the species name, there is some doubt about the trend. Data may be
inadequate, there may be some taxonomic problem, etc. 11. Where a species has clearly changed in one area, but
in one or both the others there is some doubt, no ? is marked. ii. Some species have only ever been recorded in
one or two of the study areas. iv. Nomenclature follows Stace (1991) for vascular species, Moore (1986) for
Charophytes and local sources (Appendix 1) for some infraspecific taxa.

CHANGE IN ENGLISH GRAZING MARSHES 9

TABLE 2. SPECIES SHOWN TO HAVE INCREASED IN POPULATION SIZE OR DISTRIBUTION IN
ONE OR MORE STUDY AREAS, AND TO HAVE DECREASED IN THE OTHERS FROM 1840 TO 1990

Althaea officinalis Potamogeton trichoides

Bromus commutatus Ranunculus baudotii

Carex vesicaria Ranunculus circinatus

Carex viridula subsp. viridula? Rorippa amphibia

Chara hispida subsp. hispida? Schoenoplectus tabernaemontani
Epilobium tetragonum subsp. tetragonum Symphytum officinale

Persicaria bistorta Tolypella nidifica

Potamogeton natans Wolffia arrhiza

Notes: see Table 1.

Levels whose distribution trends differ from those observed in the present study, reflecting the
importance of local factors in determining floristic change.

SPECIES SHOWING A DECREASE IN ABUNDANCE OR DISTRIBUTION

Just over half the species included in the archive search underwent an apparent contraction in range
or decline in frequency after 1840 (Tables 3 & 4). Of these, 123 species declined markedly, 94
becoming extinct in at least one, and 23 in two or all study areas (Table 4). In all areas, many more
species declined from 1840-1990 than increased (Tables 5 & 6).

The sources used note plants that decreased during the lifetime of the botanist, frequently
suggesting reasons for that decline. Most of the causes advanced involve the activities of a
burgeoning human population (especially agricultural improvement), but some natural changes
were also suggested as factors in species decline. Suggested causes include:

Drainage: the commonest factor proposed as a cause for species loss in wetlands, e.g. Myrica gale
L., Parnassia palustris L., Ranunculus lingua L., Lythrum salicaria L., Narthecium ossifragum (L.)
Hudson and Vaccinium oxycoccos L. (Gibbons 1975); Thelypteris palustris Schott, Dryopteris
cristata (L.) A. Gray, Drosera rotundifolia L., Empetrum nigrum L., Viola persicifolia Schreber and
species of the Misson “‘fenny fields” (Howitt & Howitt 1963); Valeriana dioica L. and Carex
pulicaris L. (Roe 1981); Althaea officinalis L. (Hanbury & Marshall 1899); Cyperus longus L.
(White 1912).

Peat-digging: e.g. Hypericum elodes L. (White 1912; Sandwith files; Roe files); Andromeda polifolia
L. (Roe 1981); Rhynchospora fusca (L.) Aiton f. (Murray 1896; Roe 1981).

Flooding acid peat and alkaline water: Utricularia minor L. (Storer 1985 and files; Roe, pers. comm.
1982).

Enclosure and grazing: e.g. Vaccinium myrtillus L. (Lees 1888); Juncus effusus L. (Raine 1980).

Cultivation (fertilisers, liming, ploughing or reseeding): e.g. Colchicum autumnale L., Orchis morio
L. and Primula veris L. (Roe 1981); Genista tinctoria L. (White 1912); G. anglica L. and Rhinanthus
minor L. (Lees 1888); Botrychium lunaria (L.) Sw. and Sanguisorba officinalis L. (Gibbons 1975).

Watercourse management and pollution: e.g. Hottonia palustris L. and Rumex hydrolapathum
Hudson (Gibbons 1975); Stratiotes aloides L. (Roe files); Rorippa sylvestris (L.) Besser, Valeriana
officinalis L. (Gibbons 1975) and submerged aquatics especially pondweeds — herbicide (Howitt
day-book); Ranunculus fluitans Lam. — colliery effluent (Howitt & Howitt 1963).

Sea defences: e.g. salt-tolerant species at the Midrips and Wicks, Kent (N. W. Moore, pers. comm.
1981).

Industrialisation and urbanisation: e.g. Blysmus compressus (L.) Panzer ex Link (White 1912; Roe
files); Cladium mariscus (L.) Pohl and Carex elata All. (Roe files; Sandwith files); Salix aurita L.
(Howitt & Howitt 1963).

Deliberate eradication or collection: e.g. Colchicum autumnale (Roe 1981); Lychnis flos-cuculi L.
(Willis & Jefferies 1959); Osmunda regalis L. (White 1912); Pteridophyta (Gibbons 1975).

10 J.O. MOUNTFORD

TABLE 3. SPECIES SHOWN TO HAVE DECREASED IN POPULATION SIZE OR DISTRIBUTION
FROM 1840 TO 1990

(a) In all three study areas

Alnus glutinosa

Anagallis tenella

Apium inundatum

Baldellia ranunculoides

Bidens cernua

Caltha palustris

Carex diandra

Carex pulicaris

Chara globularis subsp. globularis?
Chara vulgaris var. longibracteata?
Dactylorhiza praetermissa
Eriophorum angustifolium

Galium uliginosum

Hydrocotyle vulgaris

Juncus squarrosus

Menyanthes trifoliata
Myriophyllum verticillatum

Nymphaea alba
Ophioglossum vulgatum
Orchis morio

Osmunda vulgaris
Pedicularis palustris
Potamogeton alpinus

Potamogeton polygonifolius

Potentilla palustris
Primula veris
Ranunculus lingua
Rumex maritimus
Rumex palustris
Sium latifolium
Sparganium natans
Stellaria uliginosa?
Veronica scutellata

(b) In two of the study areas (absent/unchanged in third)

Achillea ptarmica
Acorus calamus
Apium graveolens
Briza media
Butomus umbellatus
Callitriche truncata
Carex curta
Carex disticha
Carex divisa
Carex echinata
Carex elongata
Carex hostiana
Carex pallescens
Carex viridula subsp. brachyrrhyncha
Catabrosa aquatica
Chara vulgaris

var. papillata?

var. vulgaris?
Cirsium dissectum
Cirsium palustre
Cladium mariscus
Coeloglossum viride
Colchicum autumnale
Dactylorhiza incarnata subsp. incarnata
Dactylorhiza maculata
Drosera intermedia
Eleocharis quinqueflora
Epipactis palustris
Erica cinerea
Erica tetralix
Eriophorum vaginatum
Frangula alnus
Groenlandia densa
Hippuris vulgaris
Hottonia palustris
Hydrocharis morsus-ranae
Juncus bulbosus
Juncus effusus?
Lathyrus palustris

Lotus glaber
Lychnis flos-cuculli
Mentha pulegium
Molinia caerulea
Montia fontana?
Myrica gale

Myriophyllum alterniflorum

Narcissus pseudonarcissus
Narthecium ossifragum
Nuphar lutea
Pedicularis sylvatica
Persicaria laxiflora
Persicaria minor
Peucedanum palustre
Platanthera bifolia
Potamogeton gramineus
Potamogeton lucens
Potamogeton perfoliatus
Radiola linoides
Ranunculus flammula?
Ranunculus hederaceus
Ranunculus peltatus
Ranunculus penicillatus?
Rhinanthus minor s.1.
Rhynchospora alba
Sagittaria sagittifolia
Sagina nodosa

Salix aurita

Salix repens?

Schoenus nigricans
Senecio aquaticus
Spirodela polyrhiza
Stachys X ambigua
Stachys palustris
Stellaria palustris
Succisa pratensis
Thelypteris palustris
Utricularia minor
Utricularia vulgaris s.s.

CHANGE IN ENGLISH GRAZING MARSHES

TABLE 3. cont.

Vaccinium oxycoccos
Valeriana dioica
Valeriana officinalis

Veronica anagallis-aquatica?
Viola canina
Viola palustris

(c) Clearly declined or extinct in one study area (absent or unchanged in other two)

Alopecurus aequalis +
Alopecurus pratensis
Antennaria dioica +
Aster tripolium
Bidens tripartita
Blechnum spicant
Blysmus compressus +
Blysmus rufus +
Bupleurum tenuissimum
Calluna vulgaris
Carex distans
Carex elata
Carex extensa +
Carex laevigata
Carex limosa +
Carex nigra
Carex panicea +
Carex paniculata
Carex pseudocyperus
Carex rostrata +
Ceratophyllum submersum
Chenopodium chenopodioides
Danthonia decumbens
Deschampsia cespitosa
Dryopteris carthusiana
Dryopteris cristata +
Eleocharis multicaulis +
Eleogiton fluitans
Empetrum nigrum +
Euphrasia arctica
Galeopsis segetum +
Genista anglica +
Gentiana pneumonanthe +
Geum rivale
Gnaphalium uliginosum
Gymnadenia conopsea
subsp. conopsea
subsp. densiflora
Hammarbya paludosa +
Hordeum marinum
Huperzia selago +
Hypericum maculatum
Tris pseudacorus
Isolepis setacea +
Juncus gerardi
Juncus subnodulosus
Lactuca saligna
Lepidium latifolium +
Limosella aquatica +

(d) Possibly declined or extinct (?) in one study area (absent or unchanged in other two)

Angelica sylvestris
Berula erecta
Calamagrostis canescens
Callitriche hamulata
Carex remota

Lycopodiella inundata +
Lycopodium clavatum
Lysimachia thyrsiflora +
Lysimachia vulgaris
Lythrum salicaria

Mentha X verticillata
Mimulus moschatus +
Nardus stricta

Oenanthe lachenalii +
Ononis spinosa

Orchis ustulata

Petasites hybridus
Peucedanum ostruthium +
Pinguicula lusitanica
Potamogeton compressus +
Potamogeton X lintonti +
Potamogeton X salicifolius +
Potamogeton X sparganifolius
Prunus padus

Puccinellia fasciculata
Puccinellia maritima
Pyrola minor +
Ranunculus fluitans
Ranunculus omiophyllus +
Ranunculus sardous
Ranunculus tripartitus +
Rhynchospora fusca +
Ribes rubrum

Rumex hydrolapathum
Ruppia cirrhosa

Ruppia maritima
Salicornia ramosissima
Sanguisorba officinalis
Saxifraga granulata
Scheuchzeria palustris +
Schoenoplectus lacustris
Scirpus sylvaticus +
Selaginella selaginoides +
Sparganium angustifolium +
Suaeda maritima
Taraxacum palustre +
Thalictrum flavum
Triglochin maritimum
Triglochin palustre
Utricularia australis
Veronica beccabunga
Viola persicifolia +
Wahlenbergia hederacea
Zannichellia palustris

Carex vulpina s.s.
Ceratophyllum demersum
Chara pedunculata
Chara vulgaris

var. crassicaulis

12

TABLE 3. cont.

Eupatorium cannabinum
Filipendula ulmaria
Glyceria maxima
Hypericum tetrapterum
Juncus acutiflorus
Juncus ambiguus
Juncus bufonius s.s.
Juncus conglomeratus
Juncus X diffusus
Lotus pedunculatus
Luzula pallidula +
Mentha X smithiana
Myosotis scorpioides

J. 0. MOUNTFORD

Nitella translucens
Oenanthe crocata
Persicaria hydropiper
Potamogeton praelongus
Ranunculus aquatilis
Salix purpurea
Scrophularia auriculata
Scutellaria galericulata
Sison amomum
Tolypella intricata
Tolypella prolifera
Utricularia intermedia
Viburnum opulus

Notes: Where a species has become extinct in one study area, but is unchanged or unrecorded in the others, it is
marked as +. (Species which have become extinct in more than one study area are included in Table 4.) See also
Table 1.

TABLE 4. SPECIES SHOWN TO HAVE BECOME EXTINCT IN MORE THAN ONE STUDY AREA
BEFORE 1990, WITH DATES OF LAST RECORD IN EACH STUDY AREA

(a) In all three study areas
Littorella uniflora (S 1895, R 1899, I 1876)
Myosotis secunda? (S 1960, R 1960, I 1888)
Scutellaria minor (S 1914, R 1829, I 1880)
Stratiotes aloides (S 1976, R 1875, I 1855)

(b) In the Somerset Levels and Moors and Romney Marsh
Cyperus longus (S 1896, Rc. 1830) Juncus squarrosus (S 1912, Rc. 1850)
Eleocharis acicularis (S 1907, Rc. 1900) Potamogeton friesii (S 1914, R 1909)

(c) In the Somerset Levels and Moors and Idle/Misson Levels

Andromeda polifolia (S 1920, I 1833) Parnassia palustris (S 1782, I 1847)
Botrychium lunaria (S 1955, I 1955) Pinguicula vulgaris (S 1928, I c. 1905)
Carex dioica (S 1855, I 1840) Potamogeton alpinus (S 1881, I 1944)
Cicuta virosa (S 1888, I 1850) Rhinanthus angustifolius (S 1918, I 1960)
Drosera longifolia (S 1970, I 1893) Trichophorum cespitosum (S 1972, I 1905)
Hypericum elodes (S 1914, I 1896) Vaccinium oxycoccos (S 1919, I 1815)

(d) In the Romney Marsh and Idle/Misson Levels
Anagallis tenella (R 1959, I 1855)
Drosera rotundifolia (R ?, 1 1959)

Oenanthe silaifolia (R 1839, I 1930?)

Note: Numbers in brackets after species name are the dates of the last record, annotated with the appropriate
study area: S (Somerset Levels and Moors), R (Romney Marsh) and I (Idle/Misson Levels).

Succession: “upright-leaf and floating-leaf formations” (Moss 1907); Sparganium natans L. (Storer
files).

Drought: e.g. Drosera longifolia L. (Gibbons 1975); Schoenus nigricans L. (White 1912); Stratiotes
aloides (Roe files).

For most species, contemporary accounts do not suggest the cause of observed population
changes. Table 7 groups species that have declined in at least one of the study areas according to
their habitat and the probable cause of their decline. Species are allotted to a group when archive
sources (Appendix 1) indicate a link with some environmental change or where they are typical of
the plant community affected by this change (Ellenberg 1988; Rodwell 1991a, 1991b, 1992, MSS).

CHANGE IN ENGLISH GRAZING MARSHES 13

TABLE 5. TOTAL NUMBER OF SPECIES INCREASING OR DECREASING IN POPULATION AND
DISTRIBUTION IN THREE ENGLISH GRAZING MARSHES — 1840-1990

I. Species increasing in population and distribution (35 species)

Pronounced Moderate Data

increase increase poor
Three study areas Bs 0 4
Two study areas 4 3 8
One study area 1 ii 6
Totals 7 10 18

II. Species increasing in some and decreasing in other study areas: 16

III. Species decreasing in population and distribution (269 species)

Pronounced Moderate Data

decrease decrease poor
Three study areas 19 14 4
Two study areas 53 39 8
One study area Si 47 34
Totals WB: 100 46

IV. Species with no change in population and distribution: 206

DISCUSSION

The importance of documenting the declining status of species was realised as early as the
nineteenth century (Sheail 1982) and the value of artificial habitats such as grazing marsh ditches for
“substituted plant associations” identified by Moss (1907) long before the contribution of these
modified wetlands was generally accepted. Written records, herbaria and published accounts have
been used to reconstruct the past vegetation pattern. Such sources contribute to an assessment of
the scale and speed of environmental change over the past 150 years. Broad trends can be identified
and, following comparison with knowledge of the species’ ecology and present distribution,
inferences made as to the cause for such trends (Table 7).

SPECIES INCREASING IN ABUNDANCE AND DISTRIBUTION

Aliens and denizens showed the clearest evidence of increased abundance. Such species were often
absent from the study areas in 1840. Some were deliberately introduced later as ornamentals (e.g.
Nymphoides peltata and Populus alba L.) or as crops (e.g. Rorippa X sterilis Airy Shaw and Salix
spp.). Others escaped from gardens (e.g. Impatiens glandulifera Royle and Mimulus spp.) or from
aquaria and ponds (e.g. Azolla filiculoides Lam. and Elodea nuttallii).

Amongst native species, some increased because their preferred habitat became more extensive
e.g. drainage for peat-cutting combined with burning created a dry, disturbed soil favouring
Chamerion angustifolium. Others were favoured by particular management, enabling them to
compete effectively e.g. increased dredging, resulting in more deep water, may explain the increase
of Potamogeton berchtoldii Fieber. Field survey showed P. berchtoldii was typical of newly dredged
ditches both in the study areas and in Gwent (Wade & Edwards 1980).

Changes in water quality contributed to the spread of certain species. Lemna gibba L. and
Potamogeton pectinatus L. tolerate eutrophic, polluted water (Ellenberg 1988). Once rather rare
and largely coastal in the Somerset Levels, L. gibba is now common throughout the area, especially
in managed drains with high nutrient levels. Reduced salinity (by improved sea defences and flood
control) alters water quality. Some plants may have spread into the coastal levels from which they
were previously excluded by salinity e.g. Myriophyllum spicatum L. (Scotter et al. 1977).

Locally increased salinity led to the spread of some plants. In the Romney Marsh, Puccinellia
distans was once confined to coastal salt-marshes and a few sites inland with saline ground-water in

14

J. O. MOUNTFORD

TABLE 6. NUMBERS OF SPECIES SHOWING A CHANGE IN ABUNDANCE IN EACH OF THREE
ENGLISH GRAZING MARSHES OVER THE PERIOD 1840-1990
(Modified after Mountford & Sheail 1989)

(a) Somerset Levels and Moors

Increase in

abundance
and/or
distribution
+++ + 7 Mota
6 12 18 36

(b) Romney and Walland Marshes

Increase in

abundance
and/or
distribution
+++ + ? Total
4 2 14 20

(c) Idle/Misson Levels

Increase in

abundance
and/or
distribution
+++ + ? Total
y 3 15 20

Increases: +++ = marked; + = moderate; ? = trend slight or doubtful.

31

37)

"
53

Decrease in
abundance
and/or
distribution

Decrease in
abundance
and/or
distribution

Decrease in
abundance
and/or
distribution

30 Se) 15

Total

153

Total

USV7/

Total

151

Decreases: — = marked; — = moderate; ? = trend slight or doubtful; + = extinct.

TABLE 7. SPECIES DECREASING IN POPULATION SIZE AND DISTRIBUTION IN AT LEAST ONE
OF THREE ENGLISH GRAZING MARSHES, CLASSIFIED IN TERMS OF TYPICAL HABITAT AND
SUSPECTED CAUSE OF DECLINE, 1840-1990

Preferred habitat
or vegetation type

Wet woodland

Acid bog and heath

Rich fen and tall wet meadow
Old wet grassland

Grazing marsh ditches and pools

Open water

Open water

Bare wet mud and peat
Saline habitats

Varied

Varied

Varied

Suspected cause of

decline

Felling and drainage
Peat cutting and drainage

Drainage and conversion to pasture or arable

Drainage and conversion to intensive grassland or

arable

1. Regrading conversion to trapezoidal section
2. Overgrowing, increased shade and siltation

3. Elimination due to field rationalisation

Aquatic herbicides

Pollution and increased turbidity

Fencing of ditches and control of water-levels
Tidal control and sea defence improvement
Urbanisation and industrial development
Weed control and collection by botanists

No clear cause

Total number
of species

CHANGE IN ENGLISH GRAZING MARSHES 15

peaty layers (Green 1968). In recent years it has increased along those more important roads that
receive regular de-icing salt during the winter (Philp 1982).

SPECIES DECREASING IN ABUNDANCE AND DISTRIBUTION

1. Drainage: water-table change and plant community structure. Community composition may be in
equilibrium with a particular water-regime, such that either drainage or flooding can change that
composition (Thibodeau & Nickerson 1985). Wetland drainage may influence community structure
not only through changes in water-table, but also through altered inputs of dissolved nutrients
(Meade & Blackstock 1988). Those species requiring high water tables and oligotrophic peat are the
first to disappear, followed by those needing wet pasture, fen and open water (Sheail & Wells 1983).

a. Decline due to drainage and felling of wet woodland. 21 species which prefer wet alder woodland
with birch and willow declined in the study areas. Most are typical of waterlogged soils that are
nitrogen-deficient or mesotrophic (Ellenberg 1988). Those present in the study areas are mainly
planted and were not extensive: birch and osiers on peat in Somerset (Moss 1907) or pine on sand
and gravel near the R. Idle. In recent years, woodland has been cleared and the water-table lowered
for extraction of gravel. The Idle/Misson woods on sand were vulnerable to drying out following the
upgrading of the Stockwith pump, and though the total wooded area did not diminish from 1840 to
1990, the older woods were felled and new plantations have not acquired a shade flora. There are no
woods on the Romney Marsh, though woodland plants occur on overgrown drain banks.

b. Decline due to drainage and peat cutting on acid bog. Peat-cutting for fuel or horticulture and
reclamation for farming have greatly reduced the area of raised bog (Goode 1981; Limbert 1988),
though new mire communities may develop on abandoned cuttings (Giller & Wheeler 1986; Smart
et al. 1986). Wet heath and bog species grow in permanently wet, acid sites poor in nitrogen and
many are typical of the oceanic fringe of Europe (Ellenberg 1988).

Peat cutting is the single most important cause of species extinction in the study areas, particularly
Somerset and the Idle valley. Of the 76 species typical of wet heath and bog to have declined, almost
30 have become extinct. This scale of extinction was also observed at Holme Fen in Huntingdon-
shire (Sheail & Wells 1980). All but one of the 33 wetland species to have become extinct at Holme
Fen since 1800 also show a serious decline in one or more of the study areas. The turbary peats of the
Brue valley in Somerset have been cut for many years and those species which require an actively
growing raised bog surface have been exterminated (Hope-Simpson & Willis 1955). When the peat
surface was lowered, calcifuge species were displaced as the bog became flooded by alkaline water
from the adjacent limestone uplands.

c. Decline due to drainage and cultivation/grazing of fen. Primevally, fens were extensive in all three
areas, notably Somerset (Moss 1907; Williams 1970). Fens occur on peaty soils and are
minerotrophic wetlands, dominated by tall grasses, sedges and rushes (Wheeler 1983). Fen species
are typical of moist to permanently wet sites, on weakly acid or basic soils that are nitrogen-deficient
(Ellenberg 1988). Fen peat shrinks and oxidises rapidly following drainage. The consequent
lowering of the land surface puts pressure on the capacity of pumps to remove water and may
necessitate the installation of more powerful pumps (Hutchinson 1980). 60 plants of rich fen and tall
wet meadow declined in one or more study area. In contrast to the many extinctions in acid bog,
most fen species showed only a moderate decline, surviving in fragmented wetlands or on ditch
banks in the grazing marsh (Moss 1907). Early drainage and cultivation converted much fen into
grassland. Typical tall fen species survived in wet hay meadow but were eradicated from pasture.

Fen fragments survived in Somerset in 1990, especially on Altcar soils at the edge of the turbary
moors or where cutting has removed acid peat. Fen meadows are more widespread, both on Altcar
and Midelney soils. In the nineteenth century, fens survived in the Romney Marsh by ‘fleets’
(creeks), near the south coast and, in a modified form, in the >700 ha of rough grazing. By 1980,
some fleets had been eliminated or affected by farming; improvements to sea defences and drainage
had altered coastal wetlands, and rough grazing had been reduced to under 400 ha (M.A.F.F. 1968;
Latimer 1980). Near the R. Idle, washlands, fens and fen meadows occurred widely in the nineteeth
century. The land downstream of Misson was still “*. . . a sea of Glyceria maxima . . .” in 1955

16 J. O. MOUNTFORD

(Howitt day book — 21 July). The fen meadows were drained in the 1950s, and the washes converted
to arable in the 1970s after the upgrading of the Stockwith pump (Howitt & Howitt 1963; Severn-
Trent Water Authority 1974). By 1980, most of the area was intensive arable land with fen
vegetation present largely confined to a few ditch banks.

d. Decline due to drainage of old wet grassland, followed by reseeding or conversion to arable
agriculture. Traditional management of grazing marsh grassland involved either year-round sheep-
grazing or a summer hay cut and aftermath grazing, followed by waterlogging in winter.
Agricultural improvement, with increased use of fertiliser and pesticides, resulted in the conversion
of pasture to arable or increased stocking rates which altered grassland composition. In most grazing
marshes, much grassland was converted to arable in the twentieth century. For example, in 1930, all
Broadland marshes were under grass, but by 1984, 37% of the area was tilled (Driscoll 1985) whilst
in East Essex, coastal grazing marsh declined by 82% after 1938 (Williams & Hall 1987). The
composition of the wet grassland flora overlaps with that of fens but grassland species are more light-
loving, typical of warmer conditions and soils that are somewhat more acidic (Ellenberg 1988).
Nitrogen mineralisation can increase 5—10 fold in meadows following lowering of the water-table.
Thus drainage (with no other agricultural improvement) can also reduce species richness (Grootjans
et al. 1985).

In the study areas, most of the grassland species which have declined are typical of moist
meadows, though some are associated with manured meadows, fairly dry calcareous or poor
swards. Wet grassland species may have spread during the early phase of drainage when mires were
reclaimed for grazing, but with the expansion of arable land, they became restricted to farms
practising traditional grazing. 55 species of wet grassland have declined in one or more of the study
areas, 70% showing only a slight decline. In Somerset, permanent grass is the main land use, but
much of this has been improved resulting in the loss of the old grassland flora (Bradford 1978).
Fertiliser use suppresses sedges, rushes and low-growing wetland forbs (Mountford ef al. 1993). In
the Romney Marsh, the proportion of permanent grass fell from 81% in 1939 to 35% in 1980
(M.A.F.F. parish returns). In the R. Idle area, old grassland is less extensive than in the other study
areas and much rarer than formerly, only surviving near Bawtry, locally elsewhere on the Idle
washlands and around the Misson Line Bank (Mountford & Sheail 1985).

2. Open water: grazing marsh ditches and pools. Mires naturally include areas of open water but
when they are reclaimed, grazing marshes, ditches for drainage and ponds for watering stock are
created. This open water may be further modified as the needs of agriculture change.

a. Decline due to the altered management of ditches and pools. The ability of drainage ditches to act
as refuges for wetland plants was affected by the methods used to manage them. After 1840, the
ditches and ponds of grazing marshes underwent two changes. On the one hand, with improvement
of main channels, pump and under-drainage, some field ditches became redundant and were either
filled in or allowed to silt up, losing their aquatic vegetation through succession. In contrast, the
remaining ditches were liable to more intense management to ensure effective removal of excess
water.

There are rather few quantitative data on ditch removal. In Broadland, 33.5% of dykes were lost
between 1973 and 1981 when old grassland was ploughed for arable farming (Driscoll 1983, 1985).
In similar circumstances in the Pevensey Levels, the loss of dyke habitat was 40% (Palmer 1986).
From 1908 to 1986 in Hatfield Chase, the length of ditch managed by farmers decreased by 36%
(Wingfield & Wade 1988). In Gwent, still mainly pastoral, the loss of ditch length between 1882 and
1975 was only 14% (Wade 1977).

Much of the Somerset Levels remained under grass after 1840, with little under-drainage.
Consequentiy, few ditches became redundant, overgrown or were eliminated. In contrast, there is
ample evidence of ditch elimination in the Romney Marsh, particularly where grassland has
changed to arable (Latimer 1980; Mountford & Sheail 1982). Field survey of the Idle valley in 1983
indicated that 25% of ditches had been destroyed since 1950 (Mountford & Sheail 1985).

Ditches or natural streams are often regraded during drainage schemes, to accommodate
increased run-off of water. Aquatic and bankside vegetation are removed (Hill 1976). If there is no
further dredging, however, most species will return within two years (Haslam 1978). Channel

CHANGE IN ENGLISH GRAZING MARSHES 17

maintenance and weed control can have a profound effect on the vegetation of ditches (Mitchell
1974). Ditch maintenance methods changed after 1950, with mechanical and chemical techniques
replacing manual ones (Newbold ef al. 1989). The intensity of management varies, from more than
once each year in main channels to less than once in ten years in field ditches. Floristic composition
depends both on the management method used (Beltman 1984, 1987) and the frequency and date of
the last maintenance (Wade 1978).

Changes in the extent or management of watercourses were involved in the decline of 73 species.
Most were typical of well-lit sites, but representing the wide range of aquatic and swamp
communities present in grazing marsh ditches (especially Lemnetea, Potamogetonetea and Phragmi-
tetea, but also, locally, three other classes and three alliances (Ellenberg 1988)).

A particular method of ditch maintenance (chemical weed control) may be partly responsible for
the decline of 25 flowering plants and three genera of stonewort (Chara, Nitella and Tolypella).
Aquatic herbicide use is usually confined to drains managed by the National Rivers Authority or
Internal Drainage Boards. Herbicides have short-term effects on aquatic habitats and longer term
impacts dependent on the degree of habitat destruction and herbicide persistence. Different species
have differing susceptibilities (Newbold 1975). Although implicated in the decline of some species
(Howitt, pers. comm. 1983), Wade & Edwards (1980) thought it unlikely that herbicide use had
been responsible for the extinction of any macrophyte in Gwent where there was little evidence of
long term change in the ditch flora. In contrast, all the present study areas showed a decline in
aquatic macrophytes.

b. Decline due to pollution of ditches and pools. Drainage of wetlands and farmlands alters water
chemistry through: 1. increased nitrate nitrogen derived from fertiliser; 2. increased release of
calcium, magnesium and potassium; 3. reduced phosphorus transport due to reduced surface run-
off; 4. soluble pesticides; and 5. oxidation of iron pyrites to colloidal iron hydroxide (‘iron ochre’) in
acid sulphate soils (Hill 1976; Marshall 1981, 1984; Swales 1982). Increased nutrient loads lead to an
increase in the biomass of some macrophytes, whilst others decline (Boar et al. 1989). Overall,
eutrophication reduces species diversity, alters the dominants, increases turbidity and the rate of
sedimentation and causes anoxic conditions to develop (Mason 1981).

In all study areas species whose decline is associated with pollution include those typical of
nitrogen-poor sites (N indicator value <4) (Ellenberg 1988). In addition, in the Romney Marsh and
Somerset, other submerged species have declined, with wider nutrient tolerance but vulnerable to
the reduced light levels following increased turbidity. This may explain the observed spread of
Potamogeton pusillus L. in Broadland and the decline of P. perfoliatus L. (Driscoll 1982).

c. Decline due to fencing or penning of water levels. Open wet soil occurs where water levels are
allowed to vary naturally in response to weather or where stock trample the margins. Conversion to
arable or fencing halts bank poaching (Driscoll 1984) and control of water level (penning in summer
to maintain wet fences or pumping down) eliminates wet mud or alters its seasonal occurrence. The
filling in of redundant farm ponds, once intensely grazed and trampled, has further contributed to
the reduction of the wet mud habitat.

A well defined group of 13 declining species was typical of herbaceous vegetation in this habitat
including wet, often winter-flooded, depressions, newly flooded sites and peat cuttings. The species
are mostly annuals of well-lit sites, over a wide range of soil acidity, but with some trend toward sites
richer in available nitrogen (Ellenberg 1988).

3. Coastal and other habitats. Grazing marshes are usually coastal in Britain, created as a late stage
in the impoundment and reclamation of saltmarsh. Certain species have declined in the study areas
but their reduced population has been caused by factors other than, or additional to, drainage and
watercourse management.

a. Decline due to tide control and improved sea defences. Impoundment leads to decreased salinity
and less exchange between marsh water and estuarine water, both in nutrients and biota (Montague
et al. 1987; Wade & Edwards 1980). In Gwent, main drains were much less saline than the less
frequently dredged minor channels (Wade 1978) whilst in Broadland, saline ground-water seeped
into ditches from the adjacent water-table after drainage (Driscoll 1985).

18 J. 0. MOUNTFORD

There is salt water adjacent to all the study areas, but sea-floods and tidal rivers are now much
more carefully controlled than formerly, reducing the saline input. Hence, 17 coastal species
declined from 1840 to 1990. Such species once occurred not only where coastal vegetation gave way
to grazing marsh but also further inland (Green 1968).

b. Decline due to urbanisation and industrialisation. Human habitation and industry greatly
expanded from 1840 to 1990, and within the study areas, became particularly extensive near the
coast. Certain species, often coastal or halophytic, once mainly occurred in those areas since
urbanised. Such development often cannot be distinguished from the improvement of sea defences
as the cause of their decline. Contemporary accounts record how housing and industry destroyed
notable plant localities, both in the study areas (see Results) and elsewhere (Wade & Edwards
1980).

Increased human population has had indirect effects on the flora. Disposal of urban or industrial
waste puts pressure on wetlands, e.g. pulverised fuel ash in Gordano. Improved transport systems
were built to accommodate traffic, damaging adjacent sites. Sand and gravel extraction is a feature
of the Idle valley, but was not cited as a cause of species decline, and indeed subsequent flooding of
the pits may have increased the diversity of wetlands. However, gravel dredging near Camber
destroyed a site for Ruppia cirrhosa (Petagna) Grande (Hall 1980).

c. Decline due to terrestrial weed control and collection. Some plants are identified by graziers as
poisonous weeds in pasture and removed, e.g. Colchicum autumnale. Weed control and the
drainage or destruction of grassland may be confounded as causes of species decline. A few wetland
species were also arable weeds and were the target of improved cultivation. Direct effects of
herbicides on the wet grassland were noted in Gordano (Willis & Jefferies 1959). Enthusiasm for
plants, every bit as much as antipathy, may lead to them being removed deliberately. The Victorian
fern craze led to the decline of many species, including several from the grazing marshes.

The cause of decline in seven species is unclear. These include taxonomically difficult plants that
may now be overlooked and cultivated species that could require continued reintroduction to
ensure survival in the wild.

CONCLUSIONS

Drainage and land use change modified or destroyed large areas of wetland in England. The
resultant decline and local extinction of many wetland species was recognised by Salisbury (1927)
who showed that the decrease of wetland species was both more pronounced and more
comprehensive than in any other major habitat. If data on species decline since 1930 are examined
at a national scale, the most significant decrease is observed in wetland and grassland species
(Ratcliffe 1984).

This study of three English grazing marshes not only confirms the broad trend, but demonstrates
that between 20 and 33% of declining wetland species have become locally extinct, even over as
large an area as the Somerset Levels and Moors. The decline has been most severe in mires,
particularly raised bogs. There is evidence that the watercourses have become a major refuge for
plants, and that the decline in macrophytes was less marked (Moss 1907; Mountford & Sheail 1989).
However, even in drainage channels, there is ample evidence of reduced populations and
distribution, in which respect the study areas contrast with the relative lack of long-term change in
Gwent (Wade & Edwards 1980). Despite the choice of study areas to reflect differing degrees of
agricultural impact, comparable large declines were observed in all three. Only in the case of
extinction did the intensively farmed Idle/Misson levels show the markedly greater effect that was
predicted. The dependence of the present upon the past in community structure and conservation
can clearly be observed in the flora of grazing marshes.

ACKNOWLEDGMENTS

The results presented here were gathered as part of the botanical component of a project funded by
the then Nature Conservancy Council, whose national (Dr Chris Newbold and Margaret Palmer)

CHANGE IN ENGLISH GRAZING MARSHES 19

and regional staff were a ready source of encouragement and information. My particular thanks go
to Prof. John Sheail (I.T.E.), my collaborator in the larger project. The study depended for its
success on access to a wide variety of published and manuscript information. The then vice-county
recorders of the Botanical Society of the British Isles made available their files and records of plant
distribution: the late Miss E. J. Gibbons, the late Mr R. C. L. Howitt, Mr E. G. Philp, Capt. R. G.
B. Roe and the late Dr W. A. Sledge. Other individuals who had conducted surveys of a particular
study area also provided data: Mr E. G. and Mrs L. B. Burt, Dr J. Hodgson, Dr W. Latimer, Dr F.
Rose, Mr B. Storer, Prof. A. J. Willis and Mrs P. A. Wolseley. Archive material and unpublished
data were contributed by Drs E. J. P. Marshall and P. M. Wade, as well as staff of the local
biological records centres at Bristol, Doncaster, Nottingham and Scunthorpe. Access to data at the
Biological Records Centre (Monk’s Wood) was eased by D. M. Greene and C. D. Preston. The
figures were drawn by Claire Malkowski.

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CHANGE IN ENGLISH GRAZING MARSHES 21

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(Accepted January 1993)

APPENDIX 1: FLORISTIC HISTORY — SOURCES OF INFORMATION CONSULTED

OLD COUNTY FLORAS AND THE NINETEENTH CENTURY

I. Somerset Levels and Moors

SwetE, E. H. (1854). Flora Bristoliensis. London.

CLARK, T. (1856). Catalogue of the rarer plants of the Turf Moors of Somerset. Proceedings of the Somerset
Archaeological Society 7: 64-71.

Waite, J. W. (1886). Flora of the Bristol coalfield. Bristol.

Murray, R. P. (1896). The flora of Somerset. Taunton.

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

Marsua tl, E. S. (1914). A supplement to the Flora of Somerset. Taunton.

[Marshall’s own annotated copy of the supplement (held by his great grandson, Dr E. J. P. Marshall) was

examined. |

II. Romney and Walland Marshes

ARNOLD, F. W. (1887: Ist ed.; 1907: 2nd ed.). Flora of Sussex. London.
Hansury, F. J. & MARSHALL, E. S. (1899). Flora of Kent. London.
WoL LeEy-Dop, A. H. (1937). The flora of Sussex. Hastings.

III. Idle valley and Misson Levels

DEERING, C. (1738). Catalogus stirpium, &c. or a catalogue of plants naturally growing and commonly cultivated
in divers parts of England, more especially about Nottingham. Nottingham.

Orpboyno, T. (1807). Flora Nottinghamiensis or a systematic arrangement of the plants growing naturally in the
county of Nottingham with their Linnean and English names, generic and specific characters in Latin and
English, places of growth and time of flowering. Newark.

Howrrtt, G. (1839). The Nottinghamshire flora. Nottingham.

22 J.O. MOUNTFORD

Legs, F. A. (1888). The flora of West Yorkshire. (Botanical Transactions of the Yorkshire Naturalists Union 2.)
London.

WoopruFFE-PEACOcK, E. A. (1909). A check-list of Lincolnshire plants. Lincoln.

Carr, J. W. (1909-39). Manuscript ‘Flora of Nottinghamshire’. [Copy held at Wollaton Hall Museum,
Nottingham. |

MANUSCRIPTS AND INDICES, C. 1915-1960

I. Somerset Levels and Moors |

1. J. W. White?: note-book (1918) entitled ‘Brue Plants’ [held by Prof. A. J. Willis of Sheffield University].
2. N. Y. Sandwith: Notes and files belonging to Sandwith [held by Prof. A. J. Willis]. Including: a. Complete
series of ‘Bristol Botany in 19—’ (Proceedings of the Bristol Naturalists’ Society); b. Card index for period 1912-
1965, including (all?) published records and data from other workers (e.g. I. M. Roper and H. S. Thompson); c.
Annotated copy of White (1912).

3. Capt. R. G. B. Roe: Notes and files, including all the sources of records he could trace for the period between
the first Flora writers and his own survey, e.g.: a. Card index begun by W. D. Miller (1919-1933), continued by
Dr W. Watson (1933-1952), A. D. Hallam (1952-c. 1960) and Roe (c. 1960 onward); b. Transcriptions of data
from County Museum, Taunton and other herbaria by Roe and Dr P. M. Wade.

II. Romney and Walland marshes

1. Dr F. Rose: manuscripts and field note-books including: a. Manuscript ‘Ecological Flora of Kent’, for period
from 1942 to late 1960s; b. Note-books (especially 1954-1960 when most work on the Romney Marsh was done).
2. Maidstone Museum (E. G. Philp): both pre-1970 records for Kent and herbarium specimens.

III. Idle valley and Misson Levels

1.R.C. L. & B. M. Howitt: Note-books, files and maps covering Nottinghamshire: a. Day-books: record of
botanical excursions, 1951-1978; b. Division files (essentially a manuscript Flora); c. 6-inch Ordnance Survey
maps annotated with locations of rare species.

2. E. J. Gibbons: Files and day-books for Lincolnshire, from c. 1920 onward. [Similar to Howitt. ]

ATLAS OF THE BRITISH FLORA AND CRITICAL SUPPLEMENT (1962 AND 1968)

Covers all three study areas. The maps depict the known distribution of species in the middle part of the present
century with an indication of changes from earlier times. During the preparation of the Atlas more or less
complete vice-county lists were supplied for East Kent by Dr F. Rose and North Lincolnshire by E. J. Gibbons.
Other recorders who contributed records to the Atlas were A. D. and O. M. Hallam, R. C. L. & B. M. Howitt,
Capt. and Mrs R. G. B. Roe and N. Y. Sandwith.

BIOLOGICAL RECORDS CENTRE

Data gathered during the B.S.B.I. Maps Scheme formed the nucleus of the Biological Records Centre (B.R.C.)
at Monks Wood. The original data have been supplemented with records (grid reference, locality, date,
recorder, vice-county). Printouts of B.R.C. data were searched.

MODERN COUNTY FLORAS, AFTER THE ATLAS OF THE BRITISH FLORA

I. Somerset Levels and Moors

Rog, Capt. R. G. B. (1981). The flora of Somerset. Taunton.

[The card index that formed the basis of the Flora and annotated since publication, includes tetrad maps for the
period 1960-1980. Though not published, these data are very detailed and were transcribed fully. ]

II. Romney and Walland Marshes
HALL, P. C. (1980). Sussex plant atlas. Brighton.
Pup, E. G. (1982). Atlas of the Kent flora. Maidstone.

III. Idle valley and Misson Levels

Howrrr, R. C. L. & Howrrt, B. M. (1963). A Flora of Nottinghamshire. Privately published.
Gippons, E. J. (1975). The flora of Lincolnshire. Lincoln.

Gippons, E. J. & Weston, I. (1985). Supplement to the Flora of Lincolnshire. Lincoln.

THE NOTE-BOOKS AND FILES OF CONTEMPORARY BOTANISTS

I. Somerset and Avon Levels

1. B. Storer: Data gathered during and after publication of The natural history of the Somerset Levels (1972); 2nd
ed., 1985: a. Card index for parts of National Grid square ST/3.3; b. Detailed account of Catcott and Westhay
Heaths in note-books; c. Maps, annotated with new records.

CHANGE IN ENGLISH GRAZING MARSHES 25

2. Prof. A. J. Willis: as well as data and indices inherited from earlier workers, his files include: a. Reprints of
‘Bristol Botany’ since 1965; b. Scientific papers with data on Gordano, Shapwick and Catcott Heaths (Willis &
Jefferies 1959; Hope-Simpson et al. 1963; Willis 1967; Hope-Simpson & Willis 1955, 1973).

3. Dr P. M. Wade: species lists made in 1975 in Somerset.

II. Romney and Walland marshes

1. Mrs L. B. Burt: day-books and summary of distribution of wetland plants in the area.
2. Dr F. Rose: note-books with accurate locations.

3. E. G. Philp: Manuscript maps for Ailas.

III. Idle valley and Misson Levels

1. Doncaster Museum (local B.R.C. for South Yorkshire, etc.): Site files and species maps on a 1-km square
basis.

2. Nottingham Museum (Wollaton Hall — local B.R.C.): Site files and individual species data.

3. Scunthorpe Museum (local B.R.C.): Site files, especially L.N.R.s, S.S.S.I.s, etc. Records made after 1970
were noted.

4. Nature Conservancy Council (East Midlands office - Grantham): S.S.S.I. files.

5. Dr J. G. Hodgson (Unit of Comparative Plant Ecology, University of Sheffield): rare species and site data on
detailed data base.

NATURE CONSERVANCY COUNCIL (N.C.C.) COMMISSIONED DITCH AND WETLAND STUDIES

I. Somerset Levels and Moors

SEccomBE, P. (1977). Report on the botanical evaluation of the Rhyne system. N.C.C. (Somerset Wetlands
Project [S.W.P.])

[Areas of conservation interest only. |

BrApForD, R. (1978). A report on the botanical evaluation of the meadows and other nature conservation interests
in the Somerset Levels, 1977-8. N.C.C. (S.W.P.)

RAINE, P. (1980). The flora of drainage ditches — an ecological study of the Somerset Levels. Unpublished M.Sc.
thesis, University College, London.

[North, Southlake and both Salt Moors. ]

WOLSELEY, P. A., PALMER, M. A. & WILLIAMS, R. (1984). The aquatic flora of the Somerset Levels and Moors.
Huntingdon, N.C.C.

[All S.S.S.I.s and P.S.S.S.I.s.]

II. Romney and Walland Marshes

Puituies, N. J. A. (1975). Dykes of Romney Marsh. Unpublished M.Sc. thesis, Wye College, University ot
London.

Latimer, W. (1980). A survey of the dyke flora of the Romney Marsh. Taunton, N.C.C. [S.S.S.1. oniy.]

III. Idle valley and Misson Levels

PacE, S. E. (1980). River Idle Carrs: Botanical survey of Dykes. Huntingdon, N.C.C. (East :fidlands Region).

[Areas of conservation interest only.]

WINGFIELD, M. & WapE, P. M. (1988). Hatfield Chase: the loss of drainage channel habitat. Naturalist 113: 21-
24.

[Overlaps with northern part of study area. ]

APPENDIX 2: RUMEX MARITIMUS L. — FLORISTIC HISTORY IN SOMERSET

Information given as provided in each source.

I. Nineteenth and early twentieth century records

1. White (1886)

In marshes. ““Wedmore . . . June 1843. G. H. K. Thwaites’”’, herb. Stephens. ‘“‘Mouth of the Parrett; Steart
Marsh. J. C. Collins, MSS”, New Botanist’s Guide. Frequent on the peat moors still further south.

2. Murray (1896)

Rare and very local in marshes. District 3 at the mouth of the Parrett and at Steart Marsh, J. C. Collins. District 8
“. . ina bit of marshy ground SE of Tor Hill, Wells, several plants’”’, Miss Livett; plentiful in many parts of the
peat moor and abundant near Shapwick Station, Murray. District 9 (8?) at Wedmore, Thwaites; Knowle Bridge
near Wells, Miss Livett; near Portishead, S. Rootsey.

3. White (1912)

Typical of peat. Rare and local in marshes in the southern portion of area. On the beach at Portishead, c. 1852, J.
N. Duck. “Said to have been found at Portishead by S. Rootsey”’, F/. Som. Several plants in 1884 in a marsh on

24 J. O. MOUNTFORD

Tor Hill, E. of Wells; and at Knowle Bridge, Wookey, Miss Livett. Wedmore, 1843, Thwaites in Herb. Stephens.
Salt-marshes near Highbridge, Sole, MS. Mouth of Parrett and Steart, J. C. Collins, MSS, in New Botanist’s
Guide. Plentiful on the peat moors after any fresh cutting of peat.

4. Marshall (1914)

District 3: by the towing-path of the canal below Maunsel, 1908, Marshall. District 5: Langport, W. Watson.
District 8: salt-marshes near Highbridge (Sole, MS), F/. Bristol (Wedmore locality is probably in this division).
5. White? (1918)

In marshes at Wedmore and on the moors.

II. Mid-twentieth century records

1. Sandwith files, etc.

Bank of a rhine on the Kenn side of Nailsea Moor, 1941, C. I. & N. Y. Sandwith, Bristol Botany in 1941.
Annotated copy of White (1912) confirms status on the peat-moors.

2. Roe files, card indices, etc.

District 3: on the banks of the Parrett at Bridgwater, 1915, H. S. Thompson; between the bridge and the docks,
Bridgwater, H. S. Thompson in TTN; North Newton, Marshall, herb. Druce & Hanbury; by a pool on the edge
of West Sedgemoor below Burton Pynsent, 1918, W. Watson.

District 4: Thorney Moor, 1923 — rare H. Downes in Proceedings of the Somersetshire Archaeological and
Natural History Society |Proc. SANHS}.

District 5: on banks of the Parret, Bridgwater, 1915, H. S. Thompson; small pool, close to Wills Works, right
bank of Parrett, H. S. Thompson in TTN; near Weston Zoyland, C. E. Salmon in LSR.

District 8: near Edington Junction, 1915, J. Bot.; in some quantity N. of Shapwick Station, 1926, Bristol Botany
in 1926; by the sides of a rhine near the station, 5 July 1951, Proc. SANHS; Ashcott Moor, 27 July 1928, H. J.
Gibbons in LSR; specimens from Shapwick dated 1933, herb. J. E. Lousley and H. S$. Thompson in LSR. [Roe
quotes W. Watson as suspecting that the plants at Tor Hill may have been R. palustris.|

District 9: specimen on the bank of a rhine on the Kenn side of Nailsea Moor, 1941, C. I. & N. Y. Sandwith; at
Tickenham Moor, 1930, H. J. Gibbons in LSR.

3. Atlas of the British flora (1962)

Recorded before 1930 in ST/2.4, 3.3, 3.4, 4.2, 4.3, 4.4, 4.7, 5.4. No post-1930 records.

III. Records since 1960

1. Roe files, card indices, etc.

District 8: on old peat cuttings, the Roughet — rapidly appeared in quantity in 1969 after ditches had been cut
across, J. K. Hibberd det. F. Rose [Roe noted there was very little at this site in 1970]; plentiful in some peat
cuttings in Catcott Burtle in 1970, H. W. Boon in Proc. SANHS; several in old peat cuttings, Shapwick Heath in
1980, R. S. Cropper in Bristol Botany in 1980.

2. Roe tetrad maps (notation as Philp 1982) [100-km square ST]

2.3Y (1910s); 2.4Y (1950s); 3.2S (1910s), 3.2T?; 3.3D (1910s); 3.4D (1930s); 4.2G (1920s); 4.3P (1920s); 4.4A,
4.4B, 4.4F (all 1970s onward); 4.7F (1940s). |

3. Roe (1981)

In marshes. Very rare and only seen recently in old peat cuttings on the moors near Shapwick and Westhay
(District 8). Formerly more widespread on the levels in districts 3, 4,.5 and 9 also.

4. Mountford field work (1982-88)

Not recorded.

| Watsonia, 20, 25-31 (1994) 25

Problems in reconstructing floristic change: interpreting the
sources for English grazing marshes

J.O. MOUNTFORD

Institute of Terrestrial Ecology (Natural Environment Research Council), Monks Wood, Abbots
Ripton, PEI7 2LS

ABSTRACT

In attempting to reconstruct floristic change in three English grazing marshes, a wide variety of sources, both
published and manuscript, were employed. These sources cannot be used uncritically as though they represented
points on a uniform monitoring scheme. The sources vary in 1. their objectives; 2. the scale and type of
recording, including the basic recording unit employed; 3. the evenness of the coverage achieved, both
taxonomic and geographical; and 4. in the precision or accuracy of the information included.

Keyworbs: botanical recording, county flora, manuscript source, taxonomy, wetland.

INTRODUCTION

Many authors have attempted to trace the changing status of plant species over recorded history,
using a wide variety of published and archive sources (e.g. Driscoll 1982; Sheail & Wells 1980; Wade
1983). There are fundamental problems in using data as disparate as anecdotal jottings and
systematic surveys to reconstruct change in species distribution and vegetation type. The concept of
recording area, changing understanding of the taxonomy and the rigour and motivation of the
observer all affect the quality and quantity of the data on plant distribution, posing problems in their
interpretation by modern students (Stott 1981).

The present paper describes some of the difficulties encountered in attempting to assess changes
in the flora of three English grazing marshes since 1840: 1. the Somerset Levels and Moors; 2. the
Romney and Walland Marshes; and 3. the Idle/Misson Levels (Mountford & Sheail 1989). The
distribution and abundance of 526 species of vascular plant and species of Charophyte were
investigated through a search of published and manuscript sources (Mountford 1994). Information
on species distribution extracted from these sources was ordered by species and then compiled in
chronological order. The sources employed in that study could not be regarded as representing parts
of a uniform monitoring scheme. There was great variety in the objectives and methodology
adopted in gathering the data both within and between study areas and from recorder to recorder.
Some of the apparent trends over time in species abundance may thus be discounted as artefacts. To
some extent, inconsistencies or inaccuracies could be identified by comparing different sources, but
care had to be exercised in either rejecting or accepting data. Four major problems can be
recognised in the interpretation of these data, arising from: a. the purposes of the survey; b. its unit
of recording; c. the evenness of coverage; and d. the accuracy of the information. In practice, these
problems are inter-related and strongly influenced by the motivation of the botanist which can
introduce a bias into the survey (Rich & Woodruff 1992).

PROBLEMS IN THE INTERPRETATION OF SOURCES

PURPOSES OF THE SURVEY .
Botanists have carried out surveys for widely varied reasons. When he set out to prepare an Atlas of
the Kent flora, Philp (1982) wanted to ensure that every species was fully mapped. The smallest gap
in the map of the distribution of an otherwise ubiquitous species was to be examined. He hoped to

26 J. 0. MOUNTFORD

motivate botanists to fill these gaps. However, if the gap should prove genuine, that absence might
provide an insight into the ecology of that species (E. G. Philp, pers. comm. 1981).

The motives of Victorian naturalists, keeping day-books of interesting finds, were very different.
Many considered their own personal total, or that of their vice-county, to be more important than
the fine detail of species distribution. Some Flora writers sought only to prove a species’ presence in
their county, division or parish. Other authors expected the published work to be simply a guide to
visitors intent on finding interesting plants. More recently, authors hoped to understand the ecology
of species, and gathered considerable environmental and community data to supplement those on
distribution (Graham 1988; Rose MS; Sinker et al. 1985).

THE UNIT OF RECORDING

The purpose of the survey usually determined the basic recording unit employed, if any. The nature
and size of this unit in turn affect the detail of the study and its compatibility with other studies,
particularly if the modern student seeks to make comparisons between surveys. In recent years, the
greater number of botanists, and particularly their increased mobility, has meant that more
ambitious surveys, with smaller units of recording, can be attempted.

However, Flora writing and recording schemes have often been the province of the true amateur,
and botanists have often found it easier to identify with the county or parish than the grid square. A
vice-county and parish recording scheme may result in more enthusiastic and competent recording
than the monotonous working of grid squares (Allen 1983). The parish approach may still be
justified in the 1990s, since it allows direct comparison to be made with the nineteenth century
accounts (D. A. Wells, pers. comm 1990).

The accurate reconstruction of floristic change ideally requires some measure of abundance to
accompany the distributional data. A parish or tetrad record as published may reflect a single
specimen in a precarious locality or a common species which is an important part of the local
vegetation. This detail is seldom available except in an anecdotal form, and underlines the present
need for systematic and regular site monitoring to measure environmental change (Hill & Radford
1986).

The vice-county, drainage basin and civil parish were used as recording units from the mid-
nineteenth century Floras to the more modern studies of Nottinghamshire (Howitt & Howitt 1963)
and Somerset (Roe 1981). There are weaknesses in the use of such units for recording due to their
variable size and their tendency to include several contrasting types of topography. Parish
boundaries were often originally set so as to include a variety of landscapes which could provide the
crops, livestock, fuel and fisheries that the community required. In addition, the prevalence of
winter flooding on the levels meant that villages mainly occurred on the upland fringe with the
parish extending out into the grazing marsh from there, as along the Polden Hills in Somerset
(Havinden 1981). Particularly in the grazing marshes, therefore, a parish is rarely confined to one
soil or landscape type. Interpreting the past species distribution in terms of environmental variation
may not be straightforward. For instance, the parish of Bonnington in Kent is almost equally divided
between the Romney Marsh and the undulating, often wooded land to the north. It is not always
possible to determine whether a record for ‘Bonnington’ refers to the grazing marsh, the upland, or
to both. Specifically, an ancient woodland species is almost certain to be confined to the upland and
a macrophyte of deep, still water is likely to be restricted to the grazing marsh. Species of wet
grassland, however, could occur in either type of landscape.

In some cases, greater precision in recording was achieved with the introduction of the 10-km
square and later the 2-km square ‘tetrad’ of the National Grid as standard recording units. For
example, Ranunculus flammula L. was said to be ““common and generally distributed” in Kent in the
late nineteenth century (Hanbury & Marshall i899). Modern atlases show it to be absent from the
chalk and coastal marshland (Perring & Walters 1976; Philp 1982). It is possible that R. flammula
has disappeared from the grazing marshes (and chalk) over the last century. However, it is more
likely that the authors of Flora of Kent decided to publish a summary account, rather than give a
lengthy list of parishes, thus disguising local but important gaps in the distribution of R. flammula.
Although maps of 10-km square records can reveal patterns in geology, climate or altitude, much
environmental variation is not revealed by such a coarse grid.

The 10-km square represents a large (and often varied) block of land e.g. ST/3.4, north of the
Polden Hills in Somerset, includes parts of 19 civil parishes. In an attempt to assess the impact of

PROBLEMS IN RECONSTRUCTING FLORISTIC CHANGE Dell

land use change in the Idle/Misson Levels, it was important to include only records from west of the
River Trent and south of the River Torne, since this was the area to benefit from the upgrading of
drainage pumps at West Stockwith (Mountford & Sheail 1985; Severn-Trent Water Authority
1974). The study area included parts of 10-km grid squares SE/6.0 and SK/8.9. The Atlas of the
British flora (Perring & Walters 1976) records many wetland species for these two squares which
other archive sources and modern field survey indicate are absent from the study area proper. SE/
6.0 includes parts of Potteric Carr and other well-recorded sites (Lees 1888), whilst SK/8.9 includes
Laughton and Scotton Commons east of the Trent, which have had a diverse, but now
impoverished, wet heath and bog vegetation (Gibbons 1975). Changes in these sites are irrelevant to
a study of the Idle/Misson Levels.

At present, the tetrad is the preferred unit of recording for many county Floras and smaller
projects (Crackles 1990; Hall 1980; Philp 1982). Tetrad maps are at a scale where broad soil and
land-use patterns may emerge, showing those wetland species that are now typical of the grazing
marshes e.g. Eleocharis palustris (L.) Roemer & Schultes, and those that are not, e.g. Filipendula
ulmaria (L.) Maxim. (Philp 1982). With caution, the historical record may then be re-interpreted.

EVENNESS OF COVERAGE

From early accounts of each study area, it is clear that some sites acted as ‘honey-pots’ for botanists,
whereas others, perhaps adjacent, were seldom visited. Botanists were (and still are) tempted to
pursue new species for their own personal lists at reliable localities. This was particularly the case in
the era of the Exchange Clubs when gathering material to trade with fellow botanists was an
incentive. It was comparatively rare for botanists to seek out unvisited sites especially if, as in the
instance of the levels, such areas had a reputation for monotony. For example, the Trent division of
West Yorkshire was neglected by botanists who went instead to Potteric Carr (Don division) for
wetland plants (Lees 1888). Similarly in Nottinghamshire, there were few records for the Misson
and Misterton areas until J. W. Carr began his work at the end of the nineteenth century (Howitt &
Howitt 1963). In Kent and Sussex, botanists crossed the Romney and Walland Marshes to visit and
revisit the Denge Beach with the wetlands around the Open Pits (Rose MS). Whole counties may
have been under-recorded in the past, whether due to a lack of botanists or of perceived interest
(Wells 1989).

Sometimes accessibility led to areas being recorded preferentially (Sheail 1982). Many early
records for the Somerset peat moors are in the Ashcott and Shapwick areas where the Somerset and
Dorset Joint railway made it a simple task for botanists to visit the turbary peats (Marshall 1914;
Murray 1896). Although now known to have arich flora, West Sedgemoor is rarely mentioned in the
records until the 1910s and 1920s when easier transport allowed Dr W. Watson and other botanists
to reach more remote sites (Sandwith files). Access may still influence the data gathered (Rich &
Woodruff 1992). |

Thus many ‘new’ records, produced by modern systematic mapping, reflect the discovery of
plants at sites that had not previously been visited rather than a genuine spread in range. Uneven
coverage can also result from the lack of observers needed to record a large county. There have been
many more active naturalists is Kent, Sussex and Somerset than in the counties making up the Idle/
Misson area. Tetrad atlases require considerable personnel and some species patterns shown may
simply reflect the absence of recorders in some areas. When using diverse sources to reconstruct
floristic change in grazing marshes, it had to be assumed that the description ‘widespread and
common’ implied an even and abundant distribution throughout the area, unless other sources
contradicted that interpretation (Mountford 1994).

Even within a single source, sites may be listed with uneven precision. For example, Hanbury &
Marshall (1899) both observed Sium latifolium L. in the Romney Marsh study area. Hanbury noted
it as: ““. . . abundant in trenches by the roadside, between Ham Street and Ivychurch, and by the
military canal’. This site can be located on modern maps and visited to confirm its presence.
Marshall recorded S. latifolium ‘‘near Appledore’’, implying that it grew in Appledore parish, but
making precise relocation impossible.

Herbaria present particular opportunities and problems as a source of historical plant records.
They allow the modern student to assess the accuracy of past information, by providing
corroborative evidence to the contemporary day-books (Sheail & Wells 1980). However, herbaria
frequently suffer from uneven coverage of species, focusing on rare or taxonomically interesting

28 J. O. MOUNTFORD

plants. There may be many sheets of a species from classic sites, confirming its continued presence.
There is little systematic gathering of common species, with the result that not only can their past
status rarely be demonstrated but also important infraspecific variation may remain unrecognised
(P. D. Sell, pers. comm. 1981).

ACCURACY OF INFORMATION

No two botanists will record exactly the same species in the same numbers at the same site. These
differences are exaggerated when records are made at different times (Kirby et al. 1986). Botanists
are not immune from error and opinions on taxonomy change. It is not always certain that what a
Victorian writer intended by a species or by a site name is identical with what would now be
understood.

The same population may be identified as several different taxa over a number of years. In
addition to Potamogeton pectinatus L., three species of narrow-leaved pondweed proved wide-
spread in field recording of grazing marsh ditches for the present study: Potamogeton berchtoldii
Fieber, P. pusillus L. and P. trichoides Cham. & Schldl. Botanists have been confused as to what
was meant by the Linnaean name P. pusillus, and at different times the other two species have been
named as variants of P. pusillus. The name P. panormitanus Biv. was also applied to one or more of
these pondweeds. In Somerset, a population at Baltonsborough was variously labelled P. pusillus
(vars ‘pseudotrichoides’ and tenuissimus Koch), P. trichoides and ‘P. X_ franconicus’ until
examination of material in the Cambridge University Herbarium from 1881 and modern recording
identified it as P. berchtoldii (Murray 1896; Marshall 1914 — author’s annotated copy; Roe 1981 and
Roe files).

Distinct species may be confounded and botanists record two or more species as one taxon. This is
clearly the case with the Water-Speedwells: Veronica anagallis-aquatica L. and V. catenata Pennell.
Most post-1950 records for Water-Speedwells on the grazing marshes are referred to the pink-
flowered V. catenata (Hall 1980; Howitt & Howitt 1963; Philp 1982; Roe 1981). V. catenata is hardly
mentioned in the records covering the study areas before the Second World War, although it was
described as distinct in 1921 (Pennell 1921). Marshall recorded V. anagallis-aquatica in a ‘‘pretty
form, bearing white flowers tinged with pink” at Headcorn and Westenhanger Castle in Kent
(Hanbury & Marshall 1899), which may have been V. catenata, but this assumption simply begs the
question as to why he did not record it on the Romney Marsh where it is widespread. The earlier
records all refer to one taxon, variously named V. anagallis-aquatica, V. anagallis L. or V. aquatica
Bernh. The two species do sometimes occur together and hybridise (V. x lackschewitzii Keller), but
V. catenata is confined to open, muddy sites with little or no water flow, whilst V. anagallis-aquatica
has a much wider range, often occurring by streams. Taken at face value, the historical data suggest
a huge expansion in the population of V. catenata with a proportionate decline in V. anagallis-
aquatica. Clearly this has not been the case and no assertions as to changing abundance in the
Water-Speedwells can be made from the historical record. However, not all instances of confusion
are SO apparent and real population change may be hidden.

In contrast, variation in one species may be interpreted as the presence of two or more species.
The Creeping Forget-me-not (Myosotis secunda A. Murray) was noted in several of the early
accounts of the Somerset flora:

1. Clevedon; Nailsea Moor; Yatton; Bourton, ditches nearby (Murray 1896).
2. Tickenham Moor; dykebanks near Portbury; moors near Wells (White 1912).
3. North Newton (Marshall 1914).

The only recent Somerset records for M. secunda are from the Blackdown Hills, the Bredon Hills,
Exmoor, the Quantock Hills, Chard Common and sparingly on the Mendip sandstones (Roe 1981).
None of the records for the Levels and Moors have been substantiated and it appears likely that
forms of Myosotis scorpioides L. were mistakenly identified as the upland species (Roe 1981). As
with Veronica anagallis-aquatica, the apparent decline of M. secunda is an artefact of changed
understanding of the taxonomy.

Flora writers in different periods have included a different range of taxa. Some authors included
records of Salix bushes where they may have been originally planted or derived from cultivated
osiers, whilst others rigorously excluded them (listing only clearly native sites). Some authors avidly
recorded aliens (e.g. Azolla filiculoides Lam.), others did not mention them until they became

PROBLEMS IN RECONSTRUCTING FLORISTIC CHANGE 29

thoroughly established or confine their attention to native species (Rich & Woodruff 1992). Aliens
(e.g. Elodea nuttallii (Planchon) H. St John) that resemble established species (e.g. E. canadensis
Michaux) may be overlooked until their identity is clear. In 1982, EF. nuttallii was observed in 10% of
the watercourses sampled in the Somerset Levels and Moors, particularly in larger, more nutrient-
rich rhynes and drains (Mountford & Sheail 1984). Previously it had been recorded once, in 1981,
but the assumption must be that it had been overlooked for some years.

It would be wrong to imply that present workers are more accurate or observant than those of the
past. Indeed the opposite may be true. Between 1850 and 1950, many hybrids and infraspecific taxa
were noted that probably still occur in the grazing marshes, but which many modern recorders
dismiss as specimens that do not quite fit the specification for the taxon. Reconstruction of floristic
change from 150 years of botanical recording might seem to indicate the disappearance of many
hybrids and varieties — a highly unlikely event.

The time span over which data were gathered may become a source of error. In the ‘Plant Atlases’
for Kent (Philp 1982) and Sussex (Hall 1980), records are only included from a period of ten or
twelve years, thus giving an impression of the flora at one time. In the early Floras records dated as
much as 50 years apart may be listed side by side, without distinction. In the particular case of
Lincolnshire, the absence of a nineteenth century work impelled Gibbons (1975) to take stock of the
flora from the beginnings of botanical recording. Thus records in The flora of Lincolnshire from the
Isle of Axholme (embracing the Epworth and Wroot parts of the Idle/Misson study area) include
those of Peck (1815) and those derived from her own studies. Gibbons was very careful to
distinguish such records, but where the aim of the researcher is to trace trends over time, there may
be some confounding of the data.

CONCLUSIONS

Reconstruction of floristic change from published and archive sources requires a critical approach.
Sources vary in nature, scope and accuracy. Variation in the quality and quantity of data produces
spurious trends in abundance over time, which must be identified as doubtful or discounted in any
assessment of the scale of change. Data for 526 species were compiled from three English grazing
marshes between 1840 and 1990 (Mountford 1994). Of the 34 species increasing in distribution,
there was significant doubt in the trend for 18. Similarly, where decreasing species are considered,
46 of the 269 could not be definitely said to have declined. Thirdly, there were many species among
the 526 studied where the sources provided an ambiguous or varied impression, such that the species
had to be assumed to have remained unchanged in abundance or distribution since 1840.
Problems encountered in the use of such a wide range of sources may be summarised thus:

1. Botanists gathered information for widely different reasons — approaches including both
systematic mapping and anecdotal records in day-books.

2. The method and standard unit of recording (if any) changed from 1840 to 1990. The size of the
recording unit varied, and for the first 100 years was not standard.

3. Abundance information was often not noted — a species was simply recorded as present.

4. Coverage of an area was uneven in time and space. Some areas were recorded eagerly, others
were neglected.

5. The systematic recording of 10-km or 2-km squares partly ensured a more thorough coverage of
an area.

6. The accessibility of parts of the study areas changed with time, affecting the information
gathered.

7. Where a species was recorded as ‘widespread and common’, subsequent recording cast doubt on
whether it genuinely had been ubiquitous.

8. The identification and understanding of species’ taxonomic limits and the names by which sites
are labelled varied greatly over the period of study.

9. The range of taxa recorded varied over the 150 years and between recorders. There was particular
variation in the quantity and quality of information available for alien, critical, infraspecific or
hybrid taxa.

30 J. O. MOUNTFORD

10. Particular sources may include information gathered over a long period, making the
identification of trends in time difficult.

The problems described in this historical investigation of grazing marshes are similar to those met
in contemporary studies of plant distribution (Rich & Woodruff 1992). In contrast to the A¢las of the
British flora (Perring & Walters 1976) and the B.S.B.I. Monitoring Scheme, however, the sources
used in this study could not be considered as part of a planned whole. The reconstruction of floristic
change must make use of the data available, in all its variety. The investigator cannot influence the
type or accuracy of data gathered post hoc but must be discriminating in which data are now used
and how they are interpreted.

ACKNOWLEDGMENTS

The many individuals and organisations who contributed to this study are acknowledged in
Mountford (1994). However, I would like to repeat my thanks to the B.S.B.I. recorders who were
so generous with their time and resources: the late Miss E. J. Gibbons, the late Mr R. C. L. Howitt,
Mr E. G. Philp, Capt. R. G. B. Roe and the late Dr W. A. Sledge. In writing an account of problems
with botanical data, I would not want to imply any criticism of these distinguished botanists. On the
contrary, it is partly due to their enthusiasm and critical approach to recording, that I was stimulated
to assess the pitfalls that can arise in the reconstruction of floristic change.

REFERENCES

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MarsHatt, E. S. (1914). A supplement to the Flora of Somerset. Taunton.

Mountrorp, J. O. (1994). Floristic change in English grazing marshes: the impact of 150 years of drainage and
land-use change. Watsonia 20: 3-24.

Movuntrorp, J. O. & SHEAIL, J. (1985). Vegetation and changes in farming practice on the Idle/Misson Levels
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Movuntrorp, J. O. & SHEAIL, J. (1989). The effects of agricultural land use change on the flora of three grazing
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Murray, R. P. (1896). The flora of Somerset. Taunton.

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(Accepted January 1993)

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| Watsonia, 20, 33-39 (1994) 33

Aspects of the ecology and conservation of
Damasonium alisma Miller in Western Europe

C. R. BIRKINSHAW

The Orchard, Stanedge Road, Bakewell, Derbyshire, DE45 IDG

ABSTRACT

Damasonium alisma Miller (Alismataceae) is rare and declining over at least the western part of its range. An
account is given of some aspects of its ecology and life cycle. This information is used to discuss the reasons for its
decline and to provide recommendations for its conservation.

Keyworps: Britain, France, life cycle, habitat restoration, Alismataceae.

INTRODUCTION

Damasonium alisma Miller (Alismataceae), Starfruit, is a semi-aquatic herb (Fig. 1), found in
England, France, Italy, Spain and possibly south-western Asia where there is confusion between it
and D. bourgaei Cosson.

Its populations have declined in recent decades and this species is now “‘very rare” in France
(Aymonin 1974), ‘“‘possibly nationally rare” in Spain (Smith 1988), and “‘endangered”’ in England
(Perring & Farrell 1983) where it is protected under the Wildlife and Countryside Act (1981).

This paper examines aspects of the ecology and life cycle of D. alisma and considers how best this
species may be conserved.

ECOLOGY AND LIFE CYCLE

METHODS
Between 1987 and 1990, information was collected from five habitats of D. alisma including data on
the general nature of each habitat, vegetation structure and percentage of bare ground in 1 m*
quadrats placed over D. alisma plants, associated species of vascular plants (within two metres of
each D. alisma plant), and the physical composition and pH of the substrate (top 6 cm). Three sites
were in England (one in Surrey, two in Buckinghamshire — referred to as Bucks. Sites A and B) and
two in France (in the Loire valley close to Angers and in the Sologne region close to Marcilly-en-
Gault).

Observations on its life cycle were made in the wild and in cultivation — a population was grown
outside in a large tank for three years.

Information on germination was obtained from cultivation experiments; fresh seeds collected
from cultivated plants were sown into pots of sterile loam.

HABITAT

The three English populations grow in shallow ponds, formerly used for watering livestock, on
commons on acid soil. The Sologne population grows in a shallow pond originally dug for fish
husbandry, and that at Angers grows in a periodically flooded sand pit, beside the River Loire. At
each site water levels fluctuated seasonally — they were flooded in winter, but were dry, at least in
part, during summer.

All the sites had disturbed soil, at least in the vicinity of the D. alisma plants; the causes of the
disturbance varied from site to site and included recent pond restoration work (Surrey, Bucks. Site
A), the occasional beaching of a small boat (Sologne), trampling by dogs and people (Bucks. Site B),
and sand extraction (Angers). Many of the English populations of D. alisma recorded in the past

34 C. R. BIRKINSHAW

were on grazed commons, where livestock drinking at the ponds would have caused soil disturbance.

At the sites examined, the vegetation was patchy with large areas of bare substrate where the D.
alisma plants were growing. At the Surrey and Sologne sites D. alisma grew among sparse, stunted
emergents (Sparganium erectum (Surrey) and Typha latifolia (Sologne)). Other small herbs were
present but were infrequent. Bare substrate comprised 75-90% of each 1 m? quadrat. At Bucks. Site
A, D. alisma grew among a sparse carpet of Glyceria fluitans and other small herbs, and bare
substrate comprised 60-90% of the quadrat area. At Bucks. Site B only one plant of D. alisma was
seen in an otherwise bare area. In the Angers sand pit, in the driest, sunniest part of the pit, D.
alisma occurred as tiny terrestrial plants, in small depressions, with the minute, annual Lythrum
borysthenicum and little else; bare ground amounted to 90-98% of the quadrat area. In slightly less
extreme conditions, Starfruit grew scattered amongst a sparse, patchy carpet of small, mainly
annual, herbs. In damp, shaded areas this association was enriched with perennial, creeping grasses;
bare substrate comprised 35-75% of the quadrat area. At Angers a few feeble D. alisma plants were
found in flower growing as true aquatics in shallow pools containing a dense stand of Chara vulgaris,
filamentous algae, perennial grasses and other emergents. Species associated with D. alisma are
listed in Table 1.

Pond vegetation is notoriously difficult to classify. Often it is narrowly zoned and contains several
partly developed communities in close proximity. From year to year the prominence of each
community may vary according to the water level and pond management. Moreover, as ponds are
often artificial their flora is partly the product of chance colonisation. It has not been possible to
classify satisfactorily the vegetation at the five sites as one distinct community, but elements from
several different communities, as defined by Ellenberg (1988), may be recognised.

a. Littorelletea, comprising plants which grow in shallow water at the edge of nutrient-deficient pools
of low pH on sand or gravel (indicator species for this community are marked ‘L’ on Table 1).

b. Isoeto-Nanojuncetea, comprising dwarf plants which grow on land that is inundated only in the
winter (marked ‘T’).

c. Plantaginetea, a pioneer community of pathways and flooded or damp places (marked ‘PI’).

d. Phragmitetea, a swamp community of tall reeds and sedges (marked ‘Ph’).

e. Potamogetonetea, a community of rooted aquatic plants (marked ‘Po’).

The environment preferred by Starfruit frequently resulted in the stunting of species character-
istic of the Phragmitetea and Potamogetonetea communities.

Species not marked as indicators are either aliens (e.g. Paspalum distichum), or plants which
occur in a wide range of damp or aquatic habitats (e.g. Juncus articulatus, Solanum dulcamara), or
arable weeds (e.g. Persicaria maculosa).

The composition of the top 6 cm of soil at four of the study sites (data are not available for Bucks.
Site B), and the pH range and parent material of all sites are shown in Table 2. These data support
observations by Salisbury (1970), Hess et al. (1976), Lousley Co) Rose (1981) and Meikle (1985)
(see also Sowerby 1883).

LIFE CYCLE

D. alisma seeds germinate only below water. Two lots of 120 seeds were sown; eight germinated
below water, while none germinated when sown in merely damp conditions. Most germination
occurred within three months of sowing. Dormant, submerged seeds can be stimulated to germinate
by allowing the seeds to dry and then resubmerging them. 200 seed were sown below water; 14
germinated within three months. The dormant seeds were divided into two lots, half being kept
below water, and the other half was dried and resubmerged. None of the continuously submerged
seeds germinated whereas 22% (20 out of 93) of the dried and resubmerged seeds germinated after
seven months.

Although seed sown in damp conditions but not below water will not germinate, it remains viable
for at least a year and a high proportion (93% of 150 seeds) germinate when submerged.

These observations suggest that dormant seeds remain dormant as long as they are either
constantly damp or constantly submerged.

D. alisma seeds may remain dormant for a long time. In i939, D. alisma re-appeared at Bucks.
Site B after an apparent absence of 87 years (the last report was 1902). Although it is possible that
D. alisma set seeds in intervening years, but was not seen by botanists, the pond was unsuitable for
Starfruit, because of a dense stand of emergents, for at least a decade prior to its restoration. In

DAMASONIUM ALISMA IN WESTERN EUROPE

TABLE 1. SPECIES ASSOCIATED WITH D. ALISMA AT THE FIVE STUDY SITES

(NOMENCLATURE FOLLOWS STACE (1991) AND FLORA EUROPAEA)

Community

Species code* Surrey
Agrostis stolonifera Pl xX
Alisma lanceolatum

A. plantago-aquatica Ph

Alopecurus geniculatus Pl

Apium inundatum Ie

Baldellia ranunculoides Ib,

Bidens tripartita x
Callitriche spp. Po xX

Chara vulgaris
Corrigiola litoralis
Cynodon dactylon
Elatine alsinastrum
E. hexandra
Eleocharis acicularis
E. palustris
Epilobium montanum X ciliatum
E. obscurum
Eragrostis pectinacea
Glyceria declinata
G. fluitans Ph
Gnaphalium uliginosum I
Hypericum humifusum I
Iris pseudacorus Ph
Juncus articulatus
J. bulbosus L
J. effusus
J. cf. tenageia I
Lemna minor
Limosella aquatica I
Lindernia dubia I
Lycopus europaeus
Lythrum borysthenicum I
L. hyssopifolia I
L. portula I
Mentha pulegium I
Ie,
P

ao) anlll alll lilo

~

KKK

Myriophyllum alterniflorum
Oenanthe aquatica
Paspalum distichum
Plantago major Pl
Persicaria hydropiper

P. maculosa

Potamogeton crispus Po
P. natans Po
Pulicaria vulgaris

Ranunculus flammula

R. peltatus Po
R. repens

Rorippa islandica

R. pyrenaica

Solanum dulcamara x
Sparganium erectum Ph

Stellaria uliginosa

Typha latifolia Phe

Veronica scutellata L x

KKK

* community code — see text.

Bucks. A

~ K KK

Pitas

Study Sites

Bucks. B Angers

x
x x
x x

x

Xx

xX

x

x

xX
x
x

x

xX
Xx

xX

x
xX

x

x
x

x

xX

x

ee)
N

Sologne

x
Xx

Kx x xX

KX

~ KKK

36

C. R. BIRKINSHAW

TABLE 2. PHYSICAL COMPOSITION, pH AND PARENT MATERIAL OF THE SUBSTRATE
AT THE D. ALISMA SITES

Parent material

Composition (% dry wt)
Stones (>1 cm diam.)
CaCO; (>1 cm)
Stones (0-2-1 cm)
CaCO, (0-2-1 cm)
Coarse sand
Fine sand
Silt
Clay
CaCO, (<2 cm)*
Organic matter**

pliicas

(range for five samples)

Sologne

Sand/clay
drift

0-0
0-0
2:6
0-0
72-4
12-7),
8-8
DT
0-0
0-8
4-5-5-6

* using a Rothamsted Calcimeter

** by combustion at 350 °C

KKK

Angers

Alluvial
sand

0-0
0-0
8-1
0-0
WARY
10-6
2°8
4-9
0-4
0-5
4-9-6-3

in slurry with water using Gallenkamp pH meter

Site
Surrey

Plateau
drift/
mesozoic &
tertiary
sands

4-8
0-0
10-7
0-0
ZAC2
Shey)
TS7)
21-0
0-9
7
4-0-5-2

Bucks. A Bucks. B
Plateau Gault-lined
drift/clay pond on plateau
with flints drift
0-0 Clay with
0-0 flint gravel
0-0
0-0
6-0
43-2
21:8
27-4
0-0
15
4-3-5-6 4-5-5-6

Figure 1. Damasonium alisma growing at the edge of a fish pond, Sologne, France (June 1989).

DAMASONIUM ALISMA IN WESTERN EUROPE Sy

Surrey, D. alisma reappeared in 1973 (following disturbance of the pond) after an apparent absence
of eight years (H. W. Mackworth-Praed, pers. comm. 1987).

In cultivation, germination is most prolific in early winter (November—January). The winter,
submerged state of Starfruit has short, linear leaves and forms a small ‘grassy’ tuft. The tufts can be
smothered by large pieces of sunken organic matter such as leaves; this vulnerability may explain the
disappearance of Starfruit from ponds now surrounded by woodland. In the spring, long-petioled,
floating leaves are produced. Water levels generally fall in early summer, and the young leaves of D.
alisma plants are exposed. The floating leaves die and are replaced by short-petioled leaves as plants
assume terrestrial growth form. Sometimes, during a wet summer when water levels remain high,
plants retain floating leaves and produce flowers held above water on long peduncles. Flowers
submerged by a rise in water level do not set seed. In cultivation, D. alisma grows well as a true
aquatic plant, but in the wild it is often smothered by vigorous growth of other water plants.

In England anthesis occurs between June and August. Cultivated plants were visited by small flies
(Agromyzidae), beetles (Nitidulidae) and hoverflies (Syriphidae) which feed on pollen or nectar.
These insects often had D. alisma pollen on their bodies and are probably legitimate pollinators for
this species. Starfruit is facultatively autogamous; self-pollination occurs but is infrequent (Vuille
1987).

Terrestrial plants vary in size and can bear from one to about 150 flowers or fruits. The size and
fecundity of an exposed plant is limited by the length of time its substrate remains moist; the quicker
it dries, the smaller the plant will be, and the fewer flowers it will bear. The larger, more floriferous
plants are usually found in shaded habitats or in depressions which retain rain water whereas the
smaller plants occur in exposed, sunny positions.

The fruit of D. alisma usually consists of six follicles arranged like the rays of a star, each
containing one or two seeds. The follicles remain firmly attached to the plant until the fruit is
submerged, after which they dissociate. The proximal seed in each follicle is then released but the
other seed (if present) remains trapped in the follicle — despite being trapped, this seed can
germinate and grow into a young plant.

In cultivation, individual plants, grown as aquatics in a competition-free environment, have
survived for three years, although they become progressively weaker each year. Terrestrial plants
always behave as annuals. No mature plants were seen on autumn visits to English habitats
suggesting that Starfruit behaves as an annual.

CONCLUSION
D. alisma grows in fluctuating ponds with disturbed margins on soil of low pH. These conditions are
not necessary for successful growth and regeneration but tend to reduce competition from more
vigorous species.

Any pond with fluctuating water levels is a harsh, unreliable habitat. Several adaptations assist D.
alisma to exploit this habitat; an annual life cycle, plasticity and two different life forms, and a
germination strategy that prevents total commitment to a single, possibly unfavourable year.

CONSERVATION

THE DECLINE IN ENGLAND

D. alisma has been recorded from more than one hundred localities, in 50 10-km squares, mainly in
the south-east of England. However, at any one time living plants were reported only from a
fraction of these. Starfruit has been considered rare for at least a century (e.g. Sowerby 1883;
Perring & Farrell 1983). During the present century it has become scarcer and by 1990 only three
small populations were reliably reported. This apparent decline may be illustrated using the
summary of records for this species compiled by the Biological Records Centre, Monks Wood:

Period No. of populations verified
1830-1870 21
1870-1910 DS
1910-1950 14
1950-1990 q

38 C. R. BIRKINSHAW

To identify possible reasons for the decline of D. alisma in England, the present conditions at a
number of former sites for this species were surveyed. All these sites are now unsuitable habitats for
the following reasons: ponds had been filled (e.g. Claygate (Surrey), Oxshott (Surrey), West
Molesey (Surrey)); ponds had been abandoned and now contain dense stands of emergents and are
surrounded by woodland (e.g. Whitmoor Common (Surrey), Withybed Corner Pond, Walton-on-
the-Hill (Surrey), Naphill Common (Bucks.), Littleworth Common (Bucks.)); water levels are
artificially regulated for fishing, etc. (e.g. Holmwood Common (Surrey), Wandsworth Common
(Surrey), Earlswood Common (Surrey), Brittens Pond near Guildford (Surrey), Tylers Green
(Bucks.), Coleshill (Bucks.)); ponds are habitats for large flocks of waterfowl (e.g. at Mitcham
Common (Surrey) and the Mere Pond, Walton-on-the-Hill, (Surrey)), which by their trampling,
feeding and defecation cause all but the most resilient aquatic plants to disappear.

MANAGING EXISTING POPULATIONS

The habitat of extant populations of Starfruit should be managed so that water-levels fluctuate
seasonally, open vegetation with areas of bare soil is maintained by removing invasive plants if
necessary, large quantities of smothering organic matter do not accumulate in ponds, and large
flocks of waterfowl are discouraged.

RESTORING POPULATIONS AT FORMER SITES
It is possible that viable, dormant seeds of D. alisma survive at some of its former habitats.
Appropriate restoration may allow populations to become established again through germination of
the dormant seed. Restoration may require the dredging of accumulated organic matter (being
careful not to deepen the pond or to remove the mineral mud which may contain dormant seed),
removal of dense stands of emergents, felling trees, and promoting the seasonal fluctuation of water
levels. The re-appearance of D. alisma following the restoration of a Buckinghamshire pond
provides good grounds for hope that such restoration will be successful elsewhere.

If D. alisma does not re-appear of its own accord after the restoration, a population could be re-
established by transferring seeds from elsewhere. However, this should only be done following the
approval of the national conservation authority and must be fully documented.

CONCLUSIONS

D. alisma has specific habitat requirements and is vulnerable to changes in those habitats. However,
an appreciation of its ecology suggests that extant populations can be managed and the apparent
ability of viable seed to survive, probably for several decades, in the soil suggests that former
populations may be resurrected by the restoration of former habitats.

ACKNOWLEDGMENTS

I am grateful to Dr C. D. Pigott, Ms L. Farrell and Ms J. R. Andrews for their comments on an
earlier draft of this paper and Dr A. Jarman for determining the insects visiting D. alisma flowers. I
would also like to thank the staff of Cambridge Botanic Garden, in particular Mrs J. Free, Mr P.
Freeman and Mrs B. Stace, for their support during the course of my work there. The fieldwork in
the U.K. was funded by the Nature Conservancy Council and the fieldwork in France by a British
Ecological Society’s Small Ecological Project Grant (No. 723).

REFERENCES

AyMonin, G. (1974). Rapport sur les espéces végétales endémiques francaises et sur les espéces végétales les plus
menacées en France. Direction de la Protection de la Nature, Société Botanique de France. Paris.

ELLENBERG, H. (1988). Vegetation ecology of Central Europe. Cambridge.

Hess, H. E., LANpDoLT, E. & HIRZEL, R. (1976). Flora der Schweiz. Basel.

LousLey, J. E. (1976). Flora of Surrey. London.

MEIKLE, R. D. (1985). Flora of Cyprus 2. Kew.

PERRING, F. H. & FARRELL, L. (1983). British red data books: I. Vascular plants. 2nd ed. Lincoln.

DAMASONIUM ALISMA IN WESTERN EUROPE 39

Rose, F. (1981). The wild flower key. London.

Sa.isBurY, E. (1970). Pioneer vegetation of exposed muds. Philosophical Transactions of the Royal Society
B 259: 207-255.

Situ, A. (1988). European status of rare British vascular plants. Nature Conservancy Council, Contract Survey
No. 33. Peterborough.

Sowersy, J. E. (1883). English botany. London.

Stace, C. A. (1991). The New Flora of the British Isles. Cambridge.

VUILLE, F. L. (1987). Reproductive biology of the genus Damasonium (Alismataceae). Plant Systematics and
Evolution 157: 63-71.

(Accepted May 1993)

a pe
Vien) eee
Sie

Watsonia, 20, 41-50 (1994) 4]

Ranunculus ficaria L. sensu lato

PeDs SEE

Herbarium, Department of Plant Sciences, Downing Street, The University, Cambridge,
CB2 3EA

ABSTRACT

The taxonomy, nomenclature and typification of the segregates of Ranunculus ficaria L. (Ranunculaceae) in
Europe are revised, taking particular notice of their biology. Five subspecies are recognised in Europe, four of
which occur in the British Isles. Subsp. calthifolius is confined to east-central and south-east Europe. Subsp.
ficaria is restricted to western Europe. Subsp. bulbilifer occurs throughout much of the range of the species, but
is rare in the Mediterranean region. Subsp. ficariiformis is found in the central and west Mediterranean region
and may be native north to the British Isles. Subsp. chrysocephalus 1s native in the east Mediterranean region
and is grown in British gardens where it spreads naturally. All the subspecies can spread by tubers. Subsp.
bulbilifer and subsp. ficariiformis can spread by axillary bulbils. All except subsp. bulbilifer can spread by seed.
The chromosome number of subsp. calthifolius is unknown. Subsp. ficaria is diploid. The remaining three
subspecies are tetraploid. Triploids have been recorded from the British Isles.

Keryworps: infraspecific taxonomy, reproductive biology, habitats, distribution, variation.

INTRODUCTION

In the broad sense, Ranunculus ficaria L. (Ranunculaceae), the Lesser Celandine, is a gregarious
species, easily recognised by its heart-shaped, bluntly angled or crenate, usually dark green leaves
and shining, golden yellow flowers with 7-13 petals. It is one of the heralds of spring, forming bright
patches in woods, on banks and in other damp places from March to May.

In the taxonomic elucidation of the Ranunculus ficaria aggregate, too much emphasis has been
placed on chromosome counts and not enough consideration given to morphology, ecology and
biology. Five taxa are recognised in this account, which, if cultivated, or if examined at intervals
through their flowering and fruiting periods, are easily recognised. If only seen once in the field,
however, or only a single specimen is available, it is often difficult to identify the plant with
certainty. For this reason and for the fact that intermediates occur and can spread vegetatively, I
have treated them all as subspecies. Three have small flowers and two large flowers. Two of the
small-flowered ones (subsp. calthifolius and subsp. ficaria) are without bulbils in the axils of their
leaves; one (subsp. bulbilifer) has them. Subsp. calthifolius has short stems and leaves congested in a
rosette at anthesis, whereas subsp. ficaria is more spreading with longer, leafy stems. One of the
large-flowered plants (subsp. ficariiformis) has bulbils and the other (subsp. chrysocephalus) is
without. Their ecology and their distribution appear to overlap, but tend to be different. There are
three tetraploids and one diploid. Triploids are recorded. The chromosome number of subsp.
calthifolius is unknown. ;

The nomenclature of the infraspecific taxa is in chaos. I have attempted to clarify it, but
typification of taxa is difficult as ideal specimens are rarely collected, and one needs to know what a
plant does in both flower and fruit. Only the early flowers reach full size. The best specimens are
taken late when fruit and bulbils are developed, but at this stage any flowers still open are usually
late ones, which are mostly smaller than those when the plant first comes into flower. Botanists
should check their local populations, so that we know more about the distribution and ecology of the
subspecies. On the first visit they should note flower size, checking the bulbils and fruits at a later |
date.

42 es ID), SISIUIL
NOMENCLATURE

The first problem is to deal with the typification of Ranunculus ficaria L. The typification of
Linnaean species is carried out by different authors in different ways. My method is that which I try
to apply to all typifications, that is, to try to find material which was available to the author when the
description was written, bearing in mind that the author of the taxon is not always the author of the
description. In carrying out this typification I try not to be biased by my idea of the taxon, unless
there is more than one equally applicable specimen, in which case I would choose one which would
mean continuity of application.
Linnaeus usually validated his taxa in Species plantarum (1753) in one of three different ways:

1. By copying verbatim the description or descriptive name from his earlier works or from the works
of other authors and following it by a reference in that work;

2. By altering the description slightly, but still giving the reference to the original work; or

3. By writing a completely new description with no reference after it.

In the first example the specimens available when the original description was made, or specimens
of synonyms or illustrations cited with the original description, must in my opinion have priority
over those added at a later date, which would not have been available when the description was
written. This is supported by Art. 7.15 of the /nternational code of botanical nomenclature: “When
valid publication is by reference to a pre-starting point description, the latter must be used for
purposes of typification”. From the point of view of typification, I cannot see that there is any
difference between reference to, or actually publishing again, that description. This view is
supported by Heath (1991).

In the second case, although the description is slightly altered, Linnaeus still refers to its source
and thus himself believes the alteration is not important. As the bulk of the description still applies
to the original source and especially as the alteration is often made only because additional species
are involved, I believe the type should still be chosen from the source material.

In the third case, Linnaeus has decided either the taxon is completely new, or none of the earlier
descriptions are satisfactory and he needs to write a new description. In this case all the material in
the protologue will have been seen by him before writing the description and be equally available for
selection, although I would where possible choose the one in the Linnaean herbarium, as that is the
one which is most likely to have been in front of him when writing it.

The diagnosis of Ranunculus ficaria L. (1753) reads “Ranunculus foliis cordatis angulatis
petiolatis” and is referred to Flora Suecica (1745) and there to the Hortus cliffortianus (1738), where
it occurs with the exception of ‘dentatis’ instead of ‘angulatis’. The Hortus cliffortianus is also
referred to in Species plantarum immediately following Flora Suecica. There is a single specimen in
the herbarium of the Hortus Cliffortianus (BM) labelled ‘“‘Chelidonia Minor Rotundifolia CBP
[Caspar Bauhin’s Pinax]’’, a synonym given in the Species plantarum (1753).

When I first became interested in the nomenclature of this species I entered into a correspondence
with V. H. Heywood, and a letter from him dated 1 March 1960 reads “‘As regards the typification, I
was with Tutin and Dandy when the choice of the Hort. Cliff. plant was made and for the record I
think it fair to say that Dandy did not venture a taxonomic opinion. On the occasion I agreed with
Tutin about the identity of the plant, but since then I am not sure that a conclusive determination
can be made of the Hort. Cliff. specimen. However, I think it would be wiser to regard it as the non-
bulbiliferous form so as to avoid disturbing the nomenclature again’’.

I have examined this sheet, which has on it in Dandy’s handwriting, ‘““Type of R. ficaria L.!” There
is a single plant which has three flowers, all of which are about 30 mm in diameter with the petals
probably contiguous. I can see no sign of bulbils in the leaf axils. I agree with Tutin that it is the
small-flowered plant without bulbils. Benson (1954) designated the specimen in the Linnaean
herbarium, Savage Catalogue 715/12, as the lectotype of R. ficaria. This sheet contains three
flowering stems with smaller flowers and less obviously contiguous petals. They are too young to
show any signs of bulbils. The one word changed in the diagnosis in Species plantarum ‘angulatis’, 1s,
if anything, more applicable to the Hort. Cliff. specimen. As stated above, in my opinion all
syntypes are not equal and in this case the Hort. Cliff. specimen was available when the diagnosis
was written and the Linnaean herbarium specimen was not. I am also less certain as to what the
Linnaean herbarium specimen is, but if forced to give an opinion I would say it is probably subsp.

=" ———=

i
7

|

|

Saree

em

RANUNCULUS FICARIA L. AGGREGATE 43

_ bulbilifer. | therefore reject Benson’s lectotypification and designate the Hort. Cliff. specimen as
| the lectotype of R. ficaria L. Subsp. ficaria in this paper is thus as understood by Tutin in Flora
Europaea 1: 234 (1964) and most other current works.

Ficaria verna Hudson (1762) is a new name for Ranunculus ficaria L. and therefore has the same

) type. Ficaria ranunculoides Roth (1788) is also a new name for Ranunculus ficaria L. and has the
| same type, and for that reason must be regarded as an illegitimate substitute for Ficaria verna
/ Hudson. Ranunculus praecox Salisb. (1796) is an illegitimate substitute for R. ficaria L.

R. ficaria var. aurantiacus Turrill (1954) was based on material collected by Miss Alethea Robson
near Windsor, Berks., and plants reproduced from it vegetatively at Kew. A specimen in K is
labelled ‘‘Ranunculus ficaria var., flowers Cadmium Yellow (Ridgw. Pl. III) on back suffused
Vandyke Red (PI. XIII), in a field near Windsor, as a solitary plant of this colour growing with many
ordinary yellow-flowered plants, 27 April 1932, Miss Alethea Robson’’. I designate it as the
lectotype. Another sheet labelled “Ficaria verna var. aurea, origin from near Windsor, Cult. Kew,
22 Feb. 1935” is a paralectotype, although presumably vegetatively produced from the same plant.
The colour of the petals of R. ficaria varies from pale to deep yellow, and Cadmium Yellow is only a
slight increase in the depth of colour and does not seem worthy of recognition. As the plant has no
bulbils and produces good seed, it is placed in the synonymy of subsp. ficaria. R. ficaria var. fertilis
A. R. Clapham (1952) was never described in Latin and has remained invalid.

The other small-flowered plant without bulbils was first named Ficaria calthifolia Reichenb.

_ (1832).Gussone (1844) described a different plant from Sicily as Ranunculus ficaria var. calthifolius,

but he referred back to Reichenbach’s plant and this name must be regarded as a new combination
based on that taxon. Jordan (1847) probably described the same taxon, but he referred back to

_ Gussone and thus indirectly to Reichenbach and he must be regarded as making anew combination

in Ranunculus. R. nudicaulis Kerner (1863), Ficaria intermedia Schur (1866), F. transsilvanica Schur
(1866) and F. pumila Velenovsky (1887) are all referable to this taxon. F. calthifolius was made a
subspecies of R. ficaria by Arcangeli in 1882, R. nudicaulis a subspecies of R. ficaria by Rouy &
Foucaud in 1893, and F. pumila a subspecies of F. verna by Velenovsky in 1898.

The earliest name for the small-flowered plant with axillary bulbils is Ficaria ranunculoides var.
divergens F. W. Schultz (1855). I have not designated a type, but have seen original Schultz
material. The holotype of R. ficaria var. bulbifer Albert (Albert & Jahandiez 1908) is typical of this
subspecies. The type of R. ficaria var. sinuatus Horwood (1916) is also referable to it. E. M.
Marsden-Jones when describing R. ficaria var. bulbifer (1935) apparently did not know that Albert
had used the same name for the same taxon. Marsden-Jones did not cite a type and apart from
remarking that it occurs in all British vice-counties, mentions only two localities, Potterne in
Wiltshire and Kew. I requested of K any material in their herbarium collected previous to the date
of publication of R. ficaria var. bulbifer which could be regarded as types, and received five sheets all
collected by W. B. Turrill at Kew in 1930 and all seen by Marsden-Jones. I have designated one of

_ these sheets as the lectotype of R. ficaria var. bulbifer Marsden-Jones. It is labelled “Ranunculus

ficaria, bulbifer [in pencil], Office of Works, Kew, under lime trees amongst Ivy. About 70 plants

_ more or less female with some 100’s hermaphrodite, 25 April 1930, W. B. Turrill (E. Marsden-Jones

seen on 25 April 1930)’. We shall never know which specimens the description was drawn up from,
but at least this specimen was seen by Marsden-Jones during the studies which led to his publication
of the name. The plants have rounded bulbils in the axils of the leaves and small flowers 14-16 mm in
diameter. The name R. ficaria var. bulbifer Marsden-Jones is an iilegitimate homonym of R. ficaria
var. bulbifer Albert, and R. ficaria subsp. bulbifer Lawalrée (1955), based on Marsden-Jones’
varietal name, is also illegitimate under Art. 64.4 of the /nternational code of botanical nomencla-
ture. Lambinon (1981), therefore gave it anew name, R. ficaria subsp. bulbilifer, the type of which is
that of R. ficaria var. bulbifer Marsden-Jones. Ficaria verna subsp. bulbifera A. & D. Love and
Ficaria bulbifera (A. & D. Love) Holub (1961) are both valid in that genus, and their type is that of
R. ficaria var. bulbifer Marsden-Jones.

The name of the large-flowered plant with bulbils is the one most in doubt. Ficaria grandiflora
Robert (1838) is probably it, but bulbils are not mentioned in the description and I have not seen a
type. The epithet cannot be transferred to Ranunculus owing to R. grandiflorus L. (1753). Schultz
(1858) thus gave it anew name R. ficariiformis. There are no specimens in Helsinki (H) or Geneva
(G), the two herbaria which have some Robert material. Plants from the south of France, however,
do seem to be the plant with bulbils. In view of the lack of original type material, I designate as the

44 PS DeSEEE

neotype of Ficaria grandiflora, and thus also that of R. ficariiformis, an F. Schultz specimen no. ad
407 trom Nice in CGE. It shows clearly the axillary bulbils and large hairy achenes, but the flowers,
which are late ones, are only 30 mm in diameter. R. ficaria subsp. grandiflorus (Robert) Hayek is
valid, but later than, and made illegitimate by, R. ficaria subsp. ficariiformis, which is based on the
same type. Gussone (1844) described the large-flowered plants with bulbils, but his epithet
calthifolius is referable to another taxon. Jordan (1847) said his plant does not have bulbils, but he

referred to Gussone’s plant which does. A Jordan plant from Toulon I have seen is in flower and the |.

specimen does not have bulbils, but it looks like the plant which does. The description of Arcangeli’s ,
R. ficaria subsp. calthifolius refers to this plant, but the name belongs to another taxon. |

R. ficaria forma luxurians Moss (1920) was described from Jersey, and was illustrated by E. W. |
Hunnybun from specimens sent to him from St Aubyns, Jersey, by S. Guiton on 19 February 1913.
The original drawings of Hunnybun are in the Cambridge University herbarium (CGE) and this
information is written on the drawing in Hunnybun’s hand. The drawing was at first labelled ‘“‘var.
calthaefolia vide Rouy & Foucaud Flore de France vol. 1, p. 73” and later “‘Moss says R.
ficariaeformis F. Schultz”. Both the material of Moss and of Hunnybun, used in their work on the
Cambridge British Flora, is now to be found in CGE. The only specimens of this taxon to be found

there are two sheets labelled “Ranunculus ficariaefolia, Jersey, 14 April 1913, E. W. Hunnybun’’. It |

is probable that Hunnybun did not keep the material he drew and these sheets were the only extant
material available to Moss when he described forma /uxurians. One of the sheets contains only basal
leaves, the other, which I designate as the lectotype, contains one flower and one head of achenes. It
is, in my opinion, R. ficaria subsp. ficariiformis.

I can find no name for the remaining large-flowered subspecies with no axillary bulbils, though
some of the above mentioned names have been used for it, and I have described it below as subsp.
chrysocephalus.

TAXONOMY

Ranunculus ficaria L., Sp. Pl. 550 (1753). Habitat in Europae ruderatis, umbrosis spongiosis.
LECTOTYPE: Hort. Cliff. 228 (BM).

Vernacular name: Lesser Celandine.

Description: Perennial, gregarious herb, with whitish, fibrous roots and numerous, whitish or pale
brown, fusiform or clavate root-tubers 5-50 x 3-5-6-0 mm. Stems 3—40 cm, whitish (sometimes
tinted purplish) at base, pale green above, glabrous, branched, ascending or erect, often rooting at
the decumbent base. Leaves numerous, medium to dark green above with paler veins, often
blotched or mottled whitish or purplish, paler and slightly bluish beneath with darker veins,
glabrous or nearly so, rather fleshy; the basal with lamina 0-5—8-0 x 0-5—9-0 cm, broadly ovate,
usually rounded-obtuse at apex, bluntly angled or crenate, rarely shallowly dentate, cordate at base
with basal sinus wide or with overlapping lobes, the petioles up to 28 cm, pale green, with a
sheathing base; the cauline similar to basal but smaller and with short petioles, sometimes with
whitish or pale brown axillary bulbils. Flowers 15—60 mm in diameter, solitary at the end of each
stem branch. Sepals 3 (rarely more), 5—10 X 3-7 mm, pale green, sometimes with a whitish area at
the apex or along the margin, ovate or ovate-lanceolate, concave, obtuse at apex, caducous. Petals
7-13, rarely 0, 6-26 x 3-15 mm, bright, pale to golden yellow, very rarely orange, shining on inside,
dull on outside and sometimes tinted purplish or greenish, fading to white, often contiguous,
narrowly elliptical-oblong, oblanceolate or obovate, obtuse at apex, gradually narrowed at base.
Stamens 5—72; filaments 2-0-8-5 mm, yellow; anthers yellow. Styles 5—72, 1-5—2-0 mm, greenish;
stigmas yellowish. Receptacle concave, with short, pale simple eglandular hairs. Achenes either
abortive or maturing in a globular cluster, when mature 2-5—5-0 x 1-7—3-5 mm, more or less globular
or obovoid, with a cuneate base, keeled, minutely beaked, usually with few to numerous very short
simple eglandular hairs. 2n = 16 (+0-7B), 24, 32.

1. Leaves up to 8 X 9 cm; petioles up to 28 cm; flowers up to 60 mm in diameter; achenes up to 5-0 x
Bei TT aN OU at Vr, SE I EER ETD Ck Oe Js.

RANUNCULUS FICARIA L. AGGREGATE 45

1. Leaves up to 4 x 4cm; petioles up to 15 cm; flowers up to 40 mm in diameter; achenes up to 3-5 x
’}2. Stems rather robust, but straggling; bulbils present in leaf axils after flowering ....................
| d) subsp. ficariiformis
2. Stems robust and erect; without bulbils in leaf axils after flowering ... e) subsp. chrysocephalus
ia Weawes crowded:at base) withifew on short stems. ......00.0. 2.2.00 a) subsp. calthifolius
/3. Leaves less crowded at base and more numerous on the elongated stems ....................... 4.
4. Bulbils not present in leaf axils after flowering; achenes well developed ....... b) subsp. ficaria
4. Bulbils present in leaf-axils after flowering; achenes poorly developed .... c) subsp. bulbilifera

| a. subsp. calthifolius (Reichenb.) Arcangeli, Comp. Fl. Ital. 11 (1882).

_| Synonymy

| Ficaria calthifolia |calthaefolia| Reichenb., Fl. Germ. Excurs. 718 (1832); R. ficaria var. calthifolia

| (Reichenb.) Guss., Fl. Sic. Syn. 2: 41 (1844) quoad basionym. exclud. descript.; R. calthifolius
) (Reichenb.) Jordan, Obs. Pl. Crit. 6: 2 (1847) quoad basionym. exclud. descript.; Ficaria nudicaulis

| Kerner in Oesterr. Bot. Zeitschr. 13: 188 (1863); Ficaria intermedia Schur, Enum. Pl. Transs. 14

| (1866); Ficaria transsilvanica Schur, Enum. Pl. Transs. 14 (1866); Ficaria verna subsp. calthifolia
_(Reichenb.) Nyman, Consp. 7 (1878); Ficaria pumila Velen. in Sitzb. Bohm. Gesell. Wiss. 1887: 438
| (1887); R. ficaria subsp. nudicaulis (A. Kerner) Rouy & Fouc., Fl. Fr. 1: 73 (1893); Ficaria verna
) subsp. pumila (Velen.) Velen., Fl. Bulg. Suppl. I: 6 (1898).

"Illustrations; Reichenb., Ic. Fl. Germ. Helv. 3: t. 1 (1838-1839); T. Savulescu, Fl. Rep. Pop. Rom.
2 ae plnoo. Fie. 6) (1953); M: Josifovic, Fl. Rep. Soc: Serb. 1: 253; t. 37, Fig. 2 (1970).

Description: Plant small, with short stems at anthesis. Leaves crowded at base, few on stems, up to 4
| X 4cm; petiole up to 7 cm, without bulbils in axils. Flowers up to 30 mm in diameter; petals 10-15 x
2-5-6 mm, contiguous; pollen viable. Achenes fertile, c. 2-5 x 2-0 mm, with few to numerous, short,
rigid simple eglandular hairs.

b) subsp. ficaria

Synonymy
| Ficaria verna Hudson, Fl. Angl. 214 (1762) nom. nov. pro R. ficaria L.; Ficaria ranunculoides Roth,
Tent. Fl. Germ. 1: 241 (1788) nom. nov. pro R. ficaria L., nom. superfl. illegit. pro Ficaria verna -
Hudson; R. praecox Salisb., Prodr. Stirp. Allerton 372 (1796) nom. superfl. illegit. pro R. ficaria L.;
| R. ficaria var. aurantiacus Turrill in Bot. Mag. 170: 226 (1954) (LEcToTyPE: field near Windsor,
_Berks., v.c. 23, 27 April 1932, A. Robson (K)); R. ficaria var. fertilis A. R. Clapham in Clapham, A.
R., Tutin, T. G. & Warb., E. F., Fl. Brit. Isles 101 (1952) nom. invalid. sine diagn. latin.; R. ficaria
_ subsp. fertilis Lawralrée in Robyns, Fl. Gén. Belg. (Spermat.) 2: 50 (1955) nom. invalid., sine diagn.
latin.; R. ficaria var. incumbens auct., non F. W. Schultz, Arch. Fl. Jour. Bot. 122 (1855).

Illustration: Ross-Craig, Draw. Brit. Pl. 1: pl. 35 (exclud. H) (1948).

Description: Plant rather rigid, erect and with elongated stems. Leaves up to 5 X 5 cm, numerous on
stems; petiole up to 15 cm, without axillary bulbils. Flowers usually 20-40-mm in diameter; petals
10-20 x 4-9 mm, often contiguous; pollen largely viable. Achenes mostly fertile and well-
_ developed, 2-5—3-5 x 1-7-2-2 mm, with few to numerous simple eglandular hairs. 2n = usually 16,
sometimes 16 + 1—-7B, sometimes 24.

' c) subsp. bulbilifer Lambinon in Bull. Jard. Bot. Nat. Belg. 51: 462 (1981) nom. nov. pro R. ficaria
| var. bulbifer Marsden-Jones, non Albeit.

| Synonymy
' Ficaria ranunculoides var. divergens F. W. Schultz, Arch. Fl. Jour. Bot. 122 (1855) (Described from
_ the Wissembourg area, France); R. ficaria var. bulbifer [bulbifera| Albert in Albert & Jahandiez,

46 BP. DASELE

Cat. PI. Vasc. Var 7 (1908) (HOLOTYPE: Champs inondés l’hiver, Ampus, Var, France, 27 May 1880,
A. Albert (TLON)); R. ficaria var. sinuata Horwood in Rep. botl Exch. Club. Brit. Is. 4: 312 (1916)
(LECTOTYPE: Ratcliffe, Leicester, 29 April 1909, A. R. Horwood (NMW)); R. ficaria var. divergens
(F. W. Schultz) Horwood in Horwood & Noel, Fl. Leicester 19 (1933); R. ficaria var. bulbifer
[bulbifera| Marsden-Jones in J. Linn. Soc. London (Bot.) 50: 40 (1935) (LEcToTyPE: Office of
Works, Kew, under lime trees amongst ivy, 25 April 1930, W. B. Turrill (K)), non Albert (1908); R.
ficaria var. ficaria sensu A. R. Clapham in Clapham, A. R., Tutin, T. G. and Warb., E. F., Fl. Brit.

Isles 101 (1952); R. ficaria subsp. bulbifer Lawalrée in Robyns, Fl. Gén. Belg. (Spermat.) 2: 60 |

(1955) pro R. ficaria var. bulbifer Marsden-Jones, non Albert, nom. illegit.; Ficaria verna subsp.
bulbifera A. & D. Love in Bot. Not. 114: 52 (1961) pro R. ficaria var. bulbifer Marsden-Jones, non
Albert; Ficaria bulbifera (A. & D. Léve) J. Holub in Preslia 33: 400 (1961) pro R. ficaria var.
bulbifer Marsden-Jones, non Albert.

Illustrations: J. Linn. Soc. London (Bot.) 50: 45, Figs 6, 7, 8, 10 (1935).

Description: Plant rather slender, with elongated stems and a loose spreading habit. Leaves up to 4
x 4 cm; petiole up to 15 cm with small, globular, rounded-obtuse, axillary bulbils which reproduce
the plant vegetatively. Flowers usually not more than 25 mm in diameter; petals 6-11 < 2-5 mm,
usually narrow and not contiguous; pollen largely non-viable. Achenes rarely fertile, but sometimes
yielding up to six well-developed ones per flower. 2n = usually 32, sometimes 24.

d) subsp. ficariiformis (F. W. Schultz) Rouy & Fouc., Fl. Fr. 1: 73 (1893).

Synonymy

Ficaria grandiflora Robert, Pl. Phan. Toul. 112 (1838) (NEotyPE: Lieux cultivés, prairies humides,
bords des fosses, champs a Nice, Alpes Maritimes, France, 1866, Rec. Choulette, F. Schultz, herb.
Norm. Suppl. 1, no. ad 407 (CGE)), non R. grandiflorus L., Sp. Pl. 555 (1753); R. ficaria var.
calthifolius [calthefolius| sensu Guss., Fl. Sic. Syn. 2: 41 (1844) quoad descript. exclud. syn.; R.
calthifolius sensu Jordan, Obs. Pl. Crit. 6: 2 (1847) quoad descript., non Ficaria calthifolia
Reichenb., Fl. Germ. Excurs. 718 (1832), nec R. ficaria subsp. calthifolius (Reichenb.) Arcangeli,
Comp. FI. Ital. 11 (1882) quoad basionym.; R. ficariiformis [ficariaeformis| F. W. Schultz, Arch. de
Flore 260 (1858) nom. nov. pro Ficaria grandiflora Robert, non R. grandiflorus L.; R. ficaria subsp.
grandiflora (Robert) Coutinho, Fl. Port. 232 (1913); Ficaria ranunculoides subsp. grandiflora
(Robert) Cadevall & Sallent, Fl. Catalunya 1: 42 (1913); R. ficaria forma luxurians Moss, Camb.
Brit. Fl. 3: 126 (1920) (Lectotype: Jersey, 14 April 1913, E. W. Hunnybun (CGE)); Ficaria verna
subsp. grandiflora (Robert) Hayek in Feddes Repert. 30: 327 (1927) quoad basionym, exclud.
descript.; Ficaria calthifolia subsp. grandiflora (Robert) Trinajstic in Zborn. I. Simp. Biosist.
Jugosl. 160 (1971).

Illustrations: Moss, Camb. Brit. Fl. 3: pl. 128 (1920) as forma /uxurians.

Description: Plant up to 40 cm, rather robust but stems and leaves arching and straggling. Leaves up
to 7 X 7 cm; petioles up to 28 cm, with ovoid or globular axillary bulbils. Flowers up to 50 mm in
diameter; petals 17—26 x 4-12 mm, contiguous or overlapping; pollen largely viable. Achenes 4-5 x
2-5-3-5 mm, covered with numerous very short, rigid, pale simple eglandular hairs. 2n = 32.

e) subsp. chrysocephalus P. D. Sell in Bort. J. Linn. Soc. 106: 117 (1991) (HoLoryPeE: Hort. E. A.
Bowles, Waltham Cross, 4 April 1931, W. T. Stearn (CGE).)

Illustration: Bot. Mag. 153: t. 9199 (1927) as R. ficaria grandiflorus.

Description: Plant up to 40 cm, robust and erect. Leaves up to 8 X 9 cm; petioles up to 21 cm,
without axillary bulbils. Flowers up to 60 mm in diameter; petals 18-25 x 9-15 (—18) mm,

RANUNCULUS FICARIA L. AGGREGATE 47

contiguous or overlapping; pollen largely viable. Achenes 3-4 x 2-0-2:5 mm, covered with
numerous, very short, rigid, pale simple eglandular hairs. 2n = 32.

HABITAT AND DISTRIBUTION

This is difficult to assess, as a large proportion of herbarium sheets cannot be determined with
certainty and my field experience is restricted to the British Isles. There is, however, quite a lot of
published information, which I have been able to put together with reasonable certainty. The
species, as defined here, occurs in a wide range of communities in deciduous woodlands and
hedgerows, on roadsides and ditch banks, in damp pasture, on cliff-ledges and cliff-tops and as a
weed in gardens and lawns. It is found in open habitats and in deep shade on both heavy and light
soils (especially those which are periodically wet), with a pH which varies from 4-4-6-9. Its
altitudinal range in the British Isles is from sea-level to 360 m in N. England, 730 min N. Wales, 550
m in Sutherland and 720 m in Kerry. In Continental Europe it reaches 305 min S. Norway, 1200 min
the Tatra, 1620 m in the Pre-Alps and 1900 m in the Greek mountains. It ranges throughout Europe
to approximately 60° E. in central and south-eastern parts of the former U.S.S.R., and is introduced
in N. America (cf. Meusel et al. 1965, 168; Jalas & Suominen 1989, map 1833).

Subsp. calthifolius is a plant of east-central and south-east Europe, particularly on wood margins,
roadside banks and neglected grassland. Ficaria stepporum Smirnov, from the southern area of the
former U.S.S.R., is a plant I know nothing about, but appears to be similar to this subspecies.

Subsp. ficaria is restricted to western Europe from south-west Norway southwards through
France to the westernmost Mediterranean region. It is rare in Belgium, the Netherlands and
Denmark. I question the records for the central Mediterranean (Jalas & Suominen 1989, map 1836).

Subsp. bulbilifer probably occurs throughout the range of the species, but is rare or absent in the
Mediterranean region. It is probably the only subspecies of cliffs and mountainous regions in many
areas. It is an abundant weed of lawns and gardens, where it spreads rapidly and is very difficult to
eradicate.

Subsp. ficariiformis occurs in the central and west Mediterranean region and may be native north
to the British Isles. Plants from Jersey, Guernsey, the Isles of Scilly and a small patch of low
woodland at St Ishmaels, Pembrokeshire are certainly it. It may be a native in some of these
localities, but in 1979 D. E. Coombe and C. D. Preston failed to find any large-flowered R. ficaria
outside gardens in Guernsey. Elsewhere, it may occur in and escape from gardens. In the drive from
Grange Road to Leckhampton House, Cambridge, it has survived since 1940 (J. Rishbeth in CGE).

Subsp. chrysocephalus would appear to be native of the east Mediterranean region. Hayek (1927
as Ficaria subsp. grandiflora) says the large-flowered plants of Greece and Crete are without bulbils,
and J. R. Akeroyd (pers. comm.) says this is also his experience both in the field and when
cultivating them. In the spring of 1991 he found that it was common in Crete in minimally cultivated
fields, edges of copses and by streams, especially on plateaux between 700 and 800 m; in April 1993
he observed it in S.W. Turkey. Meikle (1977) (as R. ficaria subsp. ficariiformis) says the large-
flowered plant of Cyprus is without bulbils. Turrill first drew attention to this plant when he
published a fine illustration of it in the Botanical Magazine in 1930. W. T. Stearn collected a
specimen of it (CGE) from the garden of E. A. Bowles, which Bowles said was the clone from which
the specimen was taken for this illustration. This is confirmed by an annotation on a specimen in K.
This also seems to be the clone which Marsden-Jones & Turrill (1952) discovered was tetraploid, but
without bulbils. I have used the Stearn specimens as the type of subsp. chrysocephalus.

P. D. Williams grew plants in his garden at Lanarth, Cornwall, which he obtained from Bowles.
Lavender Williams transferred some of these to St Tudy, near Camelford, some of which she also
sent to D. E. Coombe at the Cambridge Botanic Garden. Dr Coombe went to live in Chesterton
Towers, Cambridge in 1982. In the spring of 1983 he discovered it in the garden of 1, Chesterton
Towers, since when it has spread at least 100 m. It there grows with subsp. bulbilifer and appears to
produce no intermediates. In 1979 D. E. Coombe and C. D. Preston collected plants in the garden
of Dr and Mrs D. G. Jameson, Les Fontanelles, Forest, Guernsey, which had been transplanted
from Mill House, Balsham, Cambridgeshire. This suggests that subsp. chrysocephalus, which is a
handsome plant, may be fairly widespread in British gardens, where it could easily spread and
become mixed with the common weed, subsp. bulbilifer.

48 P. D. SEEE
RELATIONSHIP TO MAN, ANIMALS AND FUNGI

Ranunculus species contain the glycoside ranunculin, from which the irritant substance protoane-
monin is formed. This is recorded in R. ficaria. The highest concentration of protoanemonin is
present when the plant is flowering. All domestic animals appear to be susceptible to protoanemo-
nin poisoning, from which various symptoms develop, but it rarely kills them.

To the adherents of the Doctrine of Signatures, the tubers looked like piles and it was formerly
recommended for them both internally and externally, hence its alternative name, Pilewort. Its
acrid nature makes it more suitable as an ointment. The tuber was also likened to a cow’s udders and
hung in the byre to produce more cream in the milk. As a harbinger of spring it is a plant mentioned
by poets. The chocolate brown teleutosori of Uromyces ficariae (Alb. & Schw.) Lév. are very
common on its petioles from March to early June, and honey-coloured spermogonial and orange
aecidial stages of Uromyces dactylidis Otth are common on the undersides of leaves and on the
petioles from March until May. Entyloma ficariae Fischer & Waldh. commonly forms yellowish to
brown spots (delimited by the veins) on the leaves in April and May. Septoria ficariae Desm. is
common on the fading leaves of the plants from May to July.

REPRODUCTIVE BIOLOGY

Plants are diploid (2n = 16), triploid (2n = 24) or tetraploid (2n = 32). Gill etal. (1972) record up to
seven B-chromosomes in diploid plants, but none are recorded for triploids and tetraploids. In the
British Isles diploids are widespread, though tetraploids are more common in the east. Diploid
plants with B-chromosomes are virtually confined to southern England and the Midlands. Triploids
have been recorded from a number of widely separated localities and may be frequent (cf. Marchant
& Brighton (1974)). The diploid records for the British Isles will be referable to subsp. ficaria and
the tetraploid mostly, if not all, to subsp. bulbilifer. The triploids are possibly hybrids between these
two subspecies as they are recorded in their area and may or may not have bulbils. The flowers are
protandrous, the anthers dehiscing extrorsely. The species is normally hermaphrodite, but male
plants (usually diploids) with large petaloid perianth segments, no nectaries and numerous stamens
and carpels abortive or absent occur in some populations, while some plants of tetraploid subsp.
bulbilifer sometimes produce a few smaller female flowers as well as hermaphrodite ones. It is
entomophilous, but self-pollination occurs in the absence of visits by insects, which include
Coleoptera (particularly Meligethes sp.), Diptera, Hymenoptera (particularly Apis mellifera L.) and
Lepidoptera. For a detailed list of species visiting R. ficaria, see Marsden-Jones (1935 & 1937a). A
large proportion of the pollen of those plants found to be triploids and the tetraploid subsp.
bulbilifer is non-viable and few seeds are set, but pollen from diploids and the large-flowered
tetraploids is viable and many achenes are produced. R. ficaria is unusual among the dicotyledons in
having only one cotyledon, and Marsden-Jones (1937b) has shown that it is a single foliar organ, on
which after two or three months, a rudimentary root tuber develops. Clones of all the subspecies can
be produced by division of the root tubers. A greater and quicker spread of clones occurs in subsp.
bulbilifer and subsp. ficariiformis, where axillary bulbils separate off as the shoot system dies and
can produce flowering plants in the first year. Subsp. calthifolius, subsp. ficaria, subsp. ficariiformis
and subsp. chrysocephalus also spread by seed. Marsden-Jones (1935) says seedlings of subsp.
ficaria do not begin flowering until the second year. Flowering normally takes place between
February and May. More detailed accounts of the species occur in Taylor & Markham (1978) and
Grime ef al. (1988), but the general biology and ecology will not properly be understood until
detailed studies of the individual subspecies are made.

By the drive which runs from Grange Road to Leckhampton House at Cambridge, there is a
colony of subsp. ficariiformis which has been there since at least 1940 (fide J. Rishbeth). Subsp.
bulbilifer is also present along the whole length of the drive. Although I have searched there over a
number of years, I have never seen any plants I would call intermediate in morphology. Along a
stream in the Cambridge Botanic Garden, subsp. ficariiformis used to grow with subsp. ficaria. In
that locality there did seem to be morphological intermediates, but they were never examined
cytologically.

Plants from the Leckhampton colony of subsp. ficartiformis and the Coombe/Preston plants from

RANUNCULUS FICARIA L. AGGREGATE 49

La Gouffre, Guernsey, of subsp. chrysocephalus, were grown in pots in the Botanic Garden,
Cambridge, together with examples of subsp. ficaria and subsp. bulbilifer. | was there able to watch
them all together right through the season and R. A. Finch kindly counted their chromosomes for
me. The tetraploid count of subsp. chrysocephalus confirms that made by Marsden-Jones (Marsden
Jones & Turrill 1952). As the plants of subsp. chrysocephalus die, the stalks bend over so that the
falling seeds form a ring round the old plant and later produce a circle of seedlings.

VARIATION

As well as the variation given in the formal account of the five subspecies, there is considerable
variation in flower colour, number of petals, leaf-blotching and width of basal sinus of leaf which to
some extent is genetically controlled, but can occur in all subspecies (Marsden-Jones & Turrill
1952). Var. aurantiacus with orange petals seems to have been found only once, but you can buy it as
‘Cupreus’ from no fewer than 13 nurseries. It did not come true from seed produced either from self-
pollination or cross pollination, but was easily multiplied vegetatively. Its chromosome number was
given as 2n = 20 (presumably 16 + 4B). In Philip & Lord (1991) 25 variants are listed as being
available for sale in a wide range of nurseries. More work is needed on these garden variants to
establish to which subspecies they belong.

ACKNOWLEDGMENTS

I am much indebted to D. E. Coombe, who has continually discussed this aggregate with me
throughout the time I have been working on it, to R. A. Finch for counting the chromosomes of
plants grown in the Cambridge Botanic Garden, and to J. G. Murrell for measuring plants in several
populations and helping to prepare the manuscript. A. O. Chater, C. D. Preston and S. M. Walters
have kindly read drafts of the manuscript.

REFERENCES

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50 P.DSERL

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(Accepted April 1993)

Watsonia, 20, 51-60 (1994) 5]

Calamagrostis stricta (Timm) Koeler, C. canescens (Wigg.)
Roth and their hybrids in S. E. Yorks., v.c. 61,
northern England

F. E. CRACKLES

143 Holmgarth Drive, Bellfield Avenue, Hull, HUS& 9DX

ABSTRACT

The taxonomy of Calamagrostis Adans. (Poaceae) species occurring along Leven Canal, S.E. Yorks., v.c. 61,
has been studied. The populations examined include C. canescens (Wigg.) Roth, C. stricta (Timm) Koeler, the
hybrid C. canescens X C. stricta at two chromosome levels (2n = 28 and 2n = 56), and a strongly introgressed C.
canescens population. The two species show only slight separation in bract length/width and panicle length, but
clear separation in the length of the lowest panicle branch, glume length/width, awn length and point of awn
insertion. The hybrids are variously intermediate in all but one of these characters between those of the
presumed parents. The hybrid with 2n = 28 is sterile and produces little well-formed pollen; that with 2n = 56
produces well-formed pollen and is partially fertile (unlike Swedish Calamagrostis with high chromosome
numbers which produce empty pollen and are apomictic).

KEYworDs: introgression, taxonomy, reproductive biology, chromosome counts.

INTRODUCTION

My interest in the taxonomy, distribution and ecological requirements of Calamagrostis stricta
(Timm) Koeler (syn. C. neglecta auct., Deyeuxia neglecta auct.) was aroused in 1951 by my
discovery that this rare British grass was locally dominant along stretches of the edge of Leven
Canal, S. E. Yorks., v.c. 61 (Crackles 1953). C. canescens (Wigg.) Roth was common along the
canal bank and plants with some morphological features intermediate between those of this species
and C. stricta were present with the two species and believed to be the hybrid C. canescens X C.
stricta which had not previously been recorded for the British Isles.

The morphology of C. stricta, C. canescens and the putative hybrid at Leven was studied adning
1951 and 1952 and these studies formed an invaluable basis for later work on the taxa. Herbarium
specimens to show inflorescences at different stages of development were displayed, alongside
corresponding stages of C. stricta and C. canescens from the same locality together with drawings to
show how floral and vegetative structures of the intermediate material differed from those of the
supposed parents at the Exhibition Meeting of the Botanical Society of the British Isles in 1952
(Crackles 1953). A representative series of specimens was also deposited in K.

Calamagrostis stricta is a very variable grass in the British Isles (Hubbard 1968). At Leven, plants
of C. stricta are less than 95 cm tall and grow by the water’s edge. The leaves are narrow, firm and
rough, the bract making an acute angle with the culm. The short panicle (14 cm or less) 1s narrow
and tight except during the short flowering period. C. canescens plants-are significantly taller and
grow on the canal bank. The leaves are limp, usually with white hairs on the upper leaf surface. The
panicle is lax and nodding after flowering, the hairs surrounding the florets (callus hairs) being
conspicuous in the fruiting panicle.

Plants of the putative C. canescens X C. stricta found in 1951 were of similar height to those of C.
canescens or taller and grew on the sloping canal bank extending to the water’s edge. The leaves were
fairly firm and slightly rough and some had white hairs on the upper leaf surface as in C. canescens; the
bract was horizontal. The panicle, whichis fairly rough to touch, was conspicuous after flowering being -
tight, orange-brown in colour and significantly broader and usually longer thanin C. stricta. The ligule
of the putative hybrid was often longer than that of C. canescens (ligule of C. strictalessthan 3mm, of C.
canescens rarely greater than 4 mm, that of C. canescens X C. stricta often to 5 mm).

52 PE. CRACKEES

In April, 1952, Dr Hubbard agreed with the identification of the hybrid (pers. comm.), but some
months later seemed to have some misgivings, saying that no British hybrid grass is male fertile even
though the putative hybrid at Leven was. In 1954, Dr Nygren of Ultima Agricultural College,
Sweden, saw putative hybrid material from Leven Canal and confirmed its identity, although he
considered that this taxon was morphologically distinct from its Swedish counterpart and forecast a
high chromosome number.

In 1970, Dr D. W. Shimwell suggested that I should make a biosystematic and taxonomic study of
the Leven Calamagrostis populations. Several visits were paid to the canal in that year and the
Calamagrostis populations examined carefully.

The stretch of bank on which the putative C. canescens X C. stricta was found in 1951 was by now
overgrown with Phragmites australis (Cav.) Trin. ex Steudel and surviving plants flowered rarely.
However, morphologically similar plants to those studied in 1951 and with characters intermediate
between those of C. stricta and C. canescens and previously described in this paper were found to be
dominant along a stretch of 22 m of the north bank and approximately 450 m to the east of the Far
Fox Aqueduct. This bed occurred on the slope of the bank above the canal edge footpath, except for
the easternmost 8 m which was by the water’s edge, thus occupying a very similar position on the
canal bank to that occupied by the 1951 hybrid. In August, this population, which I shall call the H,
population, is conspicuous by virtue of its broad orange-brown panicles as was the 1951 putative
hybrid population. Pollen grains were well-formed as in that taxon.

Another population of the putative hybrid C. canescens X C. stricta was found in 1970 growing at
the edge of the water along the north bank, nearly opposite to Carr Farm. The plants had narrow,
more or less strict, spikes in the fruiting condition and could have been overlooked for C. stricta.
This taxon was also intermediate between the two parents in a number of its characters; it was
identified ‘in the field’ by the length and position of the awn. Using a hand-lens, the awn was seen to
arise about half-way up the lemma as in the H, population and the 1951 putative hybrid and did not
project beyond the lemma. This taxon differs from C. stricta in having a far longer, projecting
bottom branch to the panicle and in other respects. This population, which I shall call the H>, differs
from the H, in having empty pollen and in being conspicuous in flower as the exserted anthers are
yellow (purple in the H, and in the parental species).

A bed of C. canescens growing at the edge of a dyke, parallel to the northern bank of the canal and
a few metres to the north of it was believed to be strongly introgressed (the I, population); since, for
example, the glumes and callus hairs were of markedly different length in one spike and the panicle
was not so lax and not nodding.

In addition, a single plant found on an unusually steep stretch of the canal bank, designated as the
Leven H3, is a recombinant with both C. stricta and C. canescens characters not present in the H,;
and H, populations.

Photographs of fruiting panicles of C. canescens X C. stricta, H,; and H; individuals, are presented
together with those of the parental species for comparison (Fig. 1).

THE AREA

Leven Canal is 5-2 km in length and extends from the village of Leven, GR TA/106.453, westwards to
meet the River Hull, c. 12-5 km south of Great Driffield where spring-fed becks converge. The canal
was cut in 1802 and was sealed off from the river in 1934. The River Hull valley, in common with the
rest of Holderness, was formerly an area of extensive marshes and irregular meres with islands of
higher ground here and there. Much of the valley was flooded for several months each year and the
meres, variable in extent, represented the remains of the flood water which lingered in the lowest part
throughout the summer (Sheppard 1957). The northern part of the valley remained relatively
unchanged until the major drainage scheme of 1764 took effect (Sheppard 1958) but this caused little
or no change to Leven Carrs. A plan of the Holderness level by A. Bower, dated 1781, in the Hull City
local history library, shows irregularly shaped meres in Leven and Tickton Carrs. Comparison of this
map with the Ordnance Survey 2-5 inch to 1 mile (sheet TA/0.4), suggested that the canal was cut
through the meres. A later plan by E. Pearson, dated 1831, confirmed that the canal cut through two
meres, or their sites, and that it passed through the edge of a lake at the place where C. stricta was first
discovered (Crackles 1968). There is supporting evidence of standing water on Leven Carrs in the

CALAMAGROSTIS SPP. AND THEIR HYBRIDS IN S.E. YORKS 53

Ly
Y

y
yy,

FicureE 1. Fruiting panicles of specimens collected at Leven Canal in 1970: (a) C. stricta; (b) C. canescens X C.
stricta (H, population); (c) C. canescens X C. stricta (H2 population); and (d) C. canescens. The scale bar
represents 2 cm.

nineteenth century in a manuscript herbal compiled by the Rev. W. Whytehead while he was vicar of
Atwick, near Hornsea, S. E. Yorks. (1757-1817) in which he states that White and Yellow Water-
lilies and Water Soldier occurred in ““Leaven Carrs’. This observation was made before the
construction of the canal which is not mentioned in the manuscript.

The presence of scarce or local British species in or by the canal provides strong support for the
view that the canal derived its flora from the former meres and adjacent marshes. Scarce British
species recorded include Calamagrostis stricta, Carex appropinquata Schum., C. elata All.,
Myriophyllum verticillatum L., Potamogeton friesii Rupr. and Sium latifolium L. Other species
recorded here included Apium inundatum (L.) H. G. Reichb., Butomus umbellatus L., Carex
paniculata L., C. rostrata Stokes, C. vesicaria L., Hippuris vulgaris L., Hottonia palustris L.,
Lysimachia vulgaris L., Lythrum salicaria L., Myriophyllum spicatum L., Nuphar lutea (L.) Smith,
Nymphaea alba L., Oenanthe fistulosa L., Potamogeton alpinus Balbis, P. lucens L., P. natans L.,
Ranunculus circinatus Sibth., Sagina nodosa (L.) Fenzl, Sagittaria sagittifolia L., Samolus valerandi
L., Schoenoplectus lacustris (L.) Palla and Utricularia vulgaris L., several being uncommon or rare
vice-county species.

In addition to extensive annual flooding of the carrland, which still occurred in winter until c. 40
years ago, there were springs in the carrs. For example, in the early 1970s, I noticed a spring in the
canal, the resultant ripples reaching the north bank at the point at which beds of Phragmites australis
end and those of Schoenoplectus lacustris and Calamagrostis stricta begin.

METHOD OF STUDY

25 shoots of C. stricta, C. canescens and the putative hybrids, population H, and H; and 20 shoots of
the I, population were collected at random. Material was collected at the end of July or early
August, 1970, when the panicles were in their after-flowering condition and comparison of the
maximum number of floret characters could be made.

Measurements of bract length and width, and also of panicle length and width and length of the
bottom branch of the panicle were recorded for each individual shoot of the population samples of
each taxon. All bract measurements were obtained ‘in the field’ because of the tendency of the
leaves of some taxa to inroll on drying. In the case of the spikelet characters, five spikelets for each
individual shoot of each taxon, selected at random, were used to provide average values.

54 FB] CRACKERS

It is important to consider the variation between the different taxa, one character at a time. Such
variation is shown for six characters (Figs 2-7) with the range of variation, mean and standard
deviation given for each character.

REPRODUCTIVE BIOLOGY AND CHROMOSOME COUNTS

All the Calamagrostis material studied was perennial and all spread vegetatively by means of
rhizomes; the H, population is particularly vigorous in this respect.

C. stricta is the first of the Leven Calamagrostis populations to flower. In most years, many spikes
of C. stricta are open by mid-June, while C. canescens is not normally in good flower until the first
week in July. There may be a slight overlap of flowering times as in 1952 when a few spikes of C.
stricta were still open on 22 June when the first spikes of C. canescens were also open. When the
early part of the summer is cold, as in 1972, flowering of C. stricta may be delayed by as much as ten
days. Typically, each spike of C. stricta closes in five days, but in 1971 tightening of spikes was
delayed, only some spikes being in the closed condition by 4 July. In both these situations, there was
a significant overlap of flowering of the two species. The flowering time of the H, population is
intermediate between that of the parents, full flowering usually being attained in the fourth week of
June. Spikes of the H,; population appear to take longer to close than those of C. stricta, closing over
a period of ten days. The H> population flowers slightly earlier than C. canescens or at the same
time.

In the case of all the Leven Calamagrostis plants studied, the anthers were found to be I-shaped
and yellow, while still in the flower. The filament is coiled while the anthers are in the spikelet and
more or less straight when the anthers are protruding, so that protrusion of anthers is presumably
spontaneous.

A spike of C. stricta half-way out of the sheath was examined, while obtaining meiotic stages for a
chromosome count. Spikelets near the top of that part of the spike still enclosed in the sheath were
green and still contained pollen mother cells undergoing meiosis, while the purplish spikelets higher
up the spike contained fully-formed pollen grains. On the same date, a spike fully out of the sheath
was found to have pollen grains fully formed half-way up the spike.

Samples of ten anthers taken at random were examined. Observations concerning shape,
measurements and colour of the anthers of the various taxa being studied were made and the
percentage of well-formed pollen produced by each taxon was recorded (Table 1).

C. stricta plants from Leven Canal were found to set some seed by self-fertilisation while C.
canescens appeared to be self-sterile, a conclusion also reached by Nygren (1946) in Sweden. The I,
population, believed to be a C. canescens introgressant, is self-fertile.

The H, population produced a good percentage of fully-formed pollen grains. Isolated Hy,
panicles produced an abundance of imperfect caryopses although two plants were reared from the
contents of half an isolated panicle. Thus the evidence indicates normal sexual reproduction with
slight self-fertility. No caryopses were produced by the H), plants.

All the caryopses of the Calamagrostis taxa examined are very small and the seeds are presumably
shed enclosed in the lemma and palea to which the callus hairs are attached. C. stricta has short
stubby caryopses less than 1 mm long. The C. canescens caryopsis is c. 1 mm long with a short
narrow neck subtending a narrow platform. The caryopsis of the H, putative hybrid is 1-0-1-3 mm
long, the top of the fruit being distinctively shaped, splaying out gradually from a restriction at the
neck. The caryopses of the putative C. canescens introgressant, population I,, was very heavily
attacked by the gall-forming mite, Steneotarsonemus spinosus Schoarschmit, the effect of the attack
being to cause the lemma and palea to harden and swell. Other C. canescens populations were not
affected.

For the purpose of obtaining chromosome counts, root tips of each taxon were placed in 1%
colchicine solution for up to four hours to spread the chromosomes. Squashes were then prepared
and stained with aceto-carmine. Difficulty was experienced in perfecting the squash technique so
that the very small chromosomes were sufficiently spread to count. Meiosis was also studied in all
five taxa. Flowering spikes were fixed in 3:1 absolute alcohol to glacial acetic acid overnight and
preserved in 95% ethanol. Squashes to show the meiotic stages were stained in aceto-carmine after
pre-treatment with a drop of N HCI to aid the spreading of chromosomes.

CALAMAGROSTIS SPP. AND THEIR HYBRIDS IN S.E. YORKS 35

eee 1, population
$$ C. canescens
nn a _§ H, population
—_-) go IE — H, population
~ R$ Guwicia

2 3 4 5 6 1

Ficure 2. Bract length/bract width of Calamagrostis taxa showing mean (vertical line), range (thin line) and
standard deviation (thick line) for Leven population samples.

:
)
)

| CoRR
ner I, population
— C. canescens
ei H, population
=a H, population
—- =a — C. stricta

2 6 10 14 (Sihae DD 26 cm

Figure 3. Panicle length of Calamagrostis taxa showing mean (vertical line), range (thin line) and standard
deviation (thick line) for Leven population samples. The open horizontal bar depicts the zone mid-way between
the limits of standard deviation for C. stricta and C. canescens.

_ aa)

—____ a ———_——- I, population
$a —— C. canescens
a ee H, population
a H, population
a __ C. stricta

Figure 4. Length of bottom branch of the panicle of Calamagrostis taxa showing mean (vertical line), range (thin
line) and standard deviation (thick line) for Leven population samples. The open horizontal bar depicts the zone
mid-way between the limits of standard deviation for C. stricta and C. canescens.

56 Fos CRACKEES

sees} I, population
$$$ C. canescens
— a —$§ H, population
se H, population
ee C. stricta

8 4 5 6 7 8 9

Ficure 5. Glume length/glume width of Calamagrostis taxa showing mean (vertical line), range (thin line) and
standard deviation (thick line) for Leven population samples. The open horizontal bar depicts the zone mid-way
between the limits of standard deviation for C. stricta and C. canescens.

[a ee ee ae)

a I, population

ae C. canescens
eras — H, population
| a —§ H, population
—_—_—§ C. stricta

|
0 1 D 3 4mm

FicurE 6. Awn length of Calamagrostis taxa showing mean (vertical line), range (thin line) and standard
deviation (thick line) for Leven population samples. The open horizontal bar depicts the zone mid-way between
the limits of standard deviation for C. stricta and C. canescens.

The chromosome number of the two parental species and the H> population is approximately 2n
= 28, corresponding to counts determined by Nygren (1946) for the two species and the hybrid in
Sweden. It was established from root tip squashes that the putative Calamagrostis canescens X C.
stricta population H, had a significantly higher number. A good preparation of the second
metaphase of the H, was finally obtained and the chromosome number found to be 2n = 56.

DISCUSSION

My understanding of the species C. stricta and C. canescens as a result of the Leven studies are in
close agreement with a short description of the type specimen of C. stricta (Arundo Stricta
(Steifahrichtes Rohr.)) by Siemssen (1795) and of the two species as described by Hylander (1953)
and of that of C. stricta given by Nygren (1946). It was therefore decided to make the provisional
assumption that the range of variation in Leven C. stricta and C. canescens gives an indication of
variation usually found in populations of these species.

The hybrid population studied in 1951 was essentially the same taxon as the H, population and it

CALAMAGROSTIS SPP. AND THEIR HYBRIDS IN S.E. YORKS 57

SSS eee eens]

Ne a I, population
~ h- C. canescens
a H, population
a H, population
a C. stricta

0 20 40 60 80 100%
FicureE 7. Point of awn insertion measured from the base of the floret, as a percentage of floret height, of
Calamagrostis taxa showing mean (vertical line), range (thin line) and standard deviation (thick line) for Leven
population samples. The open horizontal bar depicts the zone mid-way between the limits of standard deviation
for C. stricta and C. canescens.

TABLE 1. ANTHER CHARACTERISTICS OF LEVEN CALAMAGROSTIS MATERIAL

Populations
Character C. stricta C. canescens H, H, I,
Length (mm) (1-4-)1-8 1:6 2-0 1-4 1-4
Shape of exserted anther I-shaped X-shaped Y-shaped I-shaped X-shaped
% fully-formed pollen 877% 85-57% 59% 4% 65%
Colour of anther Purplish Purple Purple Yellow Purple

Numerical values are means of ten anthers.

occupied a very similar position on the canal bank, the stand extending from the canal edge, but
mainly occupying a stretch of the upper part of the sloping bank. I did not have the facilities to
obtain a chromosome count in 1951, but I have no doubt that the 1951 hybrid like the H, is the
octoploid, C. canescens X C. stricta, 2n = 56. The evidence for this is that the 1951 hybrid and the H,
population are essentially similar morphologically, notably in length and width of glumes and the
broad, tight fruiting panicle and also in producing well-formed pollen grains. The 1951 hybrid
population was, however, much more morphologically uniform than the H, population. It is
thought that the stand formed by the 1951 hybrid population may have arisen from one or more F,
or F, plants. The plants were very vigorous as are those of the H, population.

In the case of both the H, and H; populations, the question arises as to whether the populations
represent a clone or a hybrid swarm. Each population forms a continuous stand which leads one to
suspect that each population is a clone; on the other hand the degree and nature of morphological
variation in each appears to rule out this possibility.

In the case of the H, and H; populations, the range of variation for a character (Figs 2-7) may
overlap markedly with the range for that character in one of the parents or be continuous or
discontinuous with it. The range of measurements for a number of characters of the H, and H,
populations were found to be intermediate when compared with the range of measurements for the
corresponding character in each species. This was true of glume shape (Fig. 5), awn length (Fig. 6)
and position of awn insertion (Fig. 7). However, the range of variation for each of these characters
in the H, and H; populations do not lie in the same intermediate position in relation to the variation
for the same character in the two species.

The range of measurements for the C. canescens I, population is seen to show a shift towards the
range for the hybrids in the case of panicle (Fig. 3) and bottom branch length (Fig. 4). The awn tends
to be longer and its insertion on the lemma lower than in typical C. canescens.

The study of the development of the anthers in C. stricta suggests a correlation between the

58 F. E. CRACKLES

change of glume colour from green to purple, the completion of meiosis and the change of anther
colour from yellow to purple. In C. canescens and the H, hybrid population, the change of glume
and anther colour is also associated with the change of anther shape from I-shape to X- or Y-shape
respectively and the production of well-formed pollen. Such a correlation could explain the
predominantly yellow colour and the I-shape of exserted anthers of the H, population which is
almost male sterile. It is interesting to note in this connection that Calamagrostis purpurea, which
produces empty pollen and is apomictic, was found by Nygren to have yellow, I-shaped anthers and
these features were used by him to separate C. purpurea from other Calamagrostis taxa in
Scandinavia (Nygren 1948). However, these are seen not to be exclusive characters of C. purpurea
which is essentially a taxon at a higher chromosome level than the Leven H> population and
morphologically distinct from it.

Observations regarding the reproductive biology of the H, and Hy; hybrid populations are at
variance with the findings of Nygren (1946, 1962) concerning their Scandinavian counterparts. The
Leven H; C. canescens X C. stricta (2n = 28) population appears to set no seed and to produce a
very low percentage of morphologically good pollen while the F; and F, generations of this hybrid in
Sweden were found to produce good pollen (Nygren 1946) and to have low fertility (Nygren 1962).
The Leven H, C. canescens X C. stricta (2n = 56) population has morphologically good pollen and
low self-fertility while all Swedish Calamagrostis taxa with a chromosome number of 2n = 46
upwards produce empty pollen grains and are apomictic (Nygren 1946) except when young and with
the exception of one population of mature plants of C. purpurea in GAllivare (Nygren 1946). It is of

interest that similar taxa may show differences in reproductive biology in different parts of their |

geographical range.

DESCRIPTION OF THE LEVEN CALAMAGROSTIS TAXA

Calamagrostis stricta (Timm) Koeler, 2n = 28

Occurs by standing water at Leven Canal. Perennial, forming tufts and having slender rhizomes.
Height: 30-94 cm. Culm erect, 2-4 noded, rough or smooth near the panicle. Leaves green; bract
narrow, stiff and rough, making an acute angle with the culm; short, more or less truncate ligule,
that of the bract 1-3-2 mm long. One or more leaf sheaths are always hairy. The panicle is short,
(6-)9-13(-15) cm long, tight and narrow before and after flowering; less than 2 cm wide, usually less
than 1-5 cm wide after flowering. The lowest branch of the panicle is short being, less than 4 cm,
average 2:7 cm. The glumes are less than 4 mm long and 1 mm wide at the broadest part; more or less
acute and rough on the keel; the outer glume one-nerved and the upper three-nerved. The lemma is
2-6-3-2 mm long and five-nerved. The callus hairs (hairs surrounding the floret) are appreciably
shorter than the lemma, 1-9—2-5 mm long, average 2-2 mm. The awn arises from one-third of the
way up the lemma or below and is long (1-9-2-8 mm, average 2-3 mm) projecting beyond the top of
the floret. The spikelet axis is prolonged as a minute bristle. The anthers are I-shaped, purplish and
produce well-formed pollen. It is usually in full flower at Leven in mid-June, with all the spikes
usually having tightened within eleven days, but the tightening of the spikes may be delayed in
certain years.

Calamagrostis canescens (Wigg.) Roth, 2n = 28

Occurs on both the slopes and the top of the canal bank. Shade tolerant. Perennial, rhizomatous,
forming beds. Usually a much taller plant than C. stricta (60-110 cm). Culm branched. Broad,
yellow-green, limp and arcuate leaves are typical. The upper surface of the leaf blade usually has
long, white hairs. The ligule of the bract is 2-6-6 mm long and tears readily. All leaf sheaths are
glabrous. Culms (4-), 5— or 6-noded, smooth near the panicle. The panicle is long, (15—)19—22(—25)
cm, lax and typically nodding after flowering. The lowest branch of the panicle is long, 5—9 cm. The
glumes are narrowly lanceolate in side view, 4-5-6 mm long and drawn out to a point which breaks
off at maturity; they are 0-6-0-7 mm wide at their broadest part. Both glumes are one-nerved. The
lemma is 2-6—-3-0 mm long and three-nerved. The callus hairs are appreciably longer (by 0-7—1-3
mm) than the floret. The awn is very delicate and short, leaving the lemma at or near the tip. The
anthers are X-shaped and purple and produce well-formed pollen. The taxon is in full flower in early
July at Leven. Self sterile.

CALAMAGROSTIS SPP. AND THEIR HYBRIDS IN S.E. YORKS 59

Calamagrostis canescens 1, population

Believed to be an introgressed C. canescens population arising by backcrossing from C. canescens X
C. stricta to C. canescens. The population occurred by the side of a drainage dyke parallel to and
near to Leven Canal. Perennial, rhizomatous, forming beds. Generally similar to C. canescens sensu
stricto, but differs by having leaves narrower and more or less erect; culm rough near panicle;
panicle not so lax and not nodding; glumes varying in length in one spike; awn insertion distinctly
below the tip of the lemma, i.e. 0-3—0-6 mm below tip; tendency to greater awn length and for callus
hairs to exceed the lemma only slightly. Self fertile.

Calamagrostis canescens X C. stricta

Both hybrid populations studied at Leven Canal show morphological features intermediate between
those of the parents, notably glume shape, awn length and point of awn insertion, the awn leaving
the lemma just below half-way. C. stricta features almost always occurring in the hybrids are culm
rough just below the panicle, one or more bottom leaf sheaths hairy, the axis prolonged as a tuft of
hairs and the panicle tightening after flowering. A C. canescens feature which commonly occurs in
the hybrids is the long white hairs on the upper leaf surface.

Calamagrostis canescens X C. stricta Leven H, population, 2n = 56

Occurs mainly on the sloping part of the canal bank, extending to the edge of the canal, but not
occurring on top of the bank. Perennial, rhizomatous, forming beds, very vigorous. Height: 62-147
cm. Leaves green, intermediate in width, with or without long white hairs on upper leaf surface;
bracts typically taking up a horizontal ‘flag flying’ position. The ligule is long, that of the bract being
3-6 mm and it does not tear. At least one lower leaf sheath is usually hairy (24 out of 25 specimens
examined). Culm 5-noded and branched, rough near the panicle. The panicle is intermediate in
length or C. canescens-like, 13-5—20-8 cm, intermediate to feel, tightening after flowering, but
conspicuously broader than in C. stricta. The lowest branch of the panicle is intermediate in length
or C. canescens-like, 3-8—6:6 cm. The glumes are intermediate in shape, being similar to those of C.
canescens in length, 4-9-5-9 mm, and to those of C. stricta in width or slightly wider, 1-0-1-2 mm
wide; the tips of the glumes do not break off at maturity. The glumes are somewhat rough, the spike
being fairly rough to feel and intermediate between the species in this respect. The lemma is notably
long, 3-2—3-8 mm, longer than in both parents. The callus hairs more or less equal the floret in length
or are very slightly shorter to 4 mm longer, the actual length of the majority of hairs being similar to
those of C. canescens. The awnis intermediate in length, 1-2—-1-9 mm, is like that of C. stricta in type,
and leaves the lemma from just below the middle, so that it falis short of the top of the lemma, by
0-3-0-8 mm. The spikelet axis is prolonged as a minute bristle as in C. stricta. The anthers are
intermediate in shape, the lobes at one end being more or less parallel and at the other end splaying
out; they are purple and at Leven produce 60% well-formed pollen. Flowering time is intermediate
between that of the parents, but there is some overlap; the population flowers typically in the third
week of June.

C. canescens X C. stricta Leven H, population, 2n = 28

Occurs by the edge of the Leven Canal. Perennial, rhizomatous, forming a narrow belt. Height: 70—
109 cm. Leaves green, intermediate in width, with or without long white hairs on the upper leaf
surface. The ligule is variable in length, but usually intermediate, 2-2-4-9 mm. At least one lower
leaf-sheath is usually hairy (24 out of 25 examined). Culm 4-noded and usually rough near the
panicle (24 out of 25 examined). The panicle is intermediate in length or C. canescens-like, 13-7—
19-2 cm, tightening after flowering except that the long bottom branch may protrude. The lowest
branch of the panicle is longer than in C. stricta, (3-5—)4-6-7-4 cm. The glumes are intermediate in
shape, similar to those of C. stricta in length to slightly longer, 3-5—4-2 mm, but narrower, 0-8—0-9
mm wide, so that the panicle has a delicate appearance. The lemma is similar in length (2-6—3-0 mm)
to that of the parents. The callus hairs usually more or less equal the floret in length or are slightly
shorter, but occasionally are appreciably shorter (0-9-0 mm less than the floret) as in C. stricta. The
awn is intermediate in length, 1-2-1-8 mm, and leaves the lemma just below the middle and usually
falls slightly short (0-0-4 mm) of the top of the floret. The anthers are I-shaped and yellow, rarely
slightly purple, when exserted. It is almost male sterile, c. 4% morphologically good pollen being
noted, while over 90% of the pollen grains are empty. No fruit has been observed. Flowering time is
intermediate between that of the parents.

C. canescens X C. stricta Leven H;

This is a recombinant represented by a single plant on an unusually steep stretch of the canal bank.

60 FEY CRACKEES

The awn length and position was similar to that of C. canescens, while in its panicle, ligule and
bottom branch of panicle length and short callus hairs it resembled C. stricta.

ACKNOWLEDGMENTS

The work described in this paper formed part of a thesis accepted for the degree of M.Sc. at the
University of Hull. Iam indebted to Dr D. W. Shimwell for advice and encouragement during the
early stages of the research and to Mr G. Nicholson for assistance with chromosome counts. My
thanks are also due to the late Mr P. Smith, Mr R. Wheeler-Osman and Miss A. Braithwaite for
technical assistance and to Dr Michael Proctor for reading the manuscript and making helpful
suggestions.

REFERENCES

CRACKLES, F. E. (1953). Calamagrostis stricta in S.E. Yorkshire. B.S.B.J. Year Book: p. 54.

CRACKLES, F. E. (1968). Some plant associations of the River Hull valley. East Yorkshire Field Studies 1: 13-24.

HuBBArD, C. E. (1968). Grasses. Harmondsworth.

HyLanper, N. (1953). Nordisk Karlvaxtflora 1: 315-317.

NyGreN, A. (1946). The genesis of some Scandinavian species of Calamagrostis. Hereditas 32: 131-262.

NyGren, A. (1948). Further studies in spontaneous and synthetic Calamagrostis purpurea. Hereditas 34: 113—
134.

NyGrEN, A. (1962). Artificial and natural hybridization in European Calamagrostis. Symbolae botanicae
upsalienses 17; 1-105.

SHEPPARD, J. A. (1957). The medieval meres of Holderness. /nstitute of British Geography: Transactions &
Papers 23: 75-85.

SHEPPARD, J. A. (1958). The draining of the Hull valley. 24 pp. East Yorkshire Local History Society, York.
(East Yorkshire Local History Series, 8.)

SteMSSEN, M. A. C. (1795). Arundo stricta (Steifahrichtes Rohr.). Magazin fiir die Naturkunde und Oekonomie
Mecklenburgs 2: 235. Schwerin & Leipzig.

(Accepted August 1993)

Watsonia, 20, 61-71 (1994) 61

Notes

RUBUS AGHADERGENSIS D. E. ALLEN (ROSACEAE): A NEW NAME FOR AN
IRISH BRAMBLE

In the 1890s W. Moyle Rogers began receiving from Canon H. W. Lett specimens of a white-
flowered, eglandular bramble new to him from various localities in Lett’s parish of Aghaderg, near
Loughbrickland, in County Down, v.c. H38. It was subsequently distributed through the Watson
Botanical Exchange Club, and in 1901 Rogers had the opportunity of studying it in the field himself.
In keeping with his preference for a hierarchy of taxa in Rubus, he decided it was best placed under
R. lindleianus Lees as a variety, for which he coined the manuscript epithet /atifolius (after the very
broad leaflets of the stem leaves). Later, after his death, H. J. Riddelsdell felt that its discovery in a
second vice-county, Armagh, v.c. H37 — though the locality in question was close by and only just
across the boundary — warranted publishing the name at last, which he accordingly did in their joint
names (Rogers & Riddelsdell 1925).

Although this taxon has been ignored in subsequent monographs of the group, it is manifestly no
mere local variant of R. lindleianus but a distinct entity that deserves to be regarded as a species in its
own right. That would not be sufficient justification, even so, for burdening the already very lengthy
list of British and Irish Rubus species with yet another name, were this bramble confined to just the
one small area from which it has so far been recorded. In 1991, however, D. A. Doogue collected in
Ravensdale (GR J/09:14), in the hill country of the north of Louth, v.c. H31, a specimen (now in
BM) of what is clearly this same plant. This extends the known range by over 25 km. It is, moreover,
suspicious that Praeger (1901) describes R. lindleianus as “frequent” in Armagh, for recent
fieldwork has shown that species to be rare over Ulster as a whole: much of what has been taken for
it in that vice-county may in reality be Lett’s bramble.

In view of this much wider range, extending into three vice-counties, raising the taxon to specific
rank now seems appropriate. That requires a new name, however, as the epithet /atifolius is
preoccupied at that level:

Rubus aghadergensis D. E. Allen, nom. et stat. nov.
R. lindleianus var. latifolius Rogers & Riddelsd., Journal of botany 63: 14 (1925).
LECTOTYPE: side of road, parish of Ballymore, near Scarva station, v.c. H37, 4 August 1894, H. W.
Lett as R. rhamnifolius, herb. Barton & Riddelsdell no. 6783 (BM), det. B. A. Miles 1964.

The species belongs in series Sylvatici (P. J. Mueller) Focke.

REFERENCES

PRAEGER, R. L. (1901). Irish topographical botany. Dublin.
Rocers, W. M. & RippELSDELL, H. J. (1925). Some varieties of Rubus. Journal of botany 63: 13-15.

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

THE NATIVE BRITISH COTONEASTER —- GREAT ORME BERRY —- RENAMED

The only Cotoneaster species (Rosaceae) native to the British Isles grows on the Great Orme’s
Head, Llandudno, North Wales. It was discovered in 1783 by John Wynne Griffith and was
subsequently rediscovered in 1821 by William Wilson of Warrington. The number of shrubs was
stated as being plentiful, but has now been reduced to only four individuals (Smith 1979). Over-
collecting and hard grazing by sheep and goats are thought to have been the main causes of its

62 NOTES

decline. By fencing and propagation of plants for re-introduction to the limestone ledges it is hoped
to save this taxon from further decline (Morris 1978).

The North Wales shrub has been included within Cotoneaster integerrimus Med., but when
Hrabétova-Uhrova (1962) studied members of the genus Cotoneaster in former Czechoslovakia, she
observed that the Great Orme taxon differed from those of Central Europe and therefore made it a
new variety of Cotoneaster integerrimus, namely var. anglicus Hrabétova.

Apomixis is common in the genus Cotoneaster, most species having been found to be tetraploids
(2n = 68) and coming true from seed. By tradition, apomictic taxa within the genus are given the
rank of species.

In France three species related to C.integerrimus have long been separated: C. intermedius Coste,
C. juranus Gandoger and C. obtusisepalus Gandoger. Two new species have also recently been
described in Scandinavia: C. scandinavicus Hylm6 and C. canescens Hylm6. The former is closely
related to C. cinnabarinus Pojarkova from the Kola Peninsula, Russia.

By courtesy of Mr Maurice Morris, Penmachno, Gwynedd, N. Wales and Dr Nigel Brown,
Treborth Botanic Garden, University of North Wales, we have had access to seed and plants of the
Great Orme taxon making it possible to study this Cotoneaster in cultivation both in Britain and
Sweden. We have concluded that the N. Wales plant clearly differs from the taxa of Central Europe
and feel it is time to raise it to the rank of species.

Cotoneaster cambricus J. Fryer & B. Hylm6, nom. et stat. nov.
(section Cotoneaster, series Cotoneaster)
Synonym: C. integerrimus Medicus var. anglicus Hrabétova, Acta Acad. Scient. Czechoslov. Basis
Brunensis 34 (6): 217, tab. 4a (1962).
HoLotypus: Llandudno, Caernarvonshire, c. 1836, L. Price no. H.899(60)4 (K).
A low deciduous shrub. Young branches greenish-brown, villous. Leaves on young growth broadly
elliptic to suborbicular, apex obtuse or acute; upper surface grey, moderately pilose, hairs persisting
or becoming subglabrous, veins slightly depressed. Inflorescence 1—2(—3) flowered, pedicels and
peduncles short 2—3(—5) mm; petals equal to or less than 1 mm longer than the calyx. Fruit small 5-8
mm, globose, clear red with orange tones; pyrenes 2-3.

Chromosome count 2n = 68, courtesy of Dr Hugh McAllister, Ness Botanic Gardens, University
of Liverpool.

DIAGNOSTIC KEY TO THE MOST CLOSELY RELATED SPECIES -

la Hypanthium and calyx pilose; upper surface of leaves persistently pilose; flowering shoots 25—45

mm with 4 leaves and 3-7 flowers; pyrenes 3—4(-5).

France: Massif Central and the Alps (Haute Savoie) ...:.................055 C. intermedius Coste
1b Hypanthium and calyx glabrous :........ 2.0.6. ..2.02 ce sise cee ce ce ects ooo le eee ey
2a Upper surface of leaves glabrous from the beginning, leaves on sterile shoots ovate, apex acute,

surface smooth with veins only slightly depressed; flowering shoots 15—25(—35) mm with 2-4

leaves and 1—3(-4) flowers; pyrenes (2—)3—-4(—5); shrub 0-1—1-0 m, arching or decumbent with

weak stems.

France: Vosges and Jura, and throughout the Alps (above zone of C. integerrimus) ............

C. juranus Gandoger

2b. Upper surface of leaves pilose when young ..........:2....0..202022.-00-0s050 = ses nee eee 3
3a Leaves on sterile shoots broadly ovate, apex acute or acuminate; upper surface pilose becoming
glabrous, rugose with depressed veins; flowering shoots 15—25(—35) mm usually with 2-3 leaves

and (1—)3—4(—7) flowers; pyrenes 2—3(—4); shrub arching, 1-0—-1-5 m.

Central Europe and the Alps (zone below C. juranus) .................. C. integerrimus Medicus
3b Leaves on sterile shoots elliptic or suborbicular, apex obtuse or acute .........................5. -
4a Leaves broadly elliptic or suborbicular, upper surface densely pilose becoming subglabrous or ©

glabrous; petals equal to or less than 1 mm longer than calyx lobes, off-white; flowering shoots —

10-20 mm with c. 2 leaves and 1—2(3) flowers; pyrenes 2—3; shrub + erect c. 1-0(—1-5) m.

Prndemic tos Wales ite tk. 20es) oes a, ee eee C. cambricus J. Fryer & B. Hylmo
4b Leaves elliptic, upper surface with long pilose hairs becoming subglabrous with only single long

NOTES 63

hairs or entirely glabrous; petals more than 1 mm longer than the calyx lobes, off-white with
reddish stripes; flowering shoots 15—25 mm with usually 2—3 leaves and 3-5 flowers; pyrenes 3-4;
shrub arching 1-1—1-5 m.

PemeinGe eNOS) (ISENG ila fyn ceeties + S8ck ob Ah RITE. ou fk vce Ds oH 3G. hiss} d. C. obtusisepalus Gandoger

ACKNOWLEDGMENTS

We are grateful to Dr David Allen for his assistance with historical research, and to David
McClintock for his insistence that we publish this work.

REFERENCES

HrasetovA-Unrova, A. (1962). Beitrag zur Taxonomie und Verbreitung der Gattung Cotoneaster in der
Tschekoslowakei. Prace Brnénské Zakladny Ceskoslovenské akademie védecké Brno 34: 197-246.

Morris, M. (1978). Cotoneaster integerrimus — a conservation exercise. B.S.B.I. Welsh Bulletin 28: 6-8.

Smitu, M. E. (1979). And then there were four. The wild Cotoneaster in Britain. Country Life, 13 September
1979, p. 788.

J. FRYER

Trooper Bottom, Froxfield, Petersfield, Hampshire, GU32 1BH
B. HyLMO

Bygatan 30, S—267 40 Bjuv, Sweden

CONTRIBUTIONS TO A CYTOLOGICAL CATALOGUE OF THE BRITISH AND
IRISH FLORA, 4

We present here chromosome counts for 121 populations of 108 vascular plant species from
documented localities in Britain. This note continues a series, the previous one of which was Al-
Bermani et al. (1993). One plant from each population was studied, except where noted. All counts
were made on squashes of root-tips, except in the case of Ophioglossum vulgatum where meiosis in
microspore mother-cells was examined. Supernumerary chromosomes are designated by the suffix
‘S’. Voucher specimens for most counts have been deposited in LTR, the remainder are at K or CGE
as indicated.

Althaea officinalis L., 2n = 42: E. Suffolk, v.c. 25, S.E. side of Benacre Broad, near Covehithe, TM/
5.8 (K).

Anagallis arvensis L. subsp. arvensis, 2n = 40: Caerns., v.c. 49, Botwnnog, SH/26.32.

Anagallis tenella (L.) L., 2n = 22: W. Lancs., v.c. 60, near Lancaster, SD/491.603; Westmorland,
v.c. 69, Lowick Common, SD/292.847 (no voucher).

Anthyllis vulneraria L. subsp. vulneraria, 2n = 12: Dorset, v.c. 9, Melbury Hill, ST/87.19.

Apium nodiflorum (L.) Lagasca, 2n = 22: Oxon, v.c. 23, Little Bourton, farm near M40 motorway,
SP/4.4.

Aquilegia vulgaris L., 2n = 14: W. Lancs., v.c. 60, Silverdale, Gait Barrows N.N.R., SD/479.774.

Artemisia absinthium L., 2n = 18: Caerns., v.c. 49, Abersoch, N. end of Porth Fawr, SH/31.27.

Aster tripolium L., 2n = 18: W. Sussex, v.c. 13, Pagham Harbour nature reserve, near Sidlesham,
SZ/8.9 (K).

Avena fatua L., 2n = 42: N. Wilts., v.c. 7, near Calne by the A4, ST/9.7 (K).

Beta vulgaris L. subsp. maritima (L.) Arcang., 2n = 18: N. E. Yorks., v.c. 62, Scarborough, TA/0.8.

64 NOTES

Brassica nigra (L.) Koch, 2n = 16: Dorset, v.c. 9, St Alban’s Head, 0-4km E.N.E. of lookout, SY/
963.756 (K); Dorset, v.c. 9, Weymouth, landward shore of Fleet Lagoon, SY/66.76; Cambs., v.c.
29, Bar Hill, TL/38.63.

Bromopsis ramosa (Hudson) Holub, 2n = 42: W. Lancs., v.c. 60, Silverdale, Gait Barrows N.N.R.,
near Hawes Water, SD/47.76 (no voucher).

Bromus interruptus (Hackel) Druce, 2n = 28: Cambs., v.c. 29, Pampisford, TL/4.4 (CGE).

Bryonia dioica Jacq., 2n = 20: E. Sussex, v.c. 14, Beachy Head, c. 1-8 km N. of Hodcombe Farm,
TV/576.968.

Buxus sempervirens L., 2n = 28: Surrey, v.c. 17, Silent Pool, near Shere, TQ/06.48.

Chenopodium vulvaria L., 2n = 18: Dorset, v.c. 9, c. 1-5 km W. of Burton Bradstock, SY/47.89.

Cirsium heterophyllum (L.) Hill, 2n = 34: Main Argyll, v.c. 98, Black Mount, off road between
Ballachulish and Tyndrum, NN/2.4 (no voucher).

Cirsium palustre (L.) Scop., 2n = 34: Caerns., v.c. 49, Garn Fadryn, SH/27.35; W. Lancs., v.c. 60,
Lower Salter, near Barkin Bridge, SD/601.636 (no voucher).

Cirsium vulgare (Savi) Ten., 2n = 68: W. Lancs., v.c. 60, Lancaster, near River Lune, SD/484.636.

Conopodium majus (Gouan) Loret, 2n = 22: W. Sutherland, v.c. 108, Nedd, NC/137.319 (no
voucher).

Crepis paludosa (L.) Moench, 2n = 12: W. Lancs., v.c. 60, near Millbeck footbridge, SD/648.638
(no voucher).

Daucus carota L. subsp. carota, 2n = 18: W. Lancs., v.c. 60, Carnforth, SD/499.711.

Dipsacus fullonum L., 2n = 18: W. Lancs., v.c. 60, near Lancaster, SD/467.624.

Drosera rotundifolia L., 2n = 20: S. Devon, v.c. 3, Dartmoor, c.2kmN.E. of Postbridge, SX/66.80.

Echium vulgare L., 2n = 32: E. Suffolk, v.c. 25, Felixstowe, Landguard Nature Reserve, TM/2.3
(K). |

Elymus caninus (L.) L., 2n = 28: Westmorland, v.c. 69, Brigsteer Park, 2:5 km S. of Brigsteer, SD/
48.87.

Elytrigia repens (L.) Desv. ex Nevski subsp. repens, 2n = 42: Oxon, v.c. 23, Little Bourton, farm
near M40 motorway, SP/4.4.

Erica tetralix L., 2n = 24: W. Sutherland, v.c. 108, near Allt na Claise, NC/174.327.

Erodium maritimum (L.) L’Hér., 2n = 20: Dorset, v.c. 9, near Moreton, SY/80.89.

Euonymus europaeus L., 2n = 32: E. Suffolk, v.c. 25, Great Wenham, TM/077.383.

Euphorbia lathyris L., 2n = 20: W. Lancs., v.c. 60, White Land Industrial Estate, near Lancaster,
SD/453.625. |

Euphorbia portlandica L., 2n = 16: Dorset, v.c. 9, Isle of Portland, Church Ope Cove, SY/69.70.

Gagea lutea (L.) Ker Gawler, 2n = 72: Westmorland, v.c. 69, Sedgwick, E. bank of R. Kent, SD/
510.878.

Geranium lucidum L., 2n = 40: Derbys., v.c. 57, Dovedale, SK/14.53 (three plants counted).

Glaucium flavum Crantz, 2n = 12: W. Lancs., v.c. 60, R. Keer estuary, S.W. of Carnforth, SD/ _
482.699. |

Gnaphalium luteo-album L., 2n = 14: Channel Is., v.c. S, Alderney, Platte Saline, WA/567.078. :

Heracleum sphondylium L. subsp. sphondylium, 2n = 22+1S: Derbys., v.c. 57, Milldale, SK/14.55.

Hippophae rhamnoides L., 2n = 24: N. Lincs., v.c. 54, Gibraltar Point, TF/56.57; W. Lancs., v.c.
60, near Heysham harbour, SD/407.604. |

Honckenya peploides (L.) Ehrh., 2n = 68: W. Lancs., v.c. 60, W. of Cockerham, near Bank _
Houses, SD/431.531. |

Hordeum marinum Hudson, 2n = 14: W. Kent, v.c. 16, Higham Marshes, near Gravesend, TO/ _
69.74. :

Hordeum murinum L. subsp. murinum, 2n = 28: Caerns., v.c. 49, Abersoch, dunes at N. end Porth
Fawr, SH/314.277; W. Lancs., v.c. 60, W. of Cockerham, near Bank Houses, SD/432.531.

Humulus lupulus L., 2n = 20: Berks., v.c. 22, Cothill Fen, SU/46.99.

Hyacinthoides non-scripta (L.) Chouard ex Rothm., 2n = 16: W. Lancs., v.c. 60, Lancaster
University grounds, SD/48.57.

Iberis amara L., 2n = 14: Surrey, v.c. 17, Box Hill, Juniper Hall nature reserve, TQ/176.528.

Lathyrus aphaca L., 2n = 14: Dorset, v.c. 9, Weymouth, landward shore of Fleet Lagoon, SY/
66.76.

Leucanthemum vulgare Lam., 2n = 18: Channel Is., v.c. S, Alderney, Essex Hill, WA/592.074.

NOTES 65

Limonium britannicum Ingrouille subsp. celticum Ingrouille var. celticum, 2n = 36: Cheshire, v.c.
58, Hilbre Island, SJ/185.878; var. pharense Ingrouille, 2n = 36: Caerns., v.c. 49, Bardsey Island,
E. of Pen Cristin, Ogof y Gaseg, SH/12.21 (150 m N.E. of site given in Hollingsworth er al.

1992)).

ante ae Miller, 2n = 12: W. Lancs., v.c. 60, near Carnforth, SD/493.714.

Lobelia dortmanna L., 2n = 14: W. Sutherland, v.c. 108, loch c. 1 km N.E. of Duartmore Bridge,
NC/211.383 (no voucher).

Lonicera periclymenum L., 2n = 54: Leics., v.c. 55, Swithland Wood, SK/5.1.

Lotus corniculatus L., 2n = 24: W. Lancs., v.c. 60, Carnforth, SD/499.711.

Luzula sylvatica (Hudson) Gaudin, 2n = 12: W. Lancs., v.c. 60, near Millbeck footbridge, SD/
648.638; Westerness, v.c. 97, Glen Nevis, along path to Stob Ban, NN/15.66.

Malva moschata L., 2n = 42: E. Sussex, v.c. 14, Ardingly, Wakehurst Place, Bloomers Valley, TQ/
3-3°(K).

Matthiola incana (L.) R. Br., 2n = 14: E. Sussex, v.c. 14, Peacehaven cliffs to Castle Hill, TQ/4.0
(K).

Medicago lupulina L., 2n = 16: Westmorland, v.c. 69, Whitbarrow, Ravens Lodge, underneath
Whitescar, SD/46.85.

Melica nutans L., 2n = 18: W. Lancs., v.c. 60, Silverdale, Gait Barrows N.N.R., SD/481.776.

Narthecium ossifragum (L.) Hudson, 2n = 26: Westmorland, v.c. 69, Lowick Common, SD/292.847
(no voucher).

Onopordum acanthium L., 2n = 34: Dorset, v.c. 9, near Bere Regis, SY/84.94.

Ophioglossum vulgatum L.,n = c. 270: Leics., v.c. 55, Leicester, Oadby, Botanic Garden meadow,
SK/61.01.

Ornithopus perpusillus L., 2n = 14: Caerns., v.c. 49, Boduan, opposite entry to Mathan Uchaf
farm, SH/319.369.

Parapholis strigosa (Dumort.) C. E. Hubb., 2n = 28: W. Kent, v.c. 16, Higham marshes, near
Gravesend, TQ/69.74.

Persicaria lapathifolia (L.) Gray, 2n = 22: W. Lancs., v.c. 60, near Heysham harbour, SD/404.599.

Phalaris arundinacea L., 2n = 28: W. Lancs., v.c. 60, River Lune, near Lancaster, SD/483.633.

Phleum phleoides (L.) Karsten, 2n = 14: W. Norfolk, v.c. 28, Brettenham, near Thetford, S. of
A1066, TL/907.844 (K).

Polypogon monspeliensis (L.) Desf., 2n = 28: S. Hants., v.c. 11, Southampton Water, Hythe,
SU/4.0.

Potentilla argentea L., 2n = 14: Dorset, v.c. 9, 1-6 km N. of Wool, SY/84.87.

Potentilla palustris (L.) Scop., 2n = 64: Warks., v.c. 38, Sutton Coldfield, Sutton Park, SP/10.97.

Primula vulgaris Hudson, 2n = 22: Westmorland, v.c. 69, Brigsteer Park, 2:5 km S. of Brigsteer,
SD/48.87.

Pulicaria dysenterica (L.) Bernh., 2n = 18: W. Lancs., v.c. 60, near Lancaster, SD/465.624.

Ranunculus flammula L. subsp. flammula, 2n = 32: S. Lancs., v.c. 59, Ainsdale dunes, SD/31.11.

Ranunculus sceleratus L.,2n = 32: Oxon, v.c. 23, Little Bourton, farm near M40 motorway, SP/4.4.

Rumex crispus L., 2n = 60: E. Sussex, v.c. 14, Cuckmere Haven, E. side of river, TV/5.9 (K).

Saponaria officinalis L., 2n = 28: W. Lancs., v.c. 60, River Lune, near Lancaster, SD/483.634.

Scilla autumnalis L., 2n = 28: Channel Is., v.c. S, Alderney, Essex Hill, WA/592.074.

Scutellaria galericulata L., 2n = 30: Salop, v.c. 40, Crose Mere, S. end, SJ/433.303.

Seriphidium maritimum (L.) Polj., 2n = 54: W. Lancs., v.c. 60, W. of Cockerham, near Bank
Houses, SD/428.532.

Serratula tinctoria L., 2n = 22: Dorset, v.c. 9, Melbury Hill, ST/87.19.

Silene noctiflora L., 2n = 24: Dorset, v.c. 9,2 km S. of Winterborne Kingston, SY/85.96.

Sison amomum L., 2n = 14: Middlesex, v.c. 21, Monken Hadley Common, S. edge, 0-25 km E. to
bridge over railway, TQ/2.9 (K).

Sonchus arvensis L., 2n = 54: Warks., v.c. 38, Ingon Grange Farm, near Snitterfield, SP/214.590.

Stachys officinalis (L.) Trev., 2n = 16: Dorset, v.c. 9, Melbury Hill, ST/87.19.

Stellaria holostea L., 2n = 26: W. Lancs., v.c. 60, near Yealand Conyers, SD/509.746.

Suaeda vera Forssk. ex J. Gmelin, 2n = 36: E. Sussex, v.c. 14, Cuckmere Haven, E. side of river,
TV/5.9 (K).

Succisa pratensis Moench, 2n = 20: W. Lancs., v.c. 60, outskirts of Lancaster, SD/491.603.

66 NOTES

Tanacetum parthenium (L.) Schultz-Bip., 2n = 18: W. Lancs., v.c. 60, River Lune estuary, near
Sunderland Point, SD/426.556.
Teucrium botrys L., 2n = 32: Surrey, v.c. 17, Box Hill, Juniper Hall nature reserve, TQ/176.528.
Thlaspi arvense L., 2n = 14: W. Lancs., v.c. 60, near Lancaster, near River Lune, SD/486.637.
Torilis nodosa (L.) Gaertner, 2n = 24: Dorset, v.c. 9, c. 1:5 km W. of Burton Bradstock, SY/47.89;
Caerns., v.c. 49, Aberdaron, 2 km E. of Aberdaen, Penrhyn-mawr Farm, SH/190.261.
Tragopogon pratensis L. subsp. minor (Miller) Wahlenb., 2n = 12: Dorset, v.c. 9, Melbury Hill, ST/
87.19.
Trientalis europaea L., 2n = c. 160: Moray, v.c. 95, Grantown-on-Spey, NJ/02.26.
Trifolium campestre Schreber, 2n = 14: W. Suffolk, v.c. 26, Lakenheath, Maidscross Hill, TL/72.82 —
(K).

Trifolium fragiferum L. subsp. fragiferum, 2n = 16: Dorset, v.c. 9, Small Mouth, ST/66.76.

Trifolium glomeratum L., 2n = 16: E. Suffolk, v.c. 25, Aldeburgh, North Warren Nature Reserve,
TM/4.5 (K).

Trifolium incarnatum L., 2n = 14: Dorset, v.c. 9, Zelstan, SY/89.98 (no voucher); subsp. molinerii,
2n = 14: W. Cornwall, v.c. 1, The Lizard, N. of Caerthillian Cove, SW/69.12.

Trifolium ochroleucon Hudson, 2n = 16: Cambs., v.c. 29, Orwell Hill, TL/36.50 (K).

Trifolium scabrum L., 2n = 10: W. Cornwall, v.c. 1, Marazion, SW/S0.31.

Trifolium squamosum L., 2n = 16: N. Essex, v.c. 19, Little Oakley, near Harwich, TM/225.278 (K).

Trifolium striatum L., 2n = 14: W. Suffolk, v.c. 26, Icklingham, Kings Forest, TL/7.7 (K).

Urtica urens L., 2n = 24: Oxon, v.c. 23, Little Bourton, farm near M40 motorway, SP/4.4.

Verbascum virgatum Stokes, 2n = c. 64: E. Cornwall, v.c. 2, Saltash, SX/4.5 (K).

Veronica chamaedrys L., 2n = 32: Dorset, v.c. 9, Wareham, Washer’s Pit, SY/86.94; W. Lancs.,
v.c. 60, Silverdale, Gait Barrows N.N.R., SD/479.776.

Veronica hederifolia L. subsp. hederifolia, 2n = 54: E. Gloucs., v.c. 33, Ashton under Hill,
SO/99.37.

Veronica polita Fries, 2n.= 14: Dorset, v.c. 9, Upton Park, SY/99.92.

Veronica serpyllifolia L. subsp. serpyllifolia, 2n = 14: N. Wilts., v.c. 7, Lockeridge, S.W. of
Marlborough, SU/150.675; Caerns., v.c. 49, Crugan, Llanbedrog, SH/33.32; W. Lancs., v.c. 60,
near Stirk Close, SD/634.650.

Vicia hirsuta (L.) Gray, 2n = 14: Dorset, v.c. 9, Weymouth, landward shore of Fleet Lagoon, SY/
66.76; Salop, v.c. 40, Sunny Hill, W. of N.-S. minor road from Brockton to Clunton, $O/3.8 (K).

Vicia sylvatica L., 2n = 14: Salop, v.c. 40, Bucknell, on N. E. edge of Tueshill Wood, SO/3.7 (K)
(two plants counted).

Vicia tetrasperma (L.) Schreber, 2n = 14: E. Sussex, v.c. 14, Ardingly Reservoir, Hanging Meadow,
TQ/3.3 (K).

ACKNOWLEDGMENTS

We should like to thank English Nature and the R.S.P.B. for permission to collect from their
reserves, and J. Terry (Seed Bank Officer at the Royal Botanic Gardens, Wakehurst), B. Bonnard,
H. J. M. Bowen, P. Evans and J. Partridge for sending us plant material.

REFERENCES

AL-BERMANI, A.-K. K. A., AL-SHAMMARY, K. I. A., BAILEY, J. P. & GoRNALL, R. J. (1993). Contributions to a
cytological catalogue of the British and Irish flora, 3. Watsonia 19: 269-271.

HOoLLincswortH, P. M., GornaLL, R. J. & BaiLey, J. P. (1992). Contributions to a cytological catalogue of the |
British and Irish flora, 2. Watsonia 19: 134-137.

R. E. Dempsey, R. J. GORNALL & J. P. BAILEY
Botany Department, University of Leicester, Leicester, LE] 7RH

NOTES 67

VARIATION AND HYBRIDISATION OF ALNUS MILLER IN IRELAND

The Common Alder, Alnus glutinosa (L.) Gaertner (Betulaceae) is known to be morphologically
variable. McVean (1953a, b) has shown that weak clinal variation in leaf and catkin dimensions
exists along a gradient running from S. W. England to N. W. Scotland.

During 1987 and 1988, samples were taken from 17 populations from as wide a range of
geographical localities in Ireland as possible. Most populations were in semi-natural habitats such as
riverside banks and carr communities, and none showed evidence of planting. At each site, from a
normal minimum of ten individual trees, I made five measurements from ten mature leaves and
three measurements from ten mature female catkins (if present), plus an assessment of leaf apex
shape and number of teeth on leaf margin, number of pairs of secondary veins and number of glands
on the leaf surface. Two ratios were calculated from these data. The following characters were
_ recorded: 1. Lamina length (measured from the top of the petiole along the main vein to the leaf
tip); 2. Maximum lamina width; 3. Lamina length to widest point; 4. Number of pairs of secondary
veins; 5. Angle of divergence of the mid-secondary veins from the main vein; 6. Angle of divergence
of the leaf-apex; 7. Angle of divergence of the leaf-base; 8. Size of the tufts of hairs in the axils of the
secondary veins (arbitary scale 0-5); 9. Length of the petiole; 10. Apex acute, subacute or rounded;
11. Number of teeth on one side of the lamina; 12. Ratio of lamina length/lamina length from base to
widest point; 13. Ratio of lamina width/(ratio of lamina length/lamina length from base to widest
point); 14. Number of glands present on a randomly chosen area of 2 mm? on the abaxial leaf
surface; 15. Mature female catkin length (following anthesis); 16. Mature female catkin width; and
17. Length of stalk of female catkin.

McVean (1953a, b) found weak clinal changes of size in leaf length and width and female catkin
length — effectively he showed that leaf and catkin dimensions were correlated, though the largest-
leaved plants did not necessarily bear the largest catkins. Unfortunately I had considerable difficulty
in sampling a sufficient number of mature female cones and was unable to confirm this finding.
However, in Fig. 1 I have plotted data for lamina length and width for Irish populations alongside
re-calculated values derived from McVean (1953a, b), all with 95% confidence limits and arranged
along a north-east/south-west axis. In Ireland there is great variation in leaf size, but it is not clinal,
nor is there any difference in leaf size between alders from different regions of Ireland. This
contrasts with Britain (McVean 1953a, b)

The differences in the variation pattern of Common Alder in Britain and Ireland may be due toa
number of causes. The first and most obvious potential cause is that the geographical and climatic
span of Ireland is much less than that found in Britain and selection pressures may be insufficient to
generate a cline. Secondly it could be that Common Alder populations in Britain are more
genetically isolated from each other than they are in Ireland. Highly isolated small populations are
likely to experience genetic drift (Levin 1981) and it is possible for clinal variation to develop under
these circumstances even in the absence of selection. Most flowering plant populations in which
clines have been found are in fact highly genetically isolated (Nic Lughadha & Parnell 1989). Alnus
pollen, however, is frequently transported over long distances (Huntley & Birks 1983), so the real
potential for genetic drift in Alnus is likely to be small. There is no evidence that there is any
difference in the degree of genetic isolation between populations of Alders in Britain and Ireland.
Thirdly it could be that the origins of A. glutinosa in Britain and Ireland are different, and that most
present-day Irish populations were derived from a few founder individuals or populations. This is
unlikely given the speed of the spread of A. glutinosa into Ireland following the last glaciation
(Huntley & Birks 1983). Fourthly it is possible that A. glutinosa is not native at all in Ireland but was
derived from a few original planted introductions. Current evidence suggests that, following the
most recent glaciation, wild Alnus reached and rapidly colonised Ireland some 1500 years later than
it did in Britain, between 6500 and 7000 B.P. (Huntley & Birks 1983), and there is no evidence that
it subsequently became extinct. So I conclude that the difference in the variation pattern in
Common Alder between Ireland and Britain is due to the relatively lesser geographical span of
Ireland.

Of the populations sampled, one (population 15) from Charlestown House, near Jamestown (Co.
Leitrim) in Co. Roscommon (v.c. H25) (see Kelly 1985) contained trees whose leaves were
unusually variable in outline and appeared intermediate between A. glutinosa and A. incana (L.)
Moench (Grey Alder). A. incana differs from A. glutinosa in its subacute to acuminate leaves

68 NOTES

3

NW se nw se

FicureE |. Means for leaf length (a,b) and breadth (c,d) (cm), with 95% confidence limits, for populations of A.
glutinosa recorded in Ireland (b,d) and Britain (a,c). The data from the populations are arranged to show a
gradient from north-west to south-east. Data for plants from Britain are plotted from revised data of McVean
(1953a, b). The code numbers identify the different populations.

Leaf length
to widest
point

(cms)

Degree of
acuteness

of
leaf apex

(score 0-2)

Number

of glands

2
per 1mm

6 Number
5 of pairs
b } of veins
3 | per leaf
2

123

O

Angle
of basal
leaf sinus

(degrees)

2:0 Number

1S H of teeth

1-0 per side

0:5 of leaf
0

123

K

NOTES

8 Maximum
6 leaf length
4 (cms)
2| f
0

A
| Angle of
10 fs primary veins
8 to midrib
6 ) (degrees)
L

E

130 Size of

120 tufts of

110 hairs in

100 veinaxils

90 (scale 0-5)

H
50 Ratio of
40 hi maximum
30 leaf length /
20 I leaf length to
10 widest point

123

L

Hl

123

44
42
40
38
36

B

F

ho Ww &

123

2-0

ae

123

Maximum
leaf breadth

(cms)

Angle of
leaf-tip

(degrees)

Petiole
length

(cms)

Ratio of leaf
breadth/
ratio of
maximum

leaf length/
leaf length to

widest point

ah

123

FiGurE 2. Character means with 95% confidence limits for all plants in population 15 (1 = A. incana,2 = A. X
pubescens, 3 = A. glutinosa).

(truncate to emarginate in A. glutinosa), the greater number of vein pairs in its leaves (10-15 versus
4-7) and in its sessile as opposed to pedunculate female catkins. The hybrid between these two
species, A. X pubescens Tausch, has only been recorded twice in Ireland (v.cc. H14, Laois & H25,
Co. Roscommon) under natural conditions (Scannell & Synnott 1988). A. x pubescens seems to be
recognisable only, with difficulty, on the basis of intermediate morphology between its parents. Fig.
2 shows the means of the measurements made on plants in population 15. For the majority of
characters, putative hybrids overlap with one or other or both of the parents, and they appear

70 NOTES

morphologically closer to A. incana (Fig. 2C,D,H,I,K,L,M,N). However, putative hybrids are
intermediate between the parental species in some characters (Fig. 2B). In other characters the
hybrid is clearly different from, and not intermediate between, its parents; for example, it has a
much higher gland density on its leaves than either of the parents (Fig. 2A). Given the difficulty in
recognising the hybrid, this previously unnoticed characteristic may well prove useful in its
identification.

ACKNOWLEDGMENTS

I thank Professor D. H. S. Richardson and Drs Q. C. B. Cronk and D. L. Kelly for helpful
comments on a manuscript draft of this paper, and to D. L. Kelly for drawing my attention to the
Alder population containing A. xX pubescens.

REFERENCES

Hunt ey, B. & Birks, H. J. B. (1983). An atlas of past and present pollen maps for Europe: 0-13,000 years ago.
Cambridge.

KELLy, D. L. (1985). Plant records from about Ireland 1965-1983. Irish Naturalists’ Journal 21: 416.

Levin, D. A. (1981). Dispersal versus gene flow in plants. Annals of the Missouri Botanical Garden 68: 233-253.

McVEANn, O. N. (1953a). Regional variation of Alnus glutinosa (L.) Gaertn. in Britain. Watsonia 3: 26-32.

McVEAn, O.N. (1953b). Biological flora of the British Isles. Alnus glutinosa (L.) Gaertn. Journal of Ecology.
41: 447-466.

Nic LucHapua, E. & PARNELL, J. (1989). Heterostyly and gene flow in Menyanthes trifoliata L. Botanical
Journal of the Linnean Society 100: 337-354. |

SCANNELL, M. J. P. & SyNnortt, D. M. (1988). Census catalogue of the flora of Ireland. Dublin.

J. PARNELL
School of Botany, Trinity College Dublin, Dublin 2, Republic of Ireland

PEDICULARIS SYLVATICA L. SUBSP. HIBERNICA D. A. WEBB
(SCROPHULARIACEAE) NEW TO WALES

- During a botanical excursion to the Gower Peninsula, Glamorgan (v.c. 41) in May 1993, Pedicularis
sylvatica subsp. hibernica was found in three sites, new to Wales. Subsequent investigation revealed
that plants with hairy calyces occur elsewhere in Wales, and it has probably been overlooked.

Pedicularis sylvatica subsp. hibernica was described by Webb (1956), and differs from subsp.
sylvatica in having the calyx and pedicels clothed in rather long white curled hairs (glabrous in subsp.
sylvatica). In Ireland it is frequent on the blanket bogs of the west coast, and has been occasionally
recorded elsewhere, mainly near the coast. In Britain it has been reported only from Kintyre and the
Outer Hebrides (Perring & Sell 1968). Slightly hairy plants (transitional to subsp. sylvatica) are not
common in Ireland, and they seem to occur chiefly near the eastern limit of subsp. hibernica (Webb
1956). Plants of subsp. hibernica on the Gower Peninsula were similar to those examined in western
Ireland.

The records seen are listed below; it is probable that the plant is widespread elsewhere in Wales,
and possibly in south-west and north-west England. I would welcome any other British records.

1. Overton Cliff, Gower, Glamorgan, v.c. 41 (SS/454.849). Two plants in coastal heath, with
subsp. sylvatica, May 1993, T. C. G. Rich et al.

2. Broad Pool, Gower, Glamorgan, v.c. 41 (SS/51.91), abundant on Molinia caerulea bog to
north-east of Broad Pool (one plant of subsp. sy/vatica was also present), especially common in
areas burnt two years previously, May 1993, T. C. G. Rich et al. (NMW).

3. Fairwood Common, Gower, Glamorgan, v.c. 41 (SS/568.922), occasional on Molinia caerulea
bog north of airport, May 1993, T. C. G. Rich et al.

4. Portmead Common, Gower, Glamorgan, v.c. 41 (SS/627.967), May 1993, Q. O. N. Kay. All
ten plants examined had hairy calyces.

——ooreee

NOTES 71

. Clyne Common, Gower, Glamorgan, v.c. 41 (SS/600.900), May 1965, Q. O. N. Kay.

. Near Caerphilly, Glamorgan, v.c. 41 (ST/1.8, tetrad B), 31 May 1920, A. W. Wade (NMW).
. Ponypriddy area, Glamorgan, v.c. 41 (ST/0.8 or ST/0.9), 16 May 1938, A. Jones (NMW).

. Glyn Neath, Glamorgan, v.c. 41 (SN/8.0, tetrad B), peaty pasture, 23 July 1941, H. A. Hyde

(NMW).

9. Between Beddau and Llantwit Fadre, Glamorgan, v.c. 41 (ST/068.848), damp ground near the
railway, 24 September 1967, A. Pinkard (NMW).
10. Ystradowen Moors, near Cowbridge, Glamorgan, v.c. 41 (ST/02.78), grass/sedge community
on damp peat, 21 May 1968, S. G. Harrison (NMW).
11. Nanthenfoel, Pont Creuddyn, Cardigan, v.c. 46 (SN/542.521), heathy pasture, scattered over
several hectares with subsp. sylvatica, 9 June 1993, A. O. Chater.

Whether this taxon merits subspecific rank is another matter, and further work on the genetics of
the expression of hairiness is required. It almost always grows with or near subsp. sylvatica, and
differs in no other character. Webb (1956) points out that a ‘“‘difference of this kind, confined to
apparently a single character, would scarcely be worth taxonomic recognition (any more than say
albinism, which is not uncommon in this species), were it not for the fact that the hairy forms occur
only in a well-defined geographical area”’.

CONAN

ACKNOWLEDGMENTS

I am grateful to Eimear Nic Lughadha, Paul A. Smith, Rebecca Warren, and Wyn, Cennydd,
Daniel Owen and Hywel Richards for their assistance in the field, and Arthur Chater, Gwynn Ellis
and Quentin Kay for assistance with other records.

REFERENCES

Wess, D. A. (1956). A new subspecies of Pedicularis sylvatica L. Watsonia 3: 239-241.
PERRING, F. H. & SELL, P. D., eds. (1968). Critical supplement to the Atlas of the British flora. London.

Ae C2Ge RICH
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Watsonia, 20, 73-83 (1994) 73

Book Reviews

The concise Oxford dictionary of botany. Edited by M. Allaby. Pp. vi + 442. Oxford University
Press, Oxford & New York. 1992. Price £18.95 (ISBN 019-866163-0 hardback); £6.99 (ISBN 019-
286094-1 paperback).

Dictionaries are keys to information; they cannot replace monographs and detailed subject text-
books, and no single-volume dictionary can ever be a complete reference work on all matters within
a subject. On the whole Allaby’s handy, pocketable dictionary succeeds in being up-to-date and, as
far as I could tell, accurate within reasonable parameters. Certainly this is likely to be a valuable
ready-reference for undergraduates and postgraduates of all ranks, serving as a source of modern
definitions and concise explanations. I do not think it is designed, as claimed, for the non-specialist
unless the person is already well-versed in the biological sciences.

For example, if you look up ““Microbiota’’, the entry leads you to “‘the smallest soil organisms,
comprising “bacteria, *fungi, “algae, and *protozoa’’, which will be a trifle confusing if you needed
to find out what the plant named Microbiota is! You can discover what the Fagaceae are, but not the
Fabaceae unless you know this is the alternative name for Leguminosae. From obscure (to this
reviewer) genera of gram-negative bacteria to world vegetation zones, by way of killing frost and
island hopping, from micro-consumers to BP — “‘Before the present (which is taken to be 1950). The
initials should not be confused with BC” — not to mention silage and Bulgaria (order Helotiales),
this is an omnium gatherum of botanical jargon, arcana and facts, but it is not a dictionary of genera
of cryptogams and phanerogams.

Criticizing a dictionary is a thankless task, and could be an endless one too. Leaving aside the
editor’s choice of entries, especially of genera, on which no agreement is likely, there is one criticism
_ that must be leve!led at Allaby’s editing. As I ‘explored’ the world’s vegetation zones, I noticed that
the explanations included such phrases as ‘“‘Part of Good’s (1974) boreal kingdom... .”, but
nowhere could I discover who Good was, nor what his 1974 work was titled. Likewise there is no
entry about boreal kingdom. Unfortunately, there is no bibliography nor even a brief list of select
references.

Apart from that criticism, I can understand that this particular dictionary will serve the academic
fraternity of botanists, but a copy is unlikely to be used frequently by an amateur botanist who Is
seeking enlightenment about native wild plants.

E. C. NELSON

Wild orchids of Scotland. B. Allan & P. J. B. Woods; photography by Sidney Clarke. Pp. vii + 135;
illustrated. H.M.S.O. Edinburgh. 1993. Price £24.95 (ISBN 0-11-494246-3).

During the 20 years [ lived in Scotland I took a very active interest in most of the wild and cultivated
flora of the country, but I carefully avoided the wild orchids as one of the groups, along with the
grasses, hawkweeds and mosses, which were difficult and best left to the specialist. This book has
changed all that and is a first rate guide which makes accurate identification easy and also makes
orchids and orchid biology accessible.

The fact that I know many of the people responsible for this book and that some of them are old
friends is not going to stop me from giving it a rave review. Having declared my interest I cannot say
anything other than that this is a marvellous book and it deserves all the praise which will, I
confidently expect, be heaped upon it.

Let me list just a few of the virtues of this book: the identification key works and is illustrated ina
way which gives confidence, even to a novice both in the use of keys and in the structure of the
orchids, that one is on the right track; the distribution maps which show the range of each species
and subspecies before and after 1970 tell interesting stories about plant colonisation and

74 BOOK REVIEWS

disappearance (or on a different level tell us about the collecting habits and knowledge of botanists);
Mary Bates’ drawings are clear, effective and well labelled; the text is informative and accessible
and the book itself is attractively but unpretentiously designed with a clarity which fully supports the
book’s purpose but which also gives it an approachable and enjoyable feel by devices such as the
simple, uncluttered, airy layout of pages which start each chapter.

This unpretentious clarity extends to the photographs which add a whole new dimension to the
book. I have long admired Sidney Clarke’s photography both for its technical professionalism and
for the way in which he shows plants, either as individuals or as elements in a habitat, in an
unembroidered and direct way. He uses his photographic art in the service of science and good
communication: there is no artiness in the pictures (though I would enjoy seeing the book’s
endpapers as a furnishing fabric or wallpaper). In a short chapter of only four pages on photography
he tells us how he does it but he doesn’t mention the lifetime of walking the Scottish hills in pursuit of
plants, his deep personal interest in photography and his commitment to the Scottish flora and its
conservation which are the essential foundation of his success. The photographs were taken in only
three seasons and one is tempted to ask if it ever rains on Scotland’s orchid habitats.

To call the text scientific is to hint that it might be scholastic, accurate but rather dull: far from it.
Here is a well-written text in the best tradition of good communication. It perfectly complements the
photographs by being concise, clear and free of any taint of jargon. Again this artlessness conceals,
and is a result of, a lifetime of enthusiasm for the study of orchids by Patrick Woods and his co-
authors.

So, is there anything I do not like about this book? No: my admiration for this initiative is
unqualified — there are no buts. Everyone associated with its production can be proud of their roles
in what is an excellent example of the way in which a variety of different talent can be brought
together to deliver an elegant product. It is an outstanding example of what botanic gardens can do
so well as ‘packagers’ of sound scientific knowledge for a wider, enthusiastic and demanding
audience. I invite the audience to join me in the applause.

R. B. BURBIDGE

The vegetation of ultramafic (serpentine) soils. Edited by A. J. M. Baker, J. Proctor & R. D. Reeves.
Pp. 509. Intercept, Andover, Hampshire. 1993. Price £47.50 (ISBN 0-946707-62-6).

The Proceedings of the First International Conference on Serpentine Ecology, held in California in
June 1991, is a well-focused collection of 36 papers from ecologists, soil scientists, plant
physiologists, taxonomists, geneticists and evolutionary and molecular biologists. Approached from
these different viewpoints, the volume contains a wide global representation of studies which
contribute towards an understanding of ultramafic ecology. Editing and production standards of the
book are well above that which we often expect from conference proceedings; this book is a valuable
reference work, broader in scope and more comprehensive than a previous excellent review edited
by B. A. Roberts & J. Proctor (The ecology of areas with serpentinized rocks (1991)).

Soils derived from ultramafic rocks such as serpentine occupy less than 1% of the earth’s surface,
being widely scattered throughout temperate and tropical regions. They are rich in ferromagnesian
minerals, with low silica content, but otherwise they are rather variable. Most evidence suggests that
different combinations of nickel toxicity, magnesium toxicity in relation to calcium deficiency, low
nutrient status and drought are responsible for an often unusual and distinctive vegetation.
Ultramafic floras are often rich in endemic and rare species. In New Caledonia, ultramafic soils
occupy about a third of the archipelago and contain 60% of the native flora with 1,844 species,
90.6% of which are endemic. In Cuba, serpentine endemic species represent 15% of the flora, with a
much higher proportion found on old serpentine soils exposed for 10-30 million years rather than on
soils exposed for 1 million years or less. The endemic content of other serpentine floras may be
considerably less, for example accounting for only 3% in the Apennines in northern Italy where they
consist mostly of neo-endemics. Plant communities on ultramafic soils range from arctic-alpine,
through meadows, communities dominated by trees and shrubs, to rain forests. However, it appears
that vegetation cannot be described as ‘serpentine’ merely as indicated by geological maps; on

BOOK REVIEWS 75

ultramafic outcrops in Ireland there is no distinctive vegetation or endemism and no occurrence of
rare species.

Hyperaccumulation of nickel by plants, to exceptionally high tissue concentrations of >1000ug
g_! dry weight (compared to normal concentrations in plant tissues of <Sug g_'), is an unusual
phenomenon that has received considerable attention and has now been identified in nearly 200
species (1-2%) on ultramafic soils. This extreme metal tolerance trait occurs surprisingly widely in
33 families of plants, although is more common in some such as the Brassicaceae and Euphorbia-
ceae. It is thought that high levels of nickel in plant tissues may confer greater resistance to attack by
insect herbivores.

Ultramafic soils and vegetation provide a valuable resource for ecological study, but many areas
remain undescribed biologically. The importance of implementing a conservation strategy for
ultramafic sites is addressed in a conference resolution. A large number of sites are substantially
disturbed by mining for iron, chrome, nickel and asbestos, or else by burning, grazing, urbanization
and agriculture. This book should appear on the shelves of all science libraries, where hopefully it
will encourage more people to become interested in ultramafic ecology.

N. M. DICKINSON

The eternal yew. T. Baxter. Pp. 192, illustrated. The Self Publishing Association Ltd, Units 7/10
Hanley Workshops, Hanley Swan, Worcestershire, in association with Trevor Baxter. 1992. Price
£19.50 (ISBN 1-85421-148-X).

The yew tree. A thousand whispers. Biography of a species. H. Hartzell Jr. Pp. xvi + 320, illustrated.
Hulogosi, Eugene, Oregon 97440. Price US$ 19.95 paperback (ISBN 0—938493-14-0).

The yew is an intriguing plant because there is excellent evidence that some individual living trees of
Taxus baccata are many centuries old. Taxus comprises perhaps seven species, including 7. baccata
(yew) native in western Europe and T. brevifolia (Californian or Pacific yew) of western U.S.A. and
Canada. Baxter’s eternal yew is strictly 7. baccata; Hartzell misunderstands the word species so his
‘biography’ is a rambling thesis about both 7. baccata and T. brevifolia, but principally the latter.

Trevor Baxter is an enthusiast, and the author of a couple of ‘poems’ and some impenetrable
prose about yew trees. His book, published in association with a self-publishing co-operative, would
have been much less painful to read had it been severely cut by a rigorous editor who could also have
corrected Baxter’s botanical orthography, and his woeful punctuation, capitalization and grammar.
On the positive side, Baxter’s chronicle is illustrated with recent black and white photographs of
many of the famous, age-less yews that grow in Great Britain and a few in Ireland; as an appendix he
has included a list of yew trees with recent measurements. In essence Baxter’s book is a modern
directory of churchyard yews, and he has not fallen into the trap of assuming that the bigger a tree’s
various dimensions — whether height, trunk girth, or canopy circumference — and the more decrepit
its appearance, the more ancient it is. Yew trees are, as Baxter points out, notoriously impossible to
age even by ring-counts due to their capacity to produce ‘compound’ trunks.

Like Baxter, Hartzell has great difficulties writing lucid prose, and his book would also have
benefited from the vigorous attentions of an editor with a good knowledge of the history of botany
and horticulture. Among his many extraordinary sentences is this one (p. 125):

‘Many of these remarkable old trees [churchyard yews in England, Ireland, Wales, Scotland and
Normandy] bear their myths and historical significance, their stories, legends and poetic
inspiration well, surviving a past so long that the entire duration of humankind on earth is only a
minute in the day to the span of the yew, whose own ancestors stretch back 200 million years in
time’.
Hartzell’s principal purpose in writing his book was to highlight the need to help protect the splendid
yew forests of the Pacific Northwest of America which are being logged so that taxol, an anti-cancer
drug, can be extracted from the bark of T. brevifolia. The figures are stark; 13-6 kg of yew bark yield
only 1 gm of taxol, and 125,000 ancient yew trees may produce about 340-500 kg of bark. The clear-
felling of the entire Pacific yew population might yield 334 kg of taxol.
Perhaps this ramshackle tome will help, but it is a confused and confusing work, interweaving

76 BOOK REVIEWS

European and American Indian folk-lore with studies of taxol and its anti-cancer properties,
uncritically mixing fact with fiction. The section headings are ‘““Old World yew’’, “Botany and
geography’, “Living witness of human history”’’, “Culture and geography of Pacific yew’’, ““The
modern dilemma” and “The metaphorical yew’, with appendices on “Yew trees larger than 20 feet
in girth in England and Wales’, ‘“‘Notable topiary and hedges in England”’, and ‘“‘Yew species and
cultivars”. Hartzell’s knowledge of the use of yews in gardens is poor — the garden at Castle Drogo,
Devon, was created in the 1920s, and the presence elsewhere of Irish yews, none of which can have
been planted before 1780, clearly indicates nineteenth century embellishments.

Concerning the Irish yew (7. baccata ‘Fastigiata’), sometimes called the Florencecourt yew, a
footnote is required to both books. This distinctive cultivar has a well-recorded history but neither
author cited the correct dates which were given in The nomenclature and history in cultivation of the
Irish yew, Taxus baccata ‘Fastigiata’, Glasra 5: 33-44 (1981).

The dust-jacket blurbs inaccurately describe Baxter’s work as ‘““comprehensive”’ and Hartzell’s as
“authoritative” — do not be deceived. For members of this Society, Baxter’s book is the more
interesting, but neither can be recommended. I suggest that you spend your pocket-money on a
well-used second-hand copy of W. Dallimore’s Holly, yew and box (1908), or J. Lowe’s The yew
trees of Great Britain and Ireland (1897); these may be antique tomes but they are well written and a
pleasure to read.

E. C. NELSON

Phylogeny and classification of the Orchid family. R. L. Dressler. Pp. 314, 16 pages (96
photographs) of colour plates, numerous line drawings. Cambridge University Press, Cambridge.
1993. Price £35 (ISBN 0—-521—45059-6).

The invitation to review this book came to me not despite my lack of specialist knowledge of the
orchids, but precisely because of it. One of the tragedies of orchidology is that, in proportion to its
verbal output, it has given rather little to botany in general. Dr Dressler’s earlier book (The orchids:
natural history and classification, 1981) was a step towards redressing the balance. I was, therefore,
happy to review a new book by the same author.

First, however, one must deal with the relationship between this book and its predecessor.
Despite the change of title, this is a revised, updated and expanded version of the earlier volume.
There has been a great output of orchid literature in the intervening years, and this has obviously
been carefully sifted and taken on board. But where there has been no new information of value,
then the treatment in the 1981 volume has been retained more or less verbatim. For instance in
chapter 2, many of the entries are scarcely changed and eleven out of 16 text figures are the same:
but the section on seed-coats has been completely revised (based on recent work by Barthlott and
his school). In the colour section at least 25 of the 96 photographs are the same. The classification
has been revised by the re-positioning of Neottieae, and by the dispersal of the ‘vandoid’ orchids:
some sections (e.g. Oncidiinae, p. 177) have undergone considerable change, others virtually none;
and many of the illustrations are the same. Title page and dust jacket make no mention of the earlier
book: a word or two there would have avoided the need for this paragraph.

The better-known aspects of orchid-pollination were well-covered in the first book. That section
has now been dropped, but there are two pages (pp. 222-223: ‘pollination’ does not occur in the
index!) with the provocative heading ‘“‘False advertisement and the evolution of efficiency”. The
first phrase refers to such devices as the display of dummy anthers (patches of yellow hairs on yellow
swellings) that attract pollen collectors but offer nothing in return; or the mimicking by orchids of
nectariferous flowers amongst which they grow, without offering nectar themselves. Insects are
quick learners: false advertisement depends on a good supply of gullible, inexperienced, insects.
The mimics may show only 2-50% pollination compared to 80-95% in related species offering
nectar. This is the ‘evolution of inefficiency’. The pay-off comes because the insects usually visit only
one flower in an unrewarding spike; thus most of the fruits set have been cross-pollinated. The
higher fruit-set of nectariferous species is due to pollination from one flower to another on the same
spike. These two pages offer no ‘false advertisement’: they give real food for thought.

Dr Dressler is not afraid to set out his views on species. They show a very sensible approach to the

BOOK REVIEWS 77

taxonomist’s problems, and lead to the conclusion that only special creation could have given us a
world with all species neatly defined, whereas under evolution “‘we should find a spectrum of not
species, almost species, just barely species, and clearly defined species, and that is exactly what
occurs in nature”. An admirably clear statement — but it is a statement of the situation. The problem
of where to apply binomial nomenclature remains.

Parallelism is a central feature of evolution, and orchids provide a treasure trove for its study.
(The index takes one only to a brief entry on p. 233: the meat of the subject is on pp. 218-222). After
listing some early straightforward examples, the author introduces two new terms: integrational and
contingent parallelism. Here only two comments are possible. His definition covers really only the
qualifying adjectives. He writes ““By ‘contingent parallelism’ I refer to features that may evolve only
after another feature is present .. . Thus feature A permits the evolution of feature B, and B
may. . .”’. This defines an evolutionary pattern (see Burtt in Transactions of the Botanical Society of
Edinburgh 42: 138, 1974); it only becomes a parallelism if it is recurrent in different lineages.
Dressler’s voice on such wide topics is not authoritative, but it is a fine stimulant for the reader to put
in some hard thinking for himself.

Don’t pass over this book because of its title. Even if you cannot tackle the orchid-classification
amidships, fore and aft, it is well worth any botanist’s time.

Babe BuRTrT

Kvétena Ceske Republiky (Flora of the Czech Republic). Edited by S. Hejny & B. Slavik. Vol. 1.
Huperziaceae — Urticaceae. Pp. 557 with numerous line drawings. 1988. Vol. 2. Fagaceae —
Empetraceae. Pp. 540 with numerous line drawings. 1990. Vol. 3. Brassicaceae — Malvaceae. Pp.
542 with numerous line drawings. 1992. Academia, Praha. Price not given (ISBN 80-200-0256-1).

These are the first three volumes of the Flora of the Czech Republic, describing the flora of a key
area in central Europe. The work covers the Czech Republic (the western half of the old
Czechoslovakia), the Slovak flora being covered in a separate series.

The work is in Czech. The first volume includes a summary and glossary in English by M.
Kovanda, and each other volume includes a brief introduction in English, and it does not take long
to make sense of the accounts. The books are hardback, in A4 format and are clearly printed with a
good binding.

Accounts of families or genera have been prepared by specialist collaborators in addition to the
editors and their assistants. Each family is given a description and a key to genera, and likewise each
genus. Relevant literature is listed. For each species, Latin and Czech names are given, with
synonymy and representative specimens where appropriate. The descriptions of each species are
very detailed. Flowering times and chromosome numbers are given. There are often notes on the
morphological variation, ecology, vegetation types and distribution, both in the Czech Republic by
the phytogeographical district and elsewhere. Hybrids are treated in full.

There are numerous black and white line drawings of the majority of species (cross referenced in
the text). These are beautifully drawn in large format and capture the look of the plants perfectly.

Such a detailed comprehensive flora will be invaluable for any British botanists visiting the Czech
Republic, which has a huge potential for botanical holidays and excursions. It will also be a valuable
reference book for armchair botanists. It is highly recommended: if only we had a flora like this in
Britain!

T. C. G. RicH

Nouvelle flore de la Belgique, du nord de la France et des régions voisines. 4th edition. J. Lambinon,
J.-E. De Langhe, L. Delvosalle & J. Duvigneaud. Pp. cxx + 1092; illustrated. Editions du
Patrimonie du Jardin botanique national de Belgique, Meise. 1992. Price BEF 1,720 (ISBN 90—
72619-07-2).

This is a concisely written excursion Flora of Belgium, Luxembourg, N.E. France, S. Netherlands
and a small part of Germany. The fourth edition is extensively revised, having drawn on the detailed

78 BOOK REVIEWS

work of the Institut floristique belgo-luxembourgeois and the Institut floristique franco-belge for
distributional data as well as on recent taxonomic and nomenclatural studies. Many more
adventives are treated than in the third edition, and a new key is presented to the trees, shrubs and
lianes using mainly vegetative characters; this allows rapid identification not only of native
arborescent species but also of cultivated and planted specimens.

One of the most valuable features of this Flora, written in French but with additional Dutch and
German vernacular names, is the attention paid to infraspecific variation. Despite the concise
format, in which almost all comparative descriptive data are confined to the keys, space is found for
detailed observations on taxa such as Parnassia palustris var. condensata Travis & Wheldon;
subspecies and interspecific hybrids of Mentha alike are treated in a single key. Special attention is
paid to the zinc-loving variants of species such as Armeria maritima and Thlaspi caerulescens, and
notes are also provided on the varied medicinal and other uses of many of the plants.

An exhaustive glossary is provided, and the Latin, French, Dutch and German indexes are
preceded by a list of nine new combinations dated “‘December 1992” but seemingly published in
April 1993. One of these establishes a new subspecies of Scabiosa columbaria (subsp. pratensis
(Jord.) Duvigneaud & Lambinon) for material with several pairs of more frequently divided cauline
leaves, inhabiting deeper alluvial soils than the typical subspecies. This taxon could perhaps occur in
old meadows in southern England, although in Belgium it is “‘en fort régression’’.

Numerous illustrations are interspersed in the text, largely of diagnostic characters; in this respect
the Flora should prove specially useful to the inexperienced botanist. In size the book is similar to ~
the first edition of Clapham, Tutin & Warburg’s Flora of the British Isles (1952); the price (which
approximates to £35 at mid-1993 exchange rates) represents good value for a work of over 1200
pages. The book deserves to be bought by private purchasers as well as by institutional libraries.

J. R. EDMONDSON

The fern guide. A field guide to the ferns, clubmosses, quillworts and horsetails of the British Isles.
J. Merryweather & M. Hill. Pp. 101-188, with numerous drawings and 29 colour plates. AIDGAP,
Field Studies Council, Preston Montford. 1992. Price £5.25 incl. p&p (ISBN 1—85153—211-0).

There are now several field guides to the ferns and fern allies of the British Isles. The fern guide is in
the usual AIDGAP format of extended, copiously illustrated keys, and is aimed at teaching field
identification to beginners. For someone with plenty of time in the field (and prolonged fine weather
as it takes a while to work through the main fern key of 48 pages) it should provide a great deal of
pleasure and names for most of what one can find. One is led efficiently step by step, and the very
good illustrations, including 29 colour photos, are integral to the process and make it generally clear
whether one is on the right lines. (When one isn’t, backtracking can be difficult as there are no back
references to the dichotomy one has come from.) The book contains many helpful hints, including
one for Equisetum X littorale that works and that was new to me. The forms of Athyrium filix-femina
in open and shaded situations, so distinct and so puzzling to the beginner, are for example
mentioned and well illustrated in colour. There are, though, some cases where the book misleads.
For example, the glands on the indusium of Dryopteris oreades are not as constant a character as 1s
implied, and the much better character of the broad, divergent teeth at the apex of the pinnules is
not mentioned and is indeed belied by the drawing.

The book has several major drawbacks. It may enable users to put names to the plants they see,
but it does little to teach them about fern classification and relationships. It thus fails to provide any
basis for a deeper understanding of the group and hence any hope of real certainty in identification.
Genera, for example, are not described, and as their constituent species often key out in different
parts of the key the reader gets no idea of how the species are grouped into genera. In several groups
of ferns, microscopic characters are essential to confirm identifications. While it is obviously
reasonable for a field key to omit these, it should surely be stated that, for example in Polypodium, |
such characters must be used if the identification is to be definite. In Jsoetes it is misleading to say |
that hand lens examination of the megaspores of /. lacustris and I. echinospora “‘may help” to
confirm the species. The vegetative characters are often completely unreliable and examination of
the megaspores, by something more powerful than a hand lens, is essential. (The AIDGAP key to

BOOK REVIEWS 79

woodlice uses microscopic characters quite extensively, and several groups of ferns are at least as
difficult to identify as are the woodlice, with or without a microscope. Why should the identification
of ferns be taken any less seriously?) It is not in the long run helpful to anyone to suggest that the
identification of ferns is easier and more foolproof than it really is.

The list of books under ‘Further reading” would have been more helpful if it had been annotated
so that the beginner had some idea of what to turn to next, and for what. Infuriatingly, we still lack a
comprehensive book for identifying ferns, that would combine the clarity and wealth of characters
(including microscopic ones) provided by Jermy & Camus in The illustrated field guide to ferns and
allied plants of the British Isles (1991), the descriptive detail and the coverage of hybrids provided by
Page in The ferns of Britain and Ireland (1982) and the taxonomic and educative framework
provided by Hyde, Wade & Harrison in Welsh ferns (1969). Of these main contenders, Jermy &
Camus is the best as a field guide but has the great drawback of having very little on hybrids. Page,
magisterial though it is, is inadequately illustrated and has only very partial keys, making it daunting
for the beginner. Hyde, Wade & Harrison is very out of date and inadequately illustrated. The book
under review can be recommended to anyone wishing to take up fern identification from scratch, but
it should not on its own be relied on to give foolproof identifications and, if it does its job, its users
will very soon need to graduate to Jermy & Camus and to Page.

A. O. CHATER

Supplement to Flora of Cheshire. A. Newton, with cover illustration by W. Young. Pp. 52. Privately
published, Leamington Spa. 1991. Price £5.75; available post free from N.M.G.M. Enterprises,
P.O. Box 33, Liverpool L69 3LA.

This nicely produced booklet brings knowledge of the Cheshire flora up-to-date (1990), and
monitors changes over the past 20 years. Aquatic species are particularly vulnerable as the
remorseless destruction of the water-filled marl pits so characteristic of the Cheshire scene
continues. On the other hand, the number of alien and adventive species grows, and many appear
here in a two-page listing. Attention might have been drawn to the misguided practice of
introducing into Nature Reserves species not native to the area (e.g. Rosa rugosa).

It is pleasant to find that some species supposed extinct or nearly so have been refound, such as
Rhamnus catharticus at Kingsmarsh. The troublesome genera Taraxacum and Rubus are well
covered with 90 species of the former and 27 of the latter.

The index ts unsatisfactory; the page references are all one page in advance of the text; thus
Salicornia 1s indexed as on p. 15 whereas it actually appears on p. 14. Nevertheless, with colourful
stiff covers the Supplement is a valuable addition to the county Flora.

N. F. McMILLAN

Marianne North at Kew Gardens. L. Ponsonby. Pp. 128, 128 colour and four black and white
illustrations. Webb & Bower, in association with the Royal Botanic Gardens, Kew. 1990. Price
£15.95 (ISBN 0-86350—309-8).

Outstanding among the women who were lone explorers and botanical painters in those Victorian
days of more difficult travel, Marianne North gave her 832 paintings of plants in many countries to
the Royal Botanic Gardens, Kew. Many visitors will have seen these in the Marianne North Gallery
at Kew, now extensively renovated in recognition of the centenary of her death. Laura Ponsonby,
an authority on Marianne North, selected 130 of the paintings, many of which have not before been
published, to illustrate her biography and extensive account of those remarkable travels. An added
interest in the book are the detailed captions to these paintings which show scenes of plant life and
natural beauty from around the world. Sadly, now a hundred years later, it would be hard to find

80 BOOK REVIEWS

such unspoilt scenery and such a profusion of flowers in many of the places visited by Marianne
North.

M. Briccs

Stearn’s dictionary of plant names for gardeners. W. T. Stearn. Pp. viii + 363. Cassell, London.
1992. Price £16.99 (ISBN 0-304-34149-5).

The latin (or latin-form) names given to plants by botanists are often a great puzzle to gardeners.
How plants got the names they have and what those names mean or represent are matters that are
dealt with cursorily, if at all, in most of the horticultural-botanical literature. The present book,
which covers these subjects, is a revision of A. W. Smith’s Dictionary of plant names for gardeners of
1963, including “‘the addition of many more entries, the expansion and emendation of others [and]
the omission of materials considered irrelevant . . .”’. These changes are so extensive that the book
is really a new one, and has been so treated by the publishers.

The main part of the book (pp. 27-315) consists of an alphabetical listing of generic names and
specific epithets used for garden plants, with explanations of their meanings and origins. Though
Professor Stearn makes no claims for completeness, I have not been able to dig up any names that
are not included. Most of the entries are short and concise, but some are more extensive, for

example, those for ‘Forsythia’ (17 lines) and ‘paxtonii’ (14 lines). Professor Stearn brings a lifetime’s.

experience to his task and the range of erudition he displays is enormous: names from many
languages are cited (including Thraco-Palasgian (pre-Greek, see the entry under ‘Hyacinthus’)) and
the amount of biographical information condensed into a small space is astonishing. In spite of his
quotation of Dr Johnson’s definition of a lexicographer on p. 316, Professor Stearn wears this
erudition lightly and many of the entries include touches of humour or even the grotesque.

Gardeners will use this’ section extensively to find out the meaning of such unlikely words as
‘aiolosalpinx’ (as in Rhododendron aiolosalpinx — it means the trumpet of Aeolus, the god of the
winds) and to discover, for instance, who was the Rudbeck after whom the well-known Composite
genus Rudbeckia was named, thus increasing the interest provided by the plants they grow.

This main index is preceded by an “Introduction to Botanical Names” (pp. 17—26) which discusses
the principles used by botanists in naming plants and the various kinds of name that are used. It also
touches on the rules of botanical nomenclature especially as they affect such thorny subjects as why
familiar names seem to be wilfully changed by botanists.in order to confuse and irritate the

gardener.
The final section (pp. 317-329) is a very sensible essay entitled ““An Introduction to Vernacular
Names’’. The advantage (‘‘. . . vernacular names should not be despised and disregarded. When

only a few plants need to be distinguished within a limited area, vernacular names can be just as
useful, precise and stable as their scientific equivalents intended for international use.” — p. 318) and
disadvantages of these are presented calmly and rationally. This is a section that could be read with
advantage by taxonomic botanists as well as gardeners.

A glossary, bibliography and index complete the volume, which is well produced and contains few
printing errors. It should be on the shelves (or, more likely, on the desk or bench) of every gardener
or botanist interested in garden plants.

J. CULLEN

Red data books of Britain and Ireland: Stoneworts. N. F. Stewart & J. M. Church. Pp. 144, 14 colour
photographs, numerous line drawings and maps. Joint Nature Conservation Committee, Peter-
borough. 1992. Price £15 (ISBN 1—873701-24-1).

This particular Red data book breaks new ground in covering the entire British Isles: the vascular
plant red data books for Great Britain and Ireland were published independently in 1983 and 1988.
In this instance, however, we have the pleasure of a single volume introduced to us by both
Ministers responsible for the environment of the U.K. and the Republic of Ireland.

BOOK REVIEWS 81

Although technically algae, stoneworts have always been treated as being within the legitimate
purview of the B.S.B.I. — their large size makes them conspicuous in the field, so that many vascular
plant botanists have been attracted to pay some attention to them. In recent years, active interest in
the group has increased. The sensitivity of stoneworts to pollution makes them of considerable value
as indicators of good water quality.

As defined by the authors, there are 33 British and Irish species of stonewort, with about 400
worldwide; two species are endemic to these islands. Problems of nomenclature and taxonomic
ranking are dealt with briefly on pp. 37-39, and in a useful table. There are considerable differences
between the species as defined in this work and those of the B.S.B.I. handbook of 1986. However,
the authors have striven to include sufficient synonymy to enable the user to locate a species in the
major earlier works.

The book is well illustrated by colour plates of habitats, and there are excellent line drawings by
Margaret Tebbs (many previously published in the B.S.B.I. handbook) of individual species. The
section on habitats stresses among other things, the difference between calcium carbonate-rich and
nutrient-rich water bodies, and the damaging effects of pollution. The chapter includes a table
where the habitat preferences of each of the 33 species are presented.

The principal criteria for listing a species as a red data book species are that for Great Britain it
should have been recorded in 15 or fewer, and for Ireland ten or fewer 10-km grid squares since
1970. Together with other qualifications this produces a total of 57% and 48% (17 and 12 species) of
the stonewort floras respectively. Table 4 tabulates the threats to each species and gives each a
Threat Status (Extinct, Endangered, Vulnerable, Rare, etc).

The species accounts give useful notes on identification (with a line drawing), the distributions in
Great Britain and Ireland (with a distribution map), world distribution, ecology, main threats,
existing protection and conservation priorities. An appendix gives a key to all British and Irish
species using field characters requiring only the use of a hand lens. It is interesting to compare the
distribution maps with those in the B.S.B.I. handbook. For those few species where there are two
exactly corresponding maps additional records appear in the Red data book.

The book is an attractively presented and well designed summary of the conservation status of
stoneworts in the British Isles and is a model of clarity. I found it readable and packed with
fascinating and useful information as well as the expected data on threat status, etc. Anyone
interested in these plants, or in plant conservation generally, should purchase a copy.

P. HACKNEY

Flora Europaea, second edition, vol. 1: Psilotaceae to Platanaceae. Edited by T. G. Tutin ef al.,
assisted by J. R. Akeroyd & M. E. Newton. Pp. xlvi + 581. Cambridge University Press,
Cambridge. 1993. Price £100 (ISBN 0-521-41007-X).

The publication of the 5-volume Flora Europaea between 1964 and 1980 was a landmark in
European botany. Various reviewers of volumes of the first edition predicted that this work would
stimulate further taxonomic research on the European flora and this has indeed been the case. The
initial volume was published in 1964 and, because it was the pioneer part, of necessity it was the one
most in need of revision since it contained more errors than subsequent volumes. All European
botanists will therefore welcome this revision, produced 29 years after the original. It is partly
financed by using the royalties from sales of the series which were lodged in a trust fund,
administered by the Linnean Society of London. This new edition continues to remind us of the
great vision of the original group of taxonomists who initiated and successfully saw the project
through to completion. Flora Europaea has received much acclaim and has been recognised as the
definitive reference for botany by the Council of Europe.

Volume 1 contains treatments of the 25 families of Pteridophyta present in Europe, the five
families of gymnosperms and the first 49 families of angiosperms from the Salicales to the Rosales
(in part). Major families include the Caryophyllaceae, Chenopodiaceae, Cruciferae and Ranuncu-
laceae. This is certainly not just a reissue of the previous edition because the editors have carried out
extensive work on updating. Any taxa new to Europe since 1964 have been incorporated and many
descriptions have been improved. The keys have also been extensively revised and, in many cases,

82 BOOK REVIEWS

made easier to use, while all synonyms are cited in the text. The fact that 250 species and 150
subspecies have been described from Europe as new to science in the 79 families covered by Volume
1 shows that there is still much taxonomic activity on the Continent and that there is still much to do
in order to have a definitive taxonomic account of the region. The new records and alien species
included means that the new edition has about 350 extra taxa; in compensation, only 20 taxa have
been deleted. The useful appendices of the first edition have been revised by R. R. Mill. It is a pity
that the author abbreviation appendix has not adopted the standard abbreviations of Brummitt &
Powell Authors of plant names (1992) or for the book citations those of Stafleu & Cowan Taxonomic
literature, 2nd ed. (1976-1988), or indeed of any other standard work. Researchers will be glad to
see the inclusion of the complete synonymy of the species which will greatly facilitate comparison
with previous works.

Seeing the result of the revision, we should be grateful that the Flora Europaea editorial
committee was not just disbanded after the publication of Volume 5, but decided to embark on a
revision of Volume 1. Given the extent to which new taxa have been added in this revision, it is a pity
that there are no current plans for revisions of the other four volumes. It is much more useful to have
the changes and additions gathered together into a revised volume than spread throughout the
literature in a large number of papers.

The principal editors of this revision of Volume 1, J. R. Akeroyd and M. E. Newton, are to be
congratulated on the thoroughness of their work. Botanists truly interested in the European flora
will need to have this new volume, rather than rely on the first edition, because it contains so much
additional information.

G. T. PRANCE

Origin and geography of cultivated plants. The late N. I. Vavilov, translated by D. Love, foreword
by A. A. Filatenko. Pp. xxxii + 498, 28 line diagrams and 33 half-tones. Cambridge University
Press, Cambridge. 1992. Price £75 (ISBN 0—521—40427-4).

This is an English translation of a work first published in Russian under the (transliterated) title
Proiskozhdenie i Geografia Kul’turnykh Rastenii by the Leningrad (now St Petersburg) branch of
‘Nauka’, the Russian science publishing house, in 1987, to commemorate the centenary of the birth
of the great Russian plant geographer, ecologist and plant breeder Nikolai Ivanovich Vavilov
(1887-1943). That volume, in turn, was a collection of 25 of Vavilov’s most important papers,
originally published in various Russian journals from 1920 onwards, some of them posthumously.
The volume under review is the first in which all these papers, many of them of seminal importance,
have appeared collectively in translation. Doris L6éve’s translation is masterly on the whole,

although there are a few idiosyncrasies.

The titles of the papers show the immense breadth of Vavilov’ s interests in cultivated plants (and
domesticated animals) and their origins. They range from ‘On the origin of plants’, “Asia — the
source of species’, ‘Plant resources of the world and the mastering thereof’ to ‘On Soviet science and
the study of the problem concerning the origin of domesticated animals’. Most papers are quite
short, but the book includes his magnum opus, ‘Centres of origin of cultivated plants’, which runs to
112 pages. This contains, among other sections, a long, detailed, and utterly absorbing account of
the origins of cultivated hemp, Cannabis sativa — of more than academic interest to British botanists
now that E.C. legislation allows Cannabis to be legally grown, for fibre, once again in the British
Isles!

Vavilov was a great explorer in his search for elusive genotypes. As well as travelling through
most of Asia, he also visited the New World, in particular Mexico and Central America. Two
papers, ‘Mexico and Central America as a basic centre of origin of cultivated plants in the New
World’ and ‘The important agricultural crops of pre-Columbian America and their mutual
relationship’ resulted from these explorations.

The sense of amazement and wonder that Vavilov so obviously had on his travels, expressed with
deep feeling in many of his papers, shines through with undiminished radiance in the translation.
The papers contain a deep, rich mine of information and theory on the origin of almost every one of
the world’s economic plants. Some of this is now outdated, if only because radical changes in

BOOK REVIEWS 83

agricultural practices in the last five decades have sadly reduced the gene pool of ancestral species to
a fraction of what it was in many of the areas where Vavilov carried out his pioneering studies. The
nomenclature has also been largely left as it was in the original papers; the correct current name is,
however, always indicated in square brackets in the index to Latin plant names, and occasionally
there are helpful insertions by Doris Love, also in square brackets, in the main text to clarify or
update Vavilov’s writings.

The book has been sumptuously produced by Cambridge University Press. As supplied for
review, it had no dust jacket; instead, there is a striking gold-embossed Oat panicle on the dark
green cover, with the title and author’s name gold-embossed on a black background. The rather
small print is nevertheless easy to read; misprints appear to be extremely few, although,
disconcertingly, Nikolai Ivanovich’s first name appears in no fewer than three other variants
(Nickolay, Nikolay and Nicolay) in the first few pages of introductory matter, pp. 1-xiv. One
irritating feature is the use of American English spellings throughout. I do not mind, indeed I
expect, “‘centers of origin” in a book published in America, but in one published by one of Britain’s
most highly respected houses I abhor it. Sadly, this appears to be a growing trend, but should be
strongly resisted. My other main criticism is the very high price. This will deter many a student or
library from acquiring this book — indeed, a colleague from Jerusalem actually told me that his
library could not afford it. That is a pity, because it is a fitting memorial, not just to an almost
legendary Russian scientist the likes of whom we are unlikely to see again, but also to the richness of
a genetic ‘landscape’ now greatly diminished. I hope that a much cheaper, durable paperback
edition will be made available so that a wider public can appreciate Vavilov’s enthusiasm and
lifetime’s energies.

R. R. MILL

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Watsonia, 20, 85-87 (1994) 85

Report

ANNUAL GENERAL MEETING, 16 MAY 1993

The Annual General Meeting of the Society was held at the Lincolnshire College of Agriculture and
Horticulture, Riseholme Hall, Riseholme, Lincoln, at 12.30 p.m. 80 members were present. Dr P.
Macpherson, retiring President, taking the Chair for the first three items on the Agenda, welcomed
the members present, then stated that his first sad duty was to report the recent death of Mr R. W.
David, C.B.E., M.A., a past President and Honorary member of the Society, and joint author of
the B.S.B.I. Handbook Sedges of the British Isles.

Apologies for absence were read and the Minutes of the 1992 Annual General Meeting, published
in Watsonia 19: 205-207 (1993), were approved and signed by the President.

REPORT OF COUNCIL

The Chairman summarised the highlights of the year in his Report, which had been circulated to
members; the adoption was proposed by Dr N. K. B. Robson, seconded by Mrs M. Lindop, and the
Report was accepted unanimously.

HON. TREASURER’S REPORT AND ACCOUNTS

The Hon. Treasurer, proposing the adoption of his Report and Accounts, which had been circulated
to members, referred to the Welch Bequest Fund from which a number of grants had been paid
during the year, including to the B.S.B.I. Database (Leicester) project which was proving to be an
asset to the Society. Mr Walpole also reported that all Handbooks destroyed in the flood had been
reprinted. He invited queries on the Accounts, and there being none the adoption of the Report and
Accounts, seconded by Mr R. G. Ellis, were accepted unanimously.

ELECTION OF PRESIDENT

Dr P. Macpherson, retiring President, commented that as President he had been given an insight
into the committee structure of the Society, and had gained an awareness of the amount of work
undertaken for the Society by members of the various committees, by the committee chairmen and
secretaries in particular, and also by the Honorary Editors. He thanked all these on behalf of the
Society. The Hon. General Secretary he thanked for keeping him in touch with Society affairs, by
phone and by many postcards, and for general communications within the Society. Mr M. Walpole,
as Hon. Treasurer and as Chairman of the Publications Committee he thanked with appreciation for
discussions and advice throughout his Presidency. Thanking the members of his President’s
Energising Panel, and of the Executive Committee, Dr Macpherson referred to the honour of being
President of the B.S.B.I., also thanking the members who had elected him, saying that he was very
appreciative of their support.

Introducing Dr F. H. Perring, O.B.E., F.L.S., F.I.Biol., Dr Macpherson commented that the
President-elect was well-known to members as an author, broadcaster and enthusiast for the
conservation of wild plants, who had many contacts in the conservation world with whom he would
aim for liaison and publicity for the Society. Dr Perring was unanimously elected, and taking the
Chair he said that he would be very happy to repay the enormous debt of gratitude that he owed to
the Society on being a member for 40 years. He hoped to contact many members and would value
highly their thoughts and comments on the Society.

On behalf of the Society he particularly thanked Dr Macpherson for his Presidential Address, for

86 REPORT

his regular attendance and firm chairmanship of meetings, undeterred by the distances involved. As
President at two Royal occasions Dr Macpherson had also represented Scotland by wearing kilt and
sporran, and Dr Perring remarked that the retiring President was leaving a hard act to follow.

ELECTION OF VICE PRESIDENT, 1993-1997

Mr A. O. Chater had been nominated by Council and was known to many members as a past
Secretary of the Publications Committee, a Referee for Carex and for nomenclature, the Recorder
for Cardiganshire and his work on the Records Committee and also as joint author of the B.S.B.I.
Handbook Sedges of the British Isles. His unanimous election was welcomed, with applause.

RE-ELECTION OF HON. GENERAL SECRETARY AND HON. TREASURER

Proposing the re-election of these Officers from the chair, the President endorsed Dr Macpherson’s
comments, and again thanked these Officers for the wonderful service which they gave to the
Society. They were unanimously re-elected, with applause.

ELECTION OF COUNCIL MEMBERS

In accordance with Rule 10 nominations had been received for Mr B. A. Gale, Mr D. J. McCosh
and Dr G. Wynne. Profiles had been published, and these members were elected unanimously by
the meeting. .

ELECTION OF HONORARY MEMBERS

Council had nominated two members: proposing Mr R. H. Roberts, M.Sc., Mr R. G. Ellis
described him as well-known to north-western botanists in Britain, and to those with special
interests in ferns and the genus Dactylorhiza as a Referee for these groups, as Recorder for
Anglesey and also as a long-serving member of the Committee for Wales and author of The
flowering plants and ferns of Anglesey.

Dr Perring proposing Mr P. J. O. Trist, O.B.E., B.A., introduced him as being based in East
Anglia for many years, and editor of An ecological Flora of Breckland, but was also well-known to
many members as a Referee for Bromus, Festuca and other grasses. Mr Trist, now retired as a
Gramineae Referee, had named 1,891 grass specimens for members during his time as Referee.

The election of Mr Roberts and Mr Trist as Hon. Members was unanimous, and with applause.

AMENDMENTS TO B.S.B.I. RULES, 1993

Re-wording Rules 3c) and 4a), under Management of the Society and the addition of Rule 10 under
The Officers, with subsequent adjustment to the numbering of Rules 11-39, was approved by the
meeting, and copies are available to members on request.

RE-ELECTION OF HON. AUDITORS

The Hon. Treasurer proposing the re-election of Grant Thornton, West Walk, Leicester, reported
that they were willing to continue as Hon. Auditors and the re-election was passed unanimously,
with applause. The President would write to express the appreciation of the Society.

REPORT 87
ANY OTHER BUSINESS

On behalf of those present the President thanked Mrs I. Weston as local organiser, and her team of
local helpers for the excellent arrangements and very good facilities provided at Riseholme Hall. He
again thanked the Speakers, and Mrs M. D. Perring and Mr J. and Mrs S. M. Atkins for bringing
new and second-hand botanical books.

There being no other business, the meeting closed at 1.25 p.m.

MAry BriGGs

B.S.B.I. 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 and R. W. David. 1982. 268 pages, with a line drawing and
distribution map for every species. Paperback. ISBN 0 901158 05 4.

No. 2. UMBELLIFERS OF THE BRITISH ISLES
T. G. Tutin. 1980. 197 pages, with a line drawing for each species. Paperback. ISBN 0
901158 02X. Out of print: new edition in preparation. 7

No.3 DOCKS AND KNOTWEEDS OF THE BRITISH ISLES
J. E. Lousley and D. H. Kent. 1981. 205 pages, with many line drawings of British native
and alien taxa. Paperback. ISBN 0 901158 04 6. Out of print: new edition in preparation.

No.4 WILLOWS AND POPLARS OF GREAT BRITAIN AND IRELAND
R. D. Meikle. 1984. 198 pages, with 63 line drawings of all species, subspecies, varieties and
hybrids. Paperback. ISBN 0 901158 07 0.

No.5 CHAROPHYTES OF GREAT BRITAIN AND IRELAND
J. A. Moore. 1986. 144 pages, with line drawings of 39 species and 17 distribution maps.
Paperback. ISBN 0 901158 16X

No. 6 CRUCIFERS OF GREAT BRITAIN AND IRELAND
T. C. G. Rich. 1991. 336 pages, with descriptions of 140 taxa, most illustrated with line
drawings and 60 with distribution maps. Paperback. ISBN 0 901158 20 8.

No. 7 ROSES OF GREAT BRITAIN AND IRELAND
G. G. Graham and A. L. Primavesi. 1993. 208 pages, with descriptions and illustrations of
twelve native and eight introduced species, and descriptions of 83 hybrids. Distribution
maps are included of 32 native species and selected hybrids. Paperback. ISBN 0901158 22 4.

No. 8 PONDWEEDS OF GREAT BRITAIN AND IRELAND
C. D. Preston. In preparation.

Available from the official agents for BSBI Publications:
F. H. & M. Perring, Green Acre, Wood Lane, Oundle, Peterborough PE8 4JQ, England.
Tel: 0832 273388 Fax: 0832 274568

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.

Manuscripts must be submitted in duplicate, typewritten on one side of the paper, with wide margins
and double-spaced throughout.

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 1991). Further details on format can be found in B.S.B.I. News
51:40—42 (1989).

Tables, figure legends & appendices should be typed on separate sheets and attached at the end of
the manuscript.

Figures should be drawn in black ink and identified in pencil on the back with their number and the
author’s name. They should be drawn 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 done with transfers or high-quality stencilling, although graph axes
and other more extensive labelling are best done in pencil and left to the printer. Photographs can be
accepted if they assist in the understanding of the article.

Contributors are advised to consult the editors before submission in cases of doubt. 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: Dr B. S. Rushton, Department of Biological and Biomedical Sciences,
University of Ulster, Coleraine, Co. Londonderry, N. Ireland, BT52 1SA.

Books for Review: Dr J. R. Edmondson, Botany Department., Liverpool Museum, William
Brown St, Liverpool, L3 8EN.

Plant Records: the appropriate vice-county recorder, who should then send them to C. D. Preston,
Biological Records Centre, Monks Wood Experimental Station, Abbots Ripton, Huntingdon,
BE? 2LS.

Back issues of Watscnia are handled by Dawson UK Limited,
Cannon House, Folkestone, Kent, CT19 5EE to whom orders for all issues prior to Volume 18
part 1 should be sent.

Recent issues (Vol. 18 part 1 onwards) are available from the Hon. Treasurer of the
B.S.B.I., 68 Outwoods Road, Loughborough, Leicestershire, LE11 3LY.

_— eee einen " =
oo ced Sea RT iA Sa 2 2

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 1993-94

Elected at the Annual General Meeting, 15th May 1993
President, Dr F. H. Perring

Vice-Presidents, Mr P. S. Green, Dr G. Halliday, Mr A. C. Jermy,
Mr A. O. Chater

Honorary General Secretary, Mrs M. Briggs
Honorary Treasurer, Mr M. Walpole

Editors of Watsonia

Papers and Notes, J. R. Edmondson, R. R. Mill, E. C. Nelson, B. S. Rushton*

Plant Records, C. D. Preston
Book Reviews, J. R. Edmondson
Obituaries, J. R. Akeroyd

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

Watsonia 20: 89-95 (1994) 89

Changes in the distribution of Erica ciliaris L. and
E. X watsonii Benth. in Dorset, 1963—1987

S. B. CHAPMAN*® and R. J. ROSE**

Furzebrook Research Station, Natural Environment Research Council, Institute of Terrestrial
Ecology, Wareham, Dorset, BH20 5AS

ABSTRACT

The distribution in south-eastern Dorset of Erica ciliaris (Dorset Heath) and FE: watsonii (Ericaceae) is
described. Between 1963 and 1987 the distribution of these taxa and their relative proportions within
populations changed. These changes have been caused mainly by the afforestation of the area whieh-forms the
centre of the plant’s distribution, and by heathland fires. Further evidence for the hypothesis that E. ciliaris is
still spreading in Dorset is provided.

Keyworps: afforestation, heathland, Erica tetralix.

INTRODUCTION

Erica ciliaris L. (Dorset Heath) has a ‘Lusitanian’ distribution, being found in Morocco, the coastal
regions of Portugal, Spain and France, and locally in southern England (Fig. 1) and western Ireland.
The plant is listed in the British red data book (Perring & Farrell 1983), and is classified as rare in
Great Britain. It occurs at a single site in Ireland (Webb 1966). In southern England, E. ciliaris is
most abundant on the heathlands of south-eastern Dorset where it was first recorded in 1848
(Mansel-Pleydell 1895). E. X watsonii is a hybrid of E. ciliaris and E. tetralix L.; backcrossing may
occur as a range of variants intermediate between the two parent species are found (Gay 1957;
McClintock 1971). The distribution of E. ciliaris has been described by a number of workers. Good
(1948) commented on the static nature of the population, stating that “local geographical limits
follow no obviously recognisable edaphic or climatic boundary’’. Gay (1957, 1960) suggested that
the population was contracting as a result of introgressive hybridisation, and Chapman (1975)
proposed that the pattern of distribution of E. ciliaris and E. X watsonii ‘‘can be explained in terms
of an expanding population”’.

Evidence that E. ciliaris was increasing its range in Dorset was provided by Haskins (1978) who
examined sub-fossil seed remains in peat deposits. The oldest sub-fossil seeds were found in peat
from Wytch Heath, near the centre of the present distribution of E. ciliaris, with more recent
remains at locations on the edge of the present range.

This paper examines the current distribution pattern of E. ciliaris and discusses changes in the
relative proportions of EF. X watsonii in certain populations against the background of changes in
heathland in Dorset over a period of 24 years.

METHODS

THE SURVEY
The distribution of E. ciliaris was mapped by field surveys in 1963, 1973 and 1987. In 1963 and 1973
the boundaries of its populations were mapped and measured by planimetry. In 1973 and 1987 the

* Present address: Hope House, Winterborne Zelston, Blandford, Dorset, DT11 9EU.
**'To whom correspondence should be addressed.

90 S. B. CHAPMAN AND R. J. ROSE

Ficure 1. The distribution of Erica ciliaris in Great Britain; 10-km square records from the Atlas of the British
flora (Perring & Walters 1962) with recent additions.

abundance of E. ciliaris in 200 m X 200 m grid squares was recorded. This size of sampling unit and a
similar method were employed in more general surveys of heathland in Dorset (Webb & Haskins
1980; Chapman, Clarke & Webb 1989; Webb 1990). Within each grid square abundance was
recorded using a four point scale of cover:

1 = present as only single or isolated plants, <1% of the grid square area.

2 = more than isolated plants, 1% to 10% of the grid square area.

3 = plants present in 10% to 50% of the grid square area.

4 = present in more than 50% of the grid square area.
Estimates of the areas of vegetation containing E. ciliaris have been derived using the mid-values of
the scores for each grid square. For example, squares with a score of category 4, which has a mid-
score value of 75%, were taken to have a mean area of 3 ha. Similarly, squares with scores of 3, 2,
and 1 were assumed to have mean areas of 1-2, 0-2 and 0-02 ha respectively.

VEGETATION CLASSIFICATION
In 1973 the dominant heathland vegetation type in each square was recorded. In 1987, data relating
to general heathland features such as classification of vegetation types, changes in land use and
heathland fires were taken from the Dorset heathland survey databases for 1978 (Webb & Haskins
1980) and 1987 (Webb 1990). These surveys were based upon the same 200 m X 200 m recording
grid used for the EF. ciliaris surveys described in this paper.

Some populations of E. ciliaris situated in areas that were formerly heathland are now within
conifer plantations. Data from these sites have been included in the distribution maps and analyses,
but omitted from the analysis of the population composition and vegetation type.

POPULATION COMPOSITION
In the field E. X watsonii can be distinguished from its parents by its more robust growth form and
the intermediate nature of the inflorescence shape. Where there was difficulty in assigning plants to

DISTRIBUTION OF ERICA CILIARIS AND E. X WATSONII 91

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Ficure 2. Distribution of Erica ciliaris populations in Dorset (10-km grid lines shown): @ indicates present
1963-1987; + new records 1973-1987; O sites lost 1963-1973; x sites lost 1973-1987.

a taxon the presence or length of the anther awns was examined. This character is the most reliable
for separating E. ciliaris, E. tetralix and E. X watsonii (Chapman 1975).

In 1973 and 1987 the grid squares were classified according to the relative proportions of E.
ciliaris, E. tetralix and E. X watsonii (Chapman 1975). This classification, and a simple four category
version in which the population was deemed to be mixed if no one taxon dominated and pure if one
of the three taxa of Erica accounted for more than 50% of the population.

The degree of change in population composition using the four category classification was
represented by a three point scale; no change, class 1 change and class 2 change. A class 1 change
was defined as a change from a pure population to a mixed population or vice versa. A class 2 change
represents a change from one pure population to another pure population.

RESULTS

DISTRIBUTION AND ABUNDANCE OF ERICA CILIARIS

The total area in which populations occurred was measured in 1963 and 1973 by planimetry and for
all three surveys using the mid-value scores for the 4 point abundance scale. The two methods were
in close agreement (Table 1) with less than 5% error in the mid-score abundance value estimate.
The changes in total area between 1963 and 1973 were very small; however, in the 14 years between
1973 and 1987 a reduction of 199 ha (27-:7%) occurred.

The distribution of E. ciliaris and E. X watsonii in south-eastern Dorset is shown in Fig. 2. E.
ciliaris populations have been recorded from 445 grid squares in the period 1963 to 1987; from 423
grid squares in 1963, 411 in 1973, and 388 in 1987 (Table 1). The loss of populations from 13 grid
squares between 1963 and 1973 (all in abundance categories 1 and 2) and the discovery of only one
new population (abundance category 1) resulted in little change in the overall abundance or

92 S. B. CHAPMAN AND R. J. ROSE

distribution. However, between 1973 and 1987 there was a loss of E. ciliaris from 58 grid squares,
and reductions in the numbers of category 3 and 4 squares by 26 and 32 squares respectively. The
smaller losses of E. ciliaris during the period 1963 to 1973 were mainly due to agricultural
reclamation, while the losses recorded between 1973 and 1987 were due to a wider range of activities
(Table 2). Of these losses eleven were from sites that were still described as heathland; such losses,
often caused by scrub invasion, are generally of small populations or isolated plants. The losses from
25 squares attributed to forestry was caused by the closure of the canopy in conifer plantations.
‘These populations were previously recorded as existing in heathland amongst young plantation
conifers.

Several of the new records in the outlying areas of heathland were the result of records obtained
from observers during the period 1973 to 1987. These were small populations or even single plants
that did not significantly increase the total area occupied by the plant but changed the known range

(Fig. 2).

VEGETATION AND LAND USE CHANGES

In 1973 a number of populations were recorded in heathland that had been planted with conifers. By
1987, 119 grid squares, which were recorded as heathland in 1973, were classified as woodland of
which 100 were conifer plantation, seven semi-natural conifer, and twelve scrub or carr. In addition
to afforestation, land use change through agricultural reclamation and the construction of homes
and roads also occurred. These activities either destroyed or modified the habitat, resulting in a shift
in the proportions of the different habitat types. In 1973, 393 squares were recorded with both E.
ciliaris or E. X watsonii; of these 373 (95%) were classified as predominantly humid heath, wet
heath or peatland. In 1987, 383 squares were recorded, but only 183 (48%) were classified as being

TABLE 1. ESTIMATES OF THE AREAS OF VEGETATION CONTAINING ERICA CILIARIS AND
E. X WATSONII IN DORSET IN 1963, 1973 & 1987

1963 1973 1987

Number of squares in each area category
Cat. 1 m5 29 22
Cat. 2 167 163 205
Cat. 3 145 145 119
Cat. 4 73 74 42
Total number of squares 423 411 388
Number of new squares - 1 35
Number of lost squares - 13 58
Area (ha) (from mid-value scores) 427 429 310
Area (ha) (from planimetry) 419 408 -

TABLE 2. LOSSES OF ERICA CILIARIS FROM NUMBERS OF 200 M x 200 M

GRID SQUARES IN DORSET, ACCORDING TO THE LAND USE, OR

CHANGES OF LAND USE, OF THOSE GRID SQUARES, IN THE PERIODS
1963-1973 AND 1973-1987

Period
Land use 1963-1973 1973-1987
Heathland 1 11
Agriculture db 21
Forestry 0 25
Other 1 1
Total ; 13 58

Area (ha) +2 —119

DISTRIBUTION OF ERICA CILIARIS AND E. X WATSONII 95

TABLE 3. PROBABILITY VALUES OF ASSOCIATIONS BETWEEN POPULATION TYPE AND

VEGETATION FROM SITES ASSESSED IN 1973, USING A 2-SIDED VERSION OF FISHER’S EXACT
SIGNIFICANCE TEST

Vegetation type

Population type Dry Humid Wet Peatland
Erica ciliaris 0-13 0-40 0-21 1-00

Erica X watsonii 1-00 0-0053(—) 0-057 0-000002(+)
Erica tetralix 0-62 1-00 0-0048(+) 0-092

Mixed 1-00 0-37 0-60 0-89

humid heath, wet heath or peatland. However, those sites which have not undergone land use
change remained remarkably constant between the 1973 and 1987 survey data.

POPULATION COMPOSITION

The distribution of the different population types, classified according to the proportions of E.
ciliaris, E. tetralix and E. X watsonii in 1973, was presented by Chapman (1975). Several
associations between vegetation type and the population composition were shown. These analyses
were repeated in 1987 but no significant associations were found. This was due to the changes that
took place in the afforested areas of central Purbeck; these reduced the number of populations and
make it impossible to make comparisons or reassessments of grid squares where a part of the
original habitat had been lost.

Using the simplified classification of populations, the same 1973 data were re-examined using a 2-
sided version of Fisher’s exact significance test. Significant associations were found between E. X
watsonii populations and humid heathland and peatland (Table 3).

When this analysis was repeated for the 1987 data no significant associations were found. This was
due to fewer squares being classified as E. X watsonii. Thus changes in population composition
between years, in different vegetation types, could not be analyzed. However, no significant
differences in the proportion of change of population composition in humid heath, wet heath and
peatland were found.

Having established that changes in population composition were not related to vegetation type,
the data were examined for changes after fire. Using the 1978 heathland survey data to identify areas
that had been burnt in 1974-1977, a significant association between burnt sites and change towards
E. X watsonii populations (p<0.001, Fisher’s exact test) was found (see Fig. 3).

In Fig. 3, in addition to the association with fire, it can be seen that a line of six grid squares
(arrowed) show two class changes in an unburnt area; these six squares are roadside sites and the
changes are probably due to increased management (mowing) of the road verge between 1973 and
L987:

DISCUSSION

Since the survey in 1973, isolated or small populations of E. ciliaris have been recorded from several
locations outside the centre of distribution. The history of origin of these populations is uncertain.
Some sites are near houses or under power lines and may have been introduced by human activity.
Whether any introduction has been made intentionally is not known. However, these isolated sites
should be monitored closely as their performance and survival should provide data to test the
hypothesis that E. ciliaris may still be establishing and colonising suitable habitats in south-east
Dorset.

The reduction in the number of E. ciliaris sites between 1973 and 1987 was not the result of
additional afforestation during this period, but due to the closure of the canopy of existing
plantations. Such losses indicate that the potential for future loss remains as plantations mature
even if further afforestation of heathland is prevented.

Apart from the loss of populations of EF. ciliaris, the continuing fragmentation and increased

94 S. B. CHAPMAN AND R. J. ROSE

FiGureE 3. Change in population composition of Erica ciliaris sites in relation to heathland fires during the period
1973 to 1987 in south-eastern Dorset: @ site with a changed hybrid status, unchanged sites are marked X; burnt
areas are shown LU.

isolation of the remaining heathlands (Chapman et al. 1989) is likely to have an important effect on
future spread, and on colonisation, of suitable sites. Colonisation and establishment of fragmented
or isolated areas will be more dependent on chance introductions than natural dispersal.

The results obtained by Chapman (1975) showed a number of significant associations between
particular population types and the dominant vegetation types present in the 200 m X 200 m grid
squares. However, within a period of 14 years the loss of sites and changes due to afforestation have
produced a situation where the examination of data and the testing of a hypothesis relating
particular types of population and habitat are no longer possible. Such a situation demonstrates the
need for detailed recording of information regarding less common species especially where they are
subject to loss of habitat or changes in land use.

Mowing and burning appeared to promote the establishment of E. X watsonii, whereas more
severe disturbance, such as ploughing on Soussons Down in Devon, and in the Purbeck Forest,
encouraged the establishment of E. ciliaris and E. tetralix. This may be related to the more vigorous
vegetative regrowth of E. X watsonii after mowing or burning, and regrowth of E. ciliaris and E.
tetralix from seed following ploughing.

In Dorset, E. ciliaris occurs in areas of peat and wet heathland and does less well in drier
heathland communities. However, in Cornwall and elsewhere in Europe the plant favours dry
heathland. More comparative work is needed on the requirements and behaviour of the plant under
different edaphic and climatic conditions.

Although E. ciliaris may be expanding its range in Dorset, concern must be expressed about the
decline in the area the plant occupies. While there was little change in the distribution of E. ciliaris
between 1963 and 1973, there was a marked loss of populations in the period 1973 to 1987.
Furthermore many of the populations that have been lost were predominantly E. < watsonii. The

DISTRIBUTION OF ERICA CILIARIS AND E. x WATSONII 95

fragmentation of the heaths and changes in population composition make it difficult to predict the
natural spread of the plant over its potential range in south-eastern Dorset.

ACKNOWLEDGMENTS

We thank English Nature and the individual landowners who have allowed access to their property
over the period of this study. Thanks are also due to Dr L. E. Haskins for site records and
permission to quote from her Ph.D. thesis, and BP Petroleum Development Ltd for funding of a
survey, the results of which provided the impetus for the remaining survey that was completed in
1987.

REFERENCES

CHAPMAN, S. B. (1975). The distribution and composition of hybrid populations of Erica ciliaris L. and Erica
tetralix L. in Dorset. Journal of ecology 63: 809-824.

CHAPMAN, S. B., CLARKE, R. T. & Wess, N. R. (1989). The survey and assessment of heathland in Dorset,
England, for conservation. Biological conservation 47: 137-152.

Gay, P. A. (1957). Aspects of the phytogeography of some Lusitanian Ericaceae in the British Isles. Unpublished
Ph.D. thesis, University of London.

Gay, P. A. (1960). A new method for the comparison of populations that contain hybrids. New phytologist 59:
218-226.

Goon, R. (1948). A geographical handbook of the Dorset flora. Dorchester, Dorset Natural History and
Archaeological Society.

Haskins, L. E. (1978). The vegetation history of south-east Dorset. Unpublished Ph.D. thesis, University of
Southampton.

MANSEL-PLEYDELL, J. C. (1895). The Flora of Dorsetshire, 2nd ed. Dorchester.

McCuintTock, D. (1971). Recent developments in the knowledge of European Ericas. Botanische Jahrbiicher 90:
509-523.

PERRING, F. H. & FARRELL, L. (1983). British red data book 1, Vascular plants, 2nd ed. Nettleham.

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

Wesp, D. A. (1966). Erica ciliaris L. in Ireland. Proceedings of the Botanical Society of the British Isles 6: 221-
DDS.

Wess, N. R. (1990). Changes on the heathlands of Dorset, England, between 1978 and 1987. Biological
conservation 51: 273-286.

Wess, N. R. & Haskins, L. E. (1980). An ecological survey of heathlands in the Poole basin, Dorset, England,
‘in 1978. Biological conservation 17: 281-96.

(Accepted October 1993)

Watsonia 20: 97-103 (1994) 97

A comparison of the growth and morphology of native and
commercially obtained continental European Crataegus
monogyna Jacq. (Hawthorn) at an upland site

A. T. JONES and P. R. EVANS

A.F.R.C. Institute of Grassland and Environmental Research, Plas Gogerddan, Aberystwyth,
Dyfed, SY23 3EB

ABSTRACT

1. The growth and morphology of native Crataegus monogyna Jacq. (Rosaceae) obtained from an upland
population in mid- Wales was compared with that of commercially obtained material of Hungarian provenance in
an upland trial.

2. Six months after planting, the native plants were 35% taller, 70% more branched and had twice the total stem
length and four times as many thorns per thorny plant than the commercial material. 91% of native plants were
thorny compared with only 20% of commercial plants. Commercial plants had larger leaves, longer petioles and
a greater severity of powdery mildew attack than the natives.

3. It was possible, using vegetative characters, to separate 88% of the natives from 80% of the commercial plants
on the basis of their growth and morphology using discriminant analysis. Native seed was significantly smaller in
size than the commercial seed which suggests that the measurement of reproductive characters would allow
further discrimination between the two groups.

KEYWORDS: invasive species, amenity plantings, alien species, hedges.

INTRODUCTION

Crataegus monogyna Jacq. (Hawthorn) is the most commonly planted shrub species in new hedges
and in the repair of derelict hedges in Britain (Brooks 1980). For the purposes of hedge-planting,
whips are bought from nurseries where they are propagated from seed. There is circumstantial
evidence that for commercial purposes seed has been obtained in many cases and from at least the
early 1970s from the continent and especially eastern Europe (Dunball 1982). This is motivated by
costs as foreign seed is often less than half the price of British collected seed.

Continental C. monogyna (henceforth referred to as alien) is a common constituent of hedges
which have been planted along new roads and motorways and following recent road-widening
operations. It can be identified by its habit of early bud-burst, often in early February when it
contrasts with the unopened buds of native C. monogyna in neighbouring old hedges. This may
indicate a phenology in the aliens which is poorly adapted to the British oceanic climate with its mild
winters and a lack of rapid and reliable temperature increases more typical of spring in a continental
climate. C. monogyna as a species has low bud dormancy (Murray et al. 1989) and is responsive to
spring temperature increases. It is possible that natives have a higher dormancy than the aliens.

There is prima facie evidence that the use of alien C. monogyna material is inappropriate in.
British upland or exposed sites as it shows poor growth, is swamped by grasses and is susceptible to
dieback both by wind scorching and powdery mildew. In contrast, it may be expected that native
upland populations of C. monogyna would have a high degree of adaptation to exposure, poor soils
and possibly grazing. A comparison of the growth and morphology of native and continental
material at a worst-scenario upland site might identify whether locally obtained material is more
suitable for upland planting than continental material.

98 A. T. JONES AND P. R. EVANS
MATERIALS AND METHODS

In November 1989, seed collections were made from a hawthorn population in mid-Wales that is
unlikely to have been planted in recent decades because of its remote location, natural appearance
and great age. This population forms a woodland scrub at an altitude of 350 m on the southern side
of Cader Idris (SH/746.713), the second highest mountain in mid-Wales. The site is extremely
exposed, as the valley runs to the coast in a south-westerly direction, the most frequent wind
direction. Near the population the valley narrows and wind is funnelled through a pass to the east.
Part of the population covers large block scree derived from a neighbouring cliff, which may have
given some protection from grazing. The hawthorn trees at the site differ in size and probably age,
some of the multi-stemmed trees appearing very old. Good et al. (1990) working in upland North
Wales found the age of single-stemmed hawthorn bushes to be from ten to 115 years and also found
suckering in some individuals. The trees at the mid-Wales site would be expected from their size to
be at least as old as the oldest at the former site although suckering was not apparent.

Native seed was cleaned of fleshy covering by fermenting in water at room temperature. It was
then placed in seed-trays at a depth of 1 cm in a 50:50 mixture of grit and compost outdoors. Alien
seed that had been chitted (treated with concentrated sulphuric acid for 30 minutes, thoroughly
rinsed in water and then mixed with compost and kept in polythene bags at 5°C for three months in
order to break dormancy) was obtained from Forestart Ltd, Hadnall, Gloucester, who stated that it
was of Hungarian provenance.

Samples of clean native seed (n = 121) and chitted alien seed (n = 25) were measured for length
and width with calipers. Following germination in March 1991, seedlings were pricked out into
individual cells of compartment seed trays. In July the plants were potted into 9 cm diameter pots
containing John Innes compost no. 2. In March 1992 the material was planted at an exposed site at
Pwllpeiran Experimental Husbandry Farm at an altitude of 330 m in well-drained soil. The site was
on a small level area on a steep (30°) south-facing slope.

A total of 192 plants, comprising 63 aliens and 129 natives, were planted at a spacing of 0.5 m
between seedlings in a rectangular plot of cultivated ground measuring 7 m by 10.5 m. In order to
minimise the effect of environmental variation on the inter-group comparison, the plot was divided
into four blocks within each of which 15-16 alien seedlings and 32-33 native seedlings were planted
in completely random order. Seedlings were planted through a landscape fabric mulch (Tensar)
which suppresses weeds but is permeable to rain. .

Measurements of morphology, growth and powdery mildew score (Table 1) were made after
shoot extension had finished in August 1992. The mildew infecting the plants is hawthorn powdery
mildew (Podosphaera clandestina Lev.), a common disease causing dieback of the growing points
which can reach epidemic status (Khairi & Preece 1978).

TABLE 1. MORPHOLOGICAL CHARACTERS, GROWTH AND DISEASE OF CRATAEGUS
MONOGYNA SEEDLINGS STUDIED
(all measurements were made to the nearest mm)

. Height
. No. of branches
. Total branch length including main stem
. Stem diameter (single measurement at base)
. Petiole length (mean of three petioles)
. Leaf width (mean of three leaves)
. Leaf length (mean of three leaves)
. Bract length (mean of three bracts which are situated at the base of the petiole)
. Total no. of thorns
. Mildew incidence score:
1) no disease,
2) < three mature leaves with infection (silver and/or red blotches),
3) mature leaves with or without infection and with at least two immature leaves and stem apex infected,
4) as 3 above but with some length (> 1 cm) of stem affected.

SToOMOANANMNFWNr

—

N.B. Characters 5, 6, 7 and 8 above were sampled on the main stem at as near as possible to half-plant height.

THE GROWTH OF NATIVE AND NON-NATIVE HAWTHORN 99

Data were analysed using the multivariate statistical technique, discriminant analysis (DA)
available in the Genstat statistical package (Genstat 5 Committee 1987; Digby 1989). DA is a
statistical technique based on canonical variate analysis (Mardia et al. 1979) and finds the best linear
combination of variables that discriminate between prior groupings. The resulting discriminant
function has the greatest variable between-group variation relative to within-group variation. DA
can be used to examine the validity of an a priori classification or to allocate new members to an
existing classification. DA has been used in taxonomy, particularly at the intraspecific level, e.g. to
examine the taxonomic groupings within Hordeum vulgare L. (Baum & Bailey 1983) and
hybridisation between native and alien cultivar forms of Lotus corniculatus L. (Bonnemaison &
Jones 1986). Using DA it is possible to determine how morphologically distinct the native and alien
hawthorn groups are from each other. The analysis also determines which characters make major
contributions to any discrimination between the two groups.

RESULTS

Alien seed was significantly larger (ANOVA, p < 0.001) than native seed with mean dimensions for
width 5-0 mm and length of 6-4 mm compared with 4-4 mm and 5-5 mm for native seed. There was
no significant difference in the shape of seeds (ratio of length to width) between the two groups.

The native and alien groups exhibited different morphology and growth characteristics (Table 2)
at the upland site. The native plants showed overall greater growth than the alien plants. Native
plants were on average 35% taller, had twice the total stem length and 70% more branches than the
alien plants. Mean stem diameter was also greater in native plants. The alien plants had larger leaves
and longer petioles than the native plants but bract size was similar in both groups.

Of the native plants, 91% were thorny compared with only 20% of alien plants. Native plants also
had a greater number of thorns per thorny plant so that mean thorn density per unit length of stem
was approximately three times greater than in the aliens. The mean powdery mildew score for the
aliens indicates a higher incidence of disease of the stem apex than the natives.

Some morphological characters were highly and significantly positively correlated (Table 3): leaf
width with leaf length (indicating a uniform leaf outline shape), leaf width with petiole length, and
leaf length with petiole length. Some growth characters were also significantly and positively
correlated: stem diameter and total stem length, stem diameter and branch number, and branch
number and total stem length. The regular spacing of thorns, where they occurred, was indicated by
the high correlation of thorn number per plant with total stem length. Powdery mildew score was
significantly positively correlated with petiole length.

In the discriminant analysis the natives and aliens were introduced as separate groups. This

TABLE 2. MEAN VALUES AND THEIR STANDARD DEVIATIONS FOR MORPHOLOGICAL
CHARACTERS AND GROWTH PARAMETERS OF NATIVE (N = 129) AND ALIEN (N = 63)
MATERIAL OF CRATAEGUS MONOGYNA
(all measurements in mm)

Native Alien
mean s.d. mean s.d
Height 107-0 37-5 79-3 34-0
No. of branches 3-1 1-7 1-8 1-4
Total stem length 235-6 134-7 129-8 84-0
Stem diameter 4-2 1-4 3-6 1-5
Petiole length 4-3 17) 6:3 3-1
Leaf width ewer 3-5 15:3 5:1
Leaf length 15-4 3-9 19-5 6-7
Bract length 3-4 1-3 35 1-8
No. of thorns 10-6 13-1 0-6 {5
Thorn number + 12-4 13-3 3-0 1-8
Mildew score 1-5 0-8 2-0 1-1

N.B. Thorn number + indicates the mean of only the thorny plants; s.d. = standard deviation.

100 A. T. JONES AND P. R. EVANS

TABLE 3. CORRELATION COEFFICIENTS BETWEEN VEGETATIVE CHARACTERS OF
CRATAEGUS MONOGYNA

Height (ht)

No. of branches (n.b.) 0-377

%* KK

Stem diameter (s.d.) 0-416 0-381

se se
Petiole length (p.1.) 0-139 0-003 0-168

Leaf width (I.w.) 0-135 0-067 0-275 0: 706

Leaf length (1.1.) 0-132 0-068 0218 0-719 0893

Bract length (b.1.) 0-306 0-147 0-264 0-386 bee ee

No. of thorns (n.t.) 0-312 0-425 0-387, 5) —0-162),4—0-037,, -—0-0745 50-071

* KK KX

Mildew score (m.s.) 0-083 —0-004 —0-182 0-194 0-113 0-186 0-017 —0-075

Total stem length 0-609 0-656 0-522 0-015 0-148 01295 0-235 0-819 0-001
; * KK * KX * KX * * KX
ht n.b. s.d. p.l. i.w. Ld. b.1. nee m.s.

N.B. significance levels for the correlations coefficients are as follows: * = p < 0.05; ** =p <0.01; ***=ps<
0.001.

analysis reclassified 88% of the natives into the initial native group and 80% of the alien plants into
the initial alien group. The natives and aliens are, therefore, reasonably distinct in terms of their
growth and morphological characteristics, but there was an overall 15% overlap of the two groups
(Fig. 1), i.e. 12% of the natives and 20% of the aliens were classified incorrectly. Some of this
overlap might have been due to the effect of the harsh environment on growth and establishment.
Most of the alien plants classified by the analysis as natives had small leaves and short petioles
compared with those classified correctly as alien plants.

Those characters which are positively correlated with the discriminant function (Table 4), i.e.
high values correlated with the native group are, in decreasing order of importance, number of
thorns per plant, total stem length and branch number per plant. Characters that are negatively
correlated with the discriminant function, 1.e. high values correlated with the alien group are, in
decreasing order of importance petiole length, leaf width, leaf length and mildew score.

DISCUSSION

In this study, the native Crataegus monogyna plants were discriminated from the aliens on the basis
of higher stem length, a higher number of branches, higher thorn number and lower powdery
mildew score. It would be expected that the relatively high mildew score and low growth rates in the
aliens would be associated with low establishment rates following hedge-planting. If sufficient
growth is not made, especially as a result of dieback due to mildew, then there is often the danger of
planted hawthorn quickly becoming overgrown and shaded by grasses in a fenced situation.

The above would suggest that the natives are more suited to hedge-planting in upland situations
than the alien material and because of their thorniness combined with a bushy habit, they may also
be more grazing tolerant where fencing is insufficient to provide protection from grazing animals.
The natives have smaller leaves and shorter petioles than the aliens and thus at nodes where thorns
are present there is less unguarded leaf to graze. Certainly, from the degree of variation present in
the natives, it would be possible to select for bushy, fast-growing, thorny plants, e.g. a plant which
had shown the most growth in the experiment also had 109 thorns at a density of one thorn every 9
mm of stem. From the significant differences’ in seed size of the two groups, the gathering of

THE GROWTH OF NATIVE AND NON-NATIVE HAWTHORN 101

50
U Aliens
he Z ZA Natives
g
pea y :
3 AV
Z 20 y) 77 Vv) -
VAVA A
a
ACY
: AZIZIZe
\AAAAr
0 -ESLES. ed fe Bey [8] Fe Ba Bd BS g y) y) A) ws
ee se a 8

Discriminant analysis scores

Figure 1. Bar chart of discriminant analysis scores for alien and native groups of Crataegus monogyna.

TABLE 4. CORRELATION BETWEEN DISCRIMINANT FUNCTION
SCORES AND CHARACTER SCORES FOR CRATAEGUS MONOGYNA

Character Correlation coefficient Probability
No. of thorns 0-594 oe
Total stem length 0-560 gis
No. of branches 0-535 oor
Height 0-494 cas
Stem diameter 0-269 Be
Petiole length —0-582 er
Leaf length —0-541 ie
Leaf width —0-440 i
Mildew score —0-362 oe
Bract length —0-083

N.B. Significance levels for correlation coefficients are as follows: ** = p< 0.01;
*** = p < 0.001.

102 A. T. JONES AND P. R. EVANS

reproductive data may enable a further separation of the native and alien material on morphological
grounds.

The escape and spread of exotic plant species following their movement by man across
geographical boundaries is a potential danger to many endemic vegetation types (Salisbury 1961).
Examples include the South American Lantana montevidensis (K. Spreng.) Briq. which is invading
natural vegetation across the tropics (Ridley 1930) and Rhododendron ponticum L. from Turkey
which is infiltrating oak woodland in the British Isles (Cross 1974). A much more insidious threat is
the spread of material which is of alien provenance though conspecific with or closely related to an
indigenous species and thus able to hybridise and potentially introgress. The planting of alien
Crataegus monogyna represents such a threat. Across Europe there are six recognised subspecies of
C. monogyna (do Amaral Franco 1968) and 22 species within the genus, at least four of which
commonly hybridise with C. monogyna. A hybrid of C. monogyna and C. laevigata (Poiret) DC.
(Midland Hawthorn) which naturally occurs in the south of England, Crataegus X media (Bradshaw
1971) and which may occur in imported seed, would probably be unsuitable for hedge planting. C.
laevigata is typically an understorey species of lowland woodland and would be expected to be
intolerant of exposed sites. There is always the possibility that importations of seed may contain
alien subspecies, hybrids and even other species of the Crataegus genus which are then sold as
common Hawthorn.

The natives sampled in this trial may represent an extreme locally adapted ecotype of Crataegus
monogyna subsp. nordica franco (do Amaral Franco 1968) but the taxonomy of this species has yet
to be completely resolved. However, as alien material continues to be planted in hedges from where
it is highly likely that bird dispersal will take place, further taxonomic investigation of the British
forms will be difficult to progress.

At upland sites in the British Isles, it can be argued that native C. monogyna should be planted for
two reasons, namely that it has a higher growth performance than the aliens and also that it is more
appropriate for conservation purposes to use native provenances as these represent a component of
local biodiversity. Grant awarding bodies should ensure that in hedge-planting, British provenance
is used at the very least.

ACKNOWLEDGMENTS

We would like to thank the Ministry of Agriculture, Fisheries and Food for funding this work as part
of a 3-year open contract and Dr Roger Haggar for his advice. We also acknowledge the help of
John Wildig at Pwllpeiran Experimental Husbandry Farm and Derek Jones (Institute of Grassland
and Environmental Research) in providing assistance with the selection and maintenance of a
suitable experimental site.

REFERENCES

Baum, B. R. & BaILey, L. G. (1983). Morphometric relationships in Hordeum vulgare (Triticeae, Poaceae). I.
H. spontaneum. Canadian journal of botany 61: 2015-2022.

BONNEMAISON, F. & Jones, D. A. (1986). Variation in alien Lotus corniculatus L. 1. Morphological differences
between alien and native British plants. Heredity 56: 129-138.

BrapsHaw, A. D. (1971). The significance of hawthorns, in Hedges and local history, pp. 20-29. Standing
conference for local history. National Council of Social Service, London.

Brooks, A. (1980). Hedging: a practical handbook. British Trust for Conservation Volunteers, Berkshire.

Cross, J. R. (1974). Rhododendron ponticum L., in Biological Flora of the British Isles. Journal of ecology 63:
345-364.

Dicsy, P. G. N. (1989). Procedure Discriminate, in PAYNE, R. W., LANE, P. W., AINSLEY, A. E., BICKNELL, K.
E., Dicsy, P. G. N., HARDING, S. A., LEECH, P. K., Simpson, H. R., Topp, A. D., VERRIER, P. J. & WHITE,
R. P. eds. Genstat procedure library manual. Release 1.3(2). Lawes Agricultural Trust, Oxford.

DO AMARAL FRANCO, J. (1968). Crataegus L., in TuTINn, T. G. et al., eds. Flora Europaea 2: 73-77. Cambridge.

DunsaLt, A. P. (1982). The management and planting of motorway verges, in PERRING, F. H. ed. The flora of a
changing Britain, pp. 84-89. London. 5

GENSTAT 5 COMMITTEE (1987). Genstat 5 reference manual. Oxford.

THE GROWTH OF NATIVE AND NON-NATIVE HAWTHORN 103

Goop, J. E. G., Bryant, R. & Car.iti, P. (1990). Distribution, longevity and survival of upland hawthorn
(Crataegus monogyna) scrub in North Wales in relation to sheep grazing. Journal of applied ecology 27:
272-283.

Kuairi, S. M. & PREEcE, T. F. (1978). Hawthorn powdery mildew: occurrence, survival and ascospore
productivity of Podosphaera clandestina cleistotheca in England. Transactions of the British Mycological
Society 71: 289-293.

Marpi, K. V., Kent, J. T. & Bippy, J. M. (1979). Multivariate analysis. New York.

Murray, M. B., CANNEL, M. G. R. & Situ, R. I. (1989). Date of budburst of fifteen tree species in Britain
following climatic warming. Journal of applied ecology 26: 693-700.

Rip ey, H. N. (1930). The dispersal of plants throughout the world. Ashford, Kent.

SaLisBurY, E. (1961). Weeds and aliens. Collins New Naturalist, London.

(Accepted September 1993)

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Watsonia 20: 105-114 (1994) 105

Bracken (Pteridium aquilinum (L.) Kuhn) infestation of rough
grazing land in the catchment of the River Tyne,
northern England

A. J. CHERRILL* and A. M. LANE+

Centre for Land Use and Water Resources, Department of Agricultural and Environmental
Science, University of Newcastle-upon-Tyne, Newcastle-upon-Tyne, NEI 7RU

ABSTRACT

The distribution of bracken (Pteridium aquilinum (L.) Kuhn)within the catchment of the River Tyne, Northern
England, was surveyed in the field by stratifying sampling effort within land classes of a national environmental
landscape classification developed by the Institute of Terrestrial Ecology (I.T.E.). A total of 182 1-km squares
were surveyed in 1991 (equivalent to 6.3% of the catchment). The areas of bracken and rough grazing land
differed significantly between land classes. Bracken was estimated to cover 0-7% of the total land surface, and
1-9% of rough grazing land, within the catchment. The results are compared with those of previous surveys of
bracken, and it is concluded that a standard survey method is needed if such comparisons are to be statistically
reliable. It is proposed that the I.T.E. land classes offer a suitable framework for the organisation of future
surveys.

KEyworbDs: ferns, agricultural weed, survey, landscape classification.

INTRODUCTION

Bracken, Pteridium aquilinum (L.) Kuhn (Dennstaedtiaceae), is generally regarded as a serious
weed of agriculturally marginal land throughout Britain (Smith & Taylor 1986; Taylor 1990). It is an
invasive species which thrives in a wide range of habitats (Grime et al. 1988). Worldwide it is said to
be the most widespread of all vascular plant species (Page 1982). The success of bracken lies in its
ability to withstand grazing, burning and chemical control measures through survival of its
rhizomes, and the ability of its spores to colonise readily disturbed habitats (Page 1976, 1982;
Kirkwood & Archibald 1986; Dyer 1990; Marrs et al. 1993; Pakeman & Marrs 1992).

Dense stands of bracken reduce the area of forage available for livestock. Its fronds are poisonous
to sheep and cattle (Evans et al. 1982; MacLeod 1982; Miller et al. 1990). Stands can also act as a
reservoir for disease-carrying sheep ticks. They make it difficult to herd and monitor the condition
of a flock. Similar problems occur on grouse moors where heather can be replaced by bracken
(Hudson 1986). Bracken may also be a source of carcinogens in supplies of water and milk destined
for human consumption (Taylor 1990).

The present paper describes the results of a sample-based field survey of the distribution of
bracken and other land cover types within the catchment of the River Tyne in northern England, an
area of almost 3,000 km?. An approach used in previous surveys of land cover has been to stratify
sampling efforts within zones (or strata) which are perceived to be relatively homogeneous in terms
of the variables to be recorded (e.g. Bunce et al. 1975). Extrapolation of the results from such
sample based surveys requires that the distribution of the sampling strata are well-defined (Cochran
1977). A number of studies have shown that the objective definition of strata can be achieved via

* Corresponding author.
+ Present address: Institute of Terrestrial Ecology, Merlewood Research Station, Cumbria, LA11 6JU.

106 A. J. CHERRILL AND A. M. LANE

numerical classification of physical attribute data derived from extant thematic databases, including
Ordnance Survey (O.S.), soil, geological and meteorological maps (Bunce et al. 1981a; Blankson &
Green 1991; Cooper & Murray 1992). Numerical classification techniques have included both
agglomerative clustering (e.g. Blankson & Green 1991) and divisive indicator species analysis
methods (e.g. Hill et al. 1975), but their applications have had the common aim of categorising grid
Squares into a number of groups. Squares which fall into each group are similar in terms of their
measured physical attributes, but different from the squares in other groups. These groups of
squares are termed ‘land classes’.

The land classification used in the stratification of sampling effort in the present study was that
developed by the Institute of Terrestrial Ecology (1.T.E.) (Bunce et al. 1981a, 1983). This
classification was selected because it has a resolution of 1 km (a suitable areal unit for field survey)
and because all 1-km squares within Britain have been assigned to one of 32 land classes. This
second feature enables the results of the survey to be placed in a national context (Bunce et al.
1981a).

A total of 182 1-km squares, stratified by I.T.E. land classes, were visited in 1991 and areas of
bracken were mapped at a scale of 1:10,000. This paper describes the distribution of bracken
between land classes, and provides an estimate of the degree of infestation of rough grazing in the
catchment.

THE I.T.E. LANDSCAPE CLASSIFICATION

The I.T.E. land classification uses the 1-km cells of the O.S. national grid as its spatial unit. Grid
squares are grouped into 32 land classes on the basis of a wide range of map attribute data. The
development of the classification was in two phases (Bunce ef al. 1981a, 1991).

Initially, the land classes were identified from a classification of a nationally distributed sample of
1,228 1-km squares, each of which was situated at the intersection of a national 15 km X 15 km grid.
For each sample square, 282 attributes were recorded from existing maps. The attributes included
environmental variables (such as altitude, slope, geology, drift and climate), semi-natural land
cover features (such woodlands and lakes) and anthropogenic features (such as roads and
buildings). The sources of these data included 1:50,000 O.S. maps, 1:1,000,000 climatological maps
and geological maps. On the basis of these data, the sample squares were classified using Indicator
Species Analysis (I.S.A.) (Hill et al. 1975). The method is a divisive polythetic method of numerical
classification which produces a hierarchical, dichotomous key. At each division in the key,
‘indicator species’ are identified. This feature enabled grid squares not included in the original
I.S.A. classification procedure to be allocated to a land class on the basis of their attributes.
Descriptions of the 32 land classes are provided by Bunce ef al. (1981a) and Benefield & Bunce
(1982).

The second phase of development in the I.T.E. land classification system was the assignment of all
230,000 grid squares in Britain to the 32 land classes. In theory this could have simply involved use of
the extant key derived from the Indicator Species Analysis. Collation of the necessary ‘indicator
species’ data for every one of the grid squares in Britain, however, was logistically impractical. A
reduced set of some 60 attributes was more readily available for all grid squares. These attributes
came under eight broad headings; physiography, coastal features, climate, geology, geological drift,
land cover/use, offshore island status and distance from coast. These data were used to create a new
classification of squares. Correspondence between the two classifications was assessed by comparing
the ‘new’ and ‘old’ land classes of the original sample of 1,228 grid squares. A range of numerical
classifications were tested. That giving the closest agreement between the ‘new’ and the ‘old’ land
classes involved application of both multivariate discriminant analysis and logistic regression
algorithms (Bunce et al. 1991). This hybrid approach was therefore used to assign all 230,000 grid
squares in Britain to one of 32 land classes.

The hierarchical nature of the I.T.E. land classification allows land classes to be aggregated into
broad landscape types. In the present study, the land classes are categorised under three types of
landscape, namely: Lowland, Marginal Upland and Upland (after Bunce 1992).

BRACKEN INFESTATION IN NORTHERN ENGLAND 107

FicurE 1. The location of the catchment of the River Tyne (shaded).

THE STUDY AREA

The study area was the catchment of the River Tyne in northern England (Fig. 1). The boundary of
the catchment was supplied by the Institute of Hydrology (Wallingford). It enclosed an area of 2,903
km’, comprising 16 I.T.E. land classes. The distributions of the I.T.E. land classes within the
catchment were provided by the Institute of Terrestrial Ecology (Merlewood) and are shown in Fig.
2. The composition of the catchment in terms of the proportions of I.T.E. land classes is shown in
Table 1. In addition, the table shows the mean altitudes and main land forms within each I.T.E. land
class. In the west the catchment is characterised by two upland blocks, separated by the River Tyne-
River Solway gap. In the east, lowland landscapes predominate.

METHODS

SELECTION OF GRID SQUARES FOR FIELD SURVEY

The field survey was intended to obtain land cover data from a representative sample of the grid
squares within the catchment. There were insufficient resources to survey a representative sample of
all 16 land classes in a single summer. The five least frequent land classes, which together
represented 0-3% of the total area of the catchment (Table 1), were therefore omitted. In total, 182 —
squares were selected for survey.

Sample squares were selected from each of the eleven most frequent I.T.E. land classes in
approximate proportion to their total areas within the catchment (Table 1). Selection of squares
from within ten of these land classes was at random. Within the most frequent I.T.E. land class (i.e.
land class 22), however, sampling was further stratified. Equal numbers of squares of land class 22
were selected at random from within each of the upland blocks north and south of O.S. Northing

108 A. J. CHERRILL AND A. M. LANE

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FiGureE 2. The distribution of I.T.E. land classes within the River Tyne catchment: land class 9— a, 10—c, 13-e,
14-f, 15—k, 17-m, 18-n, 19-0, 20-r, 21-s, 22—t, 23-u, 25 -v, 26-—w, 27 -x, 28 —z. Figures at margins are
Ordnance Survey coordinates.

564000 (Fig. 2). This was done to allow a more detailed analysis of geographical variation in land
cover within I.T.E. land class 22, which represents 43-2% of the total area of the catchment.

FIELD SURVEY METHODS

Field work was undertaken between mid-April and October 1991. Each of the survey squares was
visited and the areas of the land cover types defined by the Nature Conservancy Council (N.C.C.
1990) were mapped on to 1:10,000 O.S. maps. Only 20 ha out of the total survey area of 18,200 ha
remained unsurveyed due to problems of access. Bracken was mapped as a distinct land cover type
where its canopy extended over greater than 25% of the land surface. Dense stands were readily
delineated, while the mapping of sparse bracken was more subjective.

PROCESSING AND ANALYSIS OF FIELD DATA

The 182 field survey maps were digitised using the Geographical Information System (G.I.S.) ARC/
INFO. The total areas of each of the land cover types within each of the survey squares were
calculated from this digital database. The areas of rough grazing were calculated by aggregating the
areas of the following cover types: heath, bog, moorland, unimproved grassland and land
dominated by bracken. Certain types of unimproved grassland which were not used for livestock
grazing (e.g. sand-dunes and maritime grasslands) were excluded, as were small areas of bracken on
derelict land in industrial areas.

109

BRACKEN INFESTATION IN NORTHERN ENGLAND

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110 A. J. CHERRILL AND A. M. LANE

The mean areas of rough grazing and bracken were calculated for each land class. Estimation of
the total areas of rough grazing and bracken within the catchment as a whole, and within lowland,
marginal upland and upland landscapes, followed Cochran (1977), as did estimation of the standard
deviations for these figures. Estimation of mean percentage infestation levels of bracken within land
classes used the ratios of the means for bracken and rough grazing. Estimation of the variances for
mean percentage infestation levels used the approximation given by Colquhoun (1971: p. 41). The
same approach was used in the estimation of percentage infestation levels (and associated variance
estimates) at the landscape and catchment scales.

Statistical comparison of the areas of bracken and rough grazing between land classes used
Duncan’s multiple range test within the SAS package (SAS 1989). Prior to these analyses the data
were transformed by adding 1-0 to all observations and conversion to log;). Comparison of the mean
infestation levels between land classes used Student’s two sample t-test for samples of unequal
variance (Zar 1984). Bonferroni’s correction for multiple comparisons was used to minimise the
chance of Type I errors (Miller 1977).

RESULTS

LAND COVER WITHIN THE I.T.E. LAND CLASSES
The land cover composition of each of the I.T.E. land classes is summarised in Table 2. The lowland
land classes are dominated by arable crops, sown grassland, woodland and urban development,
although the areas of these cover types varied between the land classes. The areas of rough grazing
land were also variable, but overall this cover type was a minor component of the lowland
landscapes. Rough grazing was most frequently encountered in the form of unimproved grassland.
The marginal upland land classes had markedly smaller areas of arable crops and urban
development than the lowland land classes, and were dominated by sown grassland. Woodland and
rough grazing were consistent features of these land classes. Rough grazing was composed primarily
of unimproved grassland, but areas of heathland and bog were also important (Table 2).

TABLE 2. ESTIMATED MEAN AREAS OF (a) MAJOR LAND COVER TYPES (EXCLUDING ROUGH
GRAZING), AND (b) COVER TYPES COMPRISING ROUGH GRAZING, WITHIN EACH I.T.E.
LAND CLASS OF THE RIVER TYNE CATCHMENT

Land Classes

Lowland * Marginal Upland Upland

10 13 14 25 26 27 119 = 20) S28 22N Gee sms

(a) Land uses (excluding rough grazing):

arable 31-2, 17-6 17-7 12-9 4-7 15:3 257) G2
sown grassland! 28:6 14-1 17-7 45-3 11-3 22-1 61-6 (30-2472 2-082 ee
woodland 91, 6:2 1-7. 12-1 3-5 16:3 11-1 8-910 040- O
urban 26-7 47-7 49:9| 9-8 75-3 27-8 3:2 “2-15 3-6) 50-Sieat ome
others 371 12-6 9-6 1:8 3-1. 3-8 70:5 20-7 “G4 SUS meeley
(b) Components of rough grazing:
unimproved grassland 0-9" 1-8 "93-47 11-8 2-1 9-2 A 19 GONG Oa een
marshy grassland 0-4 Ft 256° = 13) 4550-9 88 S-Oee mee
dry heath/grassland mosaic SS RS ET SE ee Oa) 3s (One neo eC
wet heath/grassland mosaic ia ee eee KD SS
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wet heath =. = fee.01. =) 2:32, ORS OO memos
bog and flush Se ete el 2 =, 10-8, 195 Aa el meena
bracken — i. + =. 0:33. .— 10:5 0:8) 2-00 eno

‘Including sown grasslands infested with Juncus effusus L.

* Including allotments and recreation areas.

322N and 22S refer to squares of land class 22 in the North and South of the catchment.
+ = <0.1%, — = 0.0%.

|

BRACKEN INFESTATION IN NORTHERN ENGLAND ie

The upland land classes were dominated by rough grazing, with limited areas of arable crops and
urban development. In the north, squares of land class 22 had large areas of woodland, whilst in the
southern squares of this land class sown grassland was more important. Heathland, bog and
unimproved grassland were important components of rough grazing land. Land class 23 was notable
for the dominance of bog and unimproved grassland (Table 2).

DISTRIBUTION OF BRACKEN

Bracken was recorded in 10-9% of lowland sample squares, 25-0% of marginal upland squares and
22-0% of upland squares, but contributed a mean area greater than 1% of land cover per square in
only two land classes (Table 3). The mean areas of bracken differed significantly (p < 0.05) between
certain land classes (Table 3). However, although bracken was encountered in a higher proportion
of sample squares in the marginal uplands and uplands, there was no clear altitudinal pattern in the
variation of the mean areas of bracken by land class (Tables 3 & 4). The mean areas of rough
grazing, in contrast, showed a clear increase from the lowlands to the uplands (Tables 3 & 4). Mean
infestation of rough grazing land by bracken ranged from 0-:0% to 4-23%, but did not differ
significantly between any pair of land classes (Table 3).

TABLE 3. MEAN AREAS OF ROUGH GRAZING LAND AND BRACKEN IN SAMPLE SQUARES OF
EACH LAND CLASS IN THE RIVER TYNE CATCHMENT AREA

Rough grazing Bracken
(ha km”) (ha km~”) % Infestation
Land =>... - < _-SEAva
Landscape Class x s.d. x s.d. x s.d.
Lowland 10 1-275 1-21 0-08 ~ 0-0" -
13 1-82¢ BE) 0-02° 0-06 1-10" 3°59
14 3-454 655 0-08 - 0-0" —
5) 18-09°°¢ 29° 7 0-25%8 0-58 1-38" 3-90
26 2-169 3-04 0-08 - 0-0" -
Di, 14-19°¢ 27-46 0-54'2 2-00 3-81> 15-92
Marginal 19 20-822? 20-04 0-848 2-09 4-03" 10-75
20 48-207? 39-37 2-04° 4-17 4-23" 9-3
28 27-607” 31-44 0-07%8 0-21 0-25" 0-80
Upland 22N See? 38-51 1-36°! 2-93 Dee 5-74
22S [2-22 33-81 0-82'8 2:59 1-14" 3-64
a3) O-927 4-90 0-08 - 0-0" -

X = mean; s.d. = standard deviation.
Means labelled with the same letter are not significantly different at p < 0.05.

TABLE 4. ESTIMATED TOTAL AREAS OF ROUGH GRAZING LAND AND BRACKEN, AND
MEAN PERCENTAGE INFESTATION OF ROUGH GRAZING LAND, IN LOWLAND,
MARGINAL UPLAND AND UPLAND LANDSCAPES, AND IN THE WHOLE CATCHMENT OF
THE RIVER TYNE

Rough grazing Bracken % Infestation
Total area ——-AAPS sq _ Ss

Landscape’ km? ha s.d. ha s.d. x s.d.
Lowland 1202 lin 3804 333 141 OS e095
Marginal upland 359 10405 1635 205 89 OT 0-91
Upland 1327 81530 1952 1500 606 1:84 0-76
Catchment 2888 109046 8966 2038 628 1-87 0-60

1 Land classes 9, 15, 17, 18 and 21 excluded.
kX = mean; s.d. = standard deviation.

112 A. J. CHERRILL AND A. M. LANE

Extrapc ition from the mean areas of bracken and rough grazing in the land classes indicates that
bracken a ‘ounted for around 1-9% of rough grazing land in each of the lowland, marginal upland
and uplana landscapes (Table 4). The estimated total area of bracken, 2,038 ha, was equivalent to
0-7% (s.d. + 0:2%) of the land surface of the catchment (Table 4).

DISCUSSION

A number of surveys of bracken distribution have been conducted in Britain, but each has used a
different approach or has focused on a limited geographical region (Lawson et al. 1986; Miller et al.
1990). This may, in part, explain variation in the estimates of the total extent of bracken in Britain,
which range from 1-2% to 2:7% of the land surface (Lawson et al. 1986; Taylor 1986). The only
study which is directly comparable to that presented here is that of Bunce ef al. (1981a). In 1978,
these authors mapped land cover (including bracken) in a nationally distributed sample of 1-km
squares of each of the 32 I.T.E. land classes. They estimated that bracken covered 316,703 ha
(1-3%) of the land surface of Britain. Comparison with the present study suggests that bracken is
less common in the catchment of the R. Tyne than elsewhere.

Comparison of the results of the present study with those of others conducted in north-east
England is difficult. The Countryside Commission (1991) used a visual interpretation of aerial
photography to estimate that 4-1% of the Northumberland National Park (which impinges upon the
catchment of the R. Tyne) is covered by bracken. Varvarigos & Lawton (1991) conducted a postal
survey of farmers, and estimated that 14-7% of rough grazing land within the Less Favoured Areas
(L.F.A.) of Northumberland and Durham supported bracken. The L.F.A.s in the catchment of the
R. Tyne lie predominantly within land classes 19, 20, 22, 23, 25, 27 and 28. Conversely, 90% of
squares of these land classes possess L.F.A. status in the study area (unpublished). Although the
catchment does not include all L.F.A. land within either County Durham or Northumberland, the
present study suggests that Varvarigos & Lawton (1991) overestimated the extent of bracken
infestation. Only around half of farmers targeted in their survey returned questionnaires, but
whether the probability of farmers returning the questionnaires was related to the extent of bracken
infestation was not addressed. It is possible that farmers with severe infestations would have been
more likely to respond. This interpretation is supported by preliminary analyses of LANDSAT
data, which suggest that bracken covers around 1-5% of the catchment of the R. Tyne (Cherrill et al.
in press).

Use of the I.T.E. land classification system in field surveys enables estimates of land cover to be
viewed in the context of broad patterns of environmental variation summarised by the land classes
(Tables 1 & 2) (Bunce et al. 1981a, 1981b, 1983; Benefield & Bunce 1982). Thus, for example, the
absence of bracken from land class 23 can be attributed to the effects of altitude and the
predominance of waterlogged mires in this land class. Bracken is rarely found above altitudes of 600
m and is known to prefer well drained soils (Thompson et al. 1986; Grime et al. 1988; Ader 1990). In
this context, it is relevant to note that although the nature of the vegetation beneath bracken was not
investigated, bracken stands were most frequently associated with areas of dry heath and
unimproved grassland (Table 2). It is these ecologically interesting cover types which are likely to be
at greatest risk of colonisation and which have probably lost most ground to bracken within the
study area.

In conclusion, the present study has followed others in being restricted to a limited geographical
region. However, it differs from many of its predecessors in one major respect, namely the use of a
national landscape classification system in the stratification of sampling effort. The use of a diversity
of methods in previous studies has led to difficulties in the collation of national statistics and a failure
to produce consistent regional estimates of the extent of bracken. The need for a standardised
method of field survey is clear. Use of the I.T.E. land classes allows the extrapolation of sample
based survey data to give regional estimates of land cover. In the future, use of the I.T.E. land
classification system would have the additional advantage of permitting comparisons between
temporally and spatially separated field surveys. Although remote sensing offers a potential solution
to the standardised collection of land cover data (e.g. Miller et al. 1990), its capabilities are often
overstated and there will be a continuing need for ‘ground-truth’ data (Price 1986; Townshend
1992). The I.T.E. land class system has recently been used as a framework for the stratification of

BRACKEN INFESTATION IN NORTHERN ENGLAND 1S:

sampling effort in field surveys designed to aid the interpretation of satellite data (Cherrill e¢ al. in
press). The classification, therefore, offers a framework for the organisation of traditional field
surveys, but also provides a link with modern remote-sensing technologies (Bunce ef al. 1992).

ACKNOWLEDGMENTS

Miss A. Marler and Mr J. Dargie did much of the field work. Thanks are also due to the I.T.E. Land
Use Research Group, the Institute of Hydrology, Dr R. Sanderson, Dr E. Martin and the many
people who allowed access to land in their care. The work was funded jointly by the Natural
Environment Research Council and the Economic and Social Research Council.

REFERENCES

Aber, K. G. (1990). The effect of climate and soils on bracken vigour and distribution in Scotland, in THomson,
J. A. & Situ, R. T., eds. Bracken biology and management, pp. 141-151. Sydney.

BENEFIELD, C. B. & BUNCE, R. G. H. (1982). A preliminary visual presentation of land classes in Britain.
Merlewood Research and Development Paper No. 91. Grange-over-Sands, Cumbria.

BLANKSON, E. J. & GREEN, B. H. (1991). Use of landscape classification as an essential prerequisite to landscape
evaluation. Landscape and urban planning 21: 149-162.

Bunce, R. G. H. (1992). The distribution and aggregation of I.T.E. land classes. Report to the Department of
the Environment and the Natural Environment Research Council. Grange-over-Sands, Cumbria.

Bunce, R. G. H., Barr, C. J. & Futter, R. M. (1992). Integration of methods for detecting land use change,
with special reference to Countryside Survey 1990, in Wuirsy, M., ed. Land use change: the causes and
consequences, pp. 69-78. London.

Bunce, R. G. H., Barr, C. J. & Wuitraker, H. (1981a). An integrated system of land classification. Annual
report of the Institute of Terrestrial Ecology, pp. 28-33.

Bunce, R. G. H., Barr, C. J. & Wuirraker, H. (1981b). Land classes in Great Britain: preliminary descriptions
for users of the Merlewood method of land classification. Merlewood Research and Development Paper No.
86. Grange-over-Sands, Cumbria.

Bunce, R. G. H., Barr, C. J. & Wuitraker, H. (1983). A stratification system for ecological sampling, in
FULLER, R. M., ed. Ecological mapping from ground, air and space, pp. 39-46. Huntingdon.

Bunce, R. G. H., Lane, A. M. J.,. Howarp, D. C. & Crarkg, R. T. (1991)... 1.T.E. land classification:
classification of all 1 km squares in Great Britain. Report to the Department of the Environment. Grange-
over-Sands, Cumbria.

Bunce, R. G. H., Morre1t, S. K. & STEEL, H. E. (1975). The application of multivariate analysis to regional
survey. Journal of environmental management 3: 151-165.

CHERRILL, A. J., LANE, A. M. & FULLER, R. M. (in press). The use of classified LANDSAT 5 Thematic Mapper
imagery in the characterisation of landscape composition: a case study in Northern England. Journal of
environmental management.

CocHRAN, W. G. (1977). Sampling techniques. New York.

Co.LquHoun, D. (1971). Lectures on biostatistics. Oxford.

Cooper, A. & Murray, R. (1992). A structured method of landscape assessment and countryside management.
Applied geography 12: 319-338.

COUNTRYSIDE CommIssION. (1991). Landscape change in the National Parks. Manchester.

Dyer, A. F. (1990). Does bracken spread by spores?, in THomson, J. A. & SmitH, R. T., eds. Bracken biology
and management, pp. 35—42. Sydney.

Evans, W. C., PATEL, M. C. & Koony, Y. (1982). Acute bracken poisoning in homogastric and ruminant
animals. Proceedings of the Royal Society of Edinburgh 81(B): 29-64.

Grime, J. P., Hopcson, J. G. & Hunt, R. (1988). Comparative plant ecology. London.

Hii1, M. O., Bunce, R. G. H. & SHAw, M. W. (1975). Indicator species analysis, a polythetic method of
classification, and its application to a survey of native pinewoods in Scotland. Journal of ecology 63: 597-
613.

Hupson, P. J. (1986). Bracken and ticks on grouse moors in the North of England, in Smit, R. T. & TAYLOR, J.
A., eds. Bracken. Ecology, land use and control technology, pp. 161-170. Carnforth.

Kirkwoop, R. C. & ARCHIBALD, L. (1986). The rhizome as a target site for the control of bracken using foliage-
applied herbicides, in SmirH, R. T. & Taytor, J. A., eds. Bracken. Ecology, land use and control
technology, pp. 341-350. Carnforth.

114 A. J. CHERRILL AND A. M. LANE

Lawson, G. J., CALLAGHAN, T. V. & Scott, R. (1986). Bracken as an energy resource, in SmitH, R. T. &
TayLor, J. A., eds. Bracken. Ecology, land use and control technology, pp. 239-248. Carnforth.

MacLeop, A. (1982). The bracken problem in the Highlands and Islands. Proceedings of the Royal Society of
Edinburgh 81(B): 11-18. .

Marrs, R. H., PAKEMAN, R. J. & Lowpay, J. E. (1993). Control of bracken and the restoration of heathland. V.
Effects of bracken control treatment on the rhizome and its relationship with frond performance. Journal of
applied ecology 30: 107-118.

Mixer, D. R., Morrice, J. G. & Wuitworty, P. L. (1990). Bracken distribution and spread in upland Scotland:
an assessment using digital mapping techniques, in THomson, J. A. & Smit, R. T., eds. Bracken biology
and management, pp. 121-132. Sydney.

MILLER, R. G. (1977). Developments in multiple comparisons. Journal of the American Statistical Association
72: 779-788.

NATURE CONSERVANCY CoUNCIL (1990). Handbook for Phase 1 habitat survey: a technique for environmental
audit. Peterborough.

Pace, C. N. (1976). The taxonomy and phytogeography of bracken — a review. Botanical journal of the Linnean
Society 73: 1-34.

PaGE, C. N. (1982). The history and spread of bracken in Britain. Proceedings of the Royal Society of Edinburgh
81(B): 3-10.

PAKEMAN, R. J. & Marrs, R. H. (1992). Vegetation development on moorland after control of Pteridium
aquilinum with asulam. Journal of vegetation science 3: 707-710.

Price, M. (1986). The analysis of vegetation change by remote sensing. Progress in physical geography 10: 473—
491.

SAS (1989). SAS/STAT User’s guide. Volume 2, Version 6. North Carolina.

SmitH, R. T. & TayLor, J. A., eds (1986). Bracken. Ecology, land use and control techniques. Carnforth.

TayLor, J. A. (1986). The bracken problem: a local hazard and global issue, in Smit, R. T. & Tay or; J. A.,
eds. Bracken. Ecology, land use and control techniques, pp. 21-42. Carnforth.

Taytor, J. A. (1990). The bracken problem: a global perspective, in THomson, J. A. & Situ, R. T., eds.
Bracken biology and management, pp. 3-19. Sydney.

THompson, T. R. E., RUDEFoRTH, C. C., HARTNuP, R., Lea, J. W. & Wricut, P. S. (1986). Slope and soil
conditions under bracken in Wales, in Smit, R. T. & TayLor, J. A., eds. Bracken. Ecology, land use and
control technology, pp. 101-108. Carnforth.

TOWNSHEND, J. R. G. (1992). Land cover. International journal of remote sensing 13: 1319-1328.

VARVARIGOS, P. & Lawton, J. H. (1991). Farmers’ perceptions of the scale of the bracken problem on farms in
Less Favoured Areas in England and Wales. Journal of applied ecology 28: 988-1003.

ZAR, J. H. (1984). Biostatistical analysis. New Jersey.

(Accepted December 1993)

Watsonia 20: 115-117 (1994) 115

The status of Bupleurum falcatum L. (Apiaceae) in the
British flora

M. H. FIELD

Sub-department of Quaternary Research, Botany School, University of Cambridge, Downing
Street, Cambridge, CB2 3EA

ABSTRACT

The umbellifer Bupleurum falcatum L. has only been recorded from one locality in the British Isles and its status
as a native in the modern British flora is uncertain. Recent analyses of fossiliferous Pleistocene temperate stage
sediments from two English sites have yielded mericarps of B. falcatum subsp. falcatum. These finds show that
this taxon was once naturally occurring on the British mainland and had a more northerly range than at present.
It is difficult to determine whether the one extant British population has a native status as a result of natural
dispersal from the continent during the Flandrian Stage, is a relic of a Late Pleistocene temperate stage
population that survived in Britain during the last cold stage (Devensian), or alternatively has an alien status
with a relatively recent introduction by human activity. Its occurrence in the Late Pleistocene fossil record may
support the opinion that the extant population is native.

Keyworps: Fossil record, native status, Pleistocene, Umbelliferae, Sickle-leaved Hare’s-Ear.

INTRODUCTION

The first British record of Bupleurum falcatum L. (Apiaceae) was made by Thomas Corder in 1831.
He described the plant as being abundant on either side of the turnpike between Chelmsford and
Ongar, Essex (Corder 1833). A number of authorities have debated whether or not this population
is indigenous to the British Isles. Gibson (1862) said it “appears truly indigenous”, whilst Jermyn
(1974) and Tutin (1980) have expressed reservations about its native status. In 1962 hedgerow
clearance and ditch cleaning destroyed the population in southern Essex (Jermyn 1974), but Stace
(1991) says it reappeared in 1979.

Palaeobotanical investigations of fossiliferous organic sediments from two gravel-pits at Frog
Hall, Warwickshire (SP/413.734) and Somersham, Cambridgeshire (TL/373.799) yielded mericarps
of B. falcatum subsp. falcatum. These finds show that this taxon was once naturally occurring in
Britain and add to the discussion about the status of this taxon in the British flora.

The macroscopic remains of B. falcatum subsp. falcatum came from assemblages that were
dominated by waterside, damp ground and aquatic taxa. It is probable that deposition at both sites
took place in a low energy, fluviatile environment or in a pond ona floodplain. The taxa represented
in both assemblages show that contemporary conditions were temperate and were probably similar
to those in southern Britain today.

The age of the temperate stage deposits at Frog Hall is controversial. The deposits stratigraph-
ically overlie the nearby type section of the cold Wolstonian Stage at Wolston. If the accepted
British stratigraphical sequence is applied (Mitchell et al. 1973) then these deposits are last
interglacial (Ipswichian) in age. However, the age of the Wolston cold stage sediments has been
disputed (Sumbler 1983). The Somersham deposits which yielded the mericarps of B. falcatum
subsp. falcatum have been correlated with Ipswichian pollen zone II (R. G. West, pers. comm.
1993).

116 M. H. FIELD
THE FOSSIL SPECIMEN

The best preserved mericarp was recovered from Frog Hall, but three flattened and less well
preserved mericarps were found at Somersham (R. G. West, pers. comm. 1993). It is possible to
determine the fossil specimen to subspecies level because the two recognised subspecies are
distinguished by the characteristics of the fruits (Tutin 1968). B. falcatum subsp. cernuum (Ten.)
Arcangeli can be identified by its relatively large fruits (c. 5 mm in length) which have winged ridges
running longitudinally. B. falcatum subsp. falcatum has a smaller fruit (c. 3 mm in length) and the
longitudinal ridges are usually not winged. Despite the variability of this species, noted by Tutin
(1968), there is no problem in the placement of the fossil specimens into the latter subspecies. Tutin
(1980) states that the modern British population belongs to subsp. falcatum also.

The fossil mericarp from Frog Hall is 2-9 mm long and has a maximum width of 1-1 mm (Fig. 1).
There are five prominent, slender, longitudinal ridges which are not winged. It is difficult to say
whether these ridges were winged or not, as delicate features are not always preserved. The
commissure is relatively broad and has a fairly deep longitudinal groove.

DISCUSSION

The occurrence of B. falcatum subsp. falcatum in British Pleistocene deposits is of botanical interest

because today this plant only occurs in one locality in the British Isles and the native status of this
population has been questioned. Both the fossil records are found north of the present day
occurrence at North Heath, Essex and show that this taxon was naturally occurring on the British
mainland in the Late Pleistocene.

There are three possible origins of the extant British population: it may be a remnant of a Late
Pleistocene temperate stage population, it may have been naturally dispersed from the continent, or
it may have been introduced by human activity.

If the modern population is a remnant of a Late Pleistocene temperate stage population it must

NOE, —_

<4

Spx
ota
is 4

ae ED

1mm

FicurE 1. The mericarp of Bupleurum falcatum subsp. falcatum recovered from Pleistocene sediments collected
from Frog Hall, Warwickshire.

STATUS OF BUPLEURUM FALCATUM L. IN THE BRITISH FLORA vy

have survived the glacial and periglacial conditions that prevailed during the Devensian Stage. If
this taxon was absent from the British mainland in the Devensian and Early Flandrian then it may
have been transported to southern England by natural dispersal from the continent. Alternatively,
the modern population may have an alien status with a relatively recent introduction by human
activity. Jermyn (1974) did speculate that fruits may have been transported to Essex by soldiers who
were returning from the Napoleonic wars. On the continent the modern distribution of this taxon
extends north-westwards along stretches of the French coastline (Fitter 1978). The Dover Strait and
the English Channel appear to have acted as an effective barrier to a more extensive colonisation of
Britain by this taxon from continental populations in the Flandrian Stage.

The lack of knowledge about the distribution and behaviour of B. falcatum subsp. falcatum in the
Late Pleistocene, due to its poor representation in the fossil record, makes it difficult to determine
the precise time and mode of arrival in England. However, the occurrence of B. falcatum subsp.
falcatum in the Late Pleistocene fossil record adds support to the opinion that the extant population
is native.

ACKNOWLEDGMENTS

I would like to thank Mrs M. Petitt and Prof. R. G. West.

REFERENCES

CorpeEr, T. (1833). Bupleurum falcatum L., in Smit, J. E., English botany supplement 2 (35): plate 2763.
London.

Fitter; A. H. (1978). An atlas of the wild flowers of Britain and northern Europe. London.

Gipson, G. S. (1862). The Flora of Essex. London.

JERMYN, S. T. (1974). Flora of Essex. Colchester.

MiTcHELL, G. F., PENNY, L. F., SHorron, F. W. & WEsT, R. G. (1973). A correlation of Quaternary deposits in
the British Isles. Geological Society of London, Special Report No. 4: 1-99.

Stace, C. A. (1991). New Flora of the British Isles. Cambridge.

SuMBLER, M. G. (1983). A new look at the type Wolstonian glacial deposits of central England. Proceedings of
the Geologists’ Association 94: 23-31.

TuTin, T. G. (1968). Bupleurum L., in TuTin, T. G. et al., eds. Flora Europaea 2: Sa 350. Cambridge.

TuTin, T. G. (1980). Umbellifers of the British Isles. London.

(Accepted December 1993)

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Watsonia 20: 119-132 (1994) 119

Seven new species of Taraxacum Wigg. (Asteraceae), native to
the British Isles

A. A. DUDMAN
Holebeck House, Cleator Moor, Cumbria, CA25 5HD
and
A. J. RICHARDS

Department of Agricultural and Environmental Science, Ridley Building,
University of Newcastle-upon-Tyne, Newcastle-upon-Tyne, NEI 7RU

ABSTRACT

Taraxacum breconense C. C. Haw., sp. nov. (sect. Celtica A. J. Richards), T. edmondsonianum H. @illg., sp.
nov. (sect. Ruderalia Kirschner, H. Ollg. & Stepanek), T. haworthianum A. A. Dudman & A. J. Richards, sp.
nov. (sect. Erythrosperma (H. Lindb.) Dahlst.), T. hirsutissimum C. C. Haw., sp. nov. (sect. Naevosa M. P.
Christ.), T. latens H. Ollg., sp. nov. (sect. Ruderalia), T. nigridentatum T. Edm., sp. nov. (sect. Ruderalia) and
T. sahlinianum A. A. Dudman & A. J. Richards, sp. nov. (sect. Hamata H. Ollg.) are described as new species,
native to the British Isles.

KrEyworps: Denmark, Germany, Netherlands, Czech Republic, Compositae.

INTRODUCTION

Before his untimely death in 1989, C. C. Haworth was planning to publish an account of British and
Irish Taraxacum as part of the handbook series of the Botanical Society of the British Isles.
Preliminary to this, he lectotypified the Dahlstedt British species (Haworth & Richards 1990) and
described six new species (Haworth 1990).

We are proceeding with the publication of this handbook, and in order to do so we need to publish
a further seven species. They appear here under a variety of authors. 7. breconense and T.
hirsutissimum appear here under Haworth’s authorship, as he left manuscript descriptions for them.
T. nigridentatum is named and described by T. Edmondson, and appears here under his authorship.
Two species are named and described by the Danish authority Hans Ollgaard, and we are grateful to
him for his permission to publish them within this paper. The remaining two species are described by
ourselves. Haworth (and others) formerly referred to T. haworthianum as Taraxacum “British
laetum’’, and 7. sahlinianum as Taraxacum “‘British atactum’’.

THE SPECIES

1. Taraxacum breconense C. C. Haw., sp. nov.

Ho .otypus: Nant Sere, Breconshire, v.c. 42, GR SO/041.247, wall/bank of lane, 13 May 1992, M.
Porter 92/38 (OXF). (Fig. 1).

Isotypus: herb. A. A. Dudman.

120 A. A. DUDMAN AND A. J. RICHARDS

FicureE 1. Holotype of Taraxacum breconense C. C. Haw. (top specimen).

Planta magnitudine mediocris. Folia patentia viridia immaculata epiceata, fere glabra et plana, late
lanceolata, ad 150 mm longa; lobi laterales 2-4, recurvati acuti, apice angusti, ad basem latiores,
saepe margine distali sat sigmoideo vel convexo in foliis bene evolutis, lobis inferioribus margine
distali plusminusve irregulariter denticulata; lobus terminalis submagnus subhastatus, late acutus,
valde subdivisus irregulariter profundeque dentatus; petiolus viridis exalatus. Scapus sub anthesi
saepe foliis brevior, interdum purpurascens, ad apicem sat pilosus. Involucri squamae exteriores 7—
10 X 2-5 mm, patentes subrecurvataeve, supra canescentes purpureo suffusae, subtus obscure

SEVEN NEW SPECIES OF TARAXACUM WIGG. 121

virides, interdum albo roseove indistincte submarginatae. Calathium 15-30 mm in diametro,
subclausum, pallide luteum; ligulae subbreves involutae; stylus stigmataque inserti lutei; antherae
epolliniferae. Achenium stramineo-brunneum, 3-5 mm longum pyramide exclusa, superne spinulo-
sum, pyramide subconica 0-7 mm longa; rostrum 8 mm longum; pappus albus.

Plant medium-sized. Leaves spreading, mid-green, lacking spots or dark markings, flat and almost
glabrous, broadly lanceolate, to 150 mm in length; leaf-lobes 2-4, acute and recurved with a narrow
apex, broader proximally with the distal margin sigmoid or convex in shape in well-developed
leaves, the proximal lobes more or less irregularly denticulate on the distal margin; terminal lobe
rather large and somewhat hastate in shape, broadly acute, markedly subdivided and irregularly and
deeply dentate; petiole green, unwinged. Scape often shorter than the leaves at flowering,
sometimes becoming purplish, hairy above. Exterior bracts 7-10 x 2:5 mm, spreading to somewhat
recurved, greyish green suffused with purple above, dark green below, sometimes indistinctly
bordered with white or rose. Capitulum 15-30 mm in diameter, not opening fully, pale yellow, the
ligules short and involute; style and stigmas inserted, yellow, anthers lacking pollen. Achene straw-
brown, 3-5 mm long (excluding the cone), spinulose above, cone more or less conical, 0-7 mm long;
rostrum 8 mm long, pappus white.

T. breconense is a close relative of T. ostenfeldii Raunk., with similar involucres and sharing small
capitula with short yellow styles which lack pollen. However, the ligules are notably involute, a rare
condition amongst British Taraxacum species (compare T. argutum Dahlst.), and the relatively
large and complexly subdivided end-lobes are distinctive and diagnostic. T. breconense is classified
within sect. Celtica A. J. Richards.

T. breconense is so far recorded from seven sites in four 10-km squares in v.c. 42 (Brecon). It has
been recorded from limestone cliffs, walls and lane-banks. Our knowledge of this species depends
entirely on the enthusiasm and expertly prepared collections of Michael Porter.

2. Taraxacum edmondsonianum H. @ilg., sp. nov.
Ho.otypus: Denmark, Jylland [Jutland], distr. 11, Hadsund, Hornbechsvej, ved en sti [by a
footpath], 2 May 1981, H. @llgaard 81-28 2./2 (C). (Fig. 2).
Isotypus: OXF (H@ 81.28 1./2).

The type specimens were collected and the plant taken into cultivation; the achenes were
described from the cultivated plant (Birgittelyst, 1982).

Planta magnitudine mediocris, 30-40 cm alta. Folia sat laete canescenti-viridia, sparsim vel in foliis
interioribus sat manifeste araneosa, late lanceolata, plerumque uniformiter lobata, margine
proximali loborum lateralium sat crispo, minute denticulato; petiolus alatus, plus minusve amoene
roseus, non striatulus; lobi laterales patentes subfalcati, raro plus minusve pagodiformes (ut in
Taraxaco aequilobo); lobus terminalis triangularis vel triangulari-hastatus, brevis, marginibus
distalibus plus minusve concavis, integris vel interdum incisis, apice lobis ibi saepe rotundatus.
Scapus pallidus vel leviter brunnescens, sub involucro dense araneoso-pilosus alibi parce. Involuc-
rum c. 14 mm longum, c. 11 mm latum, laete vel sat obscure viride, haud pruinosum. Squamae
exteriores lanceolatae, 14 (-16) mm x 4 (—5) mm, non vel vix marginatae, retroflexae, sat regulares,
supra pallide virides. Calathtum 45-55 mm in diametro, luteum convexum radians; ligulae
marginales planae vel saltem haud canaliculatae, subtus stria cano-violacea ornatae; antherae
polliniferae, granis pollinis in diametro variantibus; stylus luteus, stigmata lutea vel in sicca
levissime virescentia. Achenium fusco-stramineum, ad 4 mm longum pyramide inclusa, superne
grosse spinulosum, alibi plus minusve verrucosum vel fere laeve, in pyramidem cylindricam (0-4-)
0-5(—0-6) mm longam abrupte abiens. Rostrum c. 11 mm longum. Pappus albus.

Plant medium-sized (30-40 cm). Leaves light greyish-green, slightly or (on inner leaves) markedly
araneose-hairy, broadly lanceolate, mostly uniformly lobate, the proximal margins of the lateral
leaf-lobes somewhat crisped, minutely denticulate; petiole narrowly winged, more or less a
beautiful rose-pink, the midrib faintly reddish or brownish, lacking a striatulate pattern. Lateral
leaf-lobes patent, subfalcate, rarely more or less pagoda-shaped (as in T. aequilobum Dahlst.).

122 A. A. DUDMAN AND A. J. RICHARDS

FLORA DANICA

TARAXACUM & <

FicureE 2. Holotype of Taraxacum edmondsonianum H. @llg.

Terminal leaf-lobe triangular to triangular-hastate, short, more or less concave on distal margins,
entire or sometimes incised, and then the apices of the lobules rounded. Scape pale or becoming
slightly brownish, densely araneose-pilose under the head, the remainder slightly pilose. Involucre
about 14 mm long and 11 mm wide, pale or somewhat darkish green, not pruinose. Exterior bracts
lanceolate, 14(—16) x 4(—5) mm, unbordered or scarcely bordered, somewhat regularly arranged,
pale green above. Capitulum 45-55 mm in diameter, yellow, convex, the marginal ligules spaced,
flat or at least not furrowed, with grey-violet stripes beneath. Anthers bearing pollen with grains of
varying diameter (suggesting agamospermous behaviour). Styles yellow, stigmas yellow or in the
dry state very slightly greenish. Achene brownish-straw coloured, to 4 mm long (including the

SEVEN NEW SPECIES OF TARAXACUM WIGG. 123

cone), strongly large-spinulose above, the rest more or less verrucose or almost smooth, abruptly
contracted into a cylindrical cone of (0-4—)0-5(—0-6) mm. Rostrum about 11 mm long. Pappus white.

T. edmondsonianum is classified within sect. Ruderalia Kirchner, H. Ollg. & Stepanek. Morpholo-
gically, it is a relative of the common and widespread T. aequilobum, having many uniform leaf-
lobes and more or less winged petioles. The latter species is recognised by the lateral leaf-lobes,
usually shaped like the roof of a pagoda, and by the conspicuously irregular and twisted exterior
bracts. In contrast, the leaf-lobes of T. edmondsonianum are more patent, less acuminate and more
straight-sided on the distal margin; the proximal margin to these lobes is convex and bent upwards
throughout. Also, the exterior bracts of T. edmondsonianum are strongly and regularly reflexed,
and are not twisted. Other good marks of recognition are the delicate and beautiful rose-pink
petiole, and the small terminal leaf-lobes with usually concave upper margin. Often, some terminal
lobes are incised, the lobules so formed being rounded. 7. edmondsonianum has a paler leaf colour
than T. aequilobum and the stigmas are yellow when fresh. The constancy of these differences has
been confirmed after cultivation in standard conditions.

T. edmondsonianum is named after T. Edmondson of Chester, who recognised it as a new species
during his very thorough investigation of the Taraxacum flora of the Chester district. In the British
Isles it is so far known from Herefordshire (v.c. 36), Cheshire (v.c. 58), South Lancashire (v.c. 59),
West Lancashire (v.c. 60) and Cumberland (v.c. 70), where it has been recorded from seven 10-km
squares. It is known from at least seven separate localities in the city of Chester. Outside England, it
is recorded from nine localities in five recording districts (TBU02, 10, 11, 13a, 49) in Jutland,
Denmark, and from a single locality in Hessen, Germany. (A full list of authentic specimens
identified up until now can be obtained from the species’ author at Lupinstien 7, Birgittelyst, Viborg
8800, Denmark.)

3. Taraxacum haworthianum A. A. Dudman & A. J. Richards, sp. nov.

Ho .otypus: Druridge Bay, Northumberland, v.c. 67, GR NZ/276.965, sand-dune grassland 100 m
from the sea, 20 May 1987, A. J. Richards 87/41 (OXF). (Fig. 3).

Isoryri: herb. A. A. Dudman, H. Ollgaard, J. Stepanek.

Planta parva. Folia prostrata viridia glabra, immaculata epiceata, nervo mediano basin versus viride
purpureove, anguste lanceolata, ad 100 mm longa; lobi laterales 6-8, subintegri subrecurvati,
regulatim triangulares vel margine distali sigmoideo, apice angusto, subacuto subobtusove vel
interdum expanso; lobus terminalis subsagittatus integer in apicem acutum protractus; petiolus
angustus exalatus ad basem vivide atropurpureus, distaliter interdum viridescens. Scapus tenuis
purpureus adscendens, superne pilosus. Involucri squamae exteriores ovatae breves, 4-6 X 2 mm,
subadpressae, margine anguste albo-scarioso vel roseo-scarioso, apice abrupte purpureo-cornicu-
lato. Calathium 25-30 mm in diametro, luteolum; ligulae breves lataeque, stria cano-violescenti
subtus notatae, dentibus apicalibus purpureis; stylus stigmataque sordide lutei; antherae pollini-
ferae. Achenium 3-0 mm longum pyramide exclusa, angustum, in stato maturo atropurpureum, in
statu juveniliore rubescens, superne acute spinulosum, alibi fere laeve, pyramide anguste cylindrica
0-8-1-1 mm longa; rostrum 6—7 mm longum; pappus albus. 2n = 24”*.

Plant small. Leaves prostrate, green, glabrous, lacking spots or dark markings, the midrib green or
purple, narrowly lanceolate, to 100 mm in length; lateral leaf-lobes 6-8, subentire, somewhat
recurved, regularly triangular, or with the distal margin sigmoid, with narrow subacute to subobtuse,
or sometimes somewhat expanded apices; terminal lobe entire and subsagittate, drawn out into an
acute apex; petiole narrow and unwinged, brilliantly purple at the base. Scapes narrow, purple,
ascending, hairy above. Exterior bracts ovate, short, 4-6 x 2 mm, subadpressed to the involucre,
narrowly white- to rose-bordered, abruptly ending in a marked purple corniculation. Capitulum 25—
30 mm in diameter, pale yellow, the ligules short and wide with a grey-violet stripe and purple teeth;
style and stigmas dirty yellow; pollen present. Achene 3-0 mm (excluding the cone), narrow, dark
purple when mature, reddish when immature, acutely spinulose above, the rest almost smooth, the
cone narrowly cylindrical, 0-8—1-1 mm; rostrum 6-7 mm; pappus white. 2n = 24%.

124 A. A. DUDMAN AND A. J. RICHARDS

Ficure 3. Holotype of Taraxacum haworthianum A. A. Dudman & A. J. Richards (top right hand specimen).

T. haworthianum is a close relative of T. laetum (Dahlst.) Raunk., a widespread sect. Erythro-
sperma (H. Lindb.) Dahlst. species of the sand-dunes and other dry grasslands of northern Europe.
It was included within this species in Richards (1972), where some material was also referred to T.
laetiforme Dahlst. In later years, it was recognised that such British and Irish material differs from
T. laetum, notably by the narrow, bright purple petioles of all leaves (these are short and green in T.
laetum), and also by the grey-violet, not purple, ligule stripes, a slightly longer achene body, and by
dirty yellow, not brilliantly gold stigmas. :

SEVEN NEW SPECIES OF TARAXACUM WIGG. 125

@Ollgaard (1986) showed that T. /aetiforme is a very localized and rare Danish species, and he gave
the name T. discretum to the widespread plant from the Netherlands and Scandinavia which had
been called ‘‘T. laetiforme’’ by Haglund and van Soest. This species has achenes that are chestnut-
brown when mature, and petioles that are green in outer leaves, but purple on inner leaves. The
stigmas are a dirty greenish colour. In 1986, it was considered that T. discretum occurred in the
British Isles, but Haworth later showed that such plants were in fact forms of what is described here
as T. haworthianum with immature achenes.

For some time, Haworth considered the possibility that the future T. haworthianum could be
equated with another relative of T. Jaetum, the Dutch T. agaurum Soest. However, that species
always has winged, green petioles and golden stigmas, and in our view T. agaurum is very closely
allied to T. laetum itself. After testing these species in cultivation, it was Haworth’s opinion, with
which we concur, that no British plants can be referred to T. Jaetum, T. agaurum or T. discretum.

Haworth himself referred to this British relative of T. Jaetum and T. discretum as Taraxacum
“British laetum’’, and also as ““T. sublaetum’”’. However, we would like to abandon these Haworth
worknames, and to take this opportunity to recognise Chris Haworth’s substantial contribution to
British taraxacology. ;

T. haworthianum is widespread in species-rich, calcareous grassland (most commonly in sand-
dunes) in Britain and Ireland, having been recorded from 83 10-km squares in the following vice-
counties: 1-4, 6, 13, 15, 25, 27-29, 41, 42, 45, 46, 48, 49, 52, 55, 57-60, 67-69, 73-75, 82, 85, 90, 93—
95, 102-104, 106, 107, 109, 110, S, H9, H12, H21, H28, H39, H40.

With its prostrate rosettes of highly dissected leaves with narrow, purple petioles, and very pretty,
rounded involucres of bordered, purple corniculate exterior bracts, T. haworthianum is a distinctive
and attractive little plant. It bears more than a passing resemblance to a sect. Obliqua Dahlst.
species, T. platyglossum, Raunk. from which its slender, deep blackish-purple achenes will
immediately distinguish it.

4. Taraxacum hirsutissimum C. C. Haw., sp. nov.
Ho .otypus: Dunrossness, Shetland, v.c. 112, GR HU/3.2, above the source of the Red Burn, near
Williamsetter, roadside verge, 20 June 1979, W. A. Scott 2618 (OXF). (Fig. 4).

Planta mediocris. Folia adscendentia canescentia, per paginam superiorem breve subdense hirsuta,
sparse puncto-maculata vel immaculata, epiceata, nervo mediano viridi vel plus minusve purpureo-
tincto, oblanceolata, ad 200 mm longa; lobi laterales 4-7, patentes subrecurvative, apice anguste
acuti, margine distali saepe angulato-concavi, fere edentati; lobus terminalis sagittatus integer,
apice subprotractus acutus; petiolus purpureus, vix alatus. Scapus purpurascens, superne pilosus.
Involucri squamae exteriores leviter recurvatae, lanceolatae, 12 x 2-5 mm, supra pallide virides,
subtus pruinoso-glaucae, vix marginatae. Calathium intense luteum, ad 30 mm in diametro; ligulae
stria obscurocanescenti subtus notatae, dentibus apicalis luteis; stylus stigmataque aurantiaci;
antherae polliniferae. Achenium stramineum, 3-5 mm pyramide exclusa, pyramide conica 0-5 mm
longa; rostrum 9-10 mm longum; pappus albus.

Plant medium-sized. Leaves ascending, greyish, shortly and rather densely hairy throughout the
upper surface, unspotted or with scattered punctate spots, lacking dark markings, the midrib green
or faintly purple, oblanceolate, to 200 mm long; lateral leaf-lobes 4~7, patent or slightly recurved,
with narrow acute apices, the distal margins concave, often with a concave angle, almost lacking
teeth; terminal leaf-lobe sagittate, entire, with an acute, somewhat extended apex; petiole purple,
scarcely winged. Scape purplish, hairy above. Exterior bracts somewhat recurved, lanceolate, 12 x
2:5 mm, pale green above, bluish pruinose green beneath, scarcely bordered. Capitulum deep
yellow, to 30 mm in diameter, ligules striped dark grey beneath, ligule teeth yellow; style and
stigmas orange; pollen present. Achene straw-coloured, 3-5 mm (excluding the cone), the cone
conical, 0-5 mm; rostrum 9-10 mm; pappus white.

Over more than two decades, Richard C. Palmer and Walter A. Scott have thoroughly examined
the Taraxacum species of Shetland. A number of their gatherings relate to a distinctive member of
sect. Naevosa that is immediately recognised by its leaves, which are rendered greyish by their dense

126 A. A. DUDMAN AND A. J. RICHARDS

W
“a

Lew N

oN

-
oo

\S
ee.

j
hp

y
ie yi ee $
Ak BEE A

Ficure 4. Holotype of Taraxacum hirsutissimum C. C. Haw.

covering of short even hairs, and by the orange stigmas. Among its British relatives, this plant is
most closely related to T. euryphyllum (Dahlst.) Hjelt, and the achenes, leaf-shapes and involucres
of the two species are quite similar, although the leaves of T. hirsutissimum are generally more
multilobate and the end-lobes more sagittate. However, apart from differing in the stigma colour
and in the hairiness of the leaves, 7. euryphyllum also virtually lacks pollen, while the sparse
punctate leaf spots of T. hirsutissimum are distinctive. Haworth cultivated this species from achenes
sent by Scott, and it retained its distinctive. features in cultivation. So far, T. hirsutissimum is only
known from v.c. 112 (Shetland), where it has been recorded from eleven sites in two 10-km squares.

SEVEN NEW SPECIES OF TARAXACUM WIGG. 72]

FLORA DANICA

TARAXACUM

FicurE 5. Holotype of Taraxacum latens H. Ollg. (top specimen).

5. Taraxacum latens H. @llig., sp. nov.
Hovortyreus: Denmark, Jylland (Jutland), distr. 14, 5 km SSW Viborg, road 13 at the 86-6 km mark,
Picea plantation, 3 May 1993, H. @llgaard 111.01 (part of exsiccata (H@ 93-111.xx)) (C). (Fig. 5).

Planta magnitudine mediocris c. 30 cm alta. Folia flavescenti-viridia vel viridia, sparsim araneosa,
nec piceata nec maculata; nervo mediano viridi vel rarius subrubescenti, colore non striatulo; lobi
laterales plerumque 6-8, deltoidei, in speciminibus tenuibus plerumque plus minusve falcati, parce

128 A. A. DUDMAN AND A. J. RICHARDS

dentati vel integri vel interdum incisione alta subfissi, margine proximali integro vel denticulato,
apice loborum subobtusi vel acuti; lobus terminalis foliorum exteriorum intermediorumque parvus,
saepe incisus, apice brevi indefinito vel gradatim sat longe protracto, in foliis interioribus conspicuo,
cordato-hastato, subacuto vel subobtuso; interlobi partis superioris folii lobis lateralibus perdensis
saepe subtecti, in medio plus minusve acutanguli, non vel parum plicati; petiolus roseo-purpureus,
anguste alatus. Scapus folia aequans vel superans, viridis vel leviter brunnescens, sub involucro
densissime araneoso, alibi sparsim araneosa. Involucrum subobscure viride, curtum, saepe fere
globosum, leviter pruinosum. Squamae exteriores lanceolatae, c. 14 X 4 mm, recurvae vel
retroflexae, regulares, apice ecorniculata, laete virides vel colore roseolo tinctae, plus minusve
manifeste marginatae. Calathium mediocre, c. 45 mm in diametro, intense luteum; ligulae
marginales planae vel parum canaliculatae, subtus stria cano-violacea ornatae, dentibus apicalibus
rubescentibus; antherae polliniferae, granis pollinis diametro variis; stigmata plus minusve
virescentia. Achenium fusco-stramineum, 3-5 mm longum pyramide subcylindrica laevi c. 0-6 mm
longa bene definita inclusa. Rostrum 11 mm longum; pappus albus.

Plant medium-sized, about 30 cm high. Leaves yellowish-green to green, slightly araneose-hairy,
neither spotted nor blotched dark, the midrib green or rarely slightly reddish, but not striatulate;
lateral leaf-lobes usually 6-8, deltoid, sometimes more or less falcate in slender specimens, sparsely
dentate to entire or sometimes with deep subdivisions, the proximal margin entire or denticulate,
the apex subobtuse to acute; terminal lobe of the outer and intermediate leaves small, often
subdivided, the apex short and obscure or varying towards somewhat long-protracted, of inner
leaves conspicuous, cordate-hastate, subacute or subobtuse; interlobes of the distal part of the leaf
partially hidden by dense, overlapping lateral lobes, in the median part of the leaf more or less
acute, not or scarcely plicate; petiole rose-purple, narrowly winged. Scape equalling or exceeding
the leaves, green or lightly brownish, very densely araneose-hairy underneath the involucre, the
remainder sparsely hairy. Involucre rather dark green, short, often almost globose in bud, lightly
pruinose. Exterior bracts lanceolate, c. 14 x 4mm, recurved or reflexed, regular in arrangement,
the apex ecorniculate, pale green or faintly suffused with rose, more or less strongly bordered.
Capitulum medium-sized, about 45 mm in diameter, deep yellow; marginal ligules flat or slightly
canaliculate, striped grey-violet beneath, the apical teeth reddish; anthers bearing pollen, the pollen
grains of varying diameter; style and stigmas more or less greenish. Achene fuscous-straw-coloured,
3-5 mm including the well-defined and smooth subcylindrical cone, about 0-6 mm long. Rostrum 11
mm long; pappus white.

T. latens is classified within sect. Ruderalia (= sect. Vulgaria nom. illegit.). Amongst the numerous,
mostly anthropochorous species in this section, it is recognized by the rose-pink, somewhat winged
petioles, the pale green, more or less densely lobate leaves bearing entire proximal margins to the
lateral lobes, and the light green, strongly recurved to reflexed, regularly arranged exterior bracts
with an obviously differentiated border. The lateral leaf-lobes tend to become falcate, with a broad
basal part and a well-differentiated linear distal process, especially in old plants, or in modifications
from exposed positions. The terminal lobes of the outer and middle leaves are usually small and
inconspicuous, more or less incised, with a gradually elongate apex, while those of inner leaves are
large and conspicuous, with gross teeth almost of the stature of confluent lateral leaf-lobes.

The closest morphological relative of 7. /atens seems to be T. crispulum G. E. Haglund, a little-
known species not recorded from the British Isles, which, however, according to the description
(Haglund 1934) and the type material, has crispate interlobes with obvious dark blotchings, and
much larger teeth. Also, T. crispulum does not produce inner leaves with large terminal lobes, the
proximal margins of the lateral leaf-lobes are more or less subulate-dentate, and the achene cone isa
little shorter.

Amongst commoner Taraxacum species, T. latens is best compared with T. aequilobum (the leaf-
shape is somewhat similar, but in 7. aequilobum the exterior bracts are very irregular and more or
less twisted), T. vastisectum Markl. ex Puol. (again, the leaf-shape is somewhat similar, but the
latter species has a darker and faintly bluish leaf colour, usually well-developed teeth on the
proximal margins of the lateral leaf-lobes, no elongation at the apex of the terminal leaf-lobe, and
less reflexed, dirty purplish exterior bracts) and T. amplum Markl. (similar in leaf and petiole

SEVEN NEW SPECIES OF TARAXACUM WIGG. 129

colour, and in the border to the exterior bracts, but with more or less patent exterior bracts, and the
sagittate, usually entire, terminal lobes to the inner leaves).

In the British Isles, 7. /atens has so far been recorded from five 10-km squares in north Essex (v.c.
19), Shropshire (v.c. 40), Denbighshire (v.c. 50) and Cheshire (v.c. 58). Its distribution here
suggests that it may be introduced. Elsewhere, it has been recorded from 17 recording districts
(TBUO9, 10, 11, 14, 15, 20, 26, 28, 33, 37, 39b, 47, 48, 49, 50, 52, 53) in Jutland, Denmark, from
three sites in the Netherlands, from two sites in Germany, and from three sites in the Czech
Republic. (A full list of authentic specimens identified up until now can be obtained from the
species’ author at Lupinstien 7, Birgittelyst, Viborg 8800, Denmark.)

6. Taraxacum nigridentatum T. Edm., sp. nov.
Ho.otyeus: Hapsford, Cheshire, v.c. 58, GR SJ/468.745, 3 May 1985, T. Edmondson 1985.57
(OXF). (Fig. 6).

Planta robusta, magnitudine mediocris vel magna. Folia adscendentia erectave, obscure viridia,
immaculata, interlobis dentibusque utrinque valde piceata, nervo mediano sordide roseo, lanceo-
lata, ad 250 mm longa, multilobata; lobi laterales 5-8, subpatentes recurvative, sat longi, acuti,
margine distali recto sigmoideo, multos dentes nigros vel unum dentem grossum ferenti; lobus
terminalis subhastatus subsagittatusve, folii partibus inferioribus angustior, 1—2- saepe unilateraliter
inaequaliterve subdivisus vel dentatus, apice protractus subacutus; petiolus alatus, interdum late
alatus, roseus vel basi albus. Scapus sub anthesi folia aequans, sordide roseus. Involucri squamae
exteriores patentes vel eae infimae recurvatae, 12-14 x 2-3 mm, emarginatae, supra pallide virides
vel purpureo suffusae, subtus obscuriores. Calathium luteum, 40-50 mm in diametro, ligulis stria
cano-brunnescenti subtus notatis; stylus stigmataque sordidi; antherae polliniferae. Achenium
stramineo-brunnescens spinulosum, 3-3 mm longum pyramide exclusa, pyramide minima, ad 0-3
mm longa; rostrum 5—6 mm; pappus albus.

| FLORA BRITANNICA
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we aieunnnananneninnne amin
“igrddentatum
DET Se he
i ove pt (the
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reQges 57

Ficure 6. Holotype of Taraxacum nigridentatum T. Edm.

130 A. A. DUDMAN AND A. J. RICHARDS

Plant robust, medium-sized to large. Leaves ascending or erect, dark green, unspotted, the
interlobes and teeth heavily marked with black on both sides, midrib dirty pink, lanceolate, to 250
mm in length, multilobate; lateral leaf-lobes 5—8, patent to recurved, rather long, acute, with many
black teeth on the distal margin or with one large tooth on distal lobes, the distal margin straight-
sided or sigmoid; terminal leaf-lobe subhastate to subsagittate, narrower than lower parts of the
leaf, with 1-2 subdivisions or teeth, often on one side only, or if on both sides, then usually
alternately arranged, with an attenuate subacute apex; petiole winged, sometimes widely so, pink,
or white at the base. Scape equalling the leaves at flowering time, dirty pink. Exterior bracts mostly
spreading, the lower recurved, 12-14 x 2-3 mm, unbordered, pale green or suffused purplish
above, darker below. Capitulum deep yellow, 40-50 mm in diameter, the ligules striped greyish-
brown beneath; style and stigmas dirty coloured; pollen present. Achene straw-brown, spinulose,
3-3 mm (excluding the cone), the cone very small, to 0-3 mm; rostrum 5—6 mm; pappus white.

T. nigridentatum is a weedy species classified within sect. Ruderalia. These plants of anthropogenic
habitats tend to be much more mobile than species of more natural habitats, and consequently they
rarely occur as localized endemics. Approximately 130 species classified within this section are
recorded from the British Isles, but of these only four have yet to be recorded in another country. So
far, T. nigridentatum is not recorded with certainty away from western Britain, where it may be
endemic, although similar plants may occur in the Netherlands. It has been recorded from 29 10-km
squares, but so far is known from only the following western vice-counties: East Gloucestershire
(v.c. 33), West Gloucestershire (v.c. 34), Monmouthshire (v.c. 35), Breconshire (v.c. 42), Flintshire
(v.c. 51), Cheshire (v. c. 58) and Cumberland (v.c. 70).

T. nigridentatum is a relative of T. pannulatum Dahlst., and morphologically it is particularly
close to T. pannulatiforme. Dahlst., although in well-developed leaves, the leaf-lobes of T.
nigridentatum tend to be narrower and more multidentate. However, T. nigridentatum is invariably
and characteristically heavily blotched with black on the interlobes and leaf-teeth, while the midrib
and the petiole (except usually the very base outside which is white) are a dirty reddish-pink. In
comparative cultivation with 7. pannulatiforme, Haworth showed that the distinctive leaf colour
and the minor differences in leaf shape of T. nigridentatum were maintained (the leaves of T.
pannulatiforme are essentially green throughout).

7. Taraxacum sahlinianum A. A. Dudman & A. J. Richards, sp. nov.
Ho otyeus: Woodside, Maryport, Cumberland, v.c. 70, GR NY/047. 343, roadside, 8 May 1991, A.
A. Dudman s.n. (OXF). (Fig. 7).

Planta magnitudine mediocris, sat robusta. Folia suberecta plana, sordide obscuroviridia, immacu-
lata epiceataque, nervo mediano ordinatione intertexto sordide purpureo notato, late oblanceolata,
ad 200 mm longa; lobi laterales 2-4, patentes subrecurvative, lati subbrevesque, triangulares
subacuti, margine distali saepe recto vel subconvexo, lobis inferioribus margine distali dentibus
filiformis ornatis; lobus terminalis submagnus, obtusus sed apice ipso mucronatus, inaequaliter
grosse 1—3-dentatus; petiolus alatus vel saepe late alatus, dentatus, ordinatione intertexto purpureo
notatus. Scapus sub anthesi folia aequans, erectus. Involucri squamae exteriores pruinosae,
patentes sed decurvatae, 10 X 3 mm, vix marginatae, supra pallide virides interdum purpureo
suffusae, subtus obscuriores. Calathium luteum, 40-45 mm in diametro, ligulis stria obscura griseo-
violacea subtus notatis; stylus stigmataque sordidi; antherae polliniferae. Achenium fulvo-
stramineum, apice rugosum alibi laeve, 3-8-4-0 mm pyramide exclusa, pyramide conica 0-6 mm
longa; rostrum 10 mm longum; pappus albus.

Plant medium-sized, somewhat robust. Leaves suberect, flat, dark and dirty green, lacking spots or
dark markings, the midrib dirty purple in an interwoven pattern, broadly oblanceolate, to 200 mm
long; lateral leaf-lobes 2-4, patent to somewhat recurved, broad and rather short, triangular,
subacute, the distal margins straight or slightly convex, filiform dentate on lower lobes; terminal
leaf-lobe rather large, obtuse but mucronate, unequally provided with 1-3 large teeth; petiole
winged, often broadly so, filiform dentate; purple in an interwoven pattern. Scape equalling the
leaves at flowering time, erect. Exterior bracts spreading but downcurved, pruinose, 10 x 3 mm,

SEVEN NEW SPECIES OF TARAXACUM WIGG. 131

FicureE 7. Holotype of Taraxacum sahlinanum A. A. Dudman & A. J. Richards.

scarcely bordered, pale green but sometimes suffused purple above, darker green below. Capitulum
deep yellow, 40-45 mm diameter, the ligules striped dark grey-violet below, style and stigmas dirty
coloured; pollen present. Achene fulvous-straw coloured, rugose above, the remainder smooth,
3-8—4-0 mm (excluding the cone), the cone conical, 0-6 mm; rostrum 10 mm; pappus white.

From the shape, size, posture and pruinosity of the exterior bracts, the colour and lobation of the
leaves, and the characteristically ‘interwoven’ pattern of the colour in the midrib and petiole, it is

132 A. A. DUDMAN AND A. J. RICHARDS

clear that T. sahlinianum should be classified with sect. Hamata H. Oilg. Within this section, T.
sahlinianum is most closely allied to 7. atactum Sahlin & Soest. In particular, the large, subrounded
end-lobes to the leaves, which are often one-sidedly provided with one or a few large teeth, are
reminiscent of this species. Although T. atactum is a widespread British species, some early records
in fact refer to T. sahlinianum.

@Mllgaard first pointed out that not all British material referred to T. atactum was typical of that
species, and this was confirmed by the late C. I. Sahlin, one of the authors of the name T. atactum.
Since then, Haworth and Richards have recognised a distinctive taxon which was given the
workname Taraxacum “British atactum’’. Now that we believe that we understand this taxon well,
we feel that the time has come to describe it formally. In doing so, we would like to commemorate
the Swedish taraxacologist Carl Ingemar Sahlin who died in 1990.

T. sahlinianum is best distinguished from T. atactum, and indeed from its other relatives in sect.
Hamata, by the broad, broadly lobed leaves with an often widely winged petiole, and by the large
achene. Apart from T. atactum, no relative has a long toothed end-lobe to the leaf. The dull, dark
and rather dirty hue of the leaf is also distinctive.

T. sahlinianum will probably prove to be a widespread and frequent species of semi-natural
grassland habitats, particularly in the west of Britain. At present it is recorded from 15 10-km
squares in the following vice-counties: West Gloucestershire (v.c. 34), Monmouthshire (v.c. 35),
Breconshire (v.c. 42), Carmarthenshire (v.c. 44), West Lancashire (v.c. 60), Westmorland (v.c. 69),
Cumberland (v.c. 70) and Wexford (v.c. H12).

So far, T. sahlinianum has not been recorded outside the British Isles.

ACKNOWLEDGMENTS

We would like to thank those British workers whose efforts have led to a fuller understanding of the
species described here, particularly Tom Edmondson, Michael Porter, Merle Marsden, Richard
Palmer and Walter Scott. As always, we are indebted to authorities in other countries for critical
comments, notably Hans @llgaard and Piet Oosterveld.

REFERENCES

HaGLunp, G. E. (1934). Nagra nya eller i Sverige nyfunna Taraxacum-arter. Botaniska notiser 1934: 1-41.

Haworty, C. C. (1990). Six native species of Taraxacum new to the British Isles. Watsonia 18: 131-138.

Hawortn, C. C. & RicHarps, A. J. (1990). The lectotypification and revision of Dahlstedt’s species of
Taraxacum Weber based on British or Irish plants. Watsonia 18: 125-130.

@LLGAARD, H. (1986). Taraxacum discretum sp. nov. (Compositae). Nordic journal of botany 6: 21-24.

Ricuarps, A. J. (1972). Taraxacum flora of the British Isles. Watsonia 9 (suppl.): 1-141.

(Accepted March 1994)

Watsonia 20: 133-141 (1994) 133

New species of Rubus L. (Rosaceae) from Wales and the
Welsh Marches

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

Aberhoywy, Cyffredin Lane, Llangynidr, Crickhowell, Powys, NP8 1LR

ABSTRACT

The relationships and distributions of five brambles (Rubus L.) in the Corylifolii section occurring in Wales and
the Welsh Marches are discussed. Three new species from this section and one from series Hystrices are
described: Rubus ariconiensis A. Newton & M. Porter, sp. nov., R. iscanus A. Newton & M. Porter, sp. nov.,
R. vagensis A. Newton & M. Porter, sp. nov. and R. segontii A. Newton & M. Porter, sp. nov.

KEYworDs: Apomictic species, distribution, brambles.

INTRODUCTION

Considering their abundance in many regions, the brambles of sect. Corylifolii Lindl. have been a
neglected component of the genus Rubus. In his conclusions to a review of the section, Edees (1975)
noted ““Much remains unknown. A great deal of fieldwork must be done and probably many more
taxa described before a comprehensive account can be written.”’ Visits to many parts of Britain have
corroborated this view. Growing in profusion in the hedges of the southern Welsh Marches are
several brambles of the Corylifolii section which have puzzled generations of batologists. Recent
research has shown that four of these brambles are widespread in South Wales and the West
Midlands; these and their allies are discussed below and three are described as new species. The
opportunity is taken to describe a new bramble of the Hystrices series from North Wales.

THE SPECIES
Rubus iscanus A. Newton & M. Porter, sp. nov.

Turio arcuatus in apricis fusco-purpureus pruinosus obtuse angulatus faciebus planis glabrescens
glandulis stipitatis sparsis aculeis crebris plerumque ad angulos dispositis aequalibus patentibus
rectis vel obfalcatis aculeisque brevibus raris armatus.

Folia (3)—5-nata pedata imbricata superne parce strigosa inferne capillis simplicibus vestita.
Foliolum terminale apice cuspidatum vel acuminatum, late ellipticum vel subrotundum basi
emarginatum vel subcordatum inaequaliter serratum vel biserrate dentatum interdum lobatum
petiolulo triplo longius; foliola infima subsessilia.

Inflorescentia cylindrica fere ad apicem foliosa inferne 2-4 foliis ternatis superne 1-3 foliis
simplicibus instructa; ramuli inferiores foliis breviores 3—7 flori. Rachis vix flexuosa pilosa
tomentosa aculeis 4-6 mm rectis vel declinatis validis glandulisque stipitatis nonnullis armata. Flores
3-4 cm diam.; sepala post anthesin reflexa mox fructum laxe amplectantia tomentosa albo-

134 A. NEWTON AND M. PORTER

marginata glandulis subsessilibus rubris ornatis. Petala rosea elliptica vel obovata interdum apice
emarginata fimbriata; stamina stylos roseos vix superantia; anthera glabra; filamenta rosea; carpella
glabra; fructus subglobosus.

Stem low-medium arching, angled with flat sides, brownish-purple on exposed sides, pruinose,
initially with short tufted or simple hairs becoming almost glabrous, with scattered sessile glands and
occasional short-stalked glands; prickles c.15—25 per 5 cm mainly on the angles but some on the
faces, + equal, S—6 (—7) mm, about equal to the stem diameter = straight or slightly upswept at the
tip, patent or slightly declining, purple with straw coloured tips, rarely one or two short prickles c. 1
mm. Leaves pedate, leaflets (3—) 5, imbricate, mid-green, slightly strigose above, soft beneath
rather thinly clothed with simple hairs; terminal leaflet c. 7.5 X 6.5 cm, broadly elliptical, ovate,
obovate or subrotund with an acuminate or cuspidate apex c. 1 cm and emarginate or cordate base,
compound serrate or unevenly biserrate with flat or slightly undulate margin, the petiole c. 1/3 as
long as the lamina; basal leaflets subsessile or with petiolules up to 2 mm; petioles about the same
length or slightly longer than basal leaflets, coloured like the stems, with rather dense spreading
simple or tufted hairs and occasional very short stalked glands, c. 12-15 = straight, slightly declining
prickles c. 2 mm. Flowering branch with 3-foliate leaves below and up to 3 simple leaves above,
usually leafy almost to apex; inflorescence cylindrical consisting of a terminal head of a few flowers
borne on widely diverging peduncles and pedicels with several + distant axillary peduncles shorter
than their leaves (bearing up to 7 flowers). Rachis slightly flexuose coloured like the stem with dense
whitish spreading and adpressed simple or tufted hairs and stellate hairs, sparse to frequent sessile
or short-stalked glands, several straight slightly declining stout-based prickles 4-6 mm, crimson with
straw coloured tips; peduncles clothed like the rachis but with denser more adpressed hairs so that
they appear greyish in colour, prickles straight slightly declining, up to 4 mm; pedicels clothed as
peduncles but with shorter hair and prickles + straight patent or slightly declining, up to 3 mm,
sparse to frequent dark red short or very short stalked glands especially just below the flowers.
Flowers c. 3.5 cm in diameter; sepals greyish-green with dense felt and short simple hairs, few short
acicular prickles, occasional short stalked glands and few or many subsessile glands, white-
margined, short pointed, reflexed at petal fall, erect to clasping in fruit; petals c. 15 x 10 mm bright
pink elliptical to obovate, sometimes notched or erose at apex, with short hairs on the margins;
stamens slightly exceeding styles, filaments pink, anthers glabrous; styles pink or pink-based;
carpels glabrous, receptacle glabrous or slightly hairy; fruit.small-medium + round sometimes
partly abortive, consisting of rather few drupelets. Flowering from June to September.

Ho.otypus: Canal bank, Llangynidr, Brecs., v.c. 42, GR SO/165.195, 20 June 1992, M. Porter
(NMW).
Isotyri: BIRM, BM.

The bramble takes its name from the Roman name (Isca) for the River Usk. Rubus iscanus is one of
the first brambles to flower in the Usk valley, its large pink blooms being conspicuous from early
June in thickets and hedges from Brecon to the Severn estuary. It has also been recorded from
localities scattered across Monmouthshire (v.c. 35), from the Usk to the Wye. A specimen at NMW,
collected from Mounton near Chepstow by W. A. Shoolbred in 1892, was determined as R. eupectus
(Sudre) W. C. R. Watson by Watson in 1949.

R. iscanus has been recorded from v.cc. 34, 35, 36 and 42. Its known distribution is shown in Fig. 1.

Rubus ariconiensis A. Newton & M. Porter, sp. nov.

Turio arcuato—decumbens in apricis rubiginosus angulatus superficiebus planis vel concavis vix
pruinosus glabrescens aciculis glandulisque stipitatis 0.54 mm longis sparsis, aculeis aculeolisque
1-7 mm longis nonnullis glanduliferis haud ad angulos limitatis e basi lata rectis vel declinatis vel
devexis copiose armatus. Folia (3-) 5- nata, pedata. Foliola contigua superne parce strigosa inferne
capillis simplicibus brevibus vestita, foliolum terminale (c. 7 X 4.5 cm) ellipticum cuspidatum basi
integra vel emarginata margine denticulata. Foliola infima sessilia vel subsessilia; petioli foliolis

NEW SPECIES OF RUBUS FROM WALES 155

Ficure 1. Distribution of Rubus tenuiarmatus (+) Ficure 2. Distribution of Rubus ariconiensis.
and R. iscanus (@).

Ficure 3. Distribution of Rubus vagensis. FicureE 4. Distribution of Rubus pictorum excluding
Scotland.

1660 A. NEWTON AND M. PORTER

longiores aculeis — 3 mm longis curvatis nonnullis muniti. Inflorescentia angustata elongata infime
foliis ternatis instructa ad apicem congestum aphylla, ramuli infimi et medii adscendentes 30—45°, 5—
10 flori. Rachis vix flexuosa, pilosa, glandulis stipitatis magnitudine variis, aculeolis glanduliferis
aculeis usque 5 mm longis inaequalibus tenuibus rectis vel leviter curvatis crebris praedita. Pedicelli
superiori parti rachidis similes dense pilosi et tomentosi, aculeis patulis, aculeolis glanduliferis et
glandulis stipitatis muniti.

Flores c. 2—2.5 cm diam. Sepala griseoviridia albo-marginata, hirsuta et tomentosa, glandulis
stipitatis brevibus numerosis aciculisque raris vestita, primo patentia mox fructum amplectantia.
Petala roseata parum distantia elliptica vel obovata fimbriata; stamina stylos roseos vix superantia;
anthera glabra, filamenta rosea; carpella glabra; fructus subglobosus.

Stem low-arching, angled with flat or slightly concave sides, green becoming brownish-red on
exposed sides, with sparse simple and tufted hairs, glabrescent, occasional sessile glands and
scattered stalked glands of various lengths grading into gland-tipped prickles to 4 mm; prickles
numerous, sometimes crowded, not confined to the angles, unequal, to 7 mm, the longest about as
long as the stem diameter, narrowed abruptly from a broad compressed base, patent or declining,
straight or curved, red with yellow tips.

Leaves pedate; leaflets usually 5, sometimes 3 or 4, contiguous or not, mid or yellowish green,
sparsely strigose above, with medium to dense short simple hairs beneath; terminal leaflet c. 7 x 4.5
cm, elliptical or slightly obovate with cuspidate apex c. 1 cm and entire or emarginate base, finely
but unevenly serrate occasionally with principal teeth retrorse, flat, the petiolule rather more than
1/3 as long as the lamina (av. c. 36%); basal leaflets sessile or subsessile; petiole normally longer
than basal leaflets, with about 16 slightly curved + equal prickles 2-3 mm and sometimes a few
smaller prickles. Flowering branch with 3—5 ternate leaves and usually one simple leaf above, not
leafy to apex. Inflorescence consisting of a compact terminal head of up to 10 flowers, 3—7 cm in
length, with distant ascending axillary peduncles up to 8 cm, usually shorter than their leaves,
bearing about 5—10 flowers mainly in a terminal cluster with one or two distant below; rachis almost
straight, becoming reddish on exposed side with numerous spreading and adpressed simple and
tufted hairs, numerous stalked glands from short to long grading into gland-tipped pricklets,
frequent straight or slightly curved patent or declining unequal prickles up to 5 mm; pedicels with
dense mainly spreading simple or tufted hairs, stalked glands of various lengths grading into gland-
tipped pricklets, prickles + straight, patent, ike 3 mm.

Flowers c. 2.5 cm in diameter; sepals greyish-green with white margins, with dense short simple
hairs, stellate hairs and numerous short or very short stalked glands and rare acicles, short or
medium pointed, loosely reflexed at petal fall, later erect to clasping; petals c. 14 x 9 mm, pink,
obovate or elliptical, + cuneate at base, notched or erose at apex, numerous very short hairs on
margin, usually not contiguous; stamens exceeding styles, filaments lilac-pink, anthers glabrous;
styles pink, carpels glabrous; receptacle glabrous or occasionally with a few hairs; fruit round with
few drupelets, sometimes partly abortive. Flowering from June until August.

Hototypus: hedge, Llangynidr, Brecs., v.c. 42, GR SO/162.193, 9 July 1992, M. Porter (NMW).
IsotyPi: BIRM, BM.

This bramble seems to have been noticed first by W. M. Rogers during his tour of Breconshire and
Radnorshire in 1898. At first he identified it as R. marshalli Focke & Rogers var. semiglaber and
recorded it at Llangorse Common and in several places in the upper Wye valley. In the report of his
visit (Rogers 1899) he notes that the bramble is ‘“‘widely spread in Wales and somewhat variable in
character. It also crosses the border into Herefordshire and is especially abundant at Cusop near
Hay.” Later, in his Handbook (1900), he comments “‘The Breconshire and West Herefordshire
plant is abundant in hedges and at times recalls R. dumetorum but its rather large drupelets are
neither glaucous nor caesian in flavour. It has very showy flowers with deep pink petals, stamens and
styles.’ The petals turn a deeper pink on drying.

From a study of A. Ley’s herbarium at BIRM, which includes several gatherings of this plant, it
appears that from about 1903 Rogers referred it to R. diversifolius Lindl., which at this time was
used as a collective name for several hystrican Corylifolii including R. tuberculatus Bab. However,

NEW SPECIES OF RUBUS FROM WALES 137)

as E. S. Edees (1975) has pointed out, R. diversifolius is anom. superf. illegit. for R. vestitus Weihe
& Nees: the Breconshire and West Herefordshire plant is therefore described above as a new
species.

A recent survey of its distribution has shown that R. ariconiensis occurs in v.cc. 35, 36, 42 and 43.
It is particularly abundant in that part of Herefordshire west of the River Wye which in post-Roman
times formed the Kingdom of Irching or Archenfield, thought to have derived its name from the
Roman settlement of Ariconium. The bramble is common in hedges but also occurs in woods and
thickets and reaches an altitude of 370 m on the limestone cliffs of Craig-y-cilau near Crickhowell.
When growing in shade, as at Mouse Castle near Cusop, the flowers remain deep pink but the stem
armature is much reduced. R. ariconiensis is allied to R. tuberculatus, R. vagensis and R.
tenuiarmatus.

The known distribution is shown in Fig. 2.

Rubus vagensis A. Newton & M. Porter, sp. nov.

During fieldwork to trace the distribution of R. ariconiensis a closely related bramble was
encountered in hedges in several places in Herefordshire. At first it was thought that this might be
the long-lost Rubus tenuiarmatus described by Edwin Lees in Botany of the Malvern Hills, 2nd ed.
(1852}, where he mentioned that it grew in hedges and thickets about Great Malvern. During the
following decade R. tenuiarmatus was recorded on several field meetings of both the Woolhope
Club (based in Hereford) and the Worcestershire Naturalists. Then suddenly it disappeared from
the batological scene, submerged by C. C. Babington under R. balfourianus Blox. (R. nemorosus
Hayne & Willd.). Later however he seems to have lost faith in this judgement (MS notes in LIV).

In 1993 specimens of the Herefordshire bramble collected during 1991—2 were compared with the
type specimens of R. tenuiarmatus at CGE, at the time thought to be the sole surviving herbarium
specimen, upon which the account in Brambles of the British Isles (Edees & Newton 1988) was
based. The type specimen was collected at Bromsgrove Lickey in Worcestershire (v.c. 37) in
October 1850 by William Mathews, a life-long friend of Edwin Lees. Comparison was made difficult
both by the age of the type and the fact that it was collected so late in the year. The two brambles are
indeed similar in many respects but there are significant differences which are noted in Table 1;
consequently the Herefordshire bramble is described below as Rubus vagensis. Vaga as a Latin
name for the River Wye was an invention of 16th century scholars (Bannister 1916).

TABLE 1. DIFFERENCES BETWEEN RUBUS TENUIARMATUS AND R. VAGENSIS

Character

T. S: stem

Number of largest prickles
per 5 cm of stem

Largest prickles

Terminal leaflet shape

Terminal leaflet tip

Leaf colour

Leaf margin

Number of petiole prickles
Petal colour

Filament colour

Carpels

Fruit size

R. tenuiarmatus
Almost round

(6-) 10 (-14)

Very slender
Ovate
Acuminate
Bright green
Jagged — sharply biserrate

(S—) 9 (-12)
Bright pink
White
+ glabrous, sometimes one
or two hairs
Small — medium

R. vagensis
Distinctly angled

(15—) 20 (25)
Stronger with wider base
Obovate — suborbicular
Cuspidate
Mid — dark green
Shallowly serrate with some
principal teeth retrorse
(10— 16 (-21)

Very pale lilac pink often fading to off-white
Usually lilac pink, darkening when dried

Hairy

Medium - large

138 A. NEWTON AND M. PORTER

Turio arcuato-decumbens in apricis purpurascens angulatus superficiebus planis capillis simplicibus
stellatisque vestitus, glandulis stipitatis numerosis, aciculis sparsis, aculeis aculeolisque usque 6 mm
longis, haud ad angulos limitatis rectis vel leviter curvatis patentibus vel obliquis armatus. Folia (3-)
5-nata, pedata. Foliola contigua vel imbricata superne parce strigosa inferne capillis simplicibus
stellatisque interdum vestita; foliolum terminale (c. 8 X 6 cm) late obovatum vel ellipticum apice
cuspidatum basi integra vel emarginata margine denticulata. Rachis vix flexuosa pilosa tomentosa
glandulis stipitatis numerosis aciculis sparsis aculeis aculeolisque 1-7 mm longis inaequalibus rectis
fere patentibus numerosis praedita. Inflorescentia angustata elongata ad apicem aphylla; ramuli
infimi et medii adscendentes foliis breviores, supremi pauciflores distantes. Flores 2.5—3 cm diam.
Sepala griseo-tomentosa glandulis aciculisque ornata primo laxe reflexa tandem fructum amplectan-
tia. Petala pallide roseo-lilacina elliptica vel late obovata fimbriata; stamina stylos virides vix
superantia; filamenta pallide roseo-lilacina vel alba; antherae glabrae; carpella hirsuta; fructus
subglobosus.

Stem low-arching, angled, becoming purple on exposed sides, with sparse to numerous short simple
and tufted hairs, numerous stellate hairs, sparse or numerous short and medium stalked glands and
occasional gland-tipped acicles; prickles 15—25 per 5 cm, not confined to the angles, mostly 5-6 mm
but with some short prickles 1-2 mm and sometimes a few prickles of intermediate length, straight
or curved, patent or declining, narrowed abruptly from a compressed base, purple with straw
coloured tip. Leaves pedate; leaflets (S—) 5, usually contiguous, sometimes imbricate, mid to dark
green with adpressed simple hairs above and dense short simple hairs beneath, with sometimes an
underlayer of stellate hairs; terminal leaflet c. 8 X 6 cm, broadly obovate or suborbicular, with a
cuspidate apex c. 1 cm and entire or slightly emarginate base, slightly unevenly but shallowly serrate
with some principal teeth retrorse, flat, the petiolule 1/4-1/3 as long as lamina; basal leaflets sessile
or subsessile; petiole about the same length or longer than basal leaflets with about 15 straight or
gently curved declining + equal prickles c. 3 mm.

Flowering branch with 2-4 ternate teaves below and 0-1 simple leaf above, not leafy to apex;
inflorescence consisting of a narrow, cylindrical, rather open, few-flowered terminal head c. 4-8
(—10) cm and distant ascending axillary peduncles rather shorter than their leaves; rachis nearly
straight, greyish-green or purple in colour, with spreading and adpressed short and medium simple
or tufted hairs and dense stellate hairs, frequent to numerous medium and long stalked glands and
rare acicles, numerous straight slightly declining unequal prickles up to 7 mm and a few shorter
prickles 1-3 mm; pedicels with dense mainly adpressed short simple or tufted hairs and stellate
hairs, stalked glands of various lengths but mostly longer than the indumentum, a few acicles and
straight slender + patent prickles 2-4 mm.

Flowers c. 2.5—3 cm in diameter; sepals greyish-green with dense stellate hairs and short or very
short simple hairs, numerous short medium stalked glands and some acicular pricklets, short or
medium pointed, weakly reflexed at petal fall, later erect to clasping; petals c. 14 x 9 mm pale lilac
pink, broadly elliptical or obovate, sometimes shallowly notched or erose with hairy margin;
stamens + same length or slightly longer than styles, filaments usually pale lilac rarely white, anthers
glabrous; styles pale green, carpels hairy; fruit round to oblong (15—20 mm) with 15-30 drupelets,
sometimes partly abortive. Flowering June to August.

Hototypus: edge of wood, Aconbury, Herefs., v.c. 36, GR SO/518.327, 4 July 1992, M. Porter
(NMW).
IsoryP1: BM, BIRM.

Field studies during the past three summers have shown R. vagensis to be widespread and locally
abundant in Monmouthshire (v.c. 35) and Herefordshire (v.c. 36). In a few places, e.g. Aconbury
Hill near Hereford, R. vagensis and R. ariconiensis grow close together, but generally R. vagensis
tends to replace its ally east of the River Wye and extends across Herefordshire to the Malvern Hills
and Worcestershire (v.c. 37). There are specimens collected by A. Ley in Herefordshire at BIRM
and NMW, usually determined as R. diversifolius Lindl. R. vagensis has been recorded from v.cc. 6,
35, 36, 37, 42 and 43. The known distribution is shown in Fig. 3.

|

NEW SPECIES OF RUBUS FROM WALES 139
Rubus tenuiarmatus Lees

After a careful study of the lectotype at CGE in 1993, localities in Worcestershire where Lees had
recorded R. tenuiarmatus were searched, resulting in its rediscovery in several of the old sites. In
addition further specimens collected by William Mathews came to light in the herbarium of the City
Museum, Worcester (WOS). An undated specimen collected by Lees from Great Malvern and
labelled R. diversifolius Lindl. was found at CGE (Babington no. 162). This is the only specimen of
R. tenuiarmatus collected by Lees we have seen and it probably dates from before 1850, since
following the publication of R. tenuiarmatus in 1852 R. dumetorum var. diversifolius was dropped
from the two later editions of Botany of the Malvern Hills (Lees 1852, 1868). Later in the 19th
century batologists used R. diversifolius as a collective name for at least four hystrican Corylifolii
and there is a specimen of R. ftenuiarmatus at BIRM, collected by A. Ley near Whitbourne in
Herefordshire in 1907 as R. dumetorum var. diversifolius Lindl. That it was so labelled by Ley was
doubtless due to the inclusion of R. tenuiarmatus under R. balfourianus by Babington (1869) and its
subsequent eclipse. Various comments by Lees after 1865 indicate that his concept of R.
tenuiarmatus was broadened in later years: “‘It is a general bramble, the blueish fruit showing its
affinity to R. caesius” (1867) and ‘““When very well developed this becomes the R. balfourianus of
Bloxam’”’ (1868), deferring to Babington’s view. However it is noteworthy that all the contemporary
gatherings labelled R. tenuiarmatus that have been found so far clearly belong to the same taxon and
closely match the lectotype.

The following exsiccatae have been seen, in addition to the lectotype: Bromsgrove Lickey,
Worcs., 3 October 1850, W. Mathews, WOS; Ufftmoor Lane, Halesowen, Worcs., 4 October 1850,
W. Mathews, WOS; Huddington, Worcs., 12 August 1853, W. Mathews, WOS; Churchill,
Kidderminster, Worcs., 23 August 1853, W. Mathews, WOS; Great Malvern, Worcs., undated, E.
Lees, CGE, as R. diversifolius Lindl; Whitbourne, Herefords., 6 August 1907, A. Ley, BIRM, as R.
dumetorum vat. diversifolius Lindl.

During 1993 R. tenuiarmatus was recorded from the following sites in Worcestershire (v.c. 37):
Lane near Clent, SO/926.784, A.N. & M.P., 19 July; Hollies Hili Lane near Belbroughton, SO/
928.777, M.P., 31 July; Bridle path near Churchill, SO/876.795, M.P., 4 August; Broughton Green,
SO/954.616, M.P., 25 August; Monk Wood near Sinton Green, SO/808.608, M.P., 25 August;
Huddington, SO/941.576, M.P., 25 August; Upper Broadheath, SO/797.557, M.P., 25 August.

Additional information, particularly on floral structure, has provided a clear picture of a hitherto
shadowy bramble. R. tenuiarmatus is more closely related to R. vagensis, R. ariconiensis and R.
tuberculatus than to R. nemorosus under which it was submerged by Babington (1869). Watson
(1958) equated R. triangularis (Ley) Edees with R. tenuiarmatus but the two brambles are very
different, as Edees pointed out (1975). Both can be seen in hedges in the Teme valley along the
boundary between Herefordshire and Worcestershire.

The known distribution of R. tenuiarmatus is shown in Fig. 1.

Rubus pictorum Edees

A white-flowered bramble with very prickly and glandular stems is common in the foothills of the
Black Mountains in Herefordshire and Breconshire. Herbarium studies revealed that this plant had
been collected on numerous occasions by A. Ley and usually determined as Rubus britannicus
Rogers. In his report of a field meeting of the Woolhope Field Naturalists’ Club in the Black
Mountains in 1897 Ley comments “... all through the district the hedges and bushy banks
produced abundantly the rare bramble Rubus britannicus.” W. M. Rogers recorded the plant as
fairly frequent during his visit to Breconshire and Radnorshire in 1898. In his Handbook (Rogers
1900) it is recorded as Rubus dumetorum Weihe var. britannicus Rogers, from eleven vice-counties
in England, Scotland and Wales.

W. C. R. Watson seems to have followed Rogers’ wide interpretation of the taxon, although he
mistakenly equated the Scottish representatives with his R. iodnephes of the Richmond (Surrey)
district, but in 1975 E. S. Edees restricted R. britannicus Rogers to the plant of southern England
with a specimen from Munstead, Surrey, collected by Rogers as lectotype. Following this
separation, in 1980, the Staffordshire representatives were described as a new species R. intensior

140 A. NEWTON AND M. PORTER

Edees, and two years later their Scottish allies were segregated as R. pictorum Edees. This
reorganisation left our Marches plant in limbo. Clearly it does not fit the restricted concept of R.
britannicus Rogers, as described in Edees & Newton (1988). At first it seemed most likely, on
geographical grounds, that it might be R. intensior Edees, but closer study showed our local
examples differed from R. intensior in precisely those features which Edees (1982) used to
distinguish R. pictorum from R. intensior. Comparison with the holotype and other specimens,
including some authenticated by Edees, confirmed that our plant matched R. pictorum Edees. The
striking violet-red colour of the stems is clearly shown and occasionally, as in Scottish R. pictorum,
the flowers are pale pink in bud. Although some of the Herefordshire gatherings are more robust
and prickly than much of the Scottish material examined, it is now evident that the plant from the
Welsh Marches, formerly referred to as R. britannicus Rogers, is identical with R. pictorum Edees.
However not all of Ley’s determinations of R. britannicus can be accepted as this taxon: one sheet
from Carmarthenshire (v.c. 44) in BIRM is R. hylocharis W. C. R. Watson and another from the
same area is a mixture of R. hylocharis and R. merlinii A. Newton & M. Porter.

R. pictorum Edees has now been recorded from the following additional v.cc. 6, 35, 36, 41-44,
adding an intriguing outlier to its previously known distribution, as shown on the map (Fig. 4).

Recent studies in North Wales, particularly the Lleyn peninsula, have found the following
bramble to be widespread in the area. A description follows:

Rubus segontii A. Newton & M. Porter, sp. nov.

R. dasyphyllo similis, a quo praecipue differt facie grisea, turione minus pilosa, aciculis densioribus
armata; foliis inferne primo griseo-tomentosis, foliolo terminali angustiori basi cuneata; inflores-
centia ad apicem latiore, densiore, magis florifera; petalis albis attenuatis, sepalis insigniter per
anthesin laxe amplectantibus. Copiose in Mona et Arvonia crescit.

Similar to Rubus dasyphyllus, from which it differs in details as follows: Plant with an overall greyish
cast, stem less densely hairy, more closely beset with long acicular prickles; leaves at least at first
grey-tomentose beneath, the terminal leaflet consistently narrow with a cuneate base. Inflorescence
with a more substantial ultra-axillary portion, broader, denser and more floriferous at the summit.
Petals white, narrow; sepals noticeably clasping in flower and fruit.

Ho ortypus: hedgebank, Penbodlas, Caernarvon, v.c. 49, GR SH/282.336, August 1991, A. Newton
(NMW).

Frequent in Caernarvon and Anglesey. North Merioneth. First found by Babington c. 1850 in the
Llanberis district: distributed by J. E. Griffith from Treffos and Pen-y-garnedd in Anglesey whence
it was named “‘aggregate R. hirtus” in 1884. Many specimens were collected by W. C. Barton in 1922
around Portmadoc and Tremadoc and given the MS name ‘portmadocensis’ in his notes (BM).
Widespread in the Lleyn peninsula (Conolly & Newton 1993).

ACKNOWLEDGMENTS

We are most grateful to J. R. Edmondson (LIV), R. G. Ellis and G. Hutchinson (NMW), N.
Gordon (WOS), R. N. Lester (BIRM) and P. D. Sell (CGE) for their courtesy and help in locating
exsiccatae and references, and to R. R. Mill (E) for checking the Latin descriptions.

REFERENCES

AMPHLETT, J. & REA, C. (1909). The botany of Worcestershire. Birmingham.
BaBINGTON, C. C. (1869). The British Rubi. London.
BANNISTER, A. T. (1916). The place-names of Herefordshire. Cambridge.

NEW SPECIES OF RUBUS FROM WALES 141

ConoL_y, A. & Newron, A. (1992). Rubus in West Lleyn 1990 and 1991. Welsh bulletin of the B.S.B.I. 54:
12-15.

Epees, E. S. (1975). Notes on British Rubi, 3. Watsonia 10: 331-343.

EpeEs, E. S. (1980). Notes on British Rubi, 6. Watsonia 13: 31-34.

EpEEs, E. S. (1982). A new bramble from Scotland. Watsonia 14: 180-182.

Epegs, E. S. & Newron, A. (1988). Brambles of the British Isles. London.

Legs, E. (1852). Botany of the Malvern Hills, 2nd ed., p. 51. London.

Legs, E. (1853). A descriptive list of the British Rubi. Phytologist 4: 818.

LEEs, E. (1867). The botany of Worcestershire. Worcester.

LEES, E. (1868). Botany of the Malvern Hills, 3rd ed., pp. 67-68. London.

Ley, A. (1898). Botanical notes. Transactions of the Woolhope Naturalists’ Field Club: 268.

Ley, A. (1902). Two fresh Rubus forms. Journal of botany (London) 40: 69.

Ley, A. (1905). Notes additional to the Flora of Herefordshire. Transactions of the Woolhope Naturalists’ Field
Club: 69-116.

MatTHeEws, W. (1899). Midland County plants observed .. . 1849-1884. Transactions of the Worcestershire
Naturalists’ Club I.

Purcuas, W. H. & Ley, A. (1889). A Flora of Herefordshire. Hereford.

Rocers, W. M. (1894). Rubi notes. Journal of botany (London) 32: 49.

Rocers, W. M. (1895). On the Rubi list in the London Catalogue 9. Journal of botany (London) 33: 103.

Rocers, W. M. (1899). Radnorshire and Breconshire Rubi. Journal of botany (London) 37: 193-198.

Rocers, W. M. (1900). Handbook of British Rubi. London.

Watson, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland. Cambridge.

(Accepted December 1993)

Watsonia 20: 143-151 (1994) 143

Four new Anglo-Sarnian species of Rubus L. (Rosaceae)

D. E. ALLEN

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

ABSTRACT

Four new species of Rubus L. (Rosaceae) are described, all with ranges extending to both sides of the English
Channel. R. transmarinus D. E. Allen, sp. nov. (sect. Corylifolii Lindley), locally common round Bournemouth
and in the Isle of Wight and found widely across Hampshire, also occurs in Jersey and Sark and on coastal heaths
in France east of Cherbourg. R. caesarius D. E. Allen, sp. nov. (ser. Vestiti (Focke) Focke), abundant in many
parts of Jersey, is also known in one English locality, between Southampton and Portsmouth. R. couchii
Rilstone ex D. E. Allen, sp. nov. (ser. Sylvatici (P. J. Mueller) Focke) is frequent in the extreme south-east of
Cornwall and common in the north-east of Jersey; it was also distributed last century from Cherbourg. R.
percrispus D. E. Allen & R. D. Randall, sp. nov. (ser. Micantes Sudre ex Bouvet) is known from eight vice-
counties in the southern half of England, with concentrations round Colchester and Bristol. There are also single
records of it for Jersey and Guernsey.

Keyworps: Channel Islands, England, France, brambles, apomictic species.

INTRODUCTION

Intensive investigation in recent years of the Rubus flora of the Channel Islands and of that part of
France most immediately adjacent to them, the Cotentin Peninsula of Normandy, has brought tc
light numerous forms that it has not proved possible to match with published taxa. This was no mor:
than to be expected, for ail areas in which Rubus fruticosus agg. occurs in any diversity tend to hav
a proportion of putative local endemics, most of them too restricted in quantity and range to warran
adding to the formidable number of species described already in this group. In order to keep the
taxonomy within manageable limits, it is now the convention among Rubus specialists to bestow
names on only those forms that reveal a ‘regional’ (as opposed to merely ‘local’) distribution. The
four described in this paper not only fall into this category, but qualify additionally by reason of their
occurrence on both sides of the English Channel. Britain and France have many Rubus species in
common, but an increasing number are being detected with ranges that (so far at least) seem to be
only narrowly trans-Channel in character. Three examples are R. boulayi Sudre (abundant round
Bournemouth), R. thyrsigeriformis (Sudre) D. E. Allen (locally abundant round Southampton) and
R. corbieri Boulay (locally abundant in the Purbeck district of Dorset). Each of these is well-
distributed in the coastal area of France more or less opposite the main English concentrations,
namely the Cotentin Peninsula, but only the first has been recorded outside that strictly limited
territory (and whether correctly needs to be confirmed). Three of the species described below have
ranges of broadly this same pattern, though one has its English concentration much further to the
west. The fourth is much more widespread in England and, so far as is known, has been found only
twice on the other side of the Channel, on both occasions seemingly transiently.

DESCRIPTIONS OF NEW SPECIES
1. Rubus transmarinus D. E. Allen, sp. nov.

Turio arcuato-decumbens, obtusangulus, rubro-purpureus, sparsim pilosus, aculeolis saepe glandu-

144 D. E. ALLEN

liferis et aciculis et glandulis stipitatis numerosis et aculeis numerosis vel densis non ad angulos
dispositis inaequalibus rectis e basi plana declinatisque vel curvatis tenuibus munitus. Folia 5-nata,
pedata, imbricata, superne sparsim pilosa, subtus molliter pubescentia, quorum petioli aculeis (2-3
mm) numerosis declinatis vel leviter falcatis tenuibus et aculeolis et aciculis et glandulis plerumque
brevibus muniti. Foliolum terminale subrotundum vel rotundo-ovatum, paullatim acuminatum,
serratum dentibus nonnullis aduncis, planum, basi emarginata, eiusdem petiolulo ter longius.
Inflorescentia usque ad apicem foliata, superne saepe condensata, inferne ramulis axillaribus
distantibus adscendentibus aucta. Rachis vix flexuosa, supra copiose pilosa, aculeis (4-6 mm)
tenuibus subaequantibus plerumque rectis vel declinatis et aciculis et glandulosis numerosis munita.
Flores 2-3 cm diametro. Sepala griseoviridia, dense pilosa, leviter aculeolata, glandulifera,
attenuata, albo-marginata, patula tandem reflexa. Petala alba vel subroseola, obovata vel
subrotunda, ad marginem sparsim pilosa, integra, plana, non contigua. Stamina alba stylos flavos
superantia. Antherae et carpella glabri vel pilis raris praediti. Receptaculum glabrum. Fructus
rotundus.

Stem low-arching, bluntly angled, reddish-purple, with sparse medium simple and tufted hairs,
numerous stalked glands and acicles and often also numerous gland-tipped pricklets of varying
lengths; prickles numerous to crowded, not confined to the angles, unequal, the longest c. 6-9 mm,
as long as the stem diameter or a little longer, patent or slanting or curved, slender, subulate from a
narrow compressed base, with yellowish points. Leaves pedate; leaflets 5, imbricate, pale green
with sparse short simple hairs above and soft beneath with a dense layer of short simple and tufted
hairs; terminal leaflet c. 6-9 x 5-7-5 cm, subrotund or roundish ovate or (rarely) obovate, with a
gradually acuminate apex c. 1-5 cm and emarginate base, unevenly serrate with some teeth retrorse,
flat, the petiolule 3 as long as the lamina; basal leaflets subsessile; petiole as long as or a little longer
than the basal leaflets, with few to many medium simple and tufted hairs, scattered short acicles and
mostly short stalked glands, and 15—25 declining or slightly curved prickles 2-3 mm together with a
varying number of pricklets. Flowering branch with 3-foliolate and simple (trifid and entire) leaves
extending nearly to the apex; inflorescence consisting of a flat or rounded usually condensed top and
distant ascending 4~7-flowered axillary peduncles 3-7 cm 3 as long as their leaves, the lowest
sometimes much longer and divaricate; rachis slightly flexuose or + straight, with numerous short to
medium simple and tufted hairs, numerous mostly short and medium stalked glands, very short to
medium acicles and numerous mostly subequal patent or declining slender prickles 4-6 (—7) mm.
Flowers 2—3 cm in diameter; sepals greyish-green, with dense short and medium tufted and simple
hairs, few to many short stalked and sessile glands and scattered short acicles, white-bordered, long-
pointed, patent at first then reflexed; petals 9-10 x 5-5-9 mm, 5 or 6, white or pinkish, obovate to
subrotund, with sparse to very sparse short simple and tufted hairs on the margin, entire, flat, not
contiguous; stamens exceeding the styles, filaments white, anthers glabrous or with an occasional
hair; styles yellow; young carpels glabrous or slightly hairy; receptable glabrous; fruit globose.
Flowering from June to August.

Ho.otypus: hedge, Whitemoor Lane, near Copythorne, SU/323.149, S. Hants., v.c. 11, 26 June
1982, D. E. Allen s.n. (BM).

This distinctive member of section Corylifolii has been known to British batologists since the 1890s,
when it was distributed by W. Moyle Rogers from Bournemouth, S. Hants., v.c. 11, as no. 49 in the
historic Set of British Rubi, under the erroneous name of ‘R. dumetorum var. ferox Weihe’. There
is, however, a specimen in BM from nearby Poole in Dorset, v.c. 9, labelled ‘R. fuscoater’ , collected
by T. Bell Salter as early as 1837. It is locally plentiful in the Bournemouth area and parts of the Isle
of Wight, v.c. 10, and extends widely across S. Hants., v.c. 11, and here and there into N. Hants.,
v.c. 12, as well, exhibiting a preference for dry, sunny heaths and commons (though not averse to
chalk near the coast). It is also present in some quantity in at least two of the Channel Islands, Jersey
and Sark, and may well be in Guernsey too, for Rogers & Rogers (1898) recorded ‘R. dumetorum
var. ferox’ as generally distributed there; Rogers, though, is known to have included other members
of section Corylifolii under that name and this statement has yet to be confirmed. It is, however,
absent for certain from Alderney, where its place is taken by R. tuberculatus Bab. In addition, I have

FOUR NEW SPECIES OF RUBUS L. 145

collected it in a chain of French localities, on and near the coastal heaths running east from
Cherbourg (Le Becquet; Lande de Carneville; Petit Bois, Parc du Chateau, St-Pierre-Eglise:
vouchers deposited in BM). Rogers (1900) claimed to have seen ‘R. dumetorum var. ferox’ also in
both Normandy and Brittany, presumably on the visit he is known to have made to small parts of
each of these in 1897; but he appears to have passed far to the south of Cherbourg en route to Rouen
and, again, his interpretation of the taxon is in any case liable to have been over-broad. That R.
transmarinus may not occur in France more widely is suggested by the fact that Sudre (1904, 1913),
after examining Rubus material from all parts of that country, regarded it as a form restricted to
Britain, combining it with R. rubriflorus Purchas (no. 131 of the Set of British Rubi), a quite
different bramble of the North Midlands, under the new name of R. dasyphylloides — which is
consequently illegitimate —as the supposed product of recent crossing between R. caesius L. and R.
dasyphyllus (Rogers) E. S. Marshall. Watson (1958) was the first to give specific rank to Rogers’
taxon as an entity in its own right, but unfortunately he referred it, erroneously, to the apparently
non-British R. scabrosus P. J. Mueller (Edees 1975). It features as ‘H39’ in my forthcoming account
of Rubus in Hampshire (Allen, in press).

Unlike most other members of section Corylifolii, R. transmarinus has a characteristically neat
appearance. When extreme, its dense-topped panicle, neat-looking flowers, crowded slender
prickles and roundish leaflets render it unmistakable.

2. Rubus caesarius D. E. Allen, sp. nov.

Turio arcuato-decumbens, atropurpureus, obtusangulus, faciebus planis vel excavatis, striatis; pilis
paucis vel numerosis, aculeolis nonnullis, glandulis stipitatis aciculisque brevibus paucis, aculeis
crebris ad angulos dispositis inaequalibus e basi lata et compressa tenuibus declinatis vel curvatis
rubro-purpureis munitus. Folia 5-nata, digitata, saepe imbricata, superne glabrescentia, subtus
molliter cinereo-albo-tomentosa, dense pilosa. Foliolum terminale ovatum vel subrotundum, apice
cuspidatum vel acuminatum, basi emarginatum vel subintegrum, inaequaliter et saepe acute
serratum vel dentatum, planum, eiusdem petiolulo ter longius; foliola infima breviter pedicellata.
Inflorescentia usque ad apicem non foliata, longa, superne angusta et thyrsoidea vel lata et
pyramidalis, ramulis axillaribus omnibus patulis vel inferioribus adscendentibus aucta. Rachis
parum flexuosa, cinerea vel cinereo-atropurpurea, copiose pilosa, aciculis et glandulis stipitatis et
aculeolis (interdum glanduliferis) paucis, aculeis e basi longa tenuibus patulis vel declinatis vel
curvatis purpureis numerosis munita. Flores c. 2-2-5 cm diametro. Sepala cinereo-tomentosa,
glandulis stipitatis aciculisque paucis praedita, semper reflexa. Petala dilute rosea, elliptico-obovata
vel subrotunda, ad marginem sparse pilosa, integra, plana, non contigua. Stamina alba stylos virides
vix superantia. Antherae glabrae. Carpella pilosa. Receptaculum glabrum. Fructus parvus rotundus.

Stem low-arching, dark purple, blunt-angled with flat or furrowed sides, striate, with few to many
very short to medium simple and tufted hairs, numerous sessile and subsessile glands, a few pricklets
and scattered or rare short stalked glands and short acicles; prickles 10-14 per 5 cm, confined to the
angles, unequal, (3—)4-6 mm, slender from a rather broad compressed base, declining or curved,
reddish purple with pale point. Leaves digitate; leaflets 5, often imbricate, pale or dark green,
glabrescent above, softly greyish-white-felted beneath with dense short simple and tufted hairs;
terminal leaflet 5-7(-10) x 4-6(—7-5) cm, ovate or subrotund, with a cuspidate or acuminate apex
0-5—1-5 cm and an emarginate or subentire base, unequally and often finely serrate or dentate with
the principal teeth sometimes patent, flat, the petiolule 3 as long as the lamina; petiolules of basal
leaflets 1-4 mm; petiole as long as or a little longer than the basal leaflets, coloured and clothed like
the stem, with 8-11 curved prickles 2-4 mm. Flowering branch with 3- foliolate leaves below and 0-4
simple leaves above, not leafy to the apex; inflorescence long, with a dense truncate top, sometimes
narrow and thyrsoid above, sometimes broad and pyramidal, the upper, middle and sometimes
even the lowest peduncles patent, more often the lower peduncles ascending and sometimes longer
than their leaves, all divided about half-way or well above the middle; rachis slightly flexuous,
greyish or dark greyish-purple, with numerous adpressed and patent short to medium simple and
tufted hairs, sparse sessile glands, scattered purple short and medium stalked glands, rarer short
acicles and (sometimes gland-tipped) pricklets, and numerous long-based, slender-pointed, patent

146 D. E. ALLEN

or declining or curved purple prickles 2-6 mm; pedicels clothed like the upper part of the rachis,
with numerous slender declining or curved prickles i-2 mm. Flowers 2-2-5 cm in diameter; sepals
grey with dense stellate hairs and few to many medium simple and tufted hairs, numerous sessile and
scattered short stalked glands and short acicles, short- or long-pointed, reflexed; petals 9-10 x 5-6
mm, pale pink, elliptical-cbovate to subrotund, entire, with sparse very short simple and tufted
hairs on the margin, flat, not contiguous; stamens equalling or slightly exceeding styles, filaments
white, anthers glabrous; styles pale green; young carpels hairy; receptacle glabrous; fruit small,
globose. Flowering in July and August.

Ho .ortypus: heathy scrub, Le Jardin d’Olivet, par. Trinity, WV/673.543, Jersey, v.c. S, 1 July 1990,
D. E. Allen s.n. (BM).

The epithet caesarius, also borne by the subspecies of Bank Vole endemic to Jersey, Clethrionomys
glareolus caesarius (Miller), is taken from the alleged Roman name for the island, Caesarea.

R. caesarius is not only a characteristic member of series Vestiti, but was actually mistaken by
Rogers for R. vestitus Weihe (MS note by him on sheet ex herb. Barton & Riddelsdell no. 9318 in
BM, where he omits the qualifying “but usually untypical” of Rogers & Rogers 1898), a species now
known to be absent from the Channel Islands, the Cornish Peninsula and extreme north-west
France. The two are in fact readily distinguished, and it would seem that Rogers played down the
differences evidently in a wish to satisfy himself that the Rubus flora of the Channel Islands is
broadly similar to that of southern England (which, however, has turned out to be anything but the
case). As he noted, this bramble is second only in abundance in Jersey to the ubiquitous R.
ulmifolius Schott, occurring in profusion on banks, roadsides and heathland in many parts (e.g. La
Gréve de Lecq; Bouley Bay; Rue de I’Aleval; Val de la Mare), though scarce or absent in others,
most noticeably in those which harbour the putatively oldest substratum of Rubus species (as at the
top of Waterworks Valley). This suggests that it may be a relatively late incomer, a possibility
rendered the more likely by its prevalence in Jersey despite its apparently total absence from all the
other Channel Islands. It can hardly, however, have come from Britain, for only a single small
population is known from there: midway between Southampton and Portsmouth, on hedgebanks
along Segensworth Road, Park Gate, SU/526.078-9, S. Hants., v.c. 11, at the north end of the one-
time Titchfield Common (a surviving fragment of which, further south, holds the only-known
Hampshire patch of R. imbricatus Hort). On the other hand, unlike R. corbieri (Allen 1992), it
would appear not to have come from the part of the French mainland that is closest either, for no
sign of it has been met with in the Cotentin Peninsula after three seasons’ extensive sampling of that
area’s Rubus florula. Should it be a species of French origin, therefore, the source must lie further to
the east or south. Although no match for it has been found among the limited French material in
British herbaria, it may yet turn out to have been among the numerous taxa described by French
_ botanists prior to 1914, type material of many of which cannot now be located.

A representative series of R. caesarius from Jersey and Hampshire has been deposited in BM. In
OXF there is a stem-piece of it in a mixed gathering by Rogers from Les Marais and Fauvic,
misdetermined as a weak form of R. radula Weihe ex Boenn. There is also in OXF an undated
specimen of G.C. Druce’s from St Aubin, variously misdetermined as R. vestitus and R. boraeanus
Genev., that may be this too. Otherwise no previous specimens have been detected in British
herbaria.

3. Rubus couchii Rilstone ex D. E. Allen, sp. nov.

Turio alte arcuatus fuscus vel nigrescens, obtuse vel acute angulatus, faciebus planis vel concavis,
striatis, glaber vel pilis paucis vel numerosis, aculeis crebris ad angulos dispositis saepe geminatis
inaequalibus e basi lata et compressa attenuatis patulis vel declinatis vel curvatis purpureis munitus.
Folia 5-nata, pedata vel digitata, vix imbricata, superne glabrescentia, subtus viridia (raro cinereo-
tomentosa), leviter pilosa. Foliolum terminale subrotundum vel elliptico-obovatum vel oblongum,
apice acuminatum, basi emarginatum vel integrum, inaequaliter acute serratum et dentatum,
planum, eiusdem petiolulo ter longius; foliola infima breviter pedicellata. Inflorescentia usque ad

FOUR NEW SPECIES OF RUBUS L. 147

apicem non foliata, foliis nonnullis quinatis ternatisque inferne instructa, saepe longa, anguste
pyramidalis truncata, ramulis axillaribus infimis distantibus adscendentibus et mediis subpatulis vel
divaricatis aucta. Rachis flexuosa vel recta, infra sicut turio vestita, ad apicem pilis densis
longioribus, glandulis stipitatis brevissimis paucis et aculeis purpureis numerosis declinatis vel
curvatis vel falcatis 2-6 mm munita. Flores c. 2-5-3 cm diametro. Sepala cinereo-tomentosa, pilis
flavescentibus numerosis vestita, aculeolata, anguste albo-marginata, omnia attenuata, patula vel
reflexa tandem fructum amplectantia. Petala rosea vel roseola, subrotunda vel late ovata vel
oblonga, subtus et ad marginem pilosa, saepe imbricata. Stamina alba stylos virides vel flavescentes,
saepe basi erubescentes, superantia. Antherae glabrae vel pilis raris praeditae. Carpella dense
pilosa. Receptaculum glabrum. Fructus rotundus.

Stem high-arching, light brown to blackish, bluntly to rather sharply angled, with flat or shallowly
furrowed sides, striate, glabrous or with few to many short and medium simple and tufted hairs,
numerous sessile glands and very rare pricklets; prickles 11-15 per 5 cm, confined to the angles,
often in pairs, unequal, the longest c. 6-8 mm, as long as or rather shorter than the stem diameter,
patent or slanting or curved, narrowing from a broad compressed base, purple with yellowish tip.
Leaves pedate or digitate; leaflets 5, not or scarcely overlapping, dark green with sparse simple hairs
above and green (rarely, greyish-white) beneath with few to many medium simple hairs mostly on
the veins; terminal leaflet c. 6-10 x 4~7 cm, subrotund or elliptical-obovate or oblong, with a
gradually acuminate apex c. 0-5—1-5 cm and entire or emarginate base, unevenly and finely serrate-
dentate with some principal teeth patent, flat, the petiolule c. 3 as long as the lamina; petiolules of
basal leaflets 1-6 mm; petiole usually well exceeding the basal leaflets, with few to many short and
medium simple and tufted hairs, rare subsessile glands and 12-20 curved prickles. Flowering branch
with 3—5-foliolate leaves below and 1-4 simple (trifid and entire) leaves above extending to usually
well short of the apex; inflorescence often large, “usually somewhat narrowly pyramidal! and
truncate” (Rilstone ms.), with distant ascending 5—6-flowered lower axillary peduncles 4-10 cm as
long as their leaves to only half as long and subpatent or divaricate middle peduncles; rachis flexuose
or straight, with many mostly adpressed simple and tufted hairs becoming dense, more spreading
and longer above together with numerous sessile and subsessile glands, a few short stalked glands
and numerous declining or curved or falcate prickles 2-6 mm. Flowers largish, 2-5—3 cm in diameter;
sepals greyish, with dense yellowish short and medium tufted hairs, numerous sessile glands and few
to many short and medium acicles, narrowly white-bordered, all long-pointed from bud, in flower
subpatent or reflexed, in fruit reflexed and ultimately erect and clasping; petals 8-12 x 5—9 mm, mid
or pale pink, entire or erose, subrotund to broadly oval or oblong, not or scarcely clawed, with
numerous hairs beneath and few to many short and medium simple and tufted hairs on the margin,
moderately concave, spaced or overlapping; stamens usually exceeding the styles, filaments white or
faintly pink, anthers glabrous or with a rare hair; styles green or yellowish, often red-based; young
carpels densely hair; receptacle glabrous; fruit globose. Flowering from June to August.

Ho orypus: Polperro, E. Cornwall, v.c. 2, 10 September 1932, F. Rilstone no. 793, herb. Barton &
Riddelsdell no. 7782 (BM).

This bramble was first brought to notice in Britain in 1920 by Rilstone, who distributed specimens of
it from around his then home village of Polperro through the Botanical Exchange Club, remarking
that it was “‘a common and constant plant over this locality’. On that occasion H. J. Riddelsdell
determined it as R. macrophylloides Genev.; after seeing it for himself in the field in 1924-25,
however, he queried it instead as R. leucanthemus P. J. Mueller before settling for R. hirtifolius P. J.
Mueller & Wirtgen. All three of these taxa were at that time mistakenly thought to be British.
Through some confusion Riddelsdeil meanwhile led Rilstone into believing that the revised
determination was R. chrysoxylon (Rogers) Rogers, a species of north-west Wales and the north
end of the Welsh Marches. Unable to accept this, Rilstone used “Cornish chrysoxylon” as a working
name for what he increasingly became convinced was an undescribed species and eventually, in
1946, annotated the sheets of it in his herbarium with the MS name “R. couchii’’ — in honour of the
celebrated nineteenth-century Polperro naturalist Jonathan Couch (1789-1870). In that same year,
in a letter to W. C. Barton (the relevant extract of which the latter subsequently attached to the

148 D..E. ALLEN

sheet which now bears the holotype), Rilstone wrote: “‘It is frequent rather than abundant, but I
have seen it in a good many places in Polperro valley and in the east and west Looe valleys, almost
always on the edge of thickets in the bottom of the valley.” Presumably because of this very local
distribution, in the far south-east corner of Cornwall, he did not include it among the several new
Cornubian species whose descriptions he was to publish four years later (Rilstone 1950).

Were that indeed the full extent of its range, R. couchii would still be undeserving of a published
name. In 1992, however, I realised that it was identical with a bramble known to me for some years
in the north-east corner of Jersey, v.c. S. Common and widespread in the district between Bouley
Bay and Flicquet Bay, it rises to abundance in Rozel Woods (WV/69.53, 69.54, 70.52). Though
evidently preferring shade, it occurs also on sunny banks and roadsides and among gorse scrub on
heathland. Representative material from four different Jersey localities has been deposited in BM,
to join the considerable number of gatherings of the species therein from Cornwall made at different
times by Rilstone, Riddelsdell, Barton and myself respectively.

Like R. caesarius, R. couchii appears not to occur in any of the other Channel Islands; it differs
from it, however, in extending to the neighbouring Cotentin Peninsula — or at least in having done so
formerly. For no. 893 of the specimens distributed late last century through the Association
Rubologique (a set unfortunately not in any British herbarium but which I have been able to
examine in Paris (P) and Geneva (G)) is clearly R. couchii too. It was collected by the Cherbourg
botanist L. Corbiére in 1889 from a shady hedge near the farm of Le Maupas on what was then the
outskirts of that city (but now largely built over). Although put out under the name of R. obvallatus
Boulay & Gillot, it does not match other material so named in various herbaria and the characters
noted by Corbiére on the label in fact fit R. couchii impressively exactly. There is a handwritten
addition to these in the sheet in G to the effect that “l’indument en brillant assez semblable celui du
vestitus.”’ If the species is still to be found in the Cotentin Peninsula, it would appear to be at best
very local, for in my own extensive sampling of the area I have failed to encounter it.

In BM there is a specimen collected by G. Didier (no. 1010) in the Forét de Verriéres, dép. Seine-
et-Oise, in 1933 and labelled “‘R. elongatispinus Sudre var.”’ which, while patently distinct from the
Batotheca Europaea material of that Pyrenean species distributed by Sudre, closely approaches R.
couchii. It differs, however, in having fewer and longer prickles and in lacking the yellowish
tomentum on the sepals and upper rachis.

With its long, narrow pyramidal inflorescence and pale pink flowers, R. couchii superficially
resembles R. phaeocarpus W. C. R. Watson. Unlike that species, however, it is more or less
eglandular; the larger flowers and finely serrate-dentate leaves additionally distinguish it from that
at once. In the numerous curved prickles, especially on the rachis and peduncles, it also recalls R.
crudelis W. C. R. Watson, though readily told from that as well by its much shorter axillary
peduncles, differently-shaped leaflets and glabrous anthers. It seems best placed in series Sylvatici,
even though the leaves of at least the inflorescence are sometimes grey-felted beneath.

4. Rubus percrispus D. E. Allen & R. D. Randall, sp. nov.

Turio arcuatus, acute angulatus, faciebus planis vel leviter excavatis, atropurpureus vel purpureo-
erubescens, parce pruinosus, sparsim pilosus vel glabrescens, aciculis glandulisque pluribus
inaequalibus, aculeis numerosis 5—20 per 5 cm inaequalibus plerumque ad angulos dispositis rectis
tenuibus e basi lata vel compressa patentibus vel declinatis, aculeolis nonnullis munitus. Folia 3—5-
nata, subdigitata undulata, inferne albo-tomentosa et pilosa. Foliolum terminale 5—8 x 4—5 cm,
obovatum vel subrotundum vel ovatum, apice cuspidatum vel acutum vel acuminatum, acutissime
et tenuissime serrulato-denticulatum; foliola infima breviter pedicellata. Inflorescentia longa
anguste pyramidalis, foliosa inferne foliis 3—-4-natis et bracteis trilobatis vel simplicibus instructa,
ramulis axillaribus longis superioribus adscendentibus aucta. Rachis + recta vel parum nutans,
superne dense tomentosa, glandulis numerosis brevibus aculeisque inaequalibus tenuibus declinatis
1-9 mm munita. Flores c. 2-2-5 cm diametro; sepala albescens-tomentosa, glandulifera, semper
valde reflexa. Petala alba, late elliptica vel oblonga vel obovata, adscendens, unguiculata, saepe
emarginata, non contigua. Stamina alba stylos flavescentes vel viridescentes superantia. Antherae
glabrae. Carpella glabra. Receptaculum + glabrum. Fructus parvus rotundus.

FOUR NEW SPECIES OF RUBUS L. 149

Stem arching, sharply angled with flat or slightly furrowed sides, dull blackish- or reddish-purple,
striate, somewhat pruinose, with a few simple and tufted hairs or sometimes glabrescent; prickles
numerous, 5—20 per 5 cm, unequal, 3-10 mm, patent or slightly declining, mostly confined to the
angles, straight, subulate from a broadish or compressed base, usually accompanied by a few
pricklets and more numerous unequal acicles and stalked glands. Leaves subdigitate; leaflets 3-5,
dark green and + glabrous above, white-felted and pilose beneath, undulate on the margins, very
deeply and finely serrate-dentate with some of the principal teeth recurved; terminal leaflet 5—8 x 4—
5 cm, varying from obovate-cuspidate or roundish- or rhomboid-acute towards the stem base to
ovate-acuminate towards the stem apex; petiolules of basal leaflets usually less than 3 mm; petiole
1—1.5 times as long as the basal leaflets with 8-18 declining and slightly curved prickles 2-5 mm;
stipules and sometimes leaf margins deep red. Flowering branch with 34-foliolate leaves below;
inflorescence long, narrowly pyramidal, leafy with trifid and simple bracts when well-developed,
cymose-racemose, peduncles long, the upper ones ascending, 2—3-flowered, overtopping the central
axis. Rachis + straight or somewhat nodding, angled or furrowed below, felted above with dense
simple and stellate hairs and abundant short stalked glands (some longer than the hairs), usually
almost unarmed towards the apex but with unequal, slender, slanting prickles 1-9 mm below.
Flowers c. 2-2-5 cm in diameter; sepals whitish-felted, usually short-pointed, dotted with very short
stalked glands, strongly reflexed throughout; petals 8-17 x 5—9 mm, faintly mauve or lilac in bud,
Opening pure white, ascending, broad elliptical or oblong or obovate, clawed, often notched, not
contiguous; stamens exceeding the styles, filaments white, anthers glabrous; styles yellowish or
greenish; young carpels glabrous; receptacle inconspicuously hirsute at the base; fruit small,
globose. Flowering in July and August.

Hotorypus: hedgebank, Thorrington, TM/095.213, N. Essex, v.c. 19, 16 July 1971, J. R. I. Wood
no. 71/99 (BM).

Representative exsiccatae:

v.c. S, Jersey: old quarry, Le Jardin d’Olivet, WV/672.544, 23 July 1978 (gone by 1990), D.E.A. s.n.
(BM).

Guernsey: Fermain Bay, WV/33.76, 30 June 1897, F. A. Rogers s.n. (K).

v.c. 4, N. Devon: hedgerow, Kingsland Barton, South Molton, SS/697.257, 26 July 1989,
R.D. Randall no. 89.46 (herb. R.D.R.).

v.c. 5, S. Somerset: by bridge, Dulverton, SS/91.27, 4 August 1976, L. J. Margetts no. 99 (NMW:
herpes. dees no: 23122),

v.c. 6, N. Somerset: Stanton Wick, ST/613.626, 16 August 1986; near Publow Wood, ST/625.640, 16
August 1986; Sandy Lane, Easton-in-Gordano, ST/512.744, 7 July 1988; junction of A368 and
A37, Chelwood, ST/624.618, 29 July 1989; Young Woods Farm, Nailsea, ST/443.694, R.D.R.
nos. 86.29, 86.32, 88.11, 89.77, 90.24 respectively (herb. R.D.R.).

v.c. 11, S. Hants.: in a holly on B3055, Brockenhurst, SU/297.018, 12 July 1986; wooded roadside,
South Baddesley, SZ/350.960, 4 August 1987; Agar’s Lane, between Sway and Hordle, SZ/
273.970, 17 June 1989, D.E.A. s.n. (BM). Roadside park, Wick, near Christchurch, S$Z/152.922,
13 July 1991, R.D.R. s.n. (herb. A. D. Marshall).

v.c. 19, N. Essex: Great Horkesley, TL/98.30, 1 September 1890, J. D. Gray s.n. (BRISTM).
Heathland below ballast pits, Stanway, TL/960.246, 27 June 1920, G..C. Brown s.n. (BM).
Hedgebank, St Osyth, TM/145.205, 17 July 1972; Stour Wood, Wrabness, TM/187.307, 1 August
1972, J. R. I. Wood nos. 72.57, 72.119 respectively (BM). Scrub, Fordham Heath, TL/94.26, 23
July 1982, A. L. Bull & J. J. Heath no. 1296 (CLR; herb. A.L.B.); 1 August 1991, R.D.R. (herb.
R.D.R.).

v.c. 25, E. Suffolk: Dodnash Wood, Bentley, TM/107.364, 19 July 1972, J. R. I. Wood no. 72/81
(BW). Lady Anne Wake-Walker’s field, East Bergholt, TM/06.35, 24 July 1973, E. S. Edees no.
20880 (NMW).

150 D. E. ALLEN

v.c. 30, Beds.: Folly Wood, near Flitwick, TL/04.35, 5 August 1956, E. S. Edees no. 12099 (NMW).

v.c. 34, W. Gloucs.: old Midland Railway, Oldland Common, ST/677.723, 13 July 1987, R.D:R. no.
87.55 (herb. R.D.R.); ST/670.710, 19 July 1990, R.D.R. no. 90.34 (herb. R.D.R.; herb. L. J.
Margetts; herb. R. W. Gould).

In the early 1970s J. R. Ironside-Wood, in preparing the account of Rubus which he contributed
to the new Flora of Essex (Jermyn 1974), came across an unfamiliar glandular bramble on
hedgebanks around Colchester, taking examples of it for his herbarium from four localities in N.
Essex, v.c. 19, and one in E. Suffolk, v.c. 25. He noted that it resembled R. raduloides (Rogers)
Sudre, but that it differed from that most noticeably in having white petals and a much less vigorous
habit. Unbeknown to him, it had in fact been collected in that same district twice previously,
respectively 52 and 82 years earlier. On the second of those occasions it had been queried by
Riddelsdell as R. griffithianus Rogers, a species at that time not well understood.

A decade later R. D. Randall began finding what was subsequently to prove to be this same
bramble in numerous, widely-scattered localities in N. Somerset, v.c. 6, and W. Gloucs., v.c. 34.
Again, R. raduloides was noted as the species which it resembled most closely (and to which
specimens from sunny spots were often referred), though some resemblance to R. echinatoides
(Rogers) Dallman was also apparent, while in semi-shade it was seen to approach R. dentatifolius
(Briggs) W. C. R. Watson.

On being shown a specimen by R.D.R. in 1990, I immediately recognised it as an unnamed
bramble that I too had known for some years (as ““H589’’), having collected it in four widely-
separated localities in S. Hants., v.c. 11, and, earlier still, in one place in Jersey. About the same
time it was established that A. L. Bull and J. J. Heath had also collected it in N. Essex, and a
Guernsey specimen was discovered in K which had been collected late last century and referred by
Rogers successively to R. anglosaxonicus Gelert (now R. micans Godron) and R. rudis Weihe.
R.D.R. and Buil have since studied the bramble round Colchester, and it has also been shown in the
field to several other Rubus specialists. The general conclusion is that it does not correspond to any
species so far described from the British Isles. Although its presence in the Channel Islands and the
pattern of its Hampshire occurrences (isolated bushes or patches scattered across the south of the
county, mainly near the coast) strongly suggest that its range may extend to France, no sign of it has
been found in the Cotentin Peninsula of Normandy and a search of Continental Rubus material in
British herbaria as well as in Geneva (G) and Leyden (L) has proved negative as well. The Belgian
specialist H. Vannerom has also been unable to come forward with any Low Countries bramble that
convincingly matches the British Isles plant.

A member of series Micantes, R. percrispus favours hedgerows and scrub on sandy or gravelly
soils, occasionally also occurring in cpen woods but there tending to keep to the margins or rides.
When growing in a dry, sunny situation the armature is well-developed and reminiscent of that of R.
raduloides and R. heterobelus W. C. R. Watson, but the panicle is at the same time small, more or
less cylindrical and lacks the characteristic leafy bracts. In moister conditions the pliant becomes
much more robust and the panicles well-developed, while in shade the prickles are less frequent and
the secondary armature and glandular development very much reduced.

The epithet percrispus refers to the strikingly undulate leaf margins, which are rendered
additionally distinctive by their very deep and fine teeth. The dark green upper side of the leaflets
stands out in marked contrast to the white under side, which in turn forms a strong contrast to the
dark purple stem and rachis.

ACKNOWLEDGMENTS

I am indebted to R. D. Randall for drawing up the description of R. percrispus and providing notes
of its habitat preferences and many of the exsiccata details. To A. L. Bull thanks are also due for
drawing attention to the material of that species collected by J. R. Ironside-Wood. I further wish to
thank A. Newton for nomenclatural advice in connection with R. transmarinus, Mrs F. Le Sueur for
hospitality and assistance in studying R. caesarius and R. couchii in Jersey, and Professor V. Nutton
for guidance on certain words in the Latin diagnoses.

FOUR NEW SPECIES OF RUBUS L. Pl
REFERENCES

ALLEN, D. E. (1992). Rubus corbieri Boulay (Rosaceae) in the British Isles. Watsonia 19: 131-133.

ALLEN, D. E. (in press). Rubus, in Bowman, R. P., Brewis, A. & Rose, F., eds. Flora of Hampshire.
Colchester.

EpEES, E. S. (1975). Notes on British Rubi, 3. Watsonia 19: 331-343.

JERMYN, S. T. (1974). Flora of Essex. Colchester.

RitsTone, F. (1950). Some Cornish Rubi. Botanical journal of the Linnean Society 53: 413-421.

RoceErs, W. M. (1900). Handbook of British Rubi. London.

Rocers, W. M. & Rocers, F. A. (1898). On the Rubi and Rosae of the Channel Islands. Journal of botany
(London) 36: 85-90.

Supre, H. (1904). Observations sur ‘Set of British Rubi’. Bulletin de la Société d’ Etudes Scientifiques d’ Angers
33: 106-145.

SuprE, H. (1913). Rubi Europae, fasc. 6. Albi.

Watson, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland. Cambridge.

(Accepted February 1994)

|
|
|
|

Watsonia 20: 153-156 (1994) 135

Notes

JUNCUS EFFUSUS VAR. SPIRALIS J. McNAB IN THE INNER HEBRIDES

Henderson (1992) drew attention to the occurrence in N.W. Scotland (W. Ross, Outer Hebrides
and Orkney) of a variant of Juncus effusus L. with suberect, gently spiral stems; he named this var.
suberectus Henderson, with a type from Big Sand, Gairloch, W. Ross. Nelson (1993) reviewed the
taxonomy of this plant in relation to the cultivated plants of J. effusus with spiral stems and
concluded that they were all referable to one taxon, for which the correct name is var. spiralis J.
McNab or forma spiralis (J. McNab) Hegi. He also cited additional herbarium specimens. In this
note I accept Nelson’s conclusion that var. suwberectus should be regarded as a synonym of var.
spiralis, but I prefer to treat the plant at varietal rank rather than as a form.

I have been familiar with the plant described by Henderson since 1987, when it was first pointed
out to me by N. F. Stewart. As Henderson (1992) stated, the most striking feature of the plant is its
open habit, with the stems prostrate to ascending but never erect. The stems are straight or slightly
spirally coiled, but I have never seen plants in the wild with stems tightly coiled like a corkscrew. At
first I thought that it might be a phenotype resulting from exposure or heavy grazing. Subsequent
field observations have led me to the conclusion that it is genetically distinct. Plants of var. spiralis
can be found in ungrazed grassland (e.g. the roadside verge along the street at Arinagour, Coll);
furthermore, there is no reason to believe that grazing pressure is more intensive in N.W. Scotland
than in other areas where only erect plants of Juncus effusus occur. I have also seen at least one
population where var. spiralis grew amongst dense upright tussocks of J. effusus, in a pasture (S. of
Loch an Duin, Coll) where both varieties would be subject to a similar degree of exposure and
grazing. Henderson (1992) reported mixed populations, and he also made direct observations on
seedling plants which demonstrated that the variant has a genetic basis.

Unfortunately I did not make a note of the locality in the Outer Hebrides (v.c. 110) where I first
saw var. spiralis. Since 1989 I have noted the sites where I have seen ‘Hebridean Juncus effusus’, as I
called the plant. These records, all of which are from the Inner Hebrides, are listed below. Most
were seen when I was looking for aquatic plants, which explains why they are from pastures near
lochs. I have never seen it growing close to water, however, as upright plants often do.

South Ebudes, v.c. 102: Noted as occasional on both Islay and Jura by N. F. Stewart from 1981
onwards, but no specific localities recorded.

Mid Ebudes, v.c. 103: N. of Loch an Eilein, Tiree, NL/98.43, 8 July 1989. Pasture on N. side of
Loch Riaghain, Tiree, NM/03.47, 9 July 1989. Pasture near sea-shore, Lon Fhadamuill, Tiree, NM/
073.493, 22 June 1990. Pasture, Vaul, Tiree, NM/04.48, 26 June 1990. Damp area in pasture,
Urvaig, Tiree, NM/07.50, 26 June 1990. S.E. coast of Gunna, NM/10.50, 6 July 1989. Pasture
between Crossapol and Caoles, Coll, NM/12.52, 5 July 1989. Near pool at Cnoc na h’Osnaiche,
N.E. of Loch Fada, Coll, NM/198.587, 29 June 1990. Verge by village street, Arinagour, Coll, NM/
22.57, 5 July 1989. Rough pasture S. of Loch an Duin, Coll, NM/21.57, 7 July 1989.

North Ebudes, v.c. 104: Bank of stream by outflow at S. end of Loch Mhor, Waterstein, Skye,
NG/142.480, 13 July 1989, CGE (Preston 89/202).

J. effusus var. spiralis is occasional on Coll and Tiree, but much less frequent than the erect
variant. I have done too little fieldwork on Skye to comment on its frequency there.

These records add to those already published by Henderson (1992) from v.cc. 105, 110 and 111
and Nelson (1993) from v.cc. 17, 44, 111, 112, H27, H28 and H34. The fact that J. effusus var.
spiralis has a distinct habit and a geographical range which differs from the widespread varieties has
led me to prefer to treat it as a variety rather than a form. This is similar to the taxonomic treatment
of coastal ecotypes, which differ from the type in their dwarf or prostrate habit and are usually
treated as varieties or subspecies.

154 NOTES

ACKNOWLEDGMENTS

I am grateful to N. F. Stewart for drawing my attention to this variant of Juncus effusus, and for
providing the records from Islay and Jura. Many of the above records were made with Nick Stewart
and other friends on memorable B.S.B.I. meetings to Coll (1989) and Tiree (1990) organised by Dr
A. Walker.

REFERENCES

HENpDERrSON, D. M. (1992). A new spiral variant of Juncus effusus L. (Juncaceae). Watsonia 19: 133-134.
NELSON, E. C. (1993). Corkscrew rush (Juncus effusus L. forma spiralis (J. McNab) Hegi) (Juncaceae) in Ireland
and Britain. Watsonia 19: 275-278.

C. D. PRESTON
Biological Records Centre, I.T.E., Monks Wood, Abbots Ripton, Huntingdon, PE17 2LS

DISTRIBUTION OF LADY ORCHID (ORCHIS PURPUREA HUDSON) IN AN.
EAST KENT WOOD

In a strip of mixed woodland in East Kent, Orchis purpurea Hudson lies in a band roughly along the
middle. This distribution invited explanation.

The wood clothes the west-facing slopes of a north-south dry valley on the upper and middle
chalk. The higher ground is capped with clay-with-flints, an acid brown earth of the Winchester
series, whereas the lower part directly over the chalk is covered by brown calcareous earth of
Coombe and Upton series. Maps show that the upper slopes have been wooded since the 1840s
(mostly coppice) and probably since the 1760s or before, whereas the lower ground has only become
woodland since the 1880s. The canopy is now fairly continuous overall but here and there gives way
to scrub; there are also small areas kept open as turf, and places where coppicing and scrub cutting
are maintained.

In the absence of a soil survey the extent of the acid cap was approximately gauged by mapping
the distribution of bluebell (Hyacinthoides non-scripta (L.) Rothm.) (Fig. 1). Fig. 2 gives the areas
shown as wooded in the Waltham Parish Tythe Map of c. 1840; the remainder of the area, at present
wooded, was shown as arable. Both figures also show the distribution of the flowering spikes of O.
purpurea counted in the plentiful year of 1986. Fig. 1 confirms that the orchid is confined to the
calcareous soil, and in Fig. 2 the orchid spikes are seen to follow more or less the boundary of the old
woodland. We infer that this was the plant’s original distribution, the old wood itself being probably
too acidic (cf. Fig. 1: Summerhayes 1968), and that it has been gradually colonising more recently
wooded chalky areas as they achieved the appropriate degree of partial shade.

The wood is an S.S.S.I. and a nature reserve belonging to the Kent Trust for Nature
Conservation; it is the site of the two abnormal forms of O. purpurea described by Ettlinger (1987)
and Ingram & Dunster (1991).

ACKNOWLEDGMENTS

We are indebted to the Librarian of the Cathedral Library, Canterbury for giving us access to the
Waltham Parish Tythe Map; to the Librarian of the Local Studies Library, Springfield (Kent County
Council) for access to other maps; to Dr S. G. McRae for laboratory measurements of soil pH; and
to the Kent Trust for Nature Conservation for permission to publish.

ee

NOTES

Ground cover
of bluebells

e Single
@ 5-9
@ 10+

Flower spikes
of lady orchid

FicurE 1. Bluebell areas (Hyacinthoides non-
scripta) in 1988. Numerals indicate sites where pH

i measurements were made in 1990 (top soil —- A

horizon): 1, pH 7-7; 2, pH 7-8; 3, pH 4-3; 4, pH 4-0.

155

200 metres

Woodland
c. 1840

e Single
@ 5-9

@10+
Flower spikes
of lady orchid

FIGURE 2. Woodland as shown on the Waltham -
Parish Tythe Map c. 1840 superimposed on the
modern outline of the Reserve; the remainder was
shown as arable.

Dots show O. purpurea flowering spikes in 1986. Sites of abnormal flower forms described by Ettlinger (1987),

E, E; by Ingram & Dunster (1991), ID.

156 NOTES
REFERENCES

ETTLINGER, D. M. T. (1987). Peloric and duplex examples of Orchis purpurea Hudson in Kent. Watsonia 16: 432.

INGRAM, G. I. C. & Dunster, C. W. (1991). A non-resupinate abnormality in Orchis purpurea Hudson.
Watsonia 18: 317-319.

SUMMERHAYES, V. S. (1968). Wild orchids of Britain, p. 253. Collins, London.

D. E. WILSON

The Keep House, Sutton Valence, Maidstone, Kent, ME17 3BS
G. I. C. INGRAM

42 Burford Lodge, Pegasus Grange, White House Road,
Oxford, OXI 4NA

Watsonia 20: 157-166 (1994) 157

Book Reviews

Mediterranean wild flowers. M. Blamey & C. Grey-Wilson. Pp. 560, with 192 pp. of colour plates
and numerous marginal line drawings. Harper Collins, London. 1993. Price £25.00 (ISBN
0-00-219901-7).

This is a comprehensive field guide to the flowers of the Mediterranean coasts and islands, including
trees and introduced species. More than 1,500 species are illustrated by Marjorie Blamey’s colourful
paintings which are bound together in the centre of the book. The format is similar to that of The
illustrated Flora of Britain and Northern Europe, published by this artist and author in 1989, but the
smaller size of the new volume makes it more easily carried and convenient for holiday field work.
This compensates for losing the pleasing coloured marginal paintings in the earlier volume, and thus
having text and illustrations together; in the new guide there are helpful line drawings illustrating
significant characters of some of the plants. The species are listed by their Latin names in the text,
followed by English names which give easy access for those not familiar with Latin; for me this is a
great improvement on the format of the earlier book, which was arranged under English names.

Alas, on the dust jacket it is stated to be a “‘complete”’ guide to the native and introduced flowers
of Mediterranean coasts, and also includes those species found growing up to 1000 m altitude. This
reads like a publisher’s description rather than the authors’, as complete coverage of this very rich
and varied area of considerable size would need a book of much larger dimensions.

The division of the region into three sections in the Introduction puts Crete and the Balkans into
the Western Mediterranean, but these floras are generally considered to be more closely allied to
those of Mediterranean countries further east than to those of the western Mediterranean and the
Iberian peninsula. From my experience the marked differences between the floras of eastern and
western Mediterranean have been insufficiently stressed in the distribution notes. We are told that
the most commonplace northern European species have been omitted; but some are included.

Over such a wide area the distribution data must have been partly written from sweeps through
the literature. As an example from Crete, where many new records have been published in recent
years, Lamium purpureum and Filago arvensis are given as “not in Crete’’, but both are listed in
Crete: checklist of the vascular plants, by Sir Colville Barclay (1986), and have been seen by the
writer in western Crete. But the Cretan tulips are clearly set out and differentiated.

To be complete for any one area would not be within the scope of this guide, but it gives excellent
general coverage. With the clear paintings, brief but diagnostic text, and the line drawings, it should
be possible to identify a good proportion of plants seen, despite the absence of keys. The book will
fulfil its aim to promote an understanding of this unique flora, and it is a reminder of the often
magically beautiful Mediterranean coastal and island flowers.

M. BRIGGS

The genera of the Umbelliferae. M. G. Pimenov & M. V. Leonov. Pp. 156, one map, one table.
Royal Botanic Gardens, Kew and Botanical Garden of Moscow University. 1993. Price £12.00
(ISBN 0-947643-58-3).

The Umbelliferae are traditionally labelled as ‘difficult’ by amateurs and professionals alike, so any
new publication which may help to unravel the taxonomic complexities is always welcome. In a
family containing over 3600 species in 455 genera the proportion growing as natives or aliens in the
British Isles is significant with a maximum of about 120 species in some 40 genera, i.e. about 3% of
the world’s species and 10% of the world’s genera are represented here.

158 BOOK REVIEWS

The published work is a listing of data gathered on the GNOM (Generis NOMenclator) computer
database which remains open for corrections and additions, thus reflecting current nomenclatural
knowledge of genera in the Umbelliferae. Adopting a modified version of Drude’s Classification
(1897-98), the bulk of the work comprises an alphabetical list of genera with authorities, place of
publication of the protologue, type, subfamily, tribe and subtribe. For each entry this is followed
where appropriate by any generic synonyms, geographical distribution, number of species and
finally any relevant literature. Unlike some works of this nature the information is easily read and
well laid out with the appropriate use of a range of typefaces and adequate spacing between entries.
About a fifth of the work is given over to a comprehensive list of references, which is more than
adequate although omitting some of the regional Floras consulted by the authors.

The authors have adopted a relatively narrow generic concept throughout but admit to difficulties
with critical genera such as Peucedanum. They recognize the genus as having perhaps only six or
eight species closely related to the type, P. officinale, a rare species in our flora being confined to
East Anglia, but much more widespread in continental Europe. However because many of the
remaining species previously placed in the genus are of uncertain position, they list Peucedanum as
having 100-120 species with the proviso that more work is needed and that this is probably the most
complex taxonomic conglomerate in the Old World Umbelliferae.

Dedicated to Professor Lincoln Constance, who has spent most of his life studying the

Umbelliferae, this is an excellent compilation of data which will be invaluable to any serious student —

of the family, but especially for those trying to grapple with the many problems of nomenclature and
generic delimitation encountered in the subfamily Apioideae.

S. G. KNEES

The Flora of Ditchley: wild flowers of an Oxfordshire estate. A. J. Dunn. Pp. xi + 68, with twelve
colour photographs and estate map. Privately published by Dr Catharine Wills, Sandford St Martin,
Oxfordshire. 1992. Price £16.45 (ISBN 0—9521310-0-S).

This is the flora of a private estate in the foothills of the Cotswolds. The 17 ha (46 acres) consist
mainly of farmland, several plantations and a few hectares of ancient woodland.

Miss Dunn gathered botanical records from 1985 until 1992, the most exciting one being a new site
for the Downy Woundwort, Stachys germanica, one of our threatened species. She has studied the
habitat requirements of this plant in great detail and published the findings in Watsonia 16: 430-431
and 18: 359-367. Thanks to the loving, careful management of the landowner, the late Mr Martin
Wills, the range of species is impressive; indeed half of the 443 recorded species are additions to the
official county records for the estate.

The book is well written and has some very readable chapters on the history and natural history of
Ditchley, but I would have welcomed a better map. I also regret that the author does not give exact
plant locations, though she mentions that index cards for each plant, carrying full details, are
available on request. Whilst I understand her concern for secrecy in the case of a few rare plants, the
general lack of precise data is bound to diminish the appeal of the book.

M. BAECKER

The Wiltshire Flora. Edited by B. Gillam. Pp. x + 386, with eight pages of colour plates and 622
maps. Pisces Publications, Newbury. 1993. Price £27.95 (ISBN 0—9508245—8-5).

Despite its large size (84 km from north to south), Wiltshire has traditionally shared the fate of
Lincolnshire and Northumberland of lying off the main botanizing routes and enjoying too slight a
reputation for rarities to lure outsiders into paying visits to it expressly. The task of investigating its
flora has consequently fallen almost wholly on the shoulders of its resident botanists, a task which
they have increasingly discharged with remarkable frequency and thoroughness. In the space of
only 36 years they have given us a county Flora of the classic type of outstanding quality, an

BOOK REVIEWS IS)

extensive supplement to that and now this further very substantial volume which is a kind of hybrid
between the two.

Most attractively produced, this latest work embodies the results of the Wiltshire Flora Mapping
Project of 1984-92, an undertaking which is shown to have had a complex gestation and to have
given rise in turn to an unusually elaborate organizational structure, with a Steering Group, a
Science Group, successive Project Coordinators and ultimately a Publication Working Party. The
whole modus operandi, including the recording methods, is described in commendable detail on
pages 3—6 and 112-118. Emphatically a team effort, in which the B.S.B.I. Recorders for the two
vice-counties, David Green and Ann Hutchison, have played a major part (with responsibility, most
notably, for the systematic list), it has ended up by giving rise, as in Surrey and Sussex, to a
permanent body dedicated to carrying on the study of the county’s flora into the future. Even had no
publication resulted, this outcome alone would surely have been justification enough for all the
work that the project has entailed.

J. D. Grose’s 1957 Flora was a product of the pre-mapping era and the 1975 Supplement was
essentially a mere appendix to that in concept. In the intervening years, however, the county’s
landscape had been drastically altered and the need had begun to be felt for a total re-survey on the
basis of the standardized mapping units offered by the National Grid. A major feature of this latest
volume is consequently over 600 tetrad distribution maps of all but the commonest and the more
critical species. In addition, it has most of the introductory sections now standard for county Floras,
35 colour plates of various shapes and sizes depicting sample habitats and some of their more
attractive individual species, and a series of short essays, by different hands, on certain plants of
more particular local interest — a welcome and valuable innovation (if slightly marred by the choice
of oppressively large type for the titles).

As usual, though, it is the systematic list which accounts for the main part of the text and to which
most readers will turn in the first instance. One thing here that will strike them very quickly is the
size of the floristic debt the county owes to the fact that its boundaries in H. C. Watson’s day took in
a tiny part of the super-rich New Forest (and ironically, if perhaps inevitably, most of the records
resulting from this appear to have been a by-product of the concurrent fieldwork for the forthcoming
Flora of Hampshire). No less eye-catching are the helpful figures at the foot of the accounts of the
species indicating the percentage of 10-km and either 2-km or 1-km squares in which each was
recorded during the project. These take the place of the inherently vaguer verbal estimates of
frequency conventional till now in works of this kind. Impressive, too, is the care given to the precise
detailing of habitats and of the extent to which species differ in quantity between one area and
another. It is a pity, though, that a wish to keep wordage and punctuation to a minimum has had the
result of making many of the lists of localities too condensed for ready comprehensibility.

Much less pardonable, by contrast, are the status categories adopted: each species is designated
either “native” or “introduction”, with no distinction made between arable weeds of age-old
presence, putative relics of former cultivation, ephemeral adventives and established horticultural
exotics. The traditional Watsonian terminology certainly has its defects but it is greatly more
informative than this kind of despairingly aggregate treatment. To discard it wholesale, instead of
seeking to improve it with modifications, is merely obscurantist and does no service to the army of
local workers on whom the task of refining our notions of status so heavily depends. Even with just
two categories in play, though, there is evidence in these pages of still some considerable
inexactness: Sinapis arvensis is classed as a native, for example, but Thlaspi arvense and Raphanus
raphanistrum are dismissed as introductions — for no reasons that are apparent. In common with
nearly all local Floras, moreover, no allowance is made for the fact that many species are of multiple
status locally: seemingly indigenous, maybe, in certain habitats, accidentally brought in in others, a
relic of deliberate cultivation in yet others. To force every species into just the one, most
‘respectable’ category (as Watson himself unfortunately did) can only be to convey a seriously
misieading idea of the more complex reality.

Lastly, the unwary should be warned that this work is, by deliberate design, a ‘snapshot’ Flora, a
record of everything found within just the period under study, not an account of all the vascular
plants known to have occurred in Wiltshire since records began. Because of this time restriction —
and it is not clear why the work had to be so hurried — pre-1984 records are excluded except where
necessary as a historical benchmark and the coverage of the larger critical groups is sketchy at best.
On the one hand this has meant that many species for which reliable past evidence of their

/

160 BOOK REVIEWS

occurrence in the county exists have no place in these pages, even though the fact that they may still
persist in the localities in question could serve as a fruitful challenge to present-day investigators.
The recent exciting disinterment in BM of E. J. Tatum’s century-old specimen of Gentianella ciliata
consequently earns no mention (indeed, the combing of herbaria formed no part of the project, by
definition). On the other hand it has seriously reduced the value of the book to anyone whose
interest lies primarily in critical plants. The account of Taraxacum does not extend beyond three
aggregates and that of Rubus is a bare list of species without localities, omitting (because observed a
few years too early) what is probably the one flowering plant taxon confined to this county in
Britain, namely R. arrhenii, as well as, with no such excuse, R. stenopetalus, the rediscovery of
which in Wiltshire and confirmation as a result as British was reported in this journal during the
duration of the project. As two Rubus specialists live in next-door counties, it seems a pity that more
effort was not made to secure a better coverage of at least that group, the more so as the duplicates

~ collected by Grose at W. C. R. Watson’s side in 1948 have mostly been destroyed by insects and the

correctness or otherwise of the determinations published in the Flora of Wiltshire badly needs to be
established. Perhaps the splendidly hyperactive botanists of this county can now be induced to make
a Critical Flora of Wiltshire the next object of their exertions?

D. E. ALLEN

Roses of Great Britain and Ireland: B.S.B.I. handbook No. 7. G. G. Graham & A. L. Primavesi.
Pp. 208. Botanical Society of the British Isles, London. 1993. Price £11.50 (ISBN 0—901158—22-4).

This long awaited addition to the B.S.B.I. Handbooks continues the high standards set by earlier
volumes in the series. The authors have produced an authoritative account which, for the first time,
makes the genus accessible to British botanists. The excellent line drawings by Margaret Gold
complement accurate and unambiguous descriptions of the twelve native and eight introduced
species. Much research in the field and in herbaria has enabled the authors to describe 83 hybrids.
Most hybrids are not mapped, but a list of v.c. records is given; I could find no explanation of “‘non-
directional records” (it means the seed parent is not known and not that the plants have grown in all
directions!).

The nomenclature and order of the species is the same as in Stace’s New Flora. English names are
given for all species though I doubt if ““Columnar-styled Dog-rose” will be very popular! I was
pleased to see “Eglantine” retained for R. rubiginosa, it would be sad to lose it to the cause of
English binomials. One other change from Stace is the addition of “Northern Dog-rose” for R.
caesia; this seems a little premature given the frequency of Leicestershire records for both
subspecies on the maps.

The introductory chapters are essential reading for anyone wishing to study wild roses. I found the
accounts on morphology and general characters particularly useful, with clear illustrations of growth
forms, prickles, leaf shapes, leaf serration, hips and shape and disposition of the sepals. This
account, together with a very useful section on collecting and pressing roses and a glossary which
puts some of the more obscure terminology into plain English, should help the amateur rhodologist
get started. The history of classification is described and the problems presented by the peculiar
reproduction and promiscuous hybridisation show why the genus has been so difficult for
taxonomists. The section on ecology and distribution is very interesting; several species appear to be
limited only by lack of suitable soils or habitat. However, I suspect the influx of new records
stimulated by this book may show up new patterns not yet visible from the paucity of accurately
determined records available to the authors.

There are two dichotomous keys, one including alien species, the other just the natives. Both are
effective and my trials with five named herbarium specimens all proved successful. Initially it seems
odd to have two keys doing the same job, but I found it reassuring to reach the same conclusion
using slightly different combinations of characters. Using the keys amply demonstrates the need for
good specimens and for the annotation of characters such as sepal disposition and bush shape whilst
in the field. The keys do not identify hybrids except for the frequent R. < dumailis.

32 distribution maps at a 10-km square scale cover the native species and selected hybrids; only
accurately determined records have been used. As the authors point out, the maps of critical species

BOOK REVIEWS 161

and hybrids are provisional. The maps for the R. canina groups, R. X dumalis and R. caesia, all show
the authors’ surveys of Leicestershire and Co. Durham, but little else. If these taxa are as frequent in
other parts of the country as they appear in these two counties then a great deal more recording is
needed before we have an idea of real distributions. The symbols used to show the two date classes
on the maps are squares and triangles rather than the black and white circles of earlier volumes. I
found this visually less effective, making it harder to pick out declines in range.

The book finishes with an extensive bibliography containing 289 entries. As 90% of these are pre-
1975 and a third are foreign I feel much of this is of limited interest to most readers.

The new ‘Demy 8vo’ format (almost A5 size) is an improvement on the cramped feel of the earlier
volumes, but there is a lot of white space and the layout does not make the best use of the larger page
size. I do not like the sombre cover design; the black edges give a funereal feel to a book about such
beautiful plants. Like Crucifers, the text has been printed from camera-ready copy, and this gives a
slightly fuzzy look to the text as opposed to the crisp typesetting of Handbooks nos. 1 to 5. The use
of hyphenation is bad, many words being split with only two letters left at the end of a line.

The fact that flowers are of little diagnostic use has meant they hardly feature in this book. This
will perhaps disappoint those who, like me, were first attracted to wild roses by the dust-jacket of
Keble Martin’s Illustrated Flora. It may also limit its sales appeal, but this is a book which all
B.S.B.I. members should buy. It has brought the study of British Roses into the twentieth century
and will stimulate me, and I am sure, others, to take much more notice of this neglected genus.

M. N. SANFORD

Atlas of the bryophytes of Britain and Ireland. Edited by M. O. Hill, C. D. Preston & A. J. E. Smith.
Volume 1: Liverworts (Hepaticae and Anthocerotae); pp. 351, including 293 distribution maps.
Volume 2: Mosses (except Diplolepideae); pp. 400, including 367 distribution maps. Harley Books,
Colchester. 1991, 1992. Price £25.00 (ISBN 0-946589-29-1, vol. 1); £30.00 (ISBN 0-946589-30-5,
vol. 2).

Using the first volume of this bryophyte distribution atlas, it took me exactly 3 minutes and 47
seconds to find out which liverwort is the most widespread in Britain and Ireland. It is Lophocolea
bidentata which is recorded from 2,421 10-km squares in the two countries (including the Channel
Islands). Of course, this fact is not one of the most interesting ones to be taken from the Adlas.
Nevertheless the example shows that the data are presented in a form that makes them readily
available.

The complete Adlas will comprise three volumes and will present distribution maps for the whole
bryophyte flora of Britain and Ireland amounting to nearly one thousand taxa. The area covered
comprises about 3,500 mapping units (10-km squares of the British and Irish national grids and of
the UTM grid in the Channel Islands).

The introductory chapters of the first volume deal with the history of bryophyte recording in the
British Isles, the mapping scheme used (including information on the availability of records from the
database), and the evenness of the survey. The first map provided shows the number of liverwort
species recorded in each of the 10-km squares. No attempt is made to distinguish between
differences in investigation effort and differences in regional species diversity. Will that problem be
discussed in the “detailed analysis of geographical coverage”’ planned for the final volume?

The liverwort volume contains 289 distribution maps for the hepatics and four for the hornworts
of Britain and Ireland. The map dots are differentiated as to the date of the last record (before or
since 1950). Accompanying the maps, short texts by various authors give additional information for
each taxon in standardized format. These texts include data on altitudinal distribution, ecology of
the species, number of grid squares in which the taxon is mapped, sexuality, fertility, means of
propagation and general distribution. Additional remarks on various subjects are given where
appropriate.

The distribution maps are followed by eight maps showing selected environmental factors,
including the mean concentration of sulphur dioxide in the atmosphere in 1987 (UK only). In a
special chapter of the second volume Crundwell considers the bryophytes of Britain and Ireland in

162 BOOK REVIEWS

a European context. 367 moss distribution maps of the same type as for liverworts follow and deal
with about half the moss flora of the archipelago.

Both published volumes conclude with bibliographic references and a “‘list of localities cited in the
text”’ with their co-ordinates in the appropriate national grids that are shown in a separate map (for
people not familiar with the grid system, an explanation of the use of it would have been useful);
each includes an index to the taxa, including selected synonyms.

No mention is made of the total number of records on which the Atlas is based. It must be a
question of hundreds of thousands. With the exception of the BeNeLux countries there is no region
in continental Europe with a comparable state of knowledge. However, Crundwell’s ‘“‘guess”’ that
“our present 10-km square records are a little over two-thirds of those we would have were our
knowledge complete” seems to be somewhat optimistic. Of 17 relevant records from two small
collections made without any special knowledge and intention in Scotland and in Southwest Ireland
four are new to their squares. From my own experience in bryophyte mapping and considering the
enormous task undertaken, I judge any degree of completeness of more than 50% an excellent
result. Of course, the data ought to be examined for less obvious biases produced by the method of
mapping. But the maps give the experienced user the decided impression that the high value of the
data for various kinds of numerical interpretation are not substantially affected by errors or
incompleteness.

I am looking forward to see the Atlas of the bryophytes of Britain and Ireland completed. It is an
invaluable tool for bryologists, biogeographers, botanists, conservationists and ecologists, and an
important work far beyond the area covered.

E. URMI

The families and genera of vascular plants. Edited by K. Kubitzski. Vol. 2: flowering plants,
dicotyledons, Magnoliid, Hamamelid and Caryophyllid families, edited by K. Kubitzski, J. G.
Rohweder & M. Bittrich. Pp. x + 653; 140 figs. Springer-Verlag, Berlin. 1993. Price DM 478 (ISBN
3—540-55509-9).

The Englerian tradition of preparing integrative botanical monographs is alive and well, to judge
from this latest volume in the series first reviewed in Watsonia 19: 44-45, 1992. It covers three of the
major subclasses of the dicotyledons, and an introductory chapter provides a chemosystematic
overview of these together with a fourth, the Ranunculidae. The main body of the volume consists
of a descriptive section for each family followed by keys to subfamilies and genera, generic
synonymy and a short description of each genus, a sketch of its geographical distribution and an
estimate of the number of species it contains. There is a selective bibliography at the end of each
family treatment, and an index to scientific names is provided at the end of the volume. The quality
of both printing and binding is excellent.

This is an encyclopaedic treatment, with one major limitation as a consequence. There was a need
to prepare complete accounts for each family, and the editors were obliged to compile some
treatments themselves when specialists either failed to deliver their promised contributions, or were
unavailable; this makes the final product somewhat uneven. In Kubitzski’s account of the
Plumbaginaceae, for example, the generic splitting carried out by Linczevski in subfam. Staticoi-
deae, resulting in around 30 species of Limonium sensu lato being transferred to 13 new genera, has
been accepted but with reservations. The revision provides no new insights into the evolution of the
family, and the key to genera uses characters such as leaf width and stigma shape to separate groups
which the author admits will not stand the test of a critical revision. Other treatments, by contrast,
such as that of the Fumariaceae, are up-to-date and definitive.

In the preface to this volume, Professor Kubitzski takes stock of the rapid development of
molecular techniques for probing the relationships between genera, families and subclasses of
dicotyledons, and the extent to which these discoveries have been incorporated into conventional
classifications. There is still enormous scope for applying these new methods to the study of taxa at
the level of genera and above, but relatively few authors have gone on to prepare monographic
treatments at this level which specify morphological characters for the newly characterised taxa.

The present editorial team, with its mixture of youth and experience, must therefore be

BOOK REVIEWS 163

congratulated for attempting such an ambitious project, albeit one which is heavily dependent on
outside contributions: 38 authors were responsible for the present work. The formidable price of
this volume may well place it beyond the reach of all but the largest botanical reference libraries, but
copies will no doubt be in demand from a wide range of users. Since the only analogous modern
work, John Hutchinson’s The genera of flowering plants, remains incomplete, one hopes that the
present series may soon be continued and eventually completed.

J. R. EDMONDSON

Algarve plants and landscape. D. J. Mabberley & P. J. Placito. Pp. xvi + 300; 279 colour and 269
black and white illustrations. Oxford University Press, Oxford. 1993. Price £45.00 (ISBN 0-19-
858702-3).

This is a difficult book to categorize. As its subtitle, “Passing tradition and ecological change”’
indicates, the authors have set out to characterize the flora of the Algarve province of Portugal
within a detailed context of the region’s geography, geology and, particularly, traditional
agriculture. There is also a balanced assessment of the impact of tourism, the region’s relatively
recent, but certainly its currently most potent, economic factor.

The natural vegetation of the Algarve is described essentially within the framework of the
province’s three main subdivisions, the coastal zone, the more inland and largely base-rich soils of
the ‘barrocal’, and the complex of acidic shales and other substrates of the upland ‘serra’. The style
is decidedly didactic, with observations for example on the derivation of plant names, their ecology
and uses, details of pollination by pseudo-copulation in some orchid species, and, consistently, with
Portuguese names for plants, communities, etc., given in parentheses. Many of these plants are
illustrated with colour plates and a fine series of line drawings by Rosemary Wise. Unfortunately,
the plates are reproduced in small format, grouped 15-18 to the page, and as a consequence are
sometimes of limited value. The drawings are on the whole effective and helpful. A further section
deals with the many exotic species encountered in streets and gardens in the province. A feature of
this book is that the plant life is not presented in isolation, but rather is interspersed with sections on,
effectively, local human geography. This extends to a survey of fishing practice, with lists of fish
species and molluscs, and other aspects of local economy, such as sea-salt production along the
coast, and above all, with an extensive exposition of traditional and more recent innovations in
agricultural practice in the region. This includes accounts of olive, carob, figs, grapes, citrus and
many other crops, each with extensive details of local cultivars. Indeed, this central section
sometimes gives the impression of a text in economic botany.

Throughout, this book is profusely illustrated with black and white photographs of plant
communities and landscapes (including a series which contrast scenes in nineteenth century
aquatints by Landmann with the same locality captured in a recent photograph) and crop cultivars,
and also with line drawings, e.g. of traditional agricultural implements, and over 20 maps. A more
judicious selection of rather fewer maps would perhaps have been more helpful: e.g. a general
introductory map with more place-names than the minimal Tavira, Faro and Silves provided in Fig.
1 would have given a useful orientation; and readers will be amazed at the extensive areas of the
Algarve where, apparently, the average age of the population is over 100 years, as depicted in Fig.
238. An omission in the legend seems to be responsible. A more serious omission is the lack of
author citation at relevant places in the text for any of the nearly 500 references listed in the
bibliography. This can only hinder the reader who wishes to pursue a particular topic further.

For plant lovers who have acquired a property in, or who regularly visit the Algarve for what the
tourist trade refers to as ‘quality’ holidays, this fact-laden book is recommended as a means of
enhancing their understanding of the region. For those visitors who prefer to toast themselves on the
beaches, with only the occasional excursion inland and minimal interaction with the local
inhabitants, the detail provided in this book is likely to be offputting, as is its price. This is a pity,
because mass-tourism, either by package-tour arrival at Faro airport, or via the new international
bridge across the river Guadiana from Spain, is changing the flora and traditional life-style of the

164 BOOK REVIEWS

Algarve at an accelerating pace. Mabberley and Placito’s enthusiasm for the region, which is evident
in this book, allows us to share something of its cultural heritage before it is too late.

P. E. GiBBs

Scandinavian ferns. B. Oligaard, with illustrations by K. Tind. Pp. 317; 103 line drawings and 114
colour plates. Rhodos, International Science and Art Publisher, Copenhagen. 1993. Price
paperback D.kr. 375 (ISBN 87—7245-530-6); hardback D.kr. 425 (ISBN 87—7245-532-2).

Scandinavian ferns is sub-titled “A natural history of ferns, clubmosses, quillworts and horsetails of
Denmark, Norway and Sweden.”’ It is natural history at its best. The text and illustrations recount
meticulous and detailed observations of Scandinavian Pteridophytes. All that has been observed of
their reproduction, variation, ecology, distribution and even the origin and meaning of their names,
is presented to the reader.

The 114 water-colour plates are painted from living subjects and breathe a life rarely found in
botanical illustrations. Clouds of spores erupting from Botrychium lunaria or shoots of Equisetum
variegatum with huge mosquitoes hanging dangerously over-head convey an impression of life for
these simpler Pteridophytes. Many of the fern illustrations are equally delightful — those of
Cystopteris for example — but my feeling was that the bold style lost some of the intricate beauty of
Athyrium and Polystichum. Whole plant illustrations invariably include rhizomes, and habitat
paintings show other species for scale and context. Frequently there are details of scales, individual
pinnae, sori, prothalli, young plants, croziers, even a droplet of water being excreted from an
Equisetum telmateia sporeling. Line drawings inserted in the text illustrate spore architecture, cells
in the annulus, stem cross-section and the fronds of hybrids and subspecies where appropriate.

There are keys to all species, and additional subspecies and varieties (eleven) and hybrids (25) are
dealt with in the text. Taxonomically there are a few differences from recent British Floras;
Diphasiastrum alpinum and D. complanatum are placed within Lycopodium for instance. As a
nomenclatural romantic I was pleased to see Asplenium viride rather than the cumbersome and
unfamiliar Asplenium trichomanes-ramosum.

The comprehensive text and illustrations might be of interest to British botanists wishing to
complement their British Floras. 56 of the 72 species described in Scandinavian ferns are members
of the British flora and all of the genera recorded from Britain bar only Adiantum and Trichomanes
are represented in the book. Diplazium sibiricum (family Woodsiaceae) was a new species for me,
and I was pleased to learn of an additional six Botrychium species not found in Britain. Taking this
large format book into the field would be cumbersome and only spoil the paintings. It is definitely a
book for the armchair where the text makes easy reading. I would recommend anyone contem-
plating a botanical excursion in Scandinavia to consider buying a copy.

G. STARK

The release of genetically-engineered organisms. Edited by B. Shorrocks & D. Coates. Pp. 48.
British Ecological Society Ecological Issues No. 4. Field Studies Council, Shrewsbury. 1993.
Price £3.50. (ISBN 1-85153-853-4).

The lack of easily obtainable and factual information on current ecological issues has led the British
Ecological Society to produce a series of booklets. Each booklet, representing the work of a group
of experts, deals with one topic and the fourth of the series is concerned with Genetically Modified
Organisms (GMOs). An organism is genetically modified if new combinations of genetic material
are achieved by techniques other than traditional breeding or other natural methods. In the first
chapter the ways in which genetic information is transmitted and manipulated are summarized.
Although these booklets are aimed at a wide audience, including teachers and policy makers, this
essential introduction on genetic engineering may be difficult to understand and possibly off-putting
to readers without a good genetic background, despite a good two-page glossary. Subsequent
chapters are easy to read and the second one discusses the risks associated with the release of GMOs

BOOK REVIEWS 165

into the environment and their potential environmental problems in relation to our knowledge of
the ecology of invasions by exotic species. The two main problems posed by GMOs are firstly the
transfer of a gene to a related species and secondly the creation of invasive organisms.

The following three chapters discuss the potential risks associated with the release of microbes,
plants and animals. The section on plants deals essentially with genetically modified crops and
arable weeds, and little mention is made of grasses and ornamental plants. Horticultural plants are
unlikely to become the target of genetical engineering in the very near future. However, because of
their economic importance, it is only a matter of time before transgenic ornamentals are produced.
Traits such as frost resistance and those related to the reproductive biology (e.g. flower colour or
structure) are likely to be targeted and these traits affect plant distribution and abundance. For
example if flower attributes are modified the pollination success and associated seed production
might be affected which could alter abundance. Since a very large proportion of invasive species are
escaped ornamentals the potential threat is great, whereas few if any terrestrial plant crops are
known to be invasive in natural or semi-natural vegetation. The problem with GMOs is that the
outcome of a particular genetic change is uncertain and like exotic species we are not able to predict
which species will become invasive in a particular environment.

The booklet is nicely produced and includes three pages of references giving readers the
opportunity to investigate particular issues in greater detail. Despite the poor treatment of
ornamental plants, this booklet, like the others in this series, is an essential source of information on
a current and potentially increasing environmental problem.

P. BINGGELI

Flora of Northumberland. G. A. Swan. Pp. 351; eight transparent overlays, 14 pages of colour
plates. Natural History Society of Northumbria, Newcastle-upon-Tyne. 1993. Price £36.00
(ISBN 0—9520782-0-1).

As children we would slide down the Willington ballast hills on pieces of cardboard, never realizing
the diversity of plant colonists that occurred there. Sadly, these white hills and their diverse floras
have disappeared, as have those of a vast number of lowland aquatic sites and many square kms of
open moorland. My grandfather, a herbalist in the Hexham area, knew the flora well and regularly
sent his children on collecting trips. Interestingly, a modern plant chemist has now produced a
comprehensive treatment of the plants of Northumberland. George A. Swan has done a
magnificent job of documenting and describing the native and naturalised flora of this large and
botanically-understudied county.

As an expatriate Northumbrian convalescing in St Louis after some serious surgery, I had hours of
pleasure reading of the plants that are now extinct, the ones I missed seeing when I lived in England
and the new occurrences of some interesting taxa, especially Ophrys apifera, Bee Orchid. The
format of the book is pleasing, with maps at the 5-km square level showing historical, recent and
current occurrences, all checked by the author before being accepted, and the text is in a clear type
on the quarto pages. A comprehensive gazetteer and a section on local habitat terms will help
visitors find their way to the various cited links, slacks, slakes, denes, hopes, cleughs, and other
areas of botanical interest. How fortunate that botanists have visited and thus ensured a mention in
print for Blakehopeburnhaugh, a strong candidate for the longest English place-name! Incidentally,
the terms Black and White in many settlement names may refer to former monastic orders, as
suggested by the village of Blanchland, Blackfriars in Newcastle and the two churches in tiny
Bywell.

The plants are listed in the Flora Europaea sequence but the names follow the recent publications
of Stace and Kent. A 50 page section by Angus G. Lunn describes the environment of vice-counties
67 and 68. The 109 species of Dandelion, Taraxacum have received an expert treatment in the hands
of John Richards. The remainder of the book is the sole effort of George Swan, although he does
fully acknowledge the companionship and active assistance of his wife Margaret. The Natural
History Society of Northumbria published the volume and Mrs Grace Hickling, our late heroine of
local natural history, apparently persuaded Mr Swan to undertake the task.

The book will surely encourage further botanizing in the region. Who will be the first to confirm

166 BOOK REVIEWS

Spring Gentian (Gentiana verna) and other plants not yet documented as county records?
Incidentally, the Bee Orchid recently found in the very grass where decades of pupils from Wallsend
Grammar School took their first ecology classes is probably a new colonisation because I am sure we
would not all have missed such a beautiful plant. Horses at the annual fair could have introduced the
seeds, and lime from the crumbling viaduct of Stephenson’s Killingworth wagonway (which
incidentally gave the world its standard railway gauge) could provide the necessary habitat. The
book also suggests numerous M.Sc. and Ph.D. thesis topics to the reviewer. Plant taxonomy is still
alive at the University of Newcastle and, hopefully, this new Flora may inspire school and college
teachers in the area. Mr Swan is only in his 70s and I wish him at least a further couple of decades of
botanising. After all, he will need them because he has recently added bryology to his list of
interests.

P. M. RICHARDSON

Watsonia 20: 167-180 (1994) 167

Obituaries

PETER HADLAND DAVIS
(1918-1992)

Born at Weston-super-Mare on 18 June 1918, Professor Peter H. Davis died in Edinburgh on
5 March 1992. My first encounter with Peter Davis was in 1946, as a raw undergraduate, attending
my first Botany I class at the University of Edinburgh, where I was planning to read Agriculture. I
was somewhat taken aback at the sight of a slender figure in a long, hairy, sheepskin coat, who
stutteringly introduced himself. Although he was ten years or more older than the rest of us, he soon
became a dominant character in our group and certainly set the pace for us as regards systematic
botany. This was easily explained by his earlier background, when he was influenced by his contacts
with the Ingwersen plant nursery which led to what became a life-long passion for plant collecting.
He had established contact with Sir William Wright Smith, Regius Keeper of the Royai Botanic
Garden and Regius Professor of Botany at the University of Edinburgh. This resulted in his going
there to read Botany, after spending most of the war years in the East Mediterranean where he was
employed by the security service (mainly dealing with screening mail) and acquired an extensive
knowledge of the flora which was to stand him in good stead later. It was, understandably,
disconcerting in our practical Botany classes at the University when faced with having to identify as
a test 100 specimens or illustrations of plants (a common practice under Sir William Wright Smith)
to find that some obscure species amongst them had been discovered for the first time by Peter
himself. Such was his disingenuous charm that we didn’t even cry foul! He was awarded First Class
Honours in Botany.

Peter Davis continued during his undergraduate years to work on his Mediterranean plant
collections which he had brought with him to Edinburgh and he soon involved me in working up the
Scrophulariaceae amongst which was a new species of Digitalis from Anatolia, D. davisii, the first
new species I ever described. In the meanwhile, I had developed an interest in Spain, having gone
there first on a plant-collecting expedition with Dr Paul Giuseppi, a Felixstowe surgeon and
renowned collector and alpine garden plant specialist, who was also a friend of Peter Davis. Peter
and I joined forces in a plant collecting tour of Spain in 1948, sponsored by the Royal Horticultural
Society. I then accompanied him on an expedition to Anatolia in 1950. Botanising with Peter was an
education in itself, for he was a real professional. Food, personal discomfort, physical exhaustion all
counted for nothing: nothing mattered but the plant collecting. He taught me how to make sketches
and accurate notes, drilled me in the meticulous preparation and numbering of the material, how to
make collections of seeds and more subtle tricks such as how not to miss plants hidden in screes.
Perhaps best of all was conversation, which ranged widely from Henry James and the use of
punctuation to Mediterranean plant life and culture, all intermingled with anecdotes about
colourful friends and acquaintances.

Thereafter, our ways separated — he remained in Edinburgh and joined the staff of the University
Botany Department, where he eventually became Professor of Taxonomic Botany (1979-85), while
I went to Cambridge and subsequently to Madrid and Liverpool. Our floristic interests too diverged
and we did not collaborate further in our respective areas although we kept in close touch and it was
as teachers of plant taxonomy that we both felt the need for a text that did justice to the subject,
rather than the old-fashioned McGraw Hill volumes that were all that were available, apart from |
manuals of plant classification systems such as Hutchinson, Warming & Potter and Lawrence (this
latter admittedly having a broader treatment of the subject matter). Thus began our collaboration
on Principles of Angiosperm Taxonomy, which was published in 1963. Although individual chapters
were drafted by one of us, we both revised and often rewrote each other’s contributions. The
intellectual clarity of the text which several reviewers commented on owed much to Peter’s fine feel
for language and style.

168 OBITUARIES

Peter was not at home with many of the ways in which plant taxonomy developed and was
happiest with the more traditional morphological aspects and with exploring the basic principles of
classification which he sought to clarify, based largely on his extensive practical experience of ‘doing
taxonomy’. Yet he enjoyed intellectual discussions with some of the leading figures in the rapidly
changing field of modern systematics and genetics such as G. Ledyard Stebbins and C. H.
Waddington. He did not engage in multivariate systematics, leaving others to master the
techniques. Not for him were molecular systematics or cladistic approaches. Nor did he, with rare
exceptions, participate in international meetings, congresses or symposia, conserving his energies
for what had become his life’s work, the preparation of the Flora of Turkey, begun in 1959. This ten-
volume Flora will undoubtedly go down in the history of taxonomy as a landmark work. Its
completion was a major achievement and very much a result of the energy he devoted to it and the
enthusiasm with which he inspired the small team of collaborators who worked with him on each
volume, although often driving them to near despair as he fussed over details.

He had a wide circle of friends in many different walks of life: when staying with him in his elegant
town house near the Water of Leith, one never knew who would knock at the door at any time of day
or night. He was a knowledgeable collector of modern paintings and drawings and an authority on
Wemyss-ware of which he built up an impressive assemblage. He showed remarkable generosity to
those he favoured and delighted in hosting parties at his home, whether for visiting Russian
botanists or some distinguished artist.

He was one of the most colourful figures in plant taxonomy and will be sadly missed. The
remembrance held for him on the occasion of his funeral was a strangely elating occasion. One could
just imagine him in some botanical Nirvana, peopled with Wemyss cats and his comfortable
furniture and objets d’art, looking down at us and giggling.

V.H. HEywoop

RECOLLECTIONS OF PETER DAVIS

Important as it is to recognise the achievements of Peter Davis in taxonomy, it would be wrong not
to try and record, both for those who knew him and those who did not, the colourful and irreverent
side of his personality which co-existed with his dedication to science.

Unlike many nine-till-five academics, who go home to the worthy sameness of domestic care, his
bachelor life enabled him to pursue wholeheartedly his interests in art, collecting, friendship and the
wilder shores of human association. He was not a career aesthete for whom these things are a pose
or an end in themselves: they were for him an essential and nourishing complement to his botany.
His marvellous parties in St Bernard’s Row, down in Stockbridge, used to bring together a great
variety of congenial spirits from many walks of life, visitors from London as well as Edinburians.
The essential requirement was that people should interest him, interest one another and never be
dull.

He had a fine collection of paintings, mainly but by no means exclusively modern. Many were by
contemporary Scottish artists, all of whom he knew and whose sensuous and assured use of colour
strongly appealed to him. He had also assembled an original and diverse collection of furniture and
bric-a-brac which came together in the pretty house to make a rich and highly distinctive whole. It
was fastidiously organised but livable, not precious. Although he knew a lot about some things,
Wemyss pottery for example, of which he was a pioneer collector, and he had a representative
collection of the bizarre products of Barvas on the western shores of the island of Lewis, a place
never reached by the potter’s wheel, his acquisitions were stimulated by passion and appetite, not
scholarship. A camp thread was discernible in some things, such as the collection of sporrans which
hung like trophies in the downstairs loo, but this was not a dominant note. Everything was
interesting and he had a wonderful eye for the strange and beautiful lurking in unexpected places.
The spoils enlarged the sensibilities of many who came to St Bernard’s Row.

On party nights the jacket of his dark suit would reveal a crimson lining, an uncharacteristic

OBITUARIES 169

excess he would probably not have continued to affect had he been to Oxford instead of Edinburgh.
In no other respect was he provincial. Sartorial effect was not otherwise something he bothered
about at all although his invariable outdoor appearance, in winter at least, did achieve a certain
idiosyncratic style. He wore a narrow cloth cap with sewn-down peak, muffler and a short, heavy
twill overcoat. His hands would be thrust into its pockets as he walked, with short steps, elbows in,
drawn up tight. With his disturbed complexion and ever-twinkling eyes behind his glasses, he looked
like some unfrocked parson who had become the faintly disreputable associate of an on-course
bookie.

It has been convenient for some to see Peter Davis as a sort of Jekyll and Hyde character: the
eminent and reputable academic on the one hand and the frivolous and slightly ‘sinister’ eccentric on
the other. I believe he was more of a piece than this suggests. The wit and judgement he exercised in
personal relations and in his enjoyment of art and the intellectual and physical stamina he brought to
his work were, in fact, a continuum. I once asked him what he thought about the fashionable trends
in genetics and molecular biology as they affected botany. It was an unkind and disingenuous
question as I was pretty sure taxonomy was thought by many at that time to have been upstaged by
these developments, more then perhaps than now. He paused for a long time, as if searching for an
unprejudiced comment on something he found uncongenial, even threatening. Then, quietly and
deliberately: “It is very interesting, and very important; but I believe it loses sight of the whole
plant, its community and its place in nature”’.

He detested pretentiousness, respected hard work, competence and above all integrity in
whomsoever he found it. His favourite novel was that cruel analysis of dishonour among thieves,
The Wings of the Dove. Looking back across the years since I was first taken to one of his parties in
early 1960; an occasion from which so many Edinburgh friendships were to spring, I realise how
keenly I used to enjoy seeing him: always acute, always interesting, always such fun. The last time I
set eyes on him, after we had been in London for some years, was from the car in the twilight of a
winter evening. He was turning out ot Moray Place into Doune Terrace in his cap and twill overcoat,
his hands in his pockets, elbows tucked in as he headed down to Stockbridge. The fading light of the
western sky was reflected in his glasses, giving him an unwontedly blind look. I wanted to stop but
we were in a hurry. Later, when I learned of his last illness, I consoled myself with the thought that
the clouds of forgetfulness had perhaps already begun to envelop him and that he would simply have
wondered whom this importunate stranger could be. I shall never know.

P. HARRISON

RONALD D’OYLEY GOOD
(1896-1992)

Ronald Good M.A., Sc.D., F.L.S., died on 11 December 1992, at the age of 96. An intelligent and
solitary man, he outlived most of his contemporaries. He was the son of a doctor who lived in High
West Street, Dorchester, close to the County Museum, which Good visited regularly from the age of
ten. What motivated his lifelong interests in botany and local history we do not know, but his
teachers at Weymouth College must have encouraged his love of science, in an age when most
intelligent boys were steered towards the classics. He was certainly recording wild flowers for the
Dorset Natural History and Archaeological Society (D.N.H.A.S.) before 1914, when he made the
outstanding record of Ranunculus ophioglossifolius near Tincleton (vide Baker, E. G., Journal of
Botany (London) 52: 277 (1914)). In 1915 the Society awarded him the Cecil Silver Medal for an
essay on Radium.

During World War I he served with the Inns of Court O.T.C. in 1916, was commissioned in the
4th Dorsets in 1917, and served in France until wounded, after which he was moved to Ireland. He
entered Downing College, Cambridge, in 1919 as an Exhibitioner, and later became a Foundation
Scholar and obtained a double First Class Honours in Botany. One of his teachers was Albert
Seward, who may have triggered his interest in plant evolution and geography. At Cambridge he
rowed for his College Eight.

On leaving Cambridge in 1922, Good joined the herbarium staff at the British Museum (Natural
History), with J. E. Dandy as a colleague. Here he built up his knowledge of the floras of the world,

170 OBITUARIES

and produced a series of papers on plant geography. He also visited Canada with the British
Association. He married in 1927.

In 1928 he joined the new University College at Hull as Head of the Botany Department, and
remained there for 31 years, becoming Professor in 1946. During this time the student population at
Hull rose from about 30 to several thousand. Among Good’s students were Eva Crackles, Professor
G. Lucas (Kew), Professor D. Read (Sheffield) and Professor W. Armstrong (Hull). Good used his
time to produce three, scholarly books, one of which, The geography of the flowering plants, was to
go through four editions and is still the standard work on the subject in English. Good preferred the
Darwinian approach to research, involving many years of thought before producing a magnum
opus, to the modern custom of writing or helping to write as many short papers as possible. Good’s
family (he had one daughter) lived at Cottingham near Hull, but every vacation was spent in his
beloved Dorset, where from 1931 to 1939 he carried out an impressive survey of old roads, lost
villages and the county’s flora. This involved visiting 7500 sites and making over a quarter of a
million records of plant species. During this time he co-operated with C. D. Day (an entomologist)
and C. Diver in ecological work, but the survey of Dorset was a one-man effort. Good joined the
D.N.H.A.S. in 1939 and won its Mansel-Pleydell silver medal. The next year he was able to publish
his work on old Dorset roads, but the war delayed other publications. During World War II he
served in the Home Guard in both Dorset and Yorkshire. He had to wait for peace before he could
see through the press his Geography of the flowering plants, a local history of Weymouth and his
Geographical handbook of the Dorset flora. Peace also allowed him to travel, notably to Bahrain
where he collected specimens, and also to Australia, South America and the Caribbean.

In 1959 Good retired from Hull and came to live in Parkstone, Dorset, where he stayed for 18
years. During this time he was an active member of the D.N.H.A.S., leading many field meetings,
editing the annual reports on botany and rainfall, acting as President between 1961 and 1964, and
then becoming a Trustee. He was also a founder member of the Dorset Trust for Nature
Conservation and was instrumental in drawing attention to one of the first Trust reserves at Brackets
Coppice, saving it from devastation. He was an accomplished artist in pastels during this part of his
retirement.

Good’s wife died in 1975, and two years later he took a flat in Albury Park near Shere in Surrey,
on the site of John Evelyn’s 17th century garden. Here I recall commenting on the tidy state of the
grounds during a visit, when he told me with his usual charm that all the able-bodied occupants
helped with gardening chores, and he himself swept the paths. Though less mobile than before, he
produced three more books at Albury, one on evolution, one on lost Dorset villages and a revised
Concise Flora of Dorset. From 1989 his left leg became paralysed and he spent his last three years in
a Henley nursing home.

Good’s contributions to British botany were mostly made in Dorset. His extensive field survey
during the 1930s resulted in many new county records for plants such as Centaurium tenuiflorum,
Epipactis leptochila and Euphrasia anglica as well as new sites for Dianthus armeria, Himantoglos-
sum hircinum, Lobelia urens and Sparganium natans (minimum). It is not clear whether he himself
saw Simethis planifolia at Branksome before its disappearance in about 1925. His extensive field
notes, species maps and some specimens are preserved at the Institute of Terrestrial Ecology,
Furzebrook. The 1948 Geographical handbook pioneered the publication of plant distribution
maps, and he not only anticipated the National Grid by producing his own Grid of Dorset, but also
subdivided each Grid square into 16 parts for mapping. All this work was done by hand, including
the preparation of maps.

In the field of world botany Good was a loner who broke new ground. His broad view of plant
geography was original and well-argued. He pointed out that affinities between the floras of
Australia and South Africa, for example, were best explained by Wegener’s theory of Continental
Drift. At this time Wegener’s theory was unpopular, but with the advent of plate tectonics it has
become accepted. Another original conclusion was that the North Temperate flora is still in a state
of flux following the catastrophe of repeated Ice Ages. He was the first botanist to suggest that the
spread of Himantoglossum hircinum in southern England in the first half of this century was due to
climatic change (New Phytologist 35: 142-170 (1936)). His two books on plant evolution express his
own ideas without reviewing the scientific literature. They emphasise our ignorance of most of the
factors controlling the evolution of plants.

Good’s herbarium specimens will be found in Dorchester Museum (DOR), the Natural History

OBITUARIES igi

Museum (BM) and I.T.E., Furzebrook, while his 1950 collection from Bahrain is at Kew (K). The
list of his publications given below may be incomplete, but shows his eclectic interests.

I am grateful to Mrs Grinsley (daughter), Kate Hebditch (County Museum, Dorchester),
Professor Gren Lucas (ex-pupil), Nigel Webb (I.T.E., Furzebrook) and others who have helped me
to compile this obituary.

MAJOR PUBLICATIONS OF R. D. GOOD

1933 Plants and human economics. Cambridge University Press.

1940 The old roads of Dorset. Reprinted 1966. H. Cummin, Bournemouth.

1945 Weyland: the story of Weymouth and its countryside. Longmans, Dorchester.

1947 The geography of the flowering plants. Longmans, London (eds 2-4 1953, 1964, 1974).

1948 A geographical handbook of the Dorset flora. Dorset Natural History and Archaeological
Society, Dorchester.

1956 Features of evolution in the flowering plants. Longmans, London (2nd ed., Dover, New Monks
1974).

1979 The lost villages of Dorset. Dovecot Press, Stanbridge.

1981 The philosophy of evolution. Dovecot Press, Stanbridge.

1984 A concise Flora of Dorset. Dorset Natural History and Archaeological Society, Dorchester.

H. J. M. BOWEN

SONIA C. HOLLAND
(1912-1993)

With the death of Sonia Holland in January 1993, the B.S.B.I. has lost one of its most accurate and
conscientious recorders and a field botanist who contributed in many ways to the study and
conservation of the British flora. Her countryside interests were wide. She had a good working
knowledge of birds and bird-song and latterly she became interested in dragonflies.

Sonia was one of the daughters of Dr C. A. Hill of Liverpool, where she was born on 4 April 1912.
On her father’s death in 1922, the family moved to Aberdovey, where her love of the countryside
began. She met her future husband, Bill Holland, at an Artillery Camp at Towyn during World War
II and they married and settled in Cheltenham in 1947. They had one son, Clive. Bill Holland died in
1972. Hers was a close-knit family; she kept in close touch with her sisters and enjoyed her visits to
them in West Somerset, where she hunted Black Poplars.

Living in Cheltenham, with the Cotswolds and Severn Vale on her doorstep, she saw the need for
an active natural history society. Thus she got together with some of her naturalist friends and in
1948 founded the Cheltenham and District Naturalists’ Society. Sadly, the established Cotteswold
Naturalists’ Field Club was no longer filling that role. The Cheltenham Society flourished and
expanded to become the North Gloucestershire Naturalists’ Society in 1957 and the Gloucestershire
Naturalists’ Society in 1974. Sonia was Vice-chairman of the Society in 1961 and she edited its
excellent bimonthly journal for many years. Sonia was a founder member of the Gloucestershire
Trust for Nature Conservation (G.T.N.C.) in 1962; she sat on its Council for 15 years and served on
its Conservation Committee. She joined the B.S.B.I. in 1969 and served on the Council (1970-74).
She was the Society’s recorder for v.cc. 33 and 34 until her death.

With the formation of the G.T.N.C., the Society for the Promotion of Nature Reserves (now
R.S.N.C.) passed the management of Badgeworth Nature Reserve from the Cotteswold Natural-
ists’ Field Club to the G.T.N.C. I had known the site, which protects the rare Adder’s-tongue
Spearwort (Ranunculus ophioglossifolius), since schooldays, before the reserve was established,
and I had seen how a fence and no management could be harmful to a rare plant. Under the
Cotteswold Naturalists there was a warden but no management was done for 28 years! I was so
worried about it that I decided to get advice from the then Nature Conservancy’s Regional Officer,
J. H. Hemsley, a sound ecologist and a personal friend. In 1961, Jim Hemsley, the Warden and I

172 OBITUARIES

visited the site in early June, flowering time of the buttercup, but none were to be seen. Jim
suggested that management be carried out in the autumn, but the Cotteswold Naturalists’ Field
Club took no notice. The following year, the G.T.N.C. took over responsibility for the reserve anda
Management Committee was formed, with Sonia Holland as Secretary and me as Chairman. So our
paths met.

Jim Hemsley’s management regime was applied in the autumn of 1962 and the buttercup flowered
well in 1963. It has been seen in flower nearly every year since then. Our official management plan
was compiled largely by Sonia and approved by the Committee. She was a conscientious and
enthusiastic Secretary. She also produced the Badgeworth Nature Reserve Handbook, full of
historical and biological information and listing the known flora and fauna of this small reserve, a
fine example of such a guide. It is now in its third, enlarged edition. As I was living in Surrey and
later in Suffolk, the task of seeing that the necessary management was carried out fell on Sonia’s
shoulders. She negotiated with the owners a small addition to the Reserve, known as Warren’s Pool,
and arranged for a movable fence to allow stock into the pool at certain times.

Sonia was keen to publicise the Reserve and got it an entry in the Guinness Book of Records as
Britain’s smallest nature reserve, 346 square yards (290 m*) together with television and national
newspaper coverage. She also encouraged local people to take an interest. The Badgeworth W.I.
produced a tie with the buttercup on it, and crowning the Buttercup Queen is the main event of the
village fete!

When Sonia was staying with us in Suffolk in 1972, we were out botanizing and I showed her some
fine trees of Black Poplar (Populus nigra), as I was starting to survey its distribution in England and
Wales for the B.S.B.I. On her return to Cheltenham she ‘got her eye in’ for this tree and began to
send me records, not only from her vice-counties, but also from other parts of the country. Sonia
visited her son in Ireland from time to time, and when driving around the countryside there spotted
several Black Poplars. These records, however, were not published, but were confirmed when
Desmond Hobson visited Ireland in 1990 (Watsonia 18: 303-305, 1991).

Sonia was interested in the development of the Gloucestershire Farming and Wildlife Advisory
Group (F.W.A.G.), and came to know the county advisor, John Hughes, who suggested that she
should be his botanical advisor and accompany him when he was asked to visit farms. This she was
pleased to do, as it gave her access to much private farmland where no botanist had previously
trodden. This resulted in Sonia discovering new sites for a number of uncommon plants and she
found several Black Poplars that could not be seen from public land. Gloucestershire F.W.A.G. was
so grateful for her help and the advice she had given to farmers on their wildlife habitats, that it
agreed to publish the results of her county Black Poplar survey as The Black Poplar in
Gloucestershire (1992). In this work she recorded and localised.355 trees, many of them pollarded,
_ all of which but one she had seen and described herself, “‘a labour of love’, as the county F.W.A.G.
president states in his foreword.

Sonia took a great interest in her local flora and discovered where most of the rarities grew and
assisted their conservation. She travelied to London specially to see a solicitor regarding the
threatened woods near Tewkesbury where the rare Cynoglossum germanicum (Green Hound’s-
tongue) grew, and succeeded in convincing him that the woods should be conserved. She personally
encouraged the commoners of Painswick Beacon to remove dozens of seedling pines which were
invading and smothering the best sites for Herminium monorchis (Musk Orchid) in the county, and
she kept an eye on Cephalanthera rubra (Red Helleborine) in all its secret sites. She was delighted
when Antennaria dioica (Mountain Everlasting) was rediscovered on Cleeve Clond, where I had
seen it in the early 1920s.

Sonia’s outstanding contribution to British botany was her work on the composition of the
Supplement to the Flora of Gloucestershire, helped by her two friends, Miss Caddick and Mrs S.
Dudley-Smith. Sonia decided on the format, selected the beautiful colour plates, edited the text and
saw through its production. It is, in my opinion, a model of what a supplement to a county Flora
should be.

Sonia was a member of the British Trust for Ornithology (B.T.O.) and she used to help the county
bird recorder, Dennis Mardle, do the monthly B.T.O. waterfowl count on the numerous flooded
gravel-pits of the Cotswold Water Park in the Upper Thames catchment area. She and Dennis
Mardle jointly published Bird watching in. the Cotswold Water Park, a booklet much in demand
amongst birdwatchers. It was in the Water Park that she developed her interest in dragonflies, which

OBITUARIES 173

resulted in her Distribution of the dragonflies of Gloucestershire (1991), a very good account, with
much information about particular habitats.

Sonia was very unhappy about the future of the splendid wild daffodil bank at Ketford in the
Leadon Valley, which had come on the market. The daffodils grew in species-rich meadows with
other interesting plants such as Gagea lutea (Yellow Star-of-Bethlehem). She tried to get the
G.T.N.C. to buy the site but the Trust was not interested, as the bank had no vehicular access. So
eventually, following the example of that pioneer conservationist G. W. Hedley, who in 1932
bought the Badgeworth buttercup site with his own money, she purchased the freehold of the
Ketford Daffodil Bank. It had been neglected for some years and Sonia, assisted by John Hughes,
returned it to a satisfactory condition. It gave Sonia great pleasure to feel that she had been able to
save this splendid wildlife site for future generations to enjoy.

Some years ago Sonia suggested to me that a very old, nearly dead, pollarded Salix fragilis
(Crack-willow) in the Badgeworth Reserve might be replaced by a pollard Black Poplar. This was
put to the Management Committee and was agreed. Sonia obtained a cutting from a local male tree,
rooted it and planted it in the Reserve in March 1973. It grew well, but nobody knew how large a
sapling had to be before its first pollarding. However, I discovered that John Evelyn in 1664 advised
cutting a sapling off at the required height when it was as thick as one’s arm: this Sonia did in March
1992. Sadly she did not live to see the development of the pollard. At the 1993 meeting of the
management Committee it was agreed that this Badgeworth Black Poplar should be treated as a
memorial to Sonia Holland, who had done so much to conserve Adder’s-tongue Spearwort and so
many other plants over the last 30 years.

E. MILNE-REDHEAD

SONIA HOLLAND: IN MEMORIAM

Sonia was cremated in Cheltenham on 21 January 1993. The service was taken by her neighbour, the
Rev. Walter Jennings, Vicar of All Saints’ Church. In his address he talked about the life-history of
a dragonfly, when the nymph, having lived at the bottom of a pond for some time, climbs a reed stem
and eventually emerges as a mature dragonfly to live thereafter in the air and sunshine but unable tc
rejoin the other nymphs underwater. He said he was telling this story for Emily and Anthony
Sonia’s grandchildren, but I think it was enjoyed by everyone else in the large congregation
particularly those who had helped in her Odonata recording.

Her son, Clive, chose Mothering Sunday as the day to scatter her ashes on her Daffodil Bank at
Ketford. A large party of family and friends met in the village to walk to the reserve on the south
bank of the River Leadon. Passers-by would have dismissed the gathering as just another field
meeting — about 50 people in wellingtons with anoraks and field glasses. Hundreds of daffodils were
flowering on this beautiful and peaceful bank. While Clive scattered the ashes over a large area of
the reserve we talked to friends or enjoyed strolling about. We saw non-flowering plants of Gagea
lutea (Yellow Star-of-Bethlehem) on the reserve and several good specimens flowering outside.
Two large Populus nigra (Black Poplar) pollards upstream reminded us of yet another of Sonia’s
study subjects. Despite the sad circumstances, this really was an enjoyable morning in a lovely
setting with the promise of spring and summer to come. I like to think that Sonia would have
enjoyed and been touched by the occasion. She would also probably have approved of the fact that
we repaired to the Rose and Crown on the way home.

S. DUDLEY-SMITH
BRIAN THOMAS STYLES
(1934-1993)
Brian Styles was born on 26 September 1934 at Chedworth in the Cotswolds and died at Oxford of a

heart attack on 27 June 1993. He was educated at Westwoods School, Northleach, Gloucs., and
Wadham College, Oxford, where he read Botany. He remained at Oxford as a research student and

174 OBITUARIES

worked at Oxford University’s Forestry Institute (formerly the Commonwealth Forestry Institute,
now part of the Department of Plant Sciences) throughout his career. His research revolved around
the study of tropical trees, but his meticulous study of Polygonum aviculare (Knotgrass) and its allies
will have made his name familiar to many members of the B.S.B.I., to whom he was a conscientious
referee for the genus.

His D.Phil. thesis, submitted in 1959, was supervised by the late E. F. (‘Heff’?) Warburg, one of
the century’s most gifted and influential British plant taxonomists and co-author, with A. R.
Clapham and T. G. Tutin, of the much loved ‘CTW’ Flora. Brian successfully elucidated the
taxonomy of the Polygonum aviculare group, that apparently intractable assemblage of annual
weeds and pauciennial plants of strandline communities. His account of them, published in
Watsonia 5: 177-214 (1962), remains the basis of that published in April 1993 in the revised first
volume of Flora Europaea. Warburg, who died in 1962, said that the thesis was the best he had seen
in Oxford and more recent revisions of the classification of Polygonum aviculare sensu lato by
botanists in northern and central Europe, Italy and North Africa have failed to add appreciably to
that elucidated by Brian. He was for more than 30 years the Society’s referee for the Polygonum
aviculare group, latterly jointly with myself.

Brian joined the staff of the Forestry Institute in 1960 as a Research Assistant sponsored by the
Colonial Welfare and Development Office (now subsumed within the Overseas Development
Administration), later becoming the Department of Forestry’s Senior Research Officer and Forest
Botanist. He was employed as a tropical forest botanist long before the current concern and
enthusiasm for the conservation of the world’s threatened tropical forests. Brian was a botanist of
the old school, a diligent researcher who published infrequently but well, a scholar, a linguist and a
field botanist. In the best tradition of Oxford and Cambridge, he threw himself vigorously into the
general work of his Department and the wider world of botany — teaching, supervising, editing and
contributing to the work of the various committees that punctuate the routine of academic life. He
gave information freely and generously to all his colleagues.

He will be remembered especially for his studies on the tropical forest trees of the family
Meliaceae, notably in collaboration with Dr T. D. Pennington, with whom he produced A generic
monograph of the Meliaceae (Blumea 22: 419-540, 1975), a conspectus of the genus Acacia in
Somalia (with A. S. Hassan) and his monographic work on tropical pines. His contribution on Pinus
to Flora Neotropica was sadly unfinished at the time of his death. A 1976 conference volume on
Variation, breeding and conservation of tropical trees, that he edited with J. Burley for the Linnean
Society of London, remains a seminal work on the study of tropical forests.

He had a particular interest in nomenclature and served for many years on the committee for the
nomenclature of cultivated plants. He was deeply concerned about the conservation of the earth’s
diminishing genetic resources, both in the wild and in cultivation. In September 1984 he organised a
most successful conference at Oxford on intraspecific variation, under the auspices of the
Systematics Association, the Royal Horticultural Society and other bodies, the proceedings of
which were published under his editorship as Infraspecific classification of wild and cultivated plants
(Systematics Association Special Volume 29, 1986).

In 1966 he married Cynthia Large, who was both wife and much valued secretary, not least in her
help with the ‘nuts-and-bolts’ organisation of successful conferences. They have a son, Jonathan.
Although not latterly connected with an Oxford College, Brian enjoyed the traditional university
life of lively conversation and good food and wine. A courteous, kindly and deeply religious man,
who rightly insisted that we began a symposium dinner with the appropriate Grace, he reminded me
of a jolly abbot. I warmed to him immediately when first I knocked cautiously on his office door in
the late 1970s to discuss our mutual polygonaceous interests. He was friendly to all those around
him, not least the countless students from Britain and overseas whom he encouraged and helped.
He was deeply sympathetic to the problems of the younger taxonomist, unable to obtain
employment in the contemporary academic world. He loved a good gossip and on my visits to
Oxford from Reading to work in the Fielding-Druce herbarium he would sidle up with a
mischievous grin to ask “‘Is it true that Professor Heywood is about to leave Reading?” or merely to
enquire about the progress of a younger colleague to the altar. He enlivened all those in his
Department with his sense of humour.

Brian’s research demonstrated the vital links between academic knowledge, economic resources
and conservation. His early death deprives the botanical community of a scholar and gentleman at a

OBITUARIES 175

time when plant taxonomy, beset by hostile forces, needs all the good people it can muster,
especially in our Universities.

J. R. AKEROYD

IRENE MARY VAUGHAN
(1889-1993)

With the death of Irene Vaughan, M.A. (Oxon), M.B.E., F.L.S., on 27 January 1993 in her 104th
year, the Botanical Society of the British Isles has lost not only its oldest member — indeed only the
second centenarian of the Society’s membership — but one of the most respected and foremost of
field botanists in Wales of recent decades. Unlike H. N. Ridley (1855-1956), the B.S.B.I.s other
centenarian, Mrs I. M. Vaughan was not a professional botanist. She had read history at St Anne’s
College, Oxford (1908-1911) but, as a woman, had not been permitted to take her degree
‘officially’. Times changed, and so, when she was 86, her degree was duly conferred.

As Irene Mary Rope, she was born and ‘raised’ in Shrewsbury, where her doctor father was in

FicureE 1. Mrs Irene M. Vaughan at Dryslwyn Castle, Carmarthenshire, 7 April 1978. (Photograph courtesy of
W. Condry.)

176 OBITUARIES

practice with a Dr Burd, one of whose nine daughters he married. Irene must have accompanied her
father on his rounds in pony and trap, and it is known that he greatly encouraged her, the youngest
of his six children, in her love and enthusiasm for flowers and all wildlife; but he died when she was
only nine. His family came from Suffolk (near Woodbridge) and much-loved visits were made to the
family home there for many years — her earliest memory is said to be of butterflies and Zinnias in this
Suffolk garden.

When World War I came, Irene joined a British team driving ambulances in Serbia. Then, joining
the newly formed W.R.N.S., she was posted to Gibraltar, rapidly promoted, awarded the M.B.E.
(Military), and was one of the first W.R.N.S. officers to serve overseas. Here in Gibraltar she met
her future husband, a naval officer, one of whose duties was to test Irene in coding; this she did
without error. Such meticulous attention to detailed exactitude stood her in good stead in later years
and was surely the foundation of her precise keeping of plant records and data of all kinds, and of
her ability to retain the voluminous character detail of the Roses, which became her speciality.

Meanwhile she had married Captain H. R. H. Vaughan R.N. whilst they were both still serving in
Gibraltar; and it was here that their infant son died. In later years there were occasional visits to her
husband during his three-year postings overseas; one was to the West Indies, and on another the
high point must have been the desert picnic with the Emir of Kuwait when uncooked sheep’s eyes
were the special delicacy on offer.

Sadly, in 1938, their daughter died on the eve of her fifteenth birthday, and it was then that the
move to Wales followed. Here, in a valley that the daughter had loved, Irene ran, single-handed
throughout the war, the small farm at Rhandirmwyn near Llandovery, Carmarthenshire, until Capt.
Vaughan’s retirement in 1945.

It was now that both the Vaughans became involved in so many aspects of conservation,
environmental ‘welfare’ and ‘wildlife’ organizations in Wales. He was closely concerned with
revitalizing the Council for the Protection of Rural Wales with her support, but foremost was their
work in rescuing the Red Kites of the upper Tywi valley from oblivion. Throughout the war, Mrs
Vaughan had ‘carried out the greater part of the work of locating nesting pairs and verifying
results’ (H. M. Salmon, in Lacey, W. S. (ed.), Welsh Wildlife in Trust, pp. 67-79, Bangor, 1970)
and gaining the co-operation of land-owners and farmers. In the early post-war years she and Capt.
Vaughan were carrying the whole burden of the organisation of Kite conservation — he was
Chairman of both an earlier and a later Kite Committee. For their immense services to Red Kite
protection over 30 years and more, they jointly received first the R.S.P.B.’s Silver Medal and then
their Gold Medal (cf. R. Lovegrove, The Kite’s Tale, Sandy, Beds., 1990).

The West Wales Field Society (later Naturalists’ Trust) was another joint concern, with Capt.
Vaughan co-editing the first few issues of their journal, Nature in Wales (1955-1956). Mrs Vaughan
had organized a short-lived ‘Botanical Association’ by then, and later was mainly responsible for
starting their Carmarthenshire Branch. She undertook various surveys on this Society’s behalf, and
in 1971 helped pioneer the Farm Nature Reserve Scheme which the Reserve Committee of this
branch had formulated. She served on the then Nature Conservancy's Committee for Wales (1958—
1960) and made a botanical survey and site quality report for a prospective National Nature Reserve
on the Castle Martin Tank Range. In 1965, with T. A. W. Davis, she visited and reported on the
prospective Nature Reserve status for Ramsey Island. Her spirited submission was largely
instrumental in saving the floristically rich Towyn Burrows from threat from the Ministry of
Defence. In 1967 she partook in the opposing, by the Council for the Protection of Rural Wales, the
Nature Conservancy and the B.S.B.I.’s Conservation Committee, of the construction of a road
through Kenfig Burrows, Glamorgan. She was much concerned with safeguarding the upper Tywi
valley from the threat of forestry and later from the Llyn Brianne reservoir project. She had worked
closely with Dafydd Davies in opposing the building of the dam, giving evidence at the inquiry on
behalf of the B.S.B.I. and the Brecknock County Naturalists’ Trust. The Quercus petraea woods at
Allt-rhyd-y-groes became a National Nature Reserve chiefly as a result of her reports, and near
Rhandirmwyn are the R.S.P.B. reserves of Gwenffrwd and Dinas; near Cilycwm a Roadside Verge
Nature Reserve protects Epipactis helleborine (Broad-leaved Helleborine).

Irene Vaughan was primarily a field botanist, with Carmarthenshire (v.c. 44) as her central
interest. Here she explored extensively, gaining a wide knowledge of the flora. She joined the
B.S.B.1. in 1952, and the start of the Maps Scheme must have been an added incentive for her. In
addition to her own contribution, as County Referee, she checked all the field cards from v.c. 44 for

OBITUARIES 177

the Atlas of the British flora (1962). She also edited the botanical notes and plant records for West
Wales in the early issues of Nature in Wales (1955-1956). Increasing involvement with the B.S.B.I.
in Wales followed: she had been local secretary for v.c. 44 (1955) and District Secretary for South
Wales (1961-2), and then became vice-county Recorder (v.c. 44) in 1961, a post she held until 1978,
just before her retirement to Suffolk in early 1979. From its inception she was on the B.S.B.I.s
Welsh Committee, for which she was Representative to the Council from 1962-1970. When the
Regional Committee for Wales was set up in 1962, Mrs Vaughan took the Chair at its first Annual
Meeting and A.G.M. in September 1963, and was Vice-chairman until 1965. Subsequently she
acted as joint and then general secretary up to 1970, and edited the Weish Region Bulletin (later
B.S.BJI. Welsh Bulletin) from 1971-1976, jointly with S. G. Harrison. She led many B.S.B.I. Field
Meetings: to Llanstephan (1967), Towyn Burrows (1969), Kidwelly (1970), and in 1967 to the
Upper Tywi Valley, visiting the Sessile Oakwood at Pen-y-rhiw-iar (by the Allt-rhyd-y-groes
N.N.R.), the site of the then proposed Llyn Brianne dam, and the Rhandirmwyn School Nature
Reserve (later submerged). Carreg Cennan was another favourite haunt and on these and other
occasions many new records were made which led to articles in the Welsh Region Bulletin, Nature in
Wales or Watsonia. These are listed in the bibliography, below. These articles were succinct and
concerned only the most outstanding finds. They were presented in their wider-than-Wales
significance and their ecological, distributional, historical or taxonomic setting. Her numerous
records, many of them new for the county, made a major contribution to R. F. May’s List of the
flowering plants and ferns of Carmarthenshire (1969); they are now in the safe-keeping of the present
vice-county recorder, Richard Pryce. Many of her records too appeared in T. A. W. Davis’ Plants of
Pembrokeshire (1970).

Mrs Vaughan was widely acknowledged in Wales and beyond for her expertise at the
determination of the wild species of Rosa, from at least the early 1960s onwards. This special
interest must have been aroused early: perhaps during the last war, when she is known to have
organized the collecting and making by the Women’s Voluntary Service of rose hip and other wild
fruit jams and jellies in a back room of her Rhandirmwyn home. She pioneered the study of Rosa in
Carmarthenshire and already in 1953 had contributed 36 specimens for Sylvester Bradley’s ill-fated
Rose Survey of 1952-1954 (R. J. Gornall & A. Primavesi, Watsonia 17: 356-359, 1989; R. J.
Gornall, pers. comm. 1993). All came from Rhandirmwyn, and comprised six or more determinable
taxa including hybrids; her R. obtusifolia specimen was one of the only five represented in the near-
thousand surviving from the country-wide survey. By 1955 she had already made at least two Rosa
records new to v.c. 44. Consulted widely long before becoming an official Referee (1975-1984), at
first jointly with R. Melville and later with G. G. Graham as well, she had published (1965) an
excellent key and synopsis on the “‘Recognition of the Roses”’ for taxa known or likely to occur in
Wales. Aithough a follower of A. H. Wolley-Dod, she understood the limitations of his system and
that many of his plethora of varieties and forms would ultimately become recognized as hybrids, and
was well aware of tre peculiarities of the genetical complications in this genus. Her penultimate
publication, in 1982, was a digest of the 13 Suffolk taxa of Rosa, three of which she had added herself
to the county fiora, with yet another later that year; all this in her three years Suffolk residence at
over 90. She was asking for the latest Rosa news just weeks before she died: what joy the Rose
Handbook would have given her!

In 1962 Mrs Vaughan was responsible for tuition in Rose identification at the Critical Plants
course held at Preston Montford Field Studies Centre. In 1962 too, Mrs Vaughan was elected a
Fellow of the Linnean Society of London, and in 1971 held office in Sect. K at the Swansea meeting
of the British Association. She was also instrumental in setting up the Llanelli Naturalists in 1971/2.

Mrs Vaughan’s collections of Rosa are in the Welsh National Herbarium at Cardiff (NMW), with
further specimens at the Ipswich Museum (IPS). Altogether she donated over a thousand flowering
plants and ferns to NMW,, as well as a number of bryophytes and lichens: most were from Wales,
both north and south. Her interests covered a wide range of living things: on perhaps her last ~
‘excursion’ — to Dunwich — in late 1991, she showed delight in the fungi there. Dogs ‘appear’
throughout her life, from those early days in Gibraltar when she had a police summons for
“unlawfully suffer[ing] a dog to be at large in a public highway’! Horses were another great love:
she would extol the advantages of botanizing from horseback. And then we hear of mid-night visits
to comfort a sick bull.

Irene Vaughan’s friends and admirers have many affectionate memories. Although outwardly

178 OBITUARIES

maybe intimidating to some, she gave unstinting encouragement and help to many a young botanist
who recall — inter alia — her agility in scrambling up (and slithering down) steep slopes more easily
than many half her age. Local legends abound — trundling bee-hives through the village at
Rhandirmwyn, with Dafydd Davies, in a cloud of angry escaping bees; ‘Mrs Vaughan’s corner’ — a
sharp turn on the way to Llandovery invariably taken on the inside in her Mini, and other
adventures in her unstable ‘Bubble-car’ in the 1960s. There are fond memories of fluttering leaves of
her precious copy of “Wolley-Dod’ escaping in the Welsh wind (retrieved by younger members); the
generosity with which she took so many young and inexperienced to see highlights of the
Carmarthenshire flora, outings with local friends and the rush back to get Capt. Vaughan’s 4 o’clock
tea. There were ‘immediate’ delicious refreshments — “Oh! the Aga does them’’. This hospitality
continued in Suffolk well into her late 90’s — three-course meals, complete with sherry, coffee (and
no washing up) for visitors from Wales. Although frustrated by increasing deafness and poor sight,
her botanical enthusiasm continued to the end. Always a keen gardener, as witness the gardens at
Rhandirmwyn and Cilycwm, and later at Woodbridge she extended her garden when 95 and planted
a special Sorbus domestica (Service-tree) when 98; her nephew observes that she was still active in
the garden up to six months before she died. By the time she reached 97 she remarked: “‘I don’t like
people to know I’m so old — they’re beginning to look at me as if I were Exhibit A”’. For her 100th
birthday she insisted on no publicity, no presents, or “‘for it to be thought that it made her more than

one day older’’. A lunch party was held, with over 80 friends and relatives; the display of documents | |

and photographs covering her life are now gathered in a commemorative album: a telegram from
the Queen, a special Apostolic Blessing from the Pope, greetings from the Director of W.R.N:S.
and many environmental and botanical organizations. The ‘locals’ from Rhandirmwyn and Cilycwm
held a party. A special issue (48) of the Welsh Bulletin had articles recalling her life in Wales and
photographs of her in the field aged 88, 89 and 96.

Her’s was a remarkable, long and active life of service to Wales, its Roses and Kites and its lovely
countryside. She gave credit to others but eschewed it for herself. It was a privilege to know her. The
wife of Sir J. E. Smith, Lady Pleasance, likewise lived to 103, alert to the last, and held ‘I am for
inquiry’: such was Irene Vaughan. Three botanical colleagues were at her funeral — the Mass in her
beloved Latin. As William Condry wrote (The Guardian, 30 January 1993) “there must still be
people living in. . . the beautiful country of the upper Tywi river who remember seeing this small
figure on a large white horse disappearing up the tracks towards the hills. Perhaps Irene Vaughan, as
a lone horse rider ever searching for her beloved kites, will pass into local legend.”

I am indebted to her nephew, Crispin Rope, and sister-in-law, Mrs L. D. Rope, for personal
details and for making available the commemorative volume, also to her many friends and admirers,
from Wales and Suffolk, who supplied information and memories.

A. P. CONOLLY

PUBLICATIONS OF IRENE VAUGHAN

1964 Preliminary notes on the distribution of roses in Carmarthenshire. B.S.B.I. Welsh Region
Bulletin 1: 6-7.

1965 Recognition of the roses. B.S.B.I. Welsh Region Bulletin 4: 1-5.

1965 [with T. A. W. Davis]. A visit to Ramsey Island, Pembrokeshire. B.S.B.I. Welsh Region
Bulletin 6: 5-6.

1967. Rosa micrantha Borrer ex Sm., R. tomentosa Sm., R. afzeliana Fr. and R. sherardii Davies
(in notes and keys of Papers read at the Local Flora Writer’s Conference, Bristol, 1966).
Proceedings of the Botanical Society of the British Isles 6: 283.

1967 Notes on the origin and distribution of Narcissus obvallaris Salisb. B.S.B.I. Welsh Region
Bulletin 10: 1-3.

1970 ‘Towyn Burrows, Carmarthenshire, in Lacey, W. S., ed., Welsh Wildlife in Trust, pp. 136—
143. Bangor.

1971 A note on Towyn Burrows. Nature in Wales 12: 1974.

1972 [with J. W. Donovan & T. A, W. Davis]. Notes on the Dune Gentian. Nature in Wales 13:
33-36.

OBITUARIES 179

1972 Lathyrus palustris L. in Carmarthenshire. Watsonia 9: 137.

1972 Hedge destruction. B.S.B.I. Welsh Region Bulletin 17: 6-7.

1973 Notes on the increase of Pyrola rotundifolia subsp. maritima (Kenyon) E. F. Warb. on Welsh
dune systems. B.S.B.I. Welsh Region Bulletin 18: 7-8.

1974 A Molinia meadow. B.S.B.I. Welsh Region Bulletin 20: 6-8.

1974 [withQ. O.N. Kay & R. H. Roserts]. The spread of Pyrola rotundifolia L. subsp. maritima
(Kenyon) E. F. Warb. in Wales. Watsonia 10: 61-67.

1976 A note on Sorbus torminalis from Silva, or a discourse of forest-trees, and the propagation of
timber in His Majesties dominions, by John Evelyn Esq. 1664. B.S.B.I. Welsh Region
Bulletin 24: 4.

1976 [ED. Mrs I. M. VAuGHAN] Sorbus torminalis in Powys, Cardigan and Carmarthenshire.
B.S.B.I. Welsh Region Bulletin 25: 4-6.

1976 Of conservation interest. B.S.B.I. Welsh Region Bulletin 25: 9-10.

1977 Lathyrus palustris sites in Somerset and in Dyfed. B.S.B.I. Welsh Bulletin 27: 15-17.

1977 Pyrola minor in Pembrey Forest, Carmarthenshire. Watsonia 11: 388-389.

1978 Lathyrus palustris L. var. pilosus (Cham.) Ledeb. in v.c. 44. B.S.B.I. News 20: 14.

1979 The decline of Liparis loeselii (L.) Rich. var. ovata on Towyn Burrows, Dyfed. B.S. B.I.
Welsh Bulletin 30: 8-10.

1981 Obituary: Thomas Arthur Warren Davis (1899-1980). Watsonia 13: 357-358.

1982 An introductory note on the native Roses of Suffolk. Transactions of the Suffolk Naturalists’
Society 18: 304-307.

1985 Notes on the identification of species in the genus Rosa in Carmarthenshire. Llanelli
Naturalists’ Newsletter, June 1985.

EDGAR DUNSTAN WIGGINS
(1911-1993)

Edgar Wiggins, an Honorary Member of the Botanical Society of the British Isles, was born on
12 September 1911 and died on 23 January 1993. A Londoner born and bred, Edgar was educated at
Latymer School, Hammersmith, and the University of Reading, where he graduated in 1932 witha
B.Sc. in Agriculture. At that time, as now, jobs in botany were hard to find so he stayed on at
Reading to take a post-graduate course in Education, after which he taught for some years at
Fairlop Upper School, Ilford. In 1940, during World War II, he joined the staff of Seal Hayne
Agricultural College, Devon, and worked subsequently for the Ministry of Agriculture on problems
of home food production — particularly potato blight. After the war, Edgar was Manager of Bayer
Agricultural’s Crawley Research Station, then Liaison Officer on the construction and development
of Fison’s new Levington Research Station, and later Manager of Fison’s Head Office Library and
Technical Information Department.

Communication, both written and verbal, was one of Edgar’s special talents, and one of his main
interests outside work was in amateur dramatics. Another interest was gardening, and he was a
founder member of Fison’s Gardening Club, for whom he edited Rakings, the Club’s newsletter, for
some 20 years. For some 35 years during the Cold War period he was involved in the scientific
aspects of Civil Defence. After taking a course at Cambridge University he became the Senior
Scientific Intelligence Officer for the East Suffolk Civil Defence Corps. Later he became directly
responsible to the Home Office as Scientific Adviser on Civil Defence for the Eastern Region. He
also became joint editor of a periodical, Fission Fragments, produced for the information of Civil
Defence scientists throughout the U.K.

Edgar joined the B.S.B.I. in 1957, and when the Society was in need of an Editor for B.S. B.T.
News his offer of editing experience in technical journalism was accepted, and he was then Hon.
Editor of 28 numbers of B.S.B.I. News, from April 1977 (No. 15) to April 1986 (No. 42). He then
became known to us as Wiggy — at the time he lived at ‘Cowpasture Farm’ and when asked how he
would like to be addressed, he replied: ““‘Wigginius pascum-vaccarum — the general usage is Wiggy,
free from any and all — an informal colloquialism, used universally; it comes naturally and trips
easily off the tongue. Edgar-— generally used by those with a professional relationship . . . Dunstan—

180 OBITUARIES

restricted, as far as possible to family and intimates. (But don’t take this too seriously. I respond to
‘Darling’ or ‘Hi yow’ from anyone.)”

During his years as Editor he furthered the scope, success and popularity with members, which
B.S.B.I. News enjoys today, and he particularly promoted the use of plant drawings as illustrations.
For these he sought and encouraged artists who donated their skills to B.S. B.I. News, contributing
to a significant collection of plant drawings which is building up through the years. We all remember
Wiggy during his time as Editor as a notable character at B.S.B.I. meetings, with his neat beard and
friendly approach to all his potential authors and contributors, and he stamped his own brand of
humour and communication on to B.S.B.I. News.

Failing eyesight forced him to resign and sadly he became blind in 1987. In his last years he was
totally chairbound — a disability which must have been extremely frustrating to such a volatile spirit.
His wife Julia nursed him throughout and we extend our sympathy to her in her loss.

M. Briccs

Watsonia 20: 181-183 (1994) 181

Report

THE ANNUAL GENERAL MEETING, 15 MAY 1993

Three papers were presented at the Annual General Meeting held at the Conference Hall,
Lincolnshire College of Agriculture and Horticulture, Riseholme, Lincoln on 15 May 1993, two in
the morning and one in the evening. The theme of the morning’s papers was Sir Joseph Banks, 1993
being the 250th Anniversary of his birth, the theme of the evening paper being “Nature
Conservation’, the papers thus linking up with the outdoor programme on Saturday afternoon and
the Sunday field excursions.

Mr Harold B. Carter, F.R.S.E., F.L.S., eminent author on Sir Joseph Banks and in charge of the
Banks Archive project at the British Museum and the Royal Society, first spoke on Banks as a
British botanist. He enlarged on some of the Banks Lincolnshire and British record sources,
regretting that much had been lost, and outlined Banks’ Welsh itinerary with connections to New
South Wales and also his Lincolnshire and Derbyshire background. Mr Carter illustrated his very
scholarly address with slides of herbarium specimens and drawings — these were of particular
interest as they had been collected and collated from a large number of sources during his
phenomenal study of Banksian papers.

Mr David N. Robinson M.Sc., Hon. Secretary for the Lincolnshire Trust for Nature Conser-
vation, next gave his address on Sir Joseph Banks — the Lincolnshire connection. Mr Robinson,
himself a native of Horncastle and very much an expert on the Lincolnshire scene, illustrated his talk
with both mounted display panels and slides. His very detailed account of the childhood
environment of the young Joseph and his family at Revesby Abbey together with activities in later
years on his Lincolnshire estates and with Lincolnshire agriculture — particularly fen drainage — was
both entertaining and significant. This was ingeniously illustrated with slides of old Revesby and
Horncastle, modern and old litho illustrations of fenland and also by slides of the portraits now at
the Usher Gallery in Lincoln and at the museum in Boston. |

The two papers were comp!ementary and gave a much appreciated sketch of the activities of this
celebrated botanist.

After the A.G.M. and lunch, two coach parties with 75 people travelled down through Lincoln to
the R.S.N.C.s headquarters at Waterside South on the River Witham where the river flows through
the celebrated Lincoln Gap. Here B.S.B.I. members were welcomed by the Chief Executive, Mr
Tim Cordy (Dr Perring’s successor at R.S.N.C.), Mr Tim Sands and the R.S.N.C. Executive
Committee whose meetings had been relocated from London to Lincoln for this event.

A sparkling elderflower champagne and cake reception was attended by over 100, the cake being
decorated splendidly with the logos of both the R.S.N.C. and B.S.B.I., with a conservation theme
throughout. The newly elected President, Dr Franklyn Perring, ceremoniously cut the cake and
replied on behalf of the Society to Mr Cordy’s welcome. Dr Perring applauded the co-operation of
the two groups in the past and looked forward to future links between them.

Two very adventurous coach drivers then proceeded to drive the party up the steep route to the
Cathedral area, belying the usual vision of flat Lincolnshire. Here at the Lawn Visitor Centre the
party was welcomed by Mr Glyn Stocker, Lincoln city landscape architect and designer of the Sir
Joseph Banks Conservatory and the John Dauber garden. Designed to contain representative plants
found on the journeys of Sir Joseph and subsequent collectors from Kew, these projects are part of
the City of Lincoln’s plan to commemorate one of Lincolnshire’s great sons. Many plants are still to
go in and both Kew and Chelsea Physic Garden are contributing relevant material. Mr Stocker led
the group through the conservatory and then out to the garden — still in glorious May sunshine, with
views down to the Trent Valley, of the Cathedral and of Lincoln City.

After the conference dinner Mr A. E. Smith, M.A., O.B.E., Chairman of the Lincolnshire Trust

182 REPORT

for Nature Conservation, took up the conservation theme with an illustrated address on Nature in
Lincolnshire. This was a masterly resumé, with links to national policy and the original vision and
formation of County Nature Reserves to which Ted Smith’s contribution was highly significant. As
Ted Smith had preceded Dr Perring at R.S.N.C. for many years the evening aptly rounded off the
mood of the meeting and was also preparatory to the Sunday excursions.

Two book displays together with sales tables were arranged in the conference hall by Dr and Mrs
Perring of B.S.B.I. Publications and Mr and Mrs Jon Atkins of Summerfield Books. These were
much appreciated by members throughout the day.

FIELD EXCURSION HELD IN CONJUNCTION WITH THE A.G.M.

CENTRAL LINCOLNSHIRE NATURE RESERVES, N. LINCS (v.c. 54), 16 May 1993

About 60 members met at the Forestry Commission car-park at Chambers Wood, Langton by |

Wragby. The woods are part of the Bardney Forest ancient Small-leaved Lime woodlands S.S.S.I.,
now covered by a Forestry Nature Reserve Agreement between the Forestry Commission, English
Nature and the Lincs. Trust. These have been worked extensively in the woodland surveys done by
Dr George Peterken. The party was split up into groups led by Pat DeLap (Forestry Commission
Officer), Keith Stephenson, a B.S.B.I. member and local Langton farmer, and I. Weston. A route
was taken through Great Scrubbs Wood, Little Scrubbs Meadow (Lincs. Trust Reserve) and Little
Scrubbs Wood. Many stages in the coppice management were observed. Notable of course were
Tilia cordata (Small-leaved Lime) and Sorbus torminalis (Wild Service-tree) but there was also a
very large Tilia platyphyllus (Large-leaved Lime). Both Crataegus monogyna (Hawthorn) and C.
laevigata (Midland Hawthorn) together with hybrid material were seen with many other woodland
plants. In the meadow Orchis morio (Green-winged Orchid), Serratula tinctoria (Saw-wort),
swarms of violets — of hybrid origin? — and surprisingly Molinia caerulea (Purple Moor-grass) —
uncommon in TF/1.7.

The party then proceeded in cars via Horncastle Market Sanene. which has the town house of Sir
Joseph Banks, to the luncheon venue on the village green at Revesby. David Robinson met the
party and pointed out features of note including the old and new Revesby Abbey sites and the Sir
Joseph Banks almhouses. Despite a rush to the afternoon venue — the Moor Farm S.S.S.I. Lincs.
Trust Reserve — all arrived on time. 54 members met here. Again the party split up into a number of
groups led by Ted Smith and Mr Terry Bailey, the Reserve Warden, another by Lincs. B.S.B.1.
members Ms R. Nickerson, Miss J. Knibb and Mrs R. Everatt, and one by I. Weston. The reserve
includes a lowland raised Sphagnum bog, a nationally scarce habitat unique to Lincolnshire, with
wet and dry heaths, acid marsh and birch and oak woodlands.

Teesdalia nudicaulis (Shepherd’s Cress) was at its most prolific for years and dominant over a
large area of dry heath. Ornithopus perpusillus (Bird’s-foot), Cerastium arvense (Field Mouse-ear),
Rumex acetosella subsp. acetosella (Sheep’s Sorrel) and Scleranthus annuus (Annual Knawel) were
noted, as were extensive swards of Ceratocapnos claviculata (Climbing Corydalis), dominating
ground cover in the woodland. On the wet heath field Montia fontana (Blinks) was plentiful (not a
common local plant) and Ophioglossum vulgatum (Adder’s-tongue) and Pedicularis sylvatica
(Lousewort) were amongst the sward. Eriophorum angustifolium (Common Cottongrass) had a
dominant spot in the meadow. Mr Clive Jermy recorded Dryopteris X deweveri (D. carthusiana X
D. expansa) — a new record for us.

Time overtook us here and only a handful of members embarked on a hasty visit to the adjacent
Kirkby Moor Reserve to see fine swards of Viola palustris (Marsh Violet) in full flower in the marshy
area — a plant seldom seen flowering well in Lincolnshire.

Thanks are due to the Principal and catering staff of the Agricultural College, the Chief Executive
and staff of the R.S.N.C. and Lincoln City officers. Special thanks are also due to our generous
guest speakers with their extra Sunday involvement — to B.S.B.I. member Mrs Thelma Brown for

I a Sas ——

REPORT 183

hospitality, to Mr Harold and Dr Mary Carter, and to Lincs. B.S.B.I. members for their support
before and during the A.G.M. weekend.

IRENE WESTON

[The above Report arrived too late for it to be incorporated with the main Report of the A.G.M. in
Watsonia 20(1) — Ed.]

8
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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
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Papers and Notes: Dr B. S. Rushton, Department of Biological and Biomedical Sciences,
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1 Si ean ae

WATSONIA

Botanical Society of the British Isles
Patron: Her Majesty Queen Elizabeth the Queen Mother

Applications for membership sheuld 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 1994-95

President, Dr F. H. Perring
President-elect, Mr D. A. Pearman

Vice-Presidents, Mr P. S. Green, Dr G. Halliday, Mr A. C. Jermy,
Mr A. O. Chater

Honorary General Secretary, Mrs M. Briggs
Honorary Treasurer, Mr M. Walpole

Editors of Watsonia

Papers and Notes, J. R. Edmondson, R. R. Mill, E. C. Nelson, B. S. Rushton*
Plant Records, C. D. Preston

Book Reviews, C. D. Preston

Obituaries, J. R. Akeroyd

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

Watsonia 20: 185-194 (1995) 185

Presidential Address, 1994

FRANKLYN PERRING
DRUCE IN NORTHAMPTONSHIRE
‘Tn rising to address you. . . , after a study of various objects of scientific interest for more than half

a century, it would seem mere hypocrisy if I told you I felt at all nervous in doing so”. Thus Miles
Berkeley began his address at the inaugural meeting of the Northamptonshire Natural History
Society in 1876. He is one of the characters in the tale I am about to tell and I could find no better
way to express my own feelings today: after 42 years as a member of the Society which has given me
so much pleasure and has for much of my life been at its centre I do not feel nervous, I feel amongst
friends and I am hopeful that you will be indulgent towards the incompleteness of what I have
prepared for this occasion.

I decided to concentrate my address on Druce in Northamptonshire when I realised that the
choice of Oxford for our A.G.M. this year would mean that, as a resident of Northamptonshire for
the last 23 years, I would be following the path which Druce took when he left the county for Oxford
in 1879. Druce’s move brought him into the mainstream of British field botany and ultimately to the
position where he was effectively the full-time, unpaid coordinator of the Society’s affairs. But by
further happy coincidence I find myself arriving in Oxford from Northamptonshire at the very
moment that the Society is taking steps to appoint a coordinator of its scientific affairs. I will touch
upon this briefly at the end of my address because I believe the consequences for the Society could
be as important as those which Druce brought about here in Oxford nearly a century ago.

George Claridge Druce (Fig. 1) was perhaps destined to be a great botanist from the day of his
birth, 23 May 1850 — the same day as Linnaeus, 143 years his senior— and, incidentally, of my
colleague and past-President Max Walters. But in other respects his birth was not so fortunate. He
was the illegitimate son of Jane Druce and no father is named on his birth certificate: there is some
speculation that his second name Claridge links him with a local family where his mother might have
been a maidservant. Druce writes later of ‘‘his patrimony” (Druce 1930), so his father’s family were
surely people of substance. He was born in Potterspury, a village in the south of the county which
had other connections with the Druce family — a Phoebe Druce married Thomas Woodward at
Potterspury on 14 October 1813. Although he made no secret of his birth elsewhere, towards the
end of his life, in writing the introduction to his Flora of Buckinghamshire (1926), he fudges the issue
a little: “‘Death early deprived me of paternal care, but in an unlimited degree there was showered
on me the devotion and love of an all too unselfish mother.”’ He added that his father’s family were
Northamptonshire people but “my mother’s family were natives of and farmers in Buckingham-
shire”. Their area appears to have been around Woughton-on-the-Green, about 14 km to the S. E.
‘where my botanical baptism was received”’.

Soon after his birth the family moved 3 km to Old Stratford, on the county boundary with only the
River Ouse between him and Buckinghamshire. There he spent the first five or six years of his life
and, as he records in both the Flora of Buckinghamshire and the Flora of Northamptonshire, the
river on the one hand and the nearby Whittlebury Forest only 3 km to the west on the other were
wonderful areas in which to become ‘turned on’ by wild life. ““Whittlebury Forest . . . with its
myriads of Primroses, and the four kinds of Violets which I grew in my tiny garden, the glossy leaved
Spurge Laurel, and Bluebells” (Druce 1926). The Ouse “‘had the White and Yellow Water Lilies;
the banks were fringed with the sweet Acorus and canary-yellow anthered Sedge, and its meads
were fragrant with Meadow Sweet or silvered with Lady’s Smock, and the hedges shaded the
beautiful Geranium pratense” (Druce 1926).

When Druce was five or six (he gives 1855 in the Flora of Buckinghamshire and 1866 (sic) in Flora
of Northamptonshire) the family moved 3 km N.W. to Yardley Gobion into 13 Chestnut Road (fide

186 F. PERRING

FicurE 1. Photograph of George Claridge Druce by Robert Chalmers F.R.P.S.

Andrew Robinson, Northamptonshire botanist who lives in the village) which runs into the rather
pleasant Chestnut Green. Here he lived with his mother and an aunt, Ann Blunt, according to
Sylvia Chandler who has looked at the 1861 Census where they are described as:

Blunt, Ann Widow 63 Fundholder born Wollstone, Bucks

Druce, Jane Niece 33 born Woughton on the Green, Bucks

Druce, George Nephew 10 Scholar born Potterspury
But, still close to Whittlebury Forest, his interest in wildlife expanded and he developed an interest
in entomology. He recalls (Druce 1926) “‘by the age of fourteen a very representative collection of
its lepidoptera was made, including an Antiopa [Camberwell Beauty] and plenty of Sinapis [Wood
White], Paniscus [Chequered Skipper] and rare clearwings. Pupae were dug for and larvae bred, so
that the various Sallows, Poplars, Buckthorn, Verbascums, etc., were familiar objects’. Wood
White is still a feature of the area and some of the finest colonies in Britain occur in Salcey Forest 5
km N.E. of Yardley Gobion. The Chequered Skipper sadly no longer occurs at Whittlebury, or in
the N.E. of Northamptonshire around its former stronghold in Rockingham Forest or indeed
anywhere in England. The only sign of it now to be seen in Northamptonshire is the pub sign at
Ashton, the estate village developed by Charles Rothschild, founder of the Society for the
Promotion of Nature Reserves (S.P.N.R.), at the beginning of the century and where his daughter
Miriam Rothschild still lives. Druce’s friendship with Charles Rothschild, which I shall return to
- later, was surely based on their mutual interest in butterflies.

Druce regarded Yardley Gobion as his home village. He named his house in Oxford Yardley
Lodge and even ‘Lodge’ may have local connections as houses at the edge of forests in

PRESIDENTIAL ADDRESS, 1994 187

Northamptonshire are often so called and the school in the village of his birth is called Potterspury
Lodge School. His mother probably kept her connections with Yardley for the rest of her life. In the
1881 Census (fide Sylvia Chandler) Jane Druce is listed as a Visitor at the Potterspury Union
Workhouse, which is in Yardley Gobion, and she is described as a housekeeper. Druce (1930) said
that his mother went to live with him in Oxford in 1879 and that “‘. . . much of my after success was
due to her energy help and cheerfulness’. Perhaps she continued to help until he was weil
established but may then have returned to Yardley from time to time, though she was living with her
son in Oxford at the time of the 1891 Census where she is described as a widow! Druce erected a
memorial tablet to her in St Leonard’s Church, Yardley and gave a burial ground to the village in her
memory in 1924.

Druce was privately educated “under my guardian. Since he had little foresight a public school
education was debarred” (Druce 1926) but at an early age ““Two ministers of the Independent
chapel at Potterspury, J. and T. B. Slye took an interest in the boy and his education”’ (Dictionary of
National Biography Suppl.). But despite his love of nature and his ability to recognise 400 species of
plants by the time he was 16 (though he did not know their names) his ‘“‘desire was to be a chemist

FiGure 2. Philadelphus Jeyes’ shop at 6, The Drapery, Northampton where Druce worked for 13 years (1866—
1879). Above: at the firm’s foundation in 1810. All Saint’s Church is to the left. Beneath its portico Druce, as a
boy, saw John Clare. Below: at the celebration of the firm’s 150th jubilee in 1960. View from beneath the
portico.

188 F. PERRING

... thanks to the generosity of my aunt, the rather heavy premium was paid and I became duly
apprenticed to a large wholesale and retail business in Northampton” (Druce 1926).

The business was Philadelphus Jeyes of 6, The Drapery in the centre of the town, close to the
market place and opposite All Saint’s Church which was largely built after the fire of 1675 (Fig. 2). It
was under the portico of the church incidentally that Druce (1930) recalls seeing, as a boy,
Northamptonshire’s treasured poet/naturalist John Clare. Clare spent the last 22 years of his life in
the Northampton Asylum but was occasionally allowed out and would sit in the portico watching
children at play or, as Druce recalls, as a little, pathetic, distraught figure, gazing at the sky.

Druce was obviously very good at his work. An assistant left after he had been there only a year
and he was promoted. At the end of his second year he became senior assistant and, when he was
still only 19, the manager left and he was given that post temporarily, but he held it for the next
eleven years.

He lived on the premises. At the time of the 1871 Census of Northampton 6, The Drapery was
recorded as a boarding house providing ‘digs’ for young pharmacists (Fitzhugh 1985; Fig. 3). They
were under the care of the housekeeper, Sarah Foll and Druce’s three companions were Frederick
Branson, Edmond Cooke and Anthony Chibnall. The first named came from Hanslope, Bucking-
hamshire, just across the River Ouse from Yardiey, and Druce was still in touch with him in 1920
when he visited him in Leeds (letter from Frederick Jeyes to ‘Betsy’ Jeyes 25 January 1920).

At this time the owner, Philadelphus Jeyes II, lived in a splendid Victorian folly he had built for
himself at Boughton 6 km to the north called Holly Lodge and Druce records (1905, 1930) how he
used to visit him rising at six o’clock, especially in the summer, and taking his Lindley’s
‘Introduction’ to read on the four miles’ [6 km] walk, have breakfast at 7.30 a.m., discuss the
business of the day and then either walk back or ride with one of the boys, who went to school in
Northampton, in time to open the shop at 8.30 a.m. By this time he had also studied Latin in the
evenings sufficiently well to pass his preliminary pharmaceutical examinations.

Mr Jeyes not only promoted Druce the chemist but promoted Druce the botanist. In 1868 he gave
£15 for one of his sons and Druce to take a 13 day walking tour through Wales where they climbed
Snowdon, Moel Eilio and Cader Idris. Though the main interest seems to have been castles, Druce
noted ferns and golden rod and they even brought back some change because he suggests (Druce
1926) hotel-keepers were over generous — the bill for two at Harlech (dinner, bed and breakfast) was
four shillings.

The next year, 1869, Mr Jeyes gave the two young men £20 and they went third class on wooden

Ficure 3 Interior of Jeyes’ shop, 6, The Drapery much as it was in Druce’s day.

PRESIDENTIAL ADDRESS, 1994 189

seats to Edinburgh where they climbed Arthur’s Seat: they then went to Blair Atholl and climbed
Ben-y-Gloe. The next stop was Kingussie from which they went by 6.00 a.m. train to Aviemore,
walked to the top of Cairngorm and back, reaching Inverness at 10.00 p.m. ‘more dead than alive,
for we had little to eat all day’’. They also climbed Ben Nevis and passed through Glasgow on their
way back to Edinburgh. The overnight train took them to Blisworth where, with sixpence left, they
bought some buns and walked the 6 km into Northampton (Druce 1926).

Strangely, although Druce refers to these important formative journeys on several occasions he
never identifies the son by name. Now, however, thanks to the help of a direct desendant of
Philadelphus Jeyes still living at Holly Lodge, Mr Anthony Jeyes, we have evidence that this was
almost certainly Theophilus Frederick Jeyes who was two years younger than Druce. Druce (1905)
says he was about his age. He was a problem child, violent at times, who terrorised the village (A.
Jeyes, pers. comm.) so perhaps Philadelphus’ generosity in sending the two boys off to Wales and
Scotland was not entirely altruistic.

Letters in Anthony Jeyes’ possession between Frederick and his elder sister Betsy show that, after
the First World War, Frederick was living in Oxford [at 97 St Aldates] and frequently met Druce
taking tea with him [at Yardley Lodge?] — so they were still on friendly terms after 50 years.

Philadelphus Jeyes not only stimulated Druce’s interest in plants by giving him the means to
explore Britain but, when his apprenticeship was complete in 1870, he gave him a £20 microscope
“selected by Henry Deane of Clapham, which is still before me” (Druce 1926).

Before following Druce during the third and last decade of his life in Northamptonshire, which
was to be his most significant scientifically, a small digression about Philadelphus Jeyes and the
family which influenced Druce so extensively seems timely: and for what follows I am much
indebted to Anthony and his nephew David Jeyes and their families.

Philadelphus Jeyes II was the son of Philadelphus I who, with a John Perrin, founded the business
in The Drapery in 1810. Philadelphus II was born in 1814, one of two surviving brothers; the other,
John, was two years younger. During their early years both boys studied botany encouraged by a Mr
Dickens, a local nurseryman and, in the late 1830s John entered into partnership with James Atkins
— a nurseryman regarded as one of the most prominent in the provinces.

Philadelphus’ father died when he was 14 and he eventually decided to follow him into the
business. Having qualified and become one of the earliest members of the newly formed
Pharmaceutical Society in 1842 he seems to have bought out John Perrin and taken over the business
in 1846. He was a successful business man and became a prominent radical non-conformist politician
and was Mayor of Northampton in 1852 when he was only 38. His knowledge of local politics was
undoubtedly invaluable to Druce in the 1870s when he was thinking of forming a local natural
history society. Druce (1918) says himself that at this time Northampton was divided into many sects
comparable with the situation in other Midlands towns described by George Eliot in Scenes of
clerical life.

John Jeyes was not a great businessman but he was a competent and inventive chemist. He left
Northampton in 1859 for London where he registered a total of 21 patents covering a variety of
subjects one of which, in December 1877, was his most famous and effective ‘Jeyes Fluid’. Though
Philadelphus may be translated as brotherly love and it is clear that they were the best of friends,
after their deaths, at the beginning of the century, the John Jeyes branch of the family considered
that a new product of the Philadelphus Jeyes branch called Carbocide was being sold to the
detriment of their Jeyes Fluid and a 13 year legal battle ensued.

The business at The Drapery continued under the name Philadelphus Jeyes until 1969 when it was
bought, gutted and run by two national multiples in succession and finally closed as a chemist’s shop
in the mid 1980s: today it is a sportswear shop though the building itself, parts of which predate the
great fire, is now protected by a conservation order.

Happily, however, the name of Jeyes was not lost to pharmacy in Northamptonshire for ever. In
1981 David Jeyes, great grandson of Philadelphus II, opened a chemist’s shop in Earls Barton 13 km
east of Northampton taking with him relics of the old shop in The Drapery.

Though as a boy Druce had been equally attracted to botany and entomology his choice of
profession now tilted the balance in favour of the former. In 1871 he helped found the
Pharmaceutical Association, “the members of which had many botanical walks. Indeed, its Field
Botany was its most successful branch” (Druce 1918).

Botany was one of the subjects for his final pharmaceutical examinations in 1872 and he clearly

190 F. PERRING

excelled in this and all the other subjects with the result that he won a special prize which included
Sowerby’s Plants of Britain, Carpenter’s Microscope, and Francis’ Ferns (Druce 1926).

So at the age of 22 he was fully qualified, acting manager of the leading chemist’s shop in town,
possessor of a microscope and some essential botanical books — what was this energetic, resourceful
young man to do next? The answer came “One night in the autumn of 1872, as in a feverish cold I lay
awake waiting for sleep that would not come, I determined that in the following year I would begin a
Herbarium, and commence a Flora of my native county. . . . These feverish thoughts, instead of
melting like a morning mist into nothing, crystallised into action, and in 1873 I collected over 700
species . . .” (Druce 1930). In the next six years he visited almost all the numerous villages in the
county and he records (Druce 1918) that he and fellow members of the Northampton Pharmaceuti-
cal Associatiun had the use of a room in Jeyes’ shop rent free: in 1879 it was a well-fitted laboratory
with a good collection of Materia Medica and specimens.

Sometime in 1875 he dined with Charles Jecks, a “‘local gentleman of independent means, and a
strong Darwin supporter” (Druce 1918), and discussed the formation of a natural history society.
The problems of dealing with the many factions in the community at that time have been referred to
earlier. However, with advice from influential people like Jecks and Jeyes and with the ability to
involve people of distinction and with titles which was to be his forte throughout his life, he went
ahead. By 7 March 1876 the rules of the Northamptonshire Natural History Society had been
formulated and Lord Lilford had agreed to become its first President. (Lilford was the county’s
leading ornithologist, publishing The birds of Northamptonshire in 1895: Lilford Hall, sadly now
empty, stands beside the River Nene 5 km south of Oundle.)

Miles Berkeley became an honorary member and President of the botanical section with Druce as
Secretary. Berkeley was born at Biggin 3 km west of Oundle (Druce 1930) and became a priest
serving as Curate of Apethorpe and Woodnewton nearby. But he also became an international
authority on fungi. He was in touch with Henslow and Darwin at Cambridge in the late 1820s and,
when Henslow retired as Professor of Botany in 1860, Berkeley was encouraged to apply for the
Chair but did not do so. Seven years later he did however apply for the Chair at Oxford but was
turned down because he was in Holy Orders (Ramsbottom 1948). In 1868 at the age of 65 he was
transferred to the more valuable living of Sibbertoft near Market Harborough. Druce first met him
at the inaugural meeting of the Northamptonshire Natural History Society on 21 April 1876 and he
later (1930) described him as “my botanical godfather, for he proposed me for the Linnean
Society’’. Druce was elected on 17 April 1879 when he was only 28 and those who recommended him
in addition to Berkeley were Lord Lilford, J. G. Baker, Henry Trimen and E. M. Holmes — a
distinguished group.

The Society had 60 members at that inaugural meeting, when. Lord Lilford spoke against
gamekeeping and in favour of birds-of-prey, and it clearly flourished (Druce 1918). In February
1877 Druce read a paper, ‘Contributions towards a Northamptonshire Flora” which was printed in
extenso in the Northampton Mercury. Druce took over as Hon. Secretary in 1878 on the resignation
of Charles Jecks who left the town. The following year at a meeting in Leicester on 20 May 1879 he
was appointed as one of the two Hon. Secretaries of the Midland Union of Natural History
Societies. On 16 June the Northamptonshire Natural History Society decided to start its own journal
and Druce, with Sanders and Scriven, became the editorial committee. Yet only two weeks later
Druce had left Jeyes and bought a chemist’s shop at 118 High Street, Oxford (Druce 1930).

The reasons for this precipitate departure when he was becoming ever more deeply involved in
local affairs in Northamptonshire and in the Midlands are recorded by Druce somewhat
ambiguously. In one place he writes “It was my wish to also save enough money from my salary to
enable me to purchase a business out of my own earnings, and not to touch my patrimony. Having
once visited Oxford and the Thames valley in search of Orchis militaris, the county so attracted me,
that in 1879 a pharmacy in the celebrated High Street of Oxford was purchased for £400” (Druce
1926). But elsewhere (Druce 1930) he confesses “It had been a great desire of mine to acquire the
business of my employer, which he had led me to believe would take place, but private matters made
him loathe to sell it, so on June 30, 1879, I left a town in which I knew “every one’’, and a county
which I dearly loved. . . .”. “The undulating, well-wooded country from which I drew my birth and
_ infant nurture and in which I spent my early years . . . will be an abiding memory” (Druce 1918).

The truth probably lies between the two. Philadelphus had eleven children all but one of whom
showed no interest in following their father into the business. However the one, Arthur Albert

|

PRESIDENTIAL ADDRESS, 1994 19]

Jeyes, younger brother of Frederick, studied at Guy’s Hospital and subsequently qualified as a
chemist. He was old enough in 1879 to join his father and, towards the end of the century, he
succeeded him as head of the firm (Fitzhugh 1985). So Druce was passed over in favour of Jeyes’ son
and, as a document in the possession of Anthony Jeyes suggests, he was probably forced to leave the
Northampton area by the terms of his employment. In an agreement with another employee dated
10 March 1876 it states that if he leaves Jeyes’ employment he may not for three years enter into
business of chemist and druggist in the town of Northampton or within 30 miles [50 km] thereof with
a fine of £100 if he did so. This explains why Druce, once he knew there was no future for him in
Northampton, looked as far afield as Reading and Oxford (Druce 1930).

Moreover there are no indications that he left with anger and cut his connections with the county
or his employers. He began to publish the Flora which he started in 1873 in the first issue of the
Journal of the Northamptonshire Natural History Society in 1880, completing it in eight parts in
1893-1894: and he contributed other articles to it until the mid-1920s. He continued to botanise in
the county: many parts were especially worked “‘some owing to the generous kindness of the Hon.
N. C. Rothschild, who was my kind host on several occasions” (Druce 1930). This friendship was
evident in 1912 when Charles Rothschild formed the S.P.N.R.: Druce was not only a member of the
Council but one of a select group on the Executive (see Rothschild 1987). And, when Rothschild
died in 1923, Druce wrote an extensive appreciation (Druce 1924). However it is also clear that he
concentrated his field work for almost all of the next 50 years in preparing Floras of other areas,
notably the three B.B.O.N.T. counties, Berks, Bucks and Oxon, and that he did not return to
serious work on Northamptonshire until he had finished the last of these, Buckinghamshire, in 1926.
Over the following four years he rewrote the ‘Flora in Parts’ and it was finally published, as a rather
colourless paperback, in 1930, in his eightieth year. Apart from the Introduction, which is
marvellous and a mine of historical information (much of it used here), it is the least satisfactory of
his great quartet of Floras and I am sure we are all looking forward eagerly to the publication of the
new Flora, in about a year’s time.

That Druce kept in touch with his home area is evident from the plaque in the church at Yardley
Gobion in memory of his mother, already referred to. The connection with the Jeyes family
continued, as we know, with periodic meetings with Frederick in Oxford. But it went further than
that and I was delighted to be shown by Mr Anthony Jeyes, grandson of Arthur Jeyes who had
stood between Druce and his ambitions for the business in Northampton, his ‘Baptismal Book’
recording his christening in 1930 where Dr Druce appears as one of his godfathers and in Druce’s
will, in the library of the Department of Plant Sciences in Oxford, appears “‘to Master Jeyes 20
guineas”’!

In preparing this part of my address I am sincerely grateful to those who have been so generous in
lending me their time and knowledge: I am especially indebted to Sylvia Chandler for notes about
Druce’s early years; to Andrew Robinson for local knowledge about Druce around Yardley
Gobion; to Ioan Thomas for access to his researches, particularly on Miles Berkeley; to Gina
Douglas in the Linnean Society library for helping locate Druceana; and finally to Anthony and
David Jeyes and their families for access to their family history and other papers and for their
hospitality in Boughton and Earls Barton.

Though Northamptonshire would undoubtedly have had a better Flora if Druce had not moved
to Oxford the botanical world and the B.S.B.I. in particular has every reason to be grateful. His
impact was immediate and decisive. Within a year, despite the demands of building up his new
business, he had helped start a natural history society which held its meetings in the Lecture Room
in the Botanic Garden (Druce 1930): already a demonstration of the energy and drive which was to
take him to the pinnacle of success, not only in Botany, but in the Pharmaceutical Society and in
local politics.

Druce was so successful in business that he was able to retire in 1905, 18 months after he had been
made Secretary of the Botanical Exchange Club, and devoted the remaining 27 years of his life to
changing it from an Exchange Club with a membership of just 42 into a national botanical field
society with a sixfold increase in numbers. He personally approached every amateur and
professional botanist. Moreover many of the new members were men and women of substance
(Allen 1986). There is little doubt that the Club was known to all those who would benefit from
membership. I fear that the same cannot be said of the B.S.B.I. today.

Clive Stace’s New Flora of the British Isles has sold over 6000 copies since it was published three

OZ F. PERRING

years ago: the Adlas of the British Flora and our Sedges Handbook have sold similar numbers, yet
the membership of the Society stands today at only 2683 and, if family members are removed and we
assume that every member has a ‘Stace’, only one in three of those who bought it (and how many
more are using it in libraries?) are members of the Society.

Mary Briggs has kindly provided some figures on the changing membership of the Society since
1955, appropriately the year after the launch of the Distribution Maps Scheme from which the Atlas
flowed. What the figures show, alarmingly (Fig. 4), is that whereas for the 21 years from 1955-1976
the annual increase in membership was 60/annum and our junior membership rose to at least 116
(7%), between 1976-1988 recruitment had dropped to 33/annum and the juniors to about 80
(3-5%), and that today our membership growth is negligible (1988-1994 mean of c.13/annum) whilst
the juniors with 43 are under 2%. These figures are reinforced by the results of the Membership
Survey we carried out in the autumn of 1993 in which we asked respondents to indicate their age
class. For the 728 who answered this section the figures were:

Under 18 1 0%
18-24 6 1%
25-34 66 9%
35—44 144 20%
45-54 167 23%
55-64 165 23%
65+ iW 24%
Don’t know! 4

Even allowing for some bias because older, retired members may have more time to fill in a
questionnaire (though I do not find that to be true for me or most of my friends!) there can be little
doubt that, taking the membership figures and the survey results together, we are a static, ageing
Society which is failing to attract the young. Yet this has been happening at a time when there has

3000 120
=)
&
2500

OQ.
s 90,
ie)
Z o
iain 2000 &
= 5
sac S00 eed ite
eal ce
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= Total B.S.B.I. 30 S
i ul junior membership es
Total B.S.B.1.
membership

0 0

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Va) eo) ~ oO N 2

O ox on an oY =

DZ = a, peng — N

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Ficure 4. Membership levels of the Botanical Society of the British Isles over the last 40 years.

PRESIDENTIAL ADDRESS, 1994 NO,

been an enormous increase in the number of posts for field naturalists in the Nature Conservancy
Council and its successor bodies, in wildlife trusts, national parks, environmental agencies and the
like, and whilst the British Trust for Ornithology has risen from 2000 in 1951 to over 10,000 today,
with more than 2000 of this increase in the last three years.

Yet you and I know that the Society is very much alive: we are involved in a range of activities of
vital importance to the botanical community of the British Isles — and further afield — with
publications like the Handbook series, the List of vascular plants and the Scarce plants in Britain. I
am sure this latter will become the most widely quoted B.S.B.I. publication since the Aflas of 1962,
and we have the new Atlas project yet to come. We also have the enormously impressive database at
Leicester which will shortly produce the long-awaited successor to Druce’s Comital Flora of 1932
(typically published in the week that he died) — and much, much more. Then there is the amazing
series of local Floras produced by members of the Society (ten in the last twelve months)
increasingly backed by computerised databases of immense and growing importance to national and
local conservation agencies which, together with up-to-date field work, are the basis for national and
county Red Data Books and lists.

The B.S.B.I. is indispensable and the need for our expertise grows rapidly. In the last few months
there have been significant developments which suggest that in future the Country Conservation
Agencies — English Nature, Scottish Natural Heritage and the Countryside Council for Wales — will
be looking increasingly to voluntary organisations like the B.S.B.I. for the information they need
about the changing flora of Britain and will be prepared to pay for it. It is in the light of this, and
three or four individual offers of contracts, that your Council has taken the decision to appoint a
coordinator. We must surely now appreciate, and make others appreciate, that we are the vascular
plant equivalent of the B.T.O. and, though I am not suggesting we need to aim for the staff of 60 and
the impressive H.Q. they inhabit at Thetford, we should at least acknowledge that in determining
what is happening to the fauna and flora of the British Isles, birds and vascular plants are the two
groups which provide the greatest opportunities for detailed study — and that the number of plant
species is an order of magnitude greater than the number of birds. We must devise a policy for
monitoring change, engage the minimum staff needed to carry it through, and aim for that
objective.

I suggest that policy shall be:

The study of the vascular plant populations of the British Isles in respect of distribution and numbers
and of the ecological factors, including those of human origin, affecting them. It shall be one of the
principal objects of this policy to answer the following questions:— In what way is contemporary man
affecting wild plants and, in particular, how are changes in forestry, agricultural and horticultural
practice influencing plant populations?

I hope this does not seem too avant-garde: it should not as it is a paraphrase of the B.T.O.’s policy
published in its Annual Report for 1962 (Hickling 1983), the year we published the Atlas and were
leaders in this field.

The functions of the coordinator were admirably summarised by the indefatigable David
Pearman in a statement sent to members in April, 1994. A primary function will be to ensure that all
the organisations which could benefit from the B.S.B.I.’s expertise, at national and local level, are
aware of the Society and act as an interface between them and the membership. But he or she cannot
be effective working alone. I am convinced that, if the Society is to achieve its full potential and
attract to its membership all those who would benefit from its resources in publications, meetings,
identification services, etc., the coordinator must be supported at the county level by voluntary
members prepared to give time to promoting the Society. For them I have coined the term ‘Link
People’. They would be additional to that remarkable, but overworked, band of county recorders in
most cases, but would work in close collaboration with them whilst having a special responsibility to
bring the Society constantly to the attention of the changing botanical staffs of universities, colleges,
wildlife trusts, local authority ecologists and so on.

One bonus of our age structure is that we must have nearly 1000 members who have retired or are
about to retire who might have time to act as ‘Link People’. Once these two elements, Coordinator
and ‘Link People’, are in place and the New Atlas project gets under way, I predict that we can, and
should, achieve a membership of 5000 by the end of the century. I strongly urge the Society to move
in that direction. Perhaps in honour of Druce and of the place from which he laid the foundation of
the Society at the beginning of the century, it might come to be known as the Oxford Movement.

194 F. PERRING
REFERENCES

ALLEN, D. E. (1986). The Botanists. St Paul’s Bibliographies, Winchester.

Druce, G. C. (1905). My apprenticeship. Typed copy of extract from Chemist & Druggist deposited in The
Library, Department of Plant Sciences, University of Oxford.

Druce, G. C. (1918). Formation of the Northamptonshire Natural History Society. Journal of the Northampton-
shire Natural History Society 19: 131-142.

Druce, G. C. (1924). Obituary. The Hon. N. Charles Rothschild. Journal of the Northamptonshire Natural
History Society 22: 135-141.

Druce, G. C. (1926). The Flora of Buckinghamshire. T. Buncle & Co., Arbroath.

Druce, G. C. (1930). The Flora of Northamptonshire. T. Buncle & Co., Arbroath.

FitzHuGH, J. (1985). Philadephus Jeyes — the firm and the family. MS in keeping of Anthony Jeyes, Boughton,
Northampton.

HICKLING, R. (1983). Enjoying ornithology. T. & A. D. Poyser, Waterhouses, Staffordshire.

RamsporTtom, J. (1948). In Phytopathological Classics 8: 5-12. American Phytopathological Society.

ROTHSCHILD, M. (1987). Changing conditions and conservation at Ashton Wold, the birthplace of the S.P.N.R.
Biological journal of the Linnean Society 32: 161-170.

20TH CENTURY DICTIONARY OF NATIONAL BIOGRAPHY SUPPLEMENT. Compact ed., p. 2607.

Watsonia 20: 195-204 (1995) 195

The Cambridge British Flora (1914-1920)

M. J. BUNTING*, D. BRIGGS AND M. BLOCK

Department of Plant Sciences, University of Cambridge, Botany School, Downing Street,
Cambridge, CB2 3EA

ABSTRACT

The reasons for the failure of The Cambridge British Flora after the publication in 1914 and 1920 of the first two
volumes have been investigated. The financial constraints imposed on Cambridge University Press by the First
World War were important, but the personal, social and financial problems of the author, Charles Edward Moss,
were crucial, since they led to his emigration to South Africa in 1917 and his subsequent loss of interest in a
project that belonged to his life in England.

Keyworbs: biography, bibliography, history, Charles Edward Moss, Edward Walter Hunnybun.

INTRODUCTION

At the British Association for the Advancement of Science meeting in Sheffield in 1910, Professor
J. W. H. Trail, Regius Professor of Botany at the University of Aberdeen, gave the Presidential
Address to the Botanical Section. In surveying the Floras of Britain then in use, he argued that there
was a pressing need for a new definitive work (Trail 1910). In his view, both teaching and research
would be stimulated by the production of an up-to-date, encyclopaedic work in which the
nomenclature was clarified. He suggested that such a Flora would probably be in the form of
monographs by specialists presented in a uniform style.

The need for a new Flora was already widely felt. In 1898, Rev. E. F. Linton’s intention to
produce just such a work was announced (Anonymous 1898), although nothing came of that
particular proposal. The Cambridge British Flora, under the editorship of C. E. Moss, was planned
as a ten-volume encyclopaedic survey of the flora of Britain, with accounts of critical genera
contributed by specialists. The two published volumes are of a very high standard, and it is therefore
of considerable interest to discover why the project was abandoned.

In considering the fate of The Cambridge British Flora, we have consulted a number of primary
sources, including documents in the archives of the University Library, Cambridge, the Department
of Botany, Natural History Museum, London, and the departments of plant sciences in the
universities of Cambridge, Oxford and Witwatersrand (South Africa). The relevant minute-books
of the meetings of both the Syndics and the Business Sub-Syndicate of Cambridge University Press
(hereafter ‘the Press’) were studied. These primary sources were used, together with contemporary
journal articles, to investigate the preparation and publication of each volume of the Flora, the
reaction of the botanical community, and the eventual abandonment of the project.

CHARLES EDWARD MOSS (1870-1930)

Moss, the youngest son of a nonconformist minister, was born in Hyde, Cheshire on 7 February
1870 (Crump 1931; Desmond 1994). His interest in botany was stimulated in 1893 when,
convalescing from a pulmonary abscess, he was ordered to spend a large amount of time out-of-
doors. He occupied himself with long walks over the Halifax moors, alone or in the company of
members of the Botanical Section of the Halifax Scientific Society, which he had joined in 1892, and
within which he soon became a prominent figure.

*Author for correspondence; present address Department of Geography, University of Waterloo, Waterloo,
Ontario, N2L 3G1, Canada.

196 M. J. BUNTING, D. BRIGGS AND M. BLOCK

In 1895, he entered Yorkshire College, Leeds (then part of the Victoria University) as a Queen’s
Scholar, and during this period was an editor of the Halifax naturalist, in which he published several
botanical papers. In 1898 he began work at Fairweather Green School, but kept up his association
with the college where he worked with W. G. Smith, mapping the vegetation associations of the
West Riding using techniques pioneered by Smith’s brother Robert in the Pentland Hills.

He became an assistant master at Sexey’s School, Bruton, Somerset in 1901, and began to study
the vegetation distribution of the area. At the end of 1902, he was appointed lecturer in biology at
Manchester Municipal Training College. This post was less well paid, but he could continue to work
for a higher degree, despite teaching every week day and several evenings.

The Central Committee for the Survey and Study of British Vegetation was set up by Tansley and
Smith in 1904 (Sheail 1987), and Moss became a prominent member. His contribution to the
committee and Types of British vegetation (Tansley 1911) was invaluable (Tansley 1931).

In 1907, Moss was awarded a doctorate by the University of Manchester for his work in Somerset
(Moss 1907), and the Back Bequest by the Royal Geographical Society for his work on the
vegetation of the Pennines (Moss 1904, 1913) which he had largely carried out at weekends and
during the vacations. At the end of the year he was offered the post of Curator of the Herbarium at
the University of Cambridge, which he took up in January 1908.

It is only at this point that his research interests turned to taxonomy. Whilst in Cambridge Moss
lectured and led field expeditions, and had a marked effect on many of his students (e.g. Levyns
1977). His lectures were described as “not brilliant ... but full of sense and philosophy”
(Ramsbottom 1931). Soon after his arrival he proposed that he should write a new ‘student’s Flora’
of the British Isles. ;

EDWARD WALTER HUNNYBUN (1848-1918)

Edward Walter Hunnybun, a Huntingdon solicitor, encouraged by the praise of his many botanical
correspondents, conceived the idea of turning his hobby of making lifelike pen-and-ink sketches of
plants into a ‘lifework’. He decided to attempt to draw all the species in the British flora, and set
about this task with single-minded determination and attention to detail. As his correspondence
with numerous collectors and field botanists testifies, he never became skilled in taxonomy, but his
network of supporters and advisors on different genera (e.g. Ley on Rosa) helped to ensure that he
drew representative, correctly identified specimens of each species and subspecies. It was important
that each specimen be representative, since he did not make-an idealised drawing, but drew the
individual specimen before him; he was fond of saying “‘I only draw what I see” (Wilmott 1920). His
great enthusiasm for this task is shown in his daily dawn bicycle rides during the collecting season in
search of specimens. Over the years, boxes of his exquisite drawings, drawn life-size in a brownish
ink over a preliminary pencil sketch, travelled around the country, to be admired and commented
on by his many correspondents.

As early as February 1901, F. J. Hanbury wrote to Hunnybun about “‘the possibility of publishing
the drawings as a separate volume, reproduced by photographic means’’.' On 4 June 1903, this
possibility became more likely, when a selection of 100 of his drawings was displayed at the Linnean
Society (Anonymous 1903). If sufficient support was forthcoming, the intention was to issue
fascicles of the drawings (Anonymous 1904). Although the pictures were much admired for their
artistic merit, G. C. Druce questioned the advisability of publishing them for scientific purposes,
since he considered the drawings to be scientifically inadequate, being “‘defective in detail” (Druce
1931).

Hunnybun considered bequeathing them to a private museum, but it is clear that he really desired
publication of at least a selection (e.g. Anonymous 1904), and in February 1908 J. Groves wrote that
‘“‘my brother and I have talked over the idea you mentioned of offering your splendid. . . drawings
.. . to the Hon. Walter Rothschild for his museum if he would agree to publish them’’ (our italics).

Sometime during the period 1908-1909 the drawings were donated to the Cambridge University
Botany School (now the Department of Plant Sciences). The actual date is unclear, but by 1909 they
were certainly in Moss’ hands, for during that year he discussed with J. Ramsbottom (a botanical
friend from his Halifax days) whether they-should produce a Flora based on the drawings

2 ON ee we

THE CAMBRIDGE BRITISH FLORA 197

(Ramsbottom 1931). Nothing seems to have come of this particular discussion, but Moss, unlike
Druce, obviously considered Hunnybun’s work worthy of publication.

PLANNING THE CAMBRIDGE BRITISH FLORA

When Hunnybun’s drawings arrived in the Botany School, Moss conceived an altogether bolder
plan than a ‘student’s Flora’, deciding to write a complete and definitive account of the British flora
classified according to Engler (1898). Hunnybun’s drawings would be used to illustrate this work. As
noted above, Moss initially considered preparing the work jointly with Ramsbottom (1931) or with
G. C. Druce, his ‘opposite number’ at Oxford University’s Fielding Herbarium (Druce 1931).

On 20 January 1911, Moss’ proposal for a Flora, to be entitled The Cambridge British Flora, was
favourably considered by the Syndics of the Press.’ He had at this point apparently changed his
plans, making it more of a solo project*, with expert contributors writing the accounts of critical
genera. Moss’ decision can partly be explained by his deep belief in and commitment to the project,
and his desire to see it completed to his own high standards. His conviction that his own views were
correct, and that those who disagreed with him were, at best, ill-informed, a characteristic even his
close friends attributed to him (e.g. Ramsbottom 1931), would have made any joint undertaking
fraught with difficulty, especially if it involved the forthright and idiosyncratic Druce. Sole control
and editorial power over the submitted manuscript would, therefore, have had great appeal to
Moss. Druce (1931) suggested that Moss made the move to sole authorship because he felt that such
a project might help him become a Fellow of one of the Cambridge colleges. Given his hardworking
and ambitious nature this is a possibility, but it should be noted that Druce made this statement after
they had quarrelled in 1915 (see below).

The contract with the Press was finally signed on 19 January 1912.4 An advance notice
(Anonymous 1911) makes much of Moss’ achievements and publications, and concludes “‘no better
choice could have been made’”’.

On 7 March 1912, a meeting of potential contributors was held in the British Museum (Natural
History). Here Moss’ habitual authoritarianism did not go down well. Moss wrote to Rendle?
(Keeper of Botany, British Museum (Natural History)), who had chaired the meeting, expressing
his thanks for the “‘perfect impartiality with which you conducted what at times threatened to
become a rather warm meeting!”’. Apparently this was a common feature of meetings involving
Moss, at which he often “‘caused an uncomfortable liveliness” (Ramsbottom 1931). During the
meeting (Anonymous 1912) there was much discussion about the citation of synonyms, etc., and
Moss “undertook to consider” the points raised. Druce “entered a formal protest” (and in doing so
appears to have spoken for the ‘establishment’) against ““Germanising our flora”’ by adopting the
Englerian system (Anonymous 1912). However, the meeting generated enthusiasm for, and interest
in, the project and showed Moss firmly in charge. E. S. Marshall® describes himself as “much
impressed by Moss’ business aptitude . . . his clearness of view and botanical ability”.

In October 1912, Moss compiled and sent out a list of 84 “Instructions to Contributors”, which
read more like an examination rubric than guidelines to acknowledged experts. Moss’ arrogance in
correcting the contributions that were eventually offered and even, on occasion, rejecting them as
inadequate, did nothing to win him friends. His editorial style was extremely high-handed. For
example, Druce’ suggested that Linton withdrew his account of Salix, intended for volume ii,
“choked off’ and found Moss “‘most difficult to work with’’. However, a letter from Moss to Linton’
was unambiguously a rejection, couched in schoolmasterly terms, describing the account as not
sufficiently up-to-date in nomenclature. Many of the older British botanists were unwilling to accept
such treatment and therefore “the editor was left with additional preparation and additional
criticism’ (Ramsbottom 1931). .

The Cambridge British Flora was never intended to be anything other than a specialist book. The
Press only intended to print 1250 copies before destroying the type.” This initial number was revised
downwards to 1000.!”

Moss had grandiose ideas about the format of the work, initially planning to include descriptions,
maps and numerous photographs, as well as the expensive and complex plates of Hunnybun’s
drawings. This led to conflict with the Press. In the spring of 1913, a debate about whether to bind
the plates with the text (as Moss desired) or in separate volumes (as the Press proposed) escalated to

198 M. J. BUNTING, D. BRIGGS AND M. BLOCK

the point where Moss threatened to take the book elsewhere. The lively correspondence between
A. C. Seward (Professor of Botany at Cambridge as well as a Syndic at the Press), attempting to
defend Moss: and the Press’ representative A. R. Waller, shows that it was indeed a major
problem.” Throughout this period Seward supported Moss, speaking for him to the Press and
excusing his abruptness and occasional outright rudeness. Moss eventually accepted the Press’s
preferred scheme, possibly under pressure from Seward.

THE ILLUSTRATIONS

In the summer of 1913, Hunnybun retired owing to increasing ill-health; he was a severe asthmatic
and until his death in 1918 moved frequently to different parts of the country trying to find a climate
that suited him. Now he could concentrate on completing and refining his drawings, replacing some
with others drawn from better or more representative specimens. This habit of redrawing was to
lead to problems with the Press.

The preliminary sketches, drawings and the copy of the London catalogue of British plants used
by Hunnybun as a checklist are full of notes stating ““Moss says” and ‘‘Moss thinks’’, generally
referring to the identification or the representativeness of a particular specimen, and showing
clearly that Moss was firmly in control of what was to be drawn as well as of what was to be written.
Hunnybun, a mild and easygoing man, knew himself to be far less expert than Moss in matters
taxonomic, and so proved to be more amenable than the contributing authors.

' THE ‘FIRST’ VOLUME AND ITS RECEPTION

In 1914, after several delays, volume ti was published. In a lengthy, controversial introduction Moss
expounded his ideas on nomenclature and classification, explaining how he wished to use the
Englerian system to bring British Floras more into line with the continental Floras, but without
using all the subspecific divisions favoured by many European taxonomists. Moss intended to use
only species, varieties and formae (e.g. in the Prospectus for the Flora). However, this approach
generated sufficient intraspecific subdivision to annoy many botanists (e.g. Anonymous 1914).
Throughout the introduction, Moss reveals a remarkable grasp of the literature, especially
considering that he had only turned to this field of botany six years earlier, and that a significant part
was written in German.

The completeness and detail of the text’s treatment of genera meant that no reviewer was inclined
to attempt a critical analysis of the entire volume. Instead most looked at the overall schemes of
nomenclature used (always a thorny subject). Moss’ use of lower case initial letters for the specific
epithet in all cases (e.g. Hieracium leyi, not Hieracium Leyi) was in direct contravention of a
recommendation of the International Botanical Congress of Vienna (held in 1905). Moss introduced
this in an attempt to simplify a confusing variety of conventions, but it was seen as unnecessary by
one reviewer (Anonymous 1914), who wrote of ‘““‘Dr Moss, who like the rest of us, likes to have his
own way and is perhaps more fortunate because more insistent in getting it’’.

Moss also insisted that a species subdivided into varieties should be fully so divided, rather than
selecting one ‘type variety’, which had only a binomial. Britten (1915), editor of Journal of botany,
considered that the logical consequence of Moss’ proposal was the “objectionable . . . American
innovation” of a trinomial system of nomenclature. He held that, for example, Populus tremula var.
glabra was essentially no different from Populus tremula glabra, and that Moss was breaking
“unwritten rules”. However, it is clear that Moss was strongly opposed to trinomials (e.g. Moss
1915), and he accused Britten of having sunk to ‘‘a Drucian level . . . of misrepresentation”’.'*

The reaction of reviewers to the text was, on the whole, positive. However, the drawings were
widely criticised. Something of the delicacy of Hunnybun’s drawings was lost in the reproduction.
Moreover, the extremely generous page size (36 X 26 cm) was not fully utilised by Hunnybun,
whose obsession with precise scale reproduction of the specimen in front of him seemed to blind him
to the space wasted in drawing small plants (e.g. Sagina boydii, Plate 27, volume 11, where less than
10% of the page was used). He also often drew twigs or sprays overlapping unnecessarily, obscuring
important features (e.g. Salix, volume ii). In’a letter to Waller at the Press’?, Hunnybun,

THE CAMBRIDGE BRITISH FLORA 199

encouraged by his many friends, hoped for a better reaction to his drawings for the next volume,
which he felt were ‘‘better and more interesting than those already published”’.

BETWEEN VOLUMES

In 1915, Moss and Druce had a serious disagreement occasioned by the internal politics of the
Botanical Exchange Club, of which Druce was then honorary secretary-treasurer. There was
widespread discontent among the members, and Moss became spokesman for the disaffected
faction. In the end he was forced to issue a formal apology to Druce, which no doubt both annoyed
and embarrassed Moss (Allen 1986).

By 1916 Moss was a bitterly frustrated man. The Press, financially stressed by the war, was
delaying the publication of further volumes of the Flora (originally supposed to be issued annually).
The minutes of a Business Sub-Syndicate meeting held on 12 January 1915 record that “‘it was
agreed to proceed slowly with the Cambridge British Flora’’.

Moss’ own finances were also strained. He had had problems with the Press over his share of the
cost of having new plates made for volume ii, to replace those he had decided were inadequate.’°
Delay in payment had culminated in a threat of legal action if he did not pay by the end of 1914.'° By
1916, all teaching staff in the Botany School had suffered a 10% pay cut in an attempt to keep the
department solvent in the face of drastically falling student numbers.'? Departmental salaries were
partly made up of payments for courses taught, at a rate of so much per lecture or demonstration,
and this element also decreased, since fewer students meant fewer classes. Moss’ pay fell by about
£40, a loss of 20% relative to his pre-war pay. Moss described himself as ‘‘almost on half pay” by
November 1915.'*

At this time Moss had a great desire to be involved in the War effort, but his hopes were frustrated
by his age (mid-40s). He managed to be assigned to training recruits in the O.T.C., and later
transferred to a munitions factory.

Moss seems to have found the writing of the Flora time-consuming, and manuscripts were
frequently late in arriving at the Press.'’ A new contract was drawn up in January 1915, under which
Moss was only to be paid the final part of the monies due to him when he delivered the relevant
manuscript.'® Prospects of promotion or of obtaining a college fellowship must have seemed remote
as problems with the Flora mounted and student numbers fell. )

In 1916 his personal problems reached their height when his marriage broke down irretrievably,
and by October 1916 divorce proceedings were under way. The severity of the scandal can scarcely
be appreciated from a late twentieth century viewpoint, but in the close, formal society of post-
Edwardian Cambridge divorce was extremely shocking. Moss appears to have been the innocent
party!’; Mrs Wedgwood (a collector and friend of Druce (Sandwith 1954)) wrote “it is difficult for us
to put ourselves into his [Moss’] position — Mrs Moss entirely deceived me, she had no marks of the
seductress about her’’.*”

Embittered, overworked and stressed, Moss decided to cut his losses. In November 1916 he
applied for a professorship at the School of Mining in Witwatersrand (later the University of
Witwatersrand).*! The School wanted him to take up the post as soon as possible, and on 3 February
1917, a week after he had given evidence in the divorce proceedings,” he and his school-age
daughter Beatrice set sail on the Balmoral Castle.”°

A letter from Mrs Wedgwood, who seems to have delighted in transmitting gossip, suggested that
Moss was considering abandoning the Flora; ‘‘he [Moss] would have ‘chucked’ the Flora had I not
been so shocked’”’.** He did not do so; instead he appointed a former student, A. J. Wilmott, then
working at the British Museum (Natural History), to see volume iii through the press.” He was
obviously still committed to the project; in a letter to Hunnybun” he proposed that if Wilmott did
well, then by volume iv or v he would be cited as co-author. In typical style Moss wrote that “‘it
would have been a mistake to take Wilmott as joint author to begin with. He is young, and he must
have something to grow up to”.*° Letters from the period provide evidence that Moss was as
authoritarian as ever, but also that Hunnybun and Wilmott dealt independently with some problems
that arose.

By April 1917 Moss appears to have been feeling more relaxed. He encouraged Wilmott to “use

your own discretion in amending or altering my manuscript” ,”° and was dismissive of Wilmott’s and

200 M. J. BUNTING, D. BRIGGS AND M. BLOCK

Hunnybun’s fears of libel over passages in the introduction to volume iii (see below), but agreed to
changes; ‘“‘with 7000 miles between us, act on your own judgement”’.”’ Moss appears to have been
dissociating himself from the project and relaxing his previous rigid control over it.

By 1918, however, the letters suggest that Moss was less happy with life in South Africa, and that
he wished to return to England before his three-year contract was completed, Ostensibly to Oversee
the Flora. A letter from Seward exists, expressing a disinclination to have him back.*® Although
Moss obviously still had allies who would support his return to Cambridge, Seward appears to have
had no desire to employ a man who had broken a contract.

VOLUME III

In 1920, after Wilmott had found that he had taken on a much larger task than he had envisaged and
after many problems with the manuscript, the Press published volume iii. Moss’ role was unclear. As
late as April 1919, Wilmott had to write a firm letter to Waller, saying ‘‘there will be no need to wait
for proof from Prof. Moss as he finished with this volume before he left the country. He left me to
finish seeing it through the press’’.*? Volume iii was issued at a markedly increased cost of £6 15s 0d
(volume ii had cost £2 5s Od), and the sum allocated to expenses for each volume had almost
quadrupled.

The introduction to volume 111 proved more controversial than that of volume ii, since it included
Moss’ final shot in the argument over nomenclature that waged between him and Britten (see
above). He wrote the introduction in January 1917, just before his departure for South Africa.
Hunnybun believed that the draft ‘“was written when [Moss] was rightly very irritated with Mr
Britten’s conduct and when he was utterly worried by the domestic troubles which have driven him
from the country. You [Wilmott] and I feel profound regret at all the trouble that has come upon
him and we realise that in his happier days he would have worded the paragraph very differently”’.°°
Hunnybun then proposed rewording the passage. The original draft introduction made by Moss is
extant, and is even more vituperative than the printed version. However, the section attacking
Britten was left exactly as Moss had worded it. Hunnybun wrote that ““Moss would never forgive me
if asked you to modify it” and adds wistfully “‘why botanical differences should occasion such bitter
strife is what I have never been able to comprehend”’.*” .

The introduction contained several direct attacks on individuals which were frowned upon by
reviewers such as Rendle (1920), who regretted the fact that “to perpetuate the differences of
opinion which have arisen in matters of very secondary importance detracts from the dignity which
such a work should possess” and suggests that the Syndics were remiss in not exerting “fatherly
censorship” over parts of the introduction.

As with volume ti, the text itself was favourably received, but the drawings were criticised, even
though Hunnybun had believed them to be far superior.

THE FATE OF THE PROJECT: 1920 ONWARDS

The Press delayed further volumes until financial restraints eased, and as early as 1918 Seward was
beginning to express doubts about the future of the series.*! In 1921 Moss married a co-worker at the
University of Witwatersrand and became head of his department, increasing his commitment to his
new life.

By 1923, no further volumes had been published. The Press decided to have the viability of the
project assessed by a committee of botanists chaired by Professor Seward, who was also a Syndic.”
In July of that year the Press decided that the work “should not proceed under the current
contract”’.°?

Moss initially expressed indignation, suggesting that the Syndics were under contractual
obligation to the subscribers to complete the series. This was investigated by the Press,* but
apparently no such obligation existed, since later meetings ignore the issue. The Press eventually
decided either to proceed under tighter rules (to prevent the delays in obtaining manuscripts from
Moss and the continual changing of plates) or to abandon the project. After consultations with their
solicitor** about Moss’ legal rights they decided to persuade Moss either to improve or take a lump

THE CAMBRIDGE BRITISH FLORA 201

sum payment in lieu of their contractual obligations.*° On 27 July 1923, “it was agreed to
recommend that Dr Moss be offered a payment in respect of work done on unpublished volumes
. . . the amount under this head not to exceed £75 in respect of any one volume; Dr Moss to be also
offered the sum of £100 (to be increased to £150, if necessary) as a solatium for the non-publication
of later volumes. It was further agreed to recommend that in the event of the offer not being
accepted . . . the Syndics proceed with the publication under the terms of the original agreement,
provided that complete copy for volume iv be delivered one year after the date of acceptance of the
offer, complete copy for subsequent volumes to be similarly delivered at annual intervals’ .*? Moss
seems to have accepted the financial offer with little protest, although he managed to extract the
maximum sum the Press were prepared to offer; “£150, plus a further sum of £80 in respect of out-
of-pocket expenses, in full discharge of any claim against the Syndics in respect of the agreement for
the publication of The Cambridge British Flora’’.*°

THE TEXT OF THE PUBLISHED VOLUMES

In the introduction to volume ii of The Cambridge British Flora, Moss listed three aims: to register
the present state of knowledge with respect to British plants, including classification, nomenclature,
characteristics and distribution; to attempt to relate British plants to allied forms in foreign
countries; and to stimulate further research, particularly in the areas of variation and distribution.
Hunnybun’’ wrote that Moss felt that ‘“‘we shall make some howlers. The Cambridge British Flora is
merely a step forward’.

Many botanists consider that The Cambridge British Flora, with its ambitious scope and attention
to detail, is one of the best taxonomic studies of the British flora. However, it has been suggested
that the volumes suffered from ‘‘elephantiasis of the format” (A. O. Chater, personal communi-
cation). Despite being incomplete, it is still invaluable for the genera treated, although modern
taxonomic ideas and conventions in nomenclature, especially the increased emphasis on the type
concept, together with Moss’ personalised methods of citing the author-attribution in the case of
hybrids, make it out-of-date for the purposes of nomenclature. Moss’ interest in, and observation
of, variation within species meant that his subdivisions often corresponded better to the situation
seen in the field than those of previous workers. For example, in Salix he subdivided S. caprea L.
into var. genuina Syme (lowland; now S. caprea var. caprea) and var. sphacelata (J. E. Sm.)
Wahlenberg (Scottish Highlands).

The analytical, formal style in which critical taxonomy was treated in The Cambridge British Flora
is even now unusual. Pioneering work on distribution mapping had been carried out on the
continent, and its use in a British Flora was also a significant step forward. Moss’ interests in
distribution mapping date back to his earliest work in the West Riding. In The Cambridge British
Flora, he simply recorded location, according to a scheme based on geographical counties rather
than the presently-preferred vice-comital system, without considering the ecology of the plant in
any detail.

The contributing authors, kept firmly in line by Moss, produced contributions of a consistently
high standard. A good example is H. W. Pugsley’s treatment of Fumaria. The account can scarcely
be bettered today.

Moss’ second aim of comparing British and continental botany was achieved in part by his use of
the Englerian classification system that was becoming commonplace in mainland Europe. Many
British botanists were parochial in outlook, but Moss was strongly influenced by several European-
minded workers such as H. Gilbert-Carter and his friend and colleague A. G. Tansley. Throughout
The Cambridge British Flora, frequent references show Moss’ familiarity with both British and
continental taxonomic writings.

THE ILLUSTRATIONS
Hunnybun’s drawings, later to be made into plates in The Cambridge British Flora, were well

received as works of art when they were exhibited at the Linnean Society in 1903 and Hunnybun’s
extensive correspondence shows that this opinion was widespread. However, Blunt (1950)

202 M. J. BUNTING, D. BRIGGS AND M. BLOCK

considers that the drawings “well demonstrate how lifeless a figure can become when sensitivity is
lacking”. Although faithful to the appearance of the living plant, they are inadequate for scientific
purposes, where more diagrammatic representation of key features is required. They often lack fine
detail and Hunnybun’s uniformly fine line and the absence of shading give no sense of texture.
Hunnybun’s emphasis on drawing living material often seems to have led to him drawing ‘weaker’
looking specimens.

As Hunnybun’s health deteriorated, he became increasingly reliant on plants sent to him by his
many correspondents, rather than collecting his own. The annual lists of desiderata which he sent to
these collectors include detailed instructions on the packing of specimens for posting, and suggest
that specimens sometimes arrived in a damaged or unusable condition. Perhaps this meant that
some of his drawings were based on less than perfect material.

Botanical illustration must always be a compromise. By accentuation of key characteristics,
illustrations should convey the essence of a species rather than an image of an individual. An
appreciation of this important principle is lacking in Hunnybun’s work. Indeed, he complained that
Pugsley wished him to produce ‘‘mental concepts of what the forms of each species should be’’** and
enlargements which were “‘absolutely diagrammatic’’.*” Hunnybun felt that his enlargements, being
based on living specimens, would be of more use to the student than a diagram. However, it appears
that even Moss thought little of Hunnybun’s enlargements, and instructed Wilmott, who was
selecting illustrations for volume iii, to “rule out any enlargements you choose; but do not, in
Heavens name, consult E.W.H. [Hunnybun] about this! It is like asking a parent which of his
children shall be cut in half!”*?

THE FAILURE OF THE CAMBRIDGE BRITISH FLORA

Tansley, in his foreword to the first edition of the Flora of the British Isles (Clapham, Tutin &
Warburg 1952), wrote that “a new British Flora has been a desideratum for the past half century and
urgently needed during the last thirty years . . . several attempts have been made to fill the gap but
none have been carried through to success’’. Although Tansley does not mention The Cambridge
British Flora by name, he is likely to be commenting on the work of his friend and former colleague
when he notes that earlier attempts “were all too ambitious, aiming at a completeness and
exhaustiveness unattainable except through years of laborious effort and the collaboration of a large
body of specialists” (our italics).

In our view Tansley was pointing to the major weakness in The Cambridge British Flora project.
To judge by the two volumes that were issued, there was no scientific reason why the project should
have failed, although Hunnybun’s drawings might have made the Flora less attractive to a potential
buyer. However, the long-term viability of the project was doubtful, as there is abundant evidence
that many of the taxonomic specialists of the day found it difficult or impossible to work with Moss,
who ‘“‘when satisfied that he had reached a sound conclusion was immovable” (Crump 1931). He was
also prone to dismiss other people’s views with “‘sweeping contempt” (Tansley 1931), although he
could be a firm friend to those who ‘‘accepted his frank expressions in the same northern spirit in
which they were given” (F.E.W. 1931). Obsessed by the Flora at its outset, Moss was unable to
sustain his interest in the project after his emigration to South Africa. Wilmott performed the task of
seeing volume iii through the press, but with his penchant for complex schemes and subtleties
(Stearn 1981) he was unsuited to the task of rallying support for continuing the project. Indeed in
the early 1920s he was involved in many projects of his own, such as editing the tenth edition of the
Manual of British botany (Babington 1922).

Thus the project died. A modern, encyclopaedic, critical Flora of Britain has yet to be written,
although an attempt was made between December 1973 and January 1985 (Stace 1991).

ACKNOWLEDGMENTS
We would like to thank C. E. Scarles of the Cambridge University Press and E. S. Leedham-Green

of the University Library, Cambridge, for access to archive material. Thanks are due to Dr D. E.
Allen for information from the Druce archives and other helpful comments, and to P. D. Sell, R. D.

THE CAMBRIDGE BRITISH FLORA 203

Meikle and Dr S. M. Walters for information on the scientific aspects of the Flora. Dr D. E. Allen,
Dr J. R. Akeroyd, D. H. Kent, Dr S. M. Walters and an anonymous reviewer commented on an
earlier version of this manuscript, for which we are grateful.

nA & WN

35
36
37
38
39

NOTES

F. J. Hanbury to E. W. Hunnybun, 12 September 1901. Hunnybun Letters (HL), Department of Plant
Sciences, Cambridge University (DPS UC).

J. Groves to E. W. Hunnybun, 7 February 1908. HL, DPS UC.

Syndicate Committee Meeting Minutes, 20 January 1911. Cambridge University Press (CUP).

Syndicate Committee Meeting Minutes, 19 January 1912. CUP.

C. E. Moss to A. B. Rendle, 11 March 1912. Rendle Correspondence in Carruthers Correspondence Box 4,
BM-G, British Museum (Natural History) (BM-NH).

E. S. Marshall to G. C. Druce, 22 February 1911. Druce papers (DP) Box 19, Department of Plant Sciences,
Oxford University (DPS OU) (D. E. Allen, personal communication).

G. C. Druce, draft of Druce 1931. DP Box 5, DPS OU.

C. E. Moss to E. F. Linton, 17 June 1912. Linton Autograph Collection, BM—B, BM—NH (D. E. Allen,
personal communication).

Syndicate Committee Meeting Minutes, 14 February 1913. CUP.

Business Sub-Syndicate Meeting Minutes, 11 February 1913. CUP.

A. C. Seward to A. R. Waller, various letters, March and April 1913. Letters from the CUP, Pr. A.,
Cambridge University Library (CUL).

E. W. Hunnybun to A. R. Waller, 14 January 1915. HL, DPS CU.

Professor A. C. Seward’s accounts book. DPS CU.

C. E. Moss to A. B. Rendle, 8 November 1915. Rendle Papers (RP), BM—R, BM-—NH.

e.g. Syndicate Committee Meeting Minutes, 21 November 1913. CUP.

Business Sub-Syndicate Meeting Minutes, 10 November 1914. CUP.

e.g. Business Sub-Syndicate Meeting Minutes, 4 November 1913. CUP.

Syndicate Committee Meeting Minutes, 15 January 1915. CUP.

Certificate of making Degree Nisi Absolute (Divorce) in the High Court of Justice (Family Division).

M. Wedgwood to E. W. Hunnybun, 20 January 1917. HL, DPS CU.

C. E. Moss to A. B. Rendle, 27 October 1916. RP, BM—R, BM-NH.

C. E. Moss to A. B. Rendle, 4 February 1917. RP, BM-R.

C. E. Moss to A. R. Waller, 4 February 1917. Letters from CUP, Pr. A., CUL.

M. Wedgwood to E. W. Hunnybun, 3 February 1917. HL, DPS CU.

Business Sub-Syndicate Meeting Minutes, 24 October 1916. CUP.

C. E. Moss to E. W. Hunnybun, 12 January 1917. HL, DPS CU.

Business Sub-Syndicate Meeting Minutes, 27 February 1917. CUP.

Syndicate Committee Meeting Minutes, 2 March 1917. CUP.

C. E. Moss to A. J. Wilmott, 18 April 1917. Wilmott Papers (WP), BM—W, BM-NH.

C. E. Moss to A. J. Wilmott, 20 April 1917. WP, BM—W, BM-NH.

A. C. Seward to A. R. Waller, 13 September 1917. Letters from the CUP, Pr. A., CUL.

A. J. Wilmott to A. R. Waller, 26 April 1919. Letters from the CUP, Pr. A., CUL.

E. W. Hunnybun to A. J. Wilmott, 11 March 1917. WP, BM—W, BM-NH.

A. C. Seward to A. B. Rendle, 17 July 1918. RP, BM—R, BM-NH.

Syndicate Committee Meeting Minutes, 2 April 1923. CUP.

Syndicate Committee Meeting Minutes, 1 July 1923. CUP.

Townsend to A. R. Waller, 1923, several letters. Letters from the CUP, Pr. A. T., CUL.

Business Sub-Syndicate Meeting Minutes, 27 July 1923. CUP.

Syndicate Committee Meeting Minutes, 21 September 1923. CUP.

E. W. Hunnybun to A. J. Wilmott, 4 March 1917. WP, BM—W, BM-NH.

E. W. Hunnybun to A. J. Wilmott, 28 November 1913. WP, BM—W, BM-NH.

C. E. Moss to A. J. Wilmott, 23 April 1917. WP, BM—W, BM-NH.

REFERENCES

ALLEN, D. E. (1986). The botanists: A history of the Botanical Society of the British Isles through 150 years. St

Paul’s Bibliographies, Winchester.

ANONYMOUS. (1898). A new British Flora. Journal of botany, London 36: 94-96.

204 M. J. BUNTING, D. BRIGGS AND M. BLOCK

Anonymous. (1903). Proceedings of the Linnean Society of London 115th session: 39.

Anonymous. (1904). Book-notes, news, etc. Journal of botany, London 42: 318-320.

Anonymous. (1911). A new British Flora. Journal of botany, London 49: 119-121.

ANonyMovws. (1912). Book-notes, news, etc. Journal of botany, London 50: 205-208.

ANONYMOUS. (1914). Reviews. Journal of botany, London §2: 131-136.

BaBINGTON, C. C. (1922). Manual of British botany. Gurney & Jackson, London.

Bunt, W. (1950). The art of botanical illustration. Collins, London.

BriTTEN, J. (1915). Reviews. Journal of botany, London 53: 252-254.

CLAPHAM, A. R., TuTin, T. G. & WarsurG, E. F. (1952). Flora of the British Isles. Cambridge University Press,
Cambridge.

Crump, W. B. (1931). In memoriam: Charles Edward Moss. The Naturalist: 55-59.

DESMOND, R. (1994). Dictionary of British and Irish botanists and horticulturists. Taylor & Francis Ltd, London.

Druce, G. C. (1931). Moss, Charles Edward (obituary). Botanical Society of the British Isles: Report: 326-327.

ENGLER, A. (1898). Syllabus der Pflanzenfamilien. Verlag von Gebriider Borntraeger, Berlin.

Levyns, M. R. (1977). Insnar’d with flow’rs. Botanical Society of South Africa, Kirstenbosch.

Moss, C. E. (1904). Peat moors of the Pennines: their age, origin and utilization. Geographical journal 23: 660—
671.

Moss, C. E. (1907). Geographical distribution of vegetation in Somerset: Bath and Bridgewater district. The
Royal Geographical Society, London.

Moss, C. E. (1913). Vegetation of the Peak District. Cambridge University Press, Cambridge.

Moss, C. E. (1914). The Cambridge British Flora. Volume 11. Cambridge University Press, Cambridge.

Moss, C. E. (1915). Trinomials. Journal of botany, London §3: 334.

Moss, C. E. (1920). The Cambridge British Flora. Volume ii. Cambridge University Press, Cambridge.

RamssortToM, J. (1931). Charles Edward Moss (obituary). Journal of botany, London 69: 20-23.

RENDLE, A. B. (1920). The Cambridge British Flora. Nature 106: 337-338.

SANDWITH, N. Y. (1954). Obituaries: M. L. Wedgwood (1854-1953). Proceedings of the Botanical Sees of the
British Isles 1954: 114-115.

SHEAIL, J. (1987). Seventy-five years in ecology. Blackwell, Oxford.

Stace, C. A. (1991). New Flora of the British Isles. Cambridge University Press, Cambridge.

STEARN, W. T. (1981). The Natural History Museum at South Kensington. Heinemann, London.

TANSLEY, A. G., ed. (1911). Types of British vegetation. Cambridge University Press, Cambridge.

TANSLEY, A. G. (1931). Charles Edward Moss. Journal of ecology 19: 210-214.

TraiL, W. H. (1910). The study of the British flora. Journal of botany, London 48: 241-250.

W[eiss], F. E. (1931). Charles Edward Moss. The North Western naturalist: 34-36.

WitmotTr, A. J. (1920). E. W. Hunnybun, in Moss, C. E. The Cambridge British Flora. Volume iii. Cambridge
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(Accepted May 1994)

Watsonia 20: 185-194 (1995) 205

Senecio vulgaris L. subsp. denticulatus (O. F. Muell.) P. D.
Sell and S. vulgaris subsp. vulgaris var. vulgaris on Jersey
(Channel Islands)

H. P. COMES

Institut f. Spezielle Botanik u. Botanischer Garten, Anselm Franz von Bentzel-Weg 9, Johannes
Gutenberg-Universitat Mainz, 55099 Mainz, Germany*

ABSTRACT

Evidence obtained from comparative cultivation confirms that Senecio vulgaris subsp. denticulatus (Asteraceae)
and its presumed derivative var. vulgaris differ consistently in a number of morphological and life history
characters. Field studies on Jersey revealed that both taxa are also ecologically distinct. Subsp. denticulatus is
only known from coastal areas of dune grassland where it occurs on soils which have large amounts of carbonate
and low amounts of mineral nutrients. Its associates include other winter-annuals and species of Mediterranean
affinity. Results obtained from soil analyses do not provide any meaningful clue to the unusual and disjunct
distribution of subsp. denticulatus and several community associates within one of its natural maritime habitats
(Les Quennevais). Some observations on the pollination strategy, seed production and predation damage of
subsp. denticulatus are reported. In contrast to subsp. denticulatus, var. vulgaris occurs in man-disturbed inland /—
and coastal localities throughout the island. Both taxa seem to interact along an extensive coastal hybrid zone,
particularly in the disturbed area north of Les Quennevais. Progeny-analyses of both short ligulate and discoid
phenotypes from Les Quennevais showed them to produce patterns of germination, phenological and leaf shape
characteristics typical of subsp. denticulatus. It is concluded that the low nutrient-supplying power of the
Quennevais area is a factor of importance in limiting the distribution of the var. vulgaris genotypes to areas
outside Les Quennevais. Information on the former distribution of subsp. denticulatus on Jersey suggests that its
habitat is seriously endangered from progressive destruction by man.

KEYworpDs: Senecio vulgaris subsp. vulgaris var. hibernicus, ecological variation, hybrid zone, adaptation.

INTRODUCTION

Senecio vulgaris L. (2n = 40) is subdivided into two subspecies, namely the type subspecies with two
varieties and subsp. denticulatus (O. F. Muell.) P. D. Sell+. Subspecies vulgaris var. hibernicus
Syme, which is distributed sympatrically with var. vulgaris and in its distribution largely limited to
the British Isles and Ireland, appears very likely to be a recently evolved stabilized introgressant
between var. vulgaris and S. squalidus (2n = 20) (e.g. Abbott 1992). This taxon will not be
considered here further. Thus all comparisons reported in this paper are between Senecio vulgaris
subsp. denticulatus and Senecio vulgaris subsp. vulgaris var. vulgaris.

Information available from cultivation experiments (Kadereit 1984a; Gilmer & Kadereit 1989)
and extensive herbarium surveys (Allen 1967; Perring & Sell 1968; Kadereit 1984a) indicates that
subsp. denticulatus and var. vulgaris differ conspicuously with regard to germination behaviour, life
cycle, morphology and reproductive output, but also geographical distribution and habitat
preference. Although no unambiguous native habitat can be delineated for the cosmopolitan weed
var. vulgaris (Kadereit 1984a), subsp. denticulatus is restricted either to natural maritime habitats
(dunes, sandy fields and cliffs) along the coasts of N.W. Europe, i.e. from W. France, the Channel
Islands, W. and S.W. Britain, to the S. and E. coasts of Sweden (Allen 1967), or can be found in the
Mediterranean area, mainly in the mountains of S. Spain and Sicily (Kadereit 1984a). In the British

*Present address: School of Biological and Medical Sciences, Sir Harold Mitchell Building, University of St
Andrews, St Andrews, Fife, KY16 9TH, Scotland, U.K.
+Nomenclature follows Stace (1991) for vascular plant species.

206 H. P. COMES

Isles only records from Ainsdale (Merseyside) and the Channel Islands have been confirmed in
recent years (Ashton 1990; Harris & Ingram 1992). Rather surprisingly, no detailed ecological study
of the natural habitat of subsp. denticulatus is available.

The present account provides some field data on the distribution, habitat and pollination strategy
of subsp. denticulatus in a maritime site on Jersey, Channel Islands. As var. vulgaris is reported to be
common on Jersey (e.g. Le Sueur 1984), another objective of the present study was to investigate
whether the two taxa are isolated through habitat differentiation, or whether they co-occur and
hybridize.

The most conspicuous morphological character of subsp. denticulatus, and certainly its most
reliable diagnostic character in the field, is the presence of ray florets 2-3—3-0 mm long. Ray florets
are absent in var. vulgaris. The presence of the ray florets is under simple genetic control of the ‘ray
floret locus’, with subsp. denticulatus homozygous for the R allele, discoid plants of var. vulgaris
homozygous for the r allele and short ligulate hybrids between both taxa heterozygous (Trow 1912;
Comes 1994). Other less reliable field characters are the rather dense arachnoid indumentum, leaf
shape characteristics (Allen 1967) and an irregular achene indumentum (Gilmer & Kadereit 1989).
Plants of subsp. denticulatus differ further from var. vulgaris in showing pronounced seed
dormancy, which also seems to be regulated by one major gene (Kadereit 1984a). Recently,

however, Ren & Abbott (1991) reported the presence of innate seed dormancy in var. vulgaris from — ;

the Mediterranean area. The authors concluded that this feature probably evolved in adaptation to
the climatic conditions of heavy winter rains and severe summer drought, and enables this taxon to
grow as a ‘typical Mediterranean’ winter annual. Given the putative Mediterranean origin of subsp.
denticulatus (Kadereit 1984b), the same explanation was previously put forward by Kadereit
(1984a) to account for the presence of both these characters, seed dormancy and winter annual life
cycle, in subsp. denticulatus. The distinctness of subsp. denticulatus from var. vulgaris is further
emphasized by its substantially prolonged time of vegetative growth, first noticed by Trow (1912,
1916) and later by Kadereit (1984a), and its potentially smaller reproductive output under standard
conditions in the greenhouse (Kadereit 1984a).

On Jersey, field studies were primarily focused on the sand dune system of St Ouen’s Bay at the
western end of the island, and in particular on the widest part of the dunes in the south, known as
Les Quennevais. In this area, initial observations revealed that subsp. denticulatus becomes
exceedingly rare c. 200 m away from the shore, but is present. again 1,000 m or more farther inland.
Since the Quennevais area is well-known for remarkable changes in soil pH and carbonate values
(Bocher 1954; Ranwell 1975), a study was designed to investigate the distribution of subsp.
denticulatus and its associated vegetation in relation to soil types along a transect, roughly coinciding
with Ranwell’s (1975) main study transect. Soil samples from outside this area were also analysed.

As I encountered discoid plants within the habitat of subsp. denticulatus which could not be
assigned to either taxon using morphological criteria, a comparative cultivation experiment was
designed to determine whether these variants exhibit (i) innate seed dormancy, (11) a substantially
slower speed of development and (ii1) a smaller reproductive capacity than typical plants of var.
vulgaris from Jersey and, as a control, var. vulgaris from the Botanic Garden, Mainz University
(Germany).

MATERIALS AND METHODS

FIELD WORK
Studies on Jersey were carried out between 13 and 19 May 1992. Following Ranwell’s terminology
(1975), the transect area on Les Quennevais (Fig. 1a), ranging from the seawall south of the slipway
at Le Braye to the inland limit of the dune system south of the enclosed valley below Ville des
Quennevais (c. 1-6 km), was classified into four zones of landscape structure according to distance
from the shore: 1. the coastal dune area (O—c. 100 m); 2. the dune plain (100-700 m); 3. the plateau
scarp (700-1000 m); and 4. the plateau dunes including the crestline (1000-1600 m). Within 300 m
distance from the shore, the number of plants of S. vulgaris was determined in 2m X 2 m sampling
areas at regular 2 m intervals. Beyond 300 m inland, only plants were counted that were found by
walking over the terrain towards the plateau scarp, and to the plateau height (c. 1025 m inland).
Along the transect, it was impossible to distinguish between subsp. denticulatus and short ligulate

SENECIO VULGARIS ON JERSEY (CHANNEL ISLANDS) 207

plants identified as probable hybrids between subsp. denticulatus and var. vulgaris. Any attempt to
quantify differences in ray floret length certainly would have resulted in confusing putative hybrids
with small individuals of subsp. denticulatus bearing small ray florets. Accordingly, all radiate plants
have been referred to as subsp. denticulatus. At various patches near the transect, however, in
particular adjacent to rabbit faeces, plants grew larger. In those places, therefore, individuals could
be classified according to their ray floret length as subsp. denticulatus (> 2 mm) and putative hybrids
(<2 mm).

Seven soil samples were taken along the transect (sites 1-7), and five samples were collected from
surrounding localities (sites 8-12) by removing soil to a depth of c. 6 cm (Fig. 1a,b). Soil samples
were allowed to air dry., and subjected to chemical analysis by the Landwirtschaftliche Unter-
suchungs- und Forschungsanstalt at Speyer (Germany). The pH, percent organic matter, free
carbonate, total nitrogen and extractable forms of magnesium (Mg) were determined for all
samples, while extractable forms of phosphorus (P,O;), potassium (K,O) and boron (B) were only
analysed for samples 1-5. At all sites along the transect records of associated vegetation were made.

OBSERVATION OF PUTATIVE POLLINATORS

Observation of insects was carried out in two natural stands at St Ouen’s Bay; stand 1 was situated in
a mobile Marram (Ammophila arenaria) dune just behind the sea wall south of Le Braye slipway
(near site 1 in Fig. 1a), and the other, stand 2, next to the bunker at La Téte du Nieér Céte (at site 10).
Stand 1 was monomorphic for Senecio vulgaris subsp. denticulatus, while stand 2 contained a few
short ligulate plants which appeared to be hybrids. Stand 1 consisted of 156 plants which were evenly
scattered throughout the observation area. Plants were (S—)9-12(—18) cm tall and mostly had one to
three open capitula. Stand 2 included much larger (up to 20 cm), often basally branched individuals
with up to 30(-40) open capitula. Approximately 70 individuals (20-30 individuals/m”) were
distributed in several patches. On 18 May 1992, the flight sequences of all insect pollinators entering
the observation area were recorded between 12.30 p.m. and 2.00 p.m. (stand 1) and 3.30 p.m. and
5.30 p.m. (stand 2).

COMPARATIVE CULTIVATION

Seed material (achenes) for comparative cultivation, including a germination experiment, was
derived from each of eight individuals of subsp. denticulatus (in the following referred to as A1—A8),
seven individuals of unidentified discoid plants (B1—B7), three individuals of short ligulate plants
identified as probable hybrids between the two taxa (C1—C3) and eight individuals cf var. vulgaris
(D1—D8) collected in various locations around the coastal plain at St Ouen’s Bay, the neighbouring
Ouaisné Common, as well as interior waste ground or agricultural sites at the eastern landward edge
of St Ouen’s Bay. Between 14 and 19 May 1992 seed material was either directly harvested in the field,
or, at the end of May, carefully removed from fruiting plants which had been pressed as herbarium
specimens. In addition, achenes were collected from eight individuals of var. vulgaris (E1—E8) in the
Botanic Garden, Mainz University, on 31 May 1992. Collecting codes of seed parents are used
throughout the text to refer to their respective greenhouse progenies and are listed in Table 1.
Collecting sites are also included, and their numbers refer to locations illustrated in Fig. 1la,b.

For a test of seed dormancy, samples of five achenes per seed parent were sown on five dates
between 31 May and 8 September 1992 in 5 cm plastic pots filled with Friihstorfer Erde, Typ P, and
were kept moist during the entire experiment. Germination was recorded daily over a period of 160
days from first sowing.

At 27, 50 and 72 days following first sowing, a variable number of seedlings (as available) from
each of the five accessions (A—E) were transplanted individually into 8 cm diameter plastic pots
filled with sterilized compost. On any one day, only samples that had germinated during the
previous 20 days were transferred. A total of 154 plants, representing between one and nine
offspring from each of the seed parents listed in Table 1 except A2, A5, A6, A8 and C3, were grown
to first fruiting under a natural light regime in an unheated greenhouse at the Botanic Garden,
Mainz University. As the experiment approached the winter period, plants were transferred to a
heated greenhouse with supplemental lighting provided by incandescent lamps (16 hour day), and
temperatures at 20 °C day / 15 °C night. Conditions were maintained this way until the end of the
experiment on 21 January 1993.

208 H. P. COMES

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2 SE Dune grassland

FIGURE 1. a. Topographic location of study sites on Les Quennevais (sites 1-7) and adjacent areas (sites 8-10).
Based on the 1:25,000 States of Jersey Official Map, 1988; additional information on contours and sand pit
outline compiled from the 1:25,000 Geological Survey Map, Jersey, 1982. Heights are in feet above M.S.L.
based on a bench mark at St Helier Harbour, Jersey, of 29-9 feet (= 9-1 m). Built-up area hatched.

b. Map of Jersey showing the location of four study sites (11-14) outside the research area (outlined). The
distribution of extant dune grassland (Le Sueur 1984) is indicated.

SENECIO VULGARIS ON JERSEY (CHANNEL ISLANDS) 209

TABLE 1. COLLECTING SITES AND COLLECTING CODES OF SEED PARENTS OF SENECIO
VULGARIS FROM WHICH EXPERIMENTAL PROGENIES WERE GROWN

Unidentified
Subsp. discoid Putative
Collecting site denticulatus plants hybrids Var. vulgaris
COASTAL PLAIN AT
ST OUEN’S BAY, JERSEY
Dune behind seawall, near Bil B24
waste ground, L’Etacq (11)
Les Quennevais, coastal Al*, A2, B3*, B4* Gi
dune and dune plain south A3*, A4*, AS

of slipway at Le Braye, 10—
100 m inland from the
shore (1/2)
Les Quennevais, plateau A6 B5*, B6* (Cy
dunes, c. 1320 m inland
from the shore (6)
On sandy soil near bunker, A7* C3 Die
La Téte du Niér Cote (10)

OUAISNE COMMON, JERSEY
Coastal dune belt, Ouaisné A& B7*
Common (13)

INTERIOR SITES AT THE
LANDWARD EDGE OF
ST OUEN’S BAY, JERSEY

Wayside in the valley below D2s
Ville des Quennevais (8)
Arable field, St Brelade (9) D3*
Waste ground, St Ouen (14) D4*-D8*
BOTANIC GARDEN, MAINZ E1*-E8*

UNIVERSITY, GERMANY

* Seed parents used both in the germination and the comparative cultivation experiment; numbers in brackets
refer to locations in Fig. 1.

STATISTICAL ANALYSES

During the experiment, plants were examined for the phenological, vegetative and reproductive
characters listed in Table 2. Means and standard errors for each character were computed. Multiple
comparisons of group means were made using Tukey’s test (VMS Version of SAS Release 6.07),
considered appropriate for unequal sample sizes (Sokal & Rohlf 1981; SAS Institute Inc. 1988), and
were evaluated at the 1% significance level. In order to represent interrelationships among groups
graphically, the data were also subjected to canonical variate analysis (C.V.A.) (Reyment et al.
1984; Reyment 1991). Character LRAY (length of ray floret) was excluded from the C.V.A.
because the character was known a priori to distinguish some of the accessions. The C.V.A. was
carried out using the SAS procedure CANDISC (VMS Version of SAS Release 6.07). This
procedure also computes both Hotelling’s multivariate T7-test, which tests the hypothesis that the
canonical means of the groups are equal, and Mahalanobis’ distances (D7) between canonical means
of groups.

RESULTS

DISTRIBUTION AND HABITAT

Senecio vulgaris subsp. denticulatus was only found in Jersey on two dune systems, namely Ouaisné
Common and the coastal stretch along St Ouen’s Bay with the largest and deepest dune system in
the south, known as Les Quennevais (Fig. 1a,b). Jersey’s third area of dune grassland, Gorey
Common, at the eastern end of the island, was not visited. All populations seen in inland areas were
var. vulgaris. By contrast, plants of subsp. denticulatus, putative hybrids, unidentified discoid plants

210 fi? ES COMBS

TABLE 2. CHARACTERS SCORED IN THE COMPARATIVE CULTIVATION EXPERIMENT OF
SENECIO VULGARIS

Character | Abbreviations

Determined by the daily inspection of all plants:

1. Days from germination to formation of first buds BUD

2. Days from first buds to first anthesis BA

3. Days from germination to first anthesis ANTH

4. Days from anthesis to first fruiting AF

5. Days from germination to first fruiting FRUIT
Measured ait the time of first fruiting:

6. Stem height (cm) from soil surface to base of apical capitulum STH

7. Number of capitula and buds NCAP

8. Length (mm) of ray floret* LRAY

9. Number of achenes per apical capitulum NACH
10. Number of unfertilized ovules per apical capitulum NUNFO
11. Number of florets per apical capitulum NFLORET

(=NACH+NUNFO)

12. Reproductive potential REPRO

(=NCAPxNACH)

* Only subsp. denticulatus and putative hybrid progenies.

and var. vulgaris grew in mixtures of varying composition in various places in the man-disturbed
area north of Les Quennevais. This area, however, was not explored extensively. Near to plants of
subsp. denticulatus found on Ouaisné Common was an extensive population of var. vulgaris growing
at a building site. However, no putative hybrids were found in the latter location.

Particular attention was paid to the Quennevais area, where subsp. denticulatus characteristically
grows either on seaward facing slopes of mobile coastal dunes dominated by Marram, and
additionally characterized by Phleum arenarium, or grows abundantly as a component of an open
turf vegetation, called fixed calcareous dune vegetation by Bocher (1954), which follows a little
farther away from the coastal dune area. Common associates here are bryophytes and small
herbaceous plants like Myosotis ramosissima and Saxifraga tridactylites, and plants of essentially
Mediterranean affinity like Lagurus ovatus, Mibora minima and Viola kitaibeliana.

RELATIONSHIP OF PLANT FREQUENCIES TO SOIL PARAMETERS AND ASSOCIATED VEGETATION

The frequency distribution of plants of Senecio vulgaris subsp. denticulatus, including six
unidentified discoid plants, in relation to the (approximate) topographic location of study sites 1—7 is
illustrated in Fig. 2. It is evident that the frequency of plants of subsp. denticulatus declines
dramatically beyond 200 m distance from the coast, with plants becoming extremely rare over most
part of the dune plain and along the plateau scarp, an area c. 800 m wide. Surprisingly, plants of
subsp. denticulatus, short ligulate hybrids and unidentified discoid plants were abundant again on
the plateau dunes (beyond c. 1025 m inland from the shore), extending as far as the inland limit of
the dune system. This observation, however, was not documented in detail. Abundance of S.
vulgaris on the plateau dunes, nonetheless, might be exemplified by 97 individuals recorded in one
subjectively chosen 1 m* quadrat at c. 1345 m distance from the shore (site 7, Fig. 1a), comprising 53
plants of subsp. denticulatus, 32 short ligulate hybrids and twelve unidentified discoid plants. This
exceeded the density of plants found in the coastal dune area, where in a3 m X 2 m sampling unit
outside the transect area, c. 50 m inland from the shore, 105 radiate, eight intermediate and two
discoid plants were counted among a total of 115 plants.

Soils of all sites from Les Quennevais have high pH and also high carbonate values (Table 3). In
general, they are low in organic matter, total nitrogen, and, as far as analysed, all minerals including
boron, magnesium, phosphorus and potassium. Also, it is evident that soil pH is largely similar in all
soil samples analysed. One of the plateau dune sites (7), where vegetation cover of subsp.
denticulatus was exceptionally high (see above), is more similar to the densely covered coastal open
turf site (2) than to any other site analysed, including the second plateau site (6) nearest to it. This

SENECIO VULGARIS ON JERSEY (CHANNEL ISLANDS) pA \Ah

Coastal Dune plain Plateau scarp Plateau dunes
dune 80

Hi. above M.S.L.in m

Celene nreSs 64 eS 6 S70 948 6 SO OTH. TZ SSLAl4e aS: 16
1 Distance landwards from H.W.M:S.T.in 100 m

FiGureE 2. Frequency distribution of subsp. denticulatus (including six unidentified discoid plants) in relation to
the approximate topographic location of study sites 1-7 on Les Quennevais. Note that the area c. 1025 m and
more farther inland from the shore (hatched) harbours large populations of subsp. denticulatus, short ligulate
hybrid, and unidentified discoid plants (not mapped in detail). Numbers correspond to sampling sites shown in
Fig. la. N = number of individuals.

relates to higher amounts of organic matter and total nitrogen, as well as to lower levels of carbonate
content, indicating a great amount of small scale variation of these edaphic factors across the
transect zone. The results from the soil analyses reveal no distinct signs of leaching on the plateau
dunes farthest east, thus confirming previous results obtained by Ranwell (1975).

Table 4 lists the species recorded in the area around the seven soil sampling sites on Les
Quennevais. Among those plant species which were found to show a disjunct pattern of distribution
along the transect similar to that observed in subsp. denticulatus, i.e. being present in the coastal
area (sites 1 and 2) and on the plateau dunes (6 and 7), but virtually absent from most parts of the
dune plain and the plateau escarpment (3-5), were Honkenya peploides, Phleum arenarium,
Saxifraga tridactylites and Viola kitaibeliana. In contrast, other taxa, for example Euphorbia
portlandica, Mibora minima and Myosotis ramosissima, were present along the entire transect.
Chemical analysis, however, makes clear that the remarkable pattern of disjunct distribution
exhibited both by Senecio vulgaris subsp. denticulatus and its four community associates is not
reflected by the soil parameters (Table 3).

As regards the surrounding localities (Fig. 1a,b), the bunker site at La Téte du Niér Cote (10),
closest to Les Quennevais, harboured plants of subsp. denticulatus and putative hybrids, but also a
single individual of var. vulgaris in a sheltered and shaded place nearby. This site is edaphically most
similar to Les Quennevais, except for the lack of carbonate (Table 3), and, apart from Armeria
maritima, has a rather similar set of species associates (not recorded in detail). In contrast, the
seawall site near waste ground at L’Etacq (11), where scattered unidentified discoid plants of
Senecio vulgaris were found, is the most unusual of all sites at St Ouen’s Bay because it has relatively
high amounts of organic matter, nitrogen and carbonate, and extremely high amounts of
magnesium. Both localities are of high pH, similar to values found on Les Quennevais. The two
interior sites (8 and 9), typical for var. vulgaris, are characterized by lower pH values than all sites at
St Ouen’s Bay, and have relatively high amounts of organic matter and nitrogen, but these values
are lower than those found at L’Etacq. Carbonate is absent from both of these interior sites. In the
maritime site at Beau Port (12) S. sylvaticus occurred profusely, but no plants of S. vulgaris were
found. This certainly reflects an edaphic composition which is virtually intolerable for the latter
species with respect to high levels of soil acidity, soil compactness, and, probably, nitrogen content
(Table 3).

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214 H. P. COMES

TABLE 4. VEGETATION ASSOCIATED WITH SENECIO VULGARIS AT SITES 1-7 IN JERSEY
(CHANNEL ISLANDS)
Records from the adjacent sites 6 and 7 are pooled for comparison.

Sites

Taxa 1 2 3 4 5 6 and 7

Ammophila arenaria a + + + at
Bellis perennis +

Calystegia soldanella “F

Cerastium sp. au
Cerastium diffusum ap ae
Cerastium semidecandrum “F

Cladonia rangiformis + 4
Claytonia perfoliata

Crataegus monogyna

Dactylis glomerata

Eryngium maritimum +

Euphorbia portlandica + + a +
Fumaria muralis a

Galium verum a
Geranium sp. i.
Geranium molle a |
Honkenya peploides+ - 4.
Juncus sp.

Lagurus ovatus* +
Mibora minima*

Myosotis ramosissima +
Oenothera stricta as

Phleum arenarium*+ + =
Plantago maritima
Pinus pinaster +

Quercus ilex* au a
Quercus robur a
Ranunculus bulbosus + +

Rosa pimpinellifolia ate aF + +
Rubus fruticosus agg.
Saxifraga tridactylitest au
Senecio vulgaris subsp. denticulatus+ + (+) +
Silene nutans a ae at
Trifolium occidentale +

Ulex europaeus + +
Vicia sativa + =- a au
Viola kitaibeliana*+ + a

+++

+4+4++
+
+
+

+

+++
+

Notes: Taxa are indicated as + = present or (+) = rare. Taxa of Mediterranean affinity are marked * and those
showing a pattern of disjunct distribution along the transect are indicated by f.

OBSERVATION OF PUTATIVE POLLINATORS
Within each of the two natural stands at St Ouen’s Bay, flies were the most frequent visitors of
subsp. denticulatus during the observation periods. Members of the family Sarcophagidae
(Sarcophaga dissimilis Meigen) were caught and identified within stand 1. These flies showed a
remarkable preference for the ray florets; systematically manipulating these florets one after the
other possibly for nectar extraction, they often moved in a circle on the capitulum, largely neglecting
the inner disc florets. Flies of the family Anthomyiidae were taken from stand 2. Though specimens
attracted by the decaying Fucus tidal litter on the nearby shore were also members of the same
family (Anthomyiidae), they were clearly a different species (P. Kirby, pers. comm., 1993).
Table 5 gives the number and the flight movements of insects in terms of number of plants visited
during a flight sequence recorded within each of the two stands. The total number of insects

SENECIO VULGARIS ON JERSEY (CHANNEL ISLANDS) 215

TABLE 5. OBSERVED NUMBER OF PUTATIVE POLLINATORS VISITING ONE TO FIVE DIFFER-

ENT PLANTS OF SENECIO VULGARIS SUBSP. DENTICULATUS IN TWO NATURAL STANDS AT
ST OUEN’S BAY, JERSEY

No. of plants visited during flight sequence

Stand 1 2 3 4 5
1 4 10 0 Z 1

2 3% 2 0 0 0

“Including one unidentified syrphid fly.
> Flights between long rayed plants.
Stand 2 harboured several short ligulate hybrids.

(including one syrphid fly) visiting stand 1 and stand 2 was 17 and 5, respectively. Mean number of
plants visited per putative pollinator was 2-2 (stand 1) and 1-4 (stand 2).

Discrimination by flies in favour of the long rayed plants of Senecio vulgaris subsp. denticulatus in
comparison to short ligulate hybrids was observed in stand 2, where both transitory flights occurred
between long rayed individuals. The observation that long rayed plants were more attractive to
insects than intermediate and discoid plants, was also made in a 12 m? mixed stand on the plateau
dunes that was observed on 14 May 1992 for a windy one-hour period (2.30 p.m.—3.30 p.m.). This
stand also comprised unequal numbers of radiate, intermediate and discoid plants of neariy equal
height ((7—)17—21 cm) in an approximate ratio of 5:3:1. During this time, one unknown insect visited
preferentially three long rayed plants during a flight sequence.

Outside observation periods, Psilothrix viridicoeruleus Fourcroy, a small beetle of the Melyridae
family, was frequently seen visiting the capitula of Senecio vulgaris subsp. denticulatus in the dunes,
not only foraging on pollen but probably also feeding on the ray florets, which often showed marked
signs of damage.

A fly seen visiting var. vulgaris in the valley below Ville des Quennevais was Siphona geniculata
De Geer (Tachinidae) which is a well-known and often abundant flower-visiting fly, with a strong
preference for yellow composites (P. Kirby, pers. comm., 1993). Evidently it does not belong to the
most frequently observed species of flies visiting Senecio vulgaris subsp. denticulatus at St Ouen’s
Bay.

SEED PRODUCTION AND PREDATION DAMAGE

The percentage seed set of subsp. denticulatus was recorded in a sample of 17 individuals chosen
randomly from the area south of the slipway at Le Braye. Care was taken to collect capitula that hac
not shed their achenes. The proportion of well-developed achenes found within one capitulum of
each plant examined averaged 88-0% and ranged between 48-8 and 98-2%. In the field, there was no
indication of any impediment for setting seed under natural conditions, including plants identified as
putative hybrids.

As indicated by decapitated, fruiting capitula which were often found lying near the base of the
parent plant, achenes of subsp. denticulatus seem to be predated even before the time of seed
dispersal. Birds are likely to be the predators, as small flocks of goldfinches have been reported to
feed on subsp. denticulatus (Le Sueur 1976). In addition, subsp. denticulatus obviously suffers from
grazing by herbivores (probably rabbits) which clip the main axis of the plants and thus affect their
branching pattern by inducing the growth of lateral shoots from near the base.

GERMINATION BEHAVIOUR AND COMPARATIVE CULTIVATION

In the course of comparative cultivation, including the germination experiment, it became strikingly
evident that in regard to the offspring of discoid plants only progeny of the unidentified discoid
parent individuals from the Quennevais area (B3—B6) showed characteristics typical of subsp.
denticulatus, while offspring of discoid plants from outside this area (B1, B2 from L’Etacq and B7
from Ouaisné Common) proved to be typical var. vulgaris. Taking account of the distinctiveness of
progenies B1, B2 and B7 on the one hand, and B3—B6 on the other, as will be demonstrated below,

216 H. P. COMES

results are presented separately. For all groups analysed, data were pooled over families for
comparison.

The results of the seed dormancy experiment are illustrated in Fig. 3. Rather unexpectedly, no
pronounced dormancy pattern was recorded for achenes of subsp. denticulatus from Les Quenne-
vais, La Téte du Niér Céte and Ouaisné Common. The minimum time required for germination was
only 23 days after first sowing in this taxon. This was followed, however, by a period of continuous
germination, which extended over 79 days. Thus, although a few achenes (all produced by parent
individuals Al and A3) germinated within four weeks from first sowing, a greater proportion of
subsp. denticulatus achenes exhibited a pattern markedly skewed towards later germination, thus
showing various degrees of dormancy. While cumulative percentages of total germination largely
remained constant over all five successive dates of sowing, time to initial germination and time to
final germination decreased from 23 (27) to 10 days and from 102 to 30 days after sowing,
respectively. Hence, partial seed dormancy, though obviously present in subsp. denticulatus, had
been completely lost by the last date of sowing (8 September 1992), i.e. after 14 week’s storage at
room temperature. There was no seed dormancy in var. vulgaris from the coastal site at La Téte du
Niér Céte and interior inland sites of Jersey, as well as from Mainz Botanic Garden. Independent of
the date of sowing, the majority of seedlings emerged within 10 days (Fig. 3e,f). Of the achenes of
unidentified discoid parent individuals, accessions from L’Etacq (B1 and B2) and the dune
grassland on Ouaisné Common (B7) showed a pattern similar to var. vulgaris, although synchroni-
zation was less apparent during the first two dates of sowing (Fig. 3b). In contrast, achenes produced
by unidentified discoid plants from Les Quennevais (B3—B6) and putative hybrids from the same
location and La Téte du Niér Céte (C1—C3) largely followed the germination pattern of subsp.
denticulatus (Fig. 3c,d). Both differed from subsp. denticulatus, however, in that the discoid plants
from the Quennevais showed a more rapid initial germination within the first four weeks following
first sowing (mainly caused by achenes produced by parent individuals B3 and B4), while, in the case
of the putative hybrids, storage at room temperature for 14 weeks was less effective in inducing
more rapid and synchronizing germination.

Turning to the results of the cultivation experiment (Table 6), it was evident that subsp.
denticulatus differed significantly (at the 1% level) from both accessions of var. vulgaris in taking a
substantially longer time for each developmental stage analysed, and producing more florets per
capitulum containing more unfertilized ovules. Number of achenes per capitulum and potential
reproductive output, however, were not significantly different for the three groups. With regard to
the two vegetative characters stem height and number of capitula (including buds), it was evident
that only plants of var. vulgaris from Jersey performed similarly to subsp. denticulatus.

The two accessions of var. vulgaris also differed from each other in several phenological and
vegetative characters. Plants from Jersey took approximately eleven days longer to complete their
life cycle due to differences in the time from germination to first buds, were taller in stature, and—as
was indicated by complete leaf collections of representative plants — produced more leaves along the
main stem than var. vulgaris from Germany. The major morphological difference between both
populations was leaf shape. Fig. 4c,d illustrates that the middle and upper cauline leaves of var.
vulgaris from Jersey were more deeply divided, with conspicuously dentate lobes.

As regards the unidentified discoid accessions, progeny from L’Etacq and Ouaisné Common (B1,
B2 and B7) were virtually indistinguishable from var. vulgaris from La Téte du Nier Cote and
interior sites in most characters, including leaf shape. The only exceptions were number of florets
and number of achenes per capitulum, which were significantly higher in the former, although no
differences were recorded for potential reproductive output. Conversely, progenies of both
unidentified discoid parent plants (B3—B6) and putative hybrid individuals (C1 and C2) from Les
Quennevais were generally closer in mean to each other and to subsp. denticulatus. In addition, all
three groups had similar leaf shape characteristics, as shown in Fig. 4a,b, and were strikingly
arachnoid. However, most developmental stages of the unidentified discoid plants from Les
Quennevais were significantly shorter than those of subsp. denticulatus , though the differences were
small.

Canonical variate analysis (C.V.A.) was conducted for the purpose of illustrating the
interrelationships among accessions, and in particular to investigate the distinctiveness of both
groups of unidentified discoid plants. Since the two groups were obtained a priori by visual
impression during cultivation, there remains the-possibility that differences between both groups

SENECIO VULGARIS ON JERSEY (CHANNEL ISLANDS) PN)

Percentage of
total germination

(a) subsp. denticulatus A1-A8

(v) 95.0
20
10
0)

(iv) 100.0
20

(iii) I 100.0

95.0

NUMBER OF GERMINATED SEEDS

- () I 92.5

10 30 50 70 90 110 130 150
Days after last date of harvest

(b) progeny of unidentified discoid parent individuals Bl, B2 and B7
(v) 73.3

(ii) 73.3

NUMBER OF GERMINATED SEEDS
5

(i) | 80.0

10 30, 50 70 90 110 130 150
Days after last date of harvest

Ficure 3. The germination behaviour of achenes of: (a) subsp. denticulatus (A 1-A8); (b) unidentified discoid
plants (B1, B2 and B7); (c) unidentified discoid plants (B3-B6); (d) putative hybrids (C1—C3); (e) var. vulgaris,
Jersey (D1—D8); and (f) var. vulgaris, Germany (E1-E8). Achenes collected between 14 and 31 May 1992 were
sown on five dates following the last date of harvest: i. 31 May, ii. 22 June, iii. 17 July, iv. 10 August, v. 8
September 1992. Arrows indicate the date of sowing. Also presented are percentages of total germination.

218 H. P. COMES

Percentage of

: aA total germination
(c) progeny of unidentified discoid parent individuals B3-B6

95.0
20
10
0 SEL 28 ee
(iv) 95.0
20-
n 10
a
w 6)
n
Fy Git) I 95.0
ze 205
Zz
S 10
a
0 ee,
\)
6 (ii) 100.0
aa 20
~
> 10
=)
Zz 0
(i) 100.0
20
10
(8) =
10 30 SO 70 90 110 130 150
Days after last date of harvest
(d) progeny of putative hybrids C!-C3
(v) 93.3
20
10
0 re
(iv) I 86.6
20
10
0
WY (iii) 86.6
Qa
a wne0
n
QA 10
WW)
2 0
LZ
S (ii) 73.3
jo4
jaa 20
o)
5 10
ra 0
~
= (i) 80.0
>
Ze

10 30 SO 70 90 110 130 150
» Days after last date of harvest

FiGuRE 3. Continued. 3

SENECIO VULGARIS ON JERSEY (CHANNEL ISLANDS) 219

Percentage of
: total germination
(e) var. vulgaris, Jersey, D1-D8

(v) 92.5
20
10
0)
(iv) 92.5
2)
Be 20
5 10
a
Wy 0
lem
< nae
Zz (iii) 95.0
=
25
es 20
) 15
10
De 5
iS) 0
a
a Ci) 97.5
oP 20
Z
10
0
(i) 95.0

10 30 50 70 90 110 130 150
Days after last date of harvest

(f) var. vulgaris, Germany, E1-E8

‘ (iv) 100.0

(11) I 95.0

NUMBER OF GERMINATED SEEDS
°

10 30 50 70 90 110 130 150
Days after last date of harvest

FIGURE 3. Continued.

H. P. COMES

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SENECIO VULGARIS ON JERSEY (CHANNEL ISLANDS) 221
b
d

Ficure 4. Silhouettes of representative middle and upper cauline leaves of cultivated progeny of: a. subsp.
denticulatus, Les Quennevais (A4); b. putative hybrid, Les Quennevais (C1); c. var. vulgaris, Jersey (D1); and
d. var. vulgaris, Germany (E2). Note that leaves of cultivated progeny of unidentified discoid plants B1, B2 and
B7, and B3—B6 were similar to (c) and (a), respectively.

10

cm

that may have been slight at the outset were ‘strengthened’ by the C.V.A., because this analysis
seeks to maximize the separation between groups (Reyment 1991). It should be emphasized that the
results of the C.V.A. were not used to investigate which characters contributed most to the
separation between groups.

Fig. 5 shows a plot of the canonical variate scores for 137 out of 154 individuals projected on to the
plane of the first two canonical axes. 17 individuals raised had missing values for some of the
characters scored, and thus were not included in the analysis, which requires equal sample sizes for
all characters (S.A.S. Institute Inc. 1988). The first two canonical variables were statistically
significant (at p<0.0001), and accounted for 86% and 9% of the total variation, respectively.

The plot of the canonical variates illustrates that all plants derived from unidentified discoid
parent individuals fall into two broad clusters. Progeny raised from individuals collected at L’Etacq
(B1 and B2) and Ouaisné Common (B7) are nearly completely intermixed with var. vulgaris from
La Téte du Nier Cote and inland sites, and to a lesser extent associated with var. vulgaris from
Germany. In contrast, offspring of four unidentified discoid plants from Les Quennevais (B3—B6)
form a distinct cluster both with subsp. denticulatus from Les Quennevais (Al, A3 and A4) and
from La Téte du Niér Céte (A7), as well as with progeny of putative hybrids from Les Quennevais
(C1 and C2). Rather interestingly, the C.V.A. shows at least moderate separation between both
populations of var. vulgaris from Jersey and Germany.

Except for progeny of putative hybrids and unidentified discoid plants from Les Quennevais,
canonical means of all six groups were significantly different according to Hotelling’s multivariate
T?-test (p<0.0001). However, the picture outlined in Fig. 5 is clearly reflected by the Mahalanobis’
distances (Table 7).

222 H. P. COMES
DISCUSSION

DISTRIBUTION, HABITAT AND ECOLOGY OF SUBSP. DENTICULATUS AND VAR. VULGARIS ON JERSEY
On Jersey, subsp. denticulatus is only known from the dune systems of the coastal plains on Ouaisné
Common and St Ouen’s Bay, where it is particularly common on Les Quennevais. The Quennevais
area in the south of St Ouen’s Bay was deliberately chosen for more detailed studies because it has
long been known to harbour extensive populations of subsp. denticulatus (Babington 1839; Trimen
1871; Lester-Garland 1903; Le Sueur 1984), and because it has suffered relatively little from soil
disturbance due to human activities in the past (Mrs F. Le Sueur, pers. comm., 1992). Thus, it can
safely be taken as representing the specific ecological requirements of subsp. denticulatus in its
natural maritime habitat. This dune system, which is obviously susceptible to various destructive
and catastrophic agencies, e.g. salt-laden winds and severe winter storms, sand mobility, water
shortage and grazing by rabbits, has mild temperatures in winter, high levels of insolation and even
rainfall throughout the year (Ranwell 1976). Here, subsp. denticulatus most characteristically grows
on mobile sand dunes or fixed calcareous dune grassland, where it is commonly associated with
other winter-annuals and species of Mediterranean affinity (Table 4). It seems likely that subsp.
denticulatus has gone unnoticed in the past as a member of the association Tortuleto-Phleetum
arenarii of the calcareous dune communities, investigated in detail by Tiixen (1937) and Westhoff
(1947). According to the latter author the main occurrence of this association is in S.W. Europe with
its main period of growth in the winter. The obviously close phytosociological relationship of subsp.
denticulatus to this association on Jersey is all the more interesting, as subsp. denticulatus has been
postulated to have migrated from the Mediterranean northward along the coasts of W. Europe at
the end of the latest glacial period (Kadereit 1984b). Similar to other community associates, it thus
would seem to have colonized in the north of Europe one of the regions mildest in winter.

The results obtained from soil analyses (Table 3) do not provide any meaningful clue to the

CANONICAL VARIATE 2

|
ro Sait lina augaiee relia (O) Dee, AG. Or sO
CANONICAL VARIATE 1
Figure 5. Canonical variate analysis for individual progeny plants of Senecio vulgaris. @ = subsp. denticulatus
(Al, A3, A4, and A7), 0 = unidentified discoid plants (B1, B2 and B7), O = unidentified discoid plants (B3—

B6), X = putative hybrids (C1, C2), @ = var. vulgaris, Jersey (D1—-D8), @ = var. vulgaris, Germany (E1—-E8).
Analysis includes characters listed in Table 2 except LRAY (length of ray floret).

SENECIO VULGARIS ON JERSEY (CHANNEL ISLANDS) 2D

TABLE 7. MAHALANOBIS’ SQUARED DISTANCES AMONG GROUPS ANALYSED IN THE
CANONICAL VARIATE ANALYSIS OF PLANTS OF SENECIO VULGARIS

Groups

1 D 3 4 5 6
1 #
2 133:465"* -
3 D209 x §3:23%2* a
4 16-71* 84-24*** 10-11 N.S. -
5 144.42*** 4-15*** 90-81*** 92-55*** 2
6 US Sekoic 15212 eee 123 -69°°7* 134.275 ** 13-0077" =

Notes: See Table 6 for explanation of group numbers. F value significance: * p<0-05, ***p<0-001, N.S. = not
significant.

unusual and disjunct distribution of subsp. denticulatus and several community associates (Honk-
enya peploides, Phleum arenarium, Saxifraga tridactylites and Viola kitaibeliana) within the
Quennevais area, as detected in the present study (Fig. 2; Table 4). Bocher (1954) has recorded a
pH value of 5-8 farther inland, thereby noting the disappearance of Phleum arenarium, Koeleria
macrantha and Tortula ruraliformis (Besch.) Ingham, and Ranwell (1975) encountered both a low
pH of 4.8, and a minimum value of carbonate content at a dune plain site c. 550 m inland from the
shore. The present results, however, show neither a drop in carbonate nor in pH value at the
landward dune plain (3), and the plateau scarp site (4). Taking account of both the close proximity
of Ranwell’s 550 m site to site 3, and the fact that the vegetation around these sites is characterized
by Cladonia rangiformis Hoffm. (Table 4), a well-known acidophilous species (Bécher 1954), it
seems reasonable to assume a rather patchy variation in edaphic composition in the region. Quite
clearly, more fine-grained estimates of edaphic factors, especially pH, as weil as availability of water
and mineral nutrients, will be necessary to settle the problem of disjunct species distribution on Les
Quennevais. Nevertheless, soil analysis does indicate that Senecio vulgaris subsp. denticulatus
occurs on soils which have large amounts of carbonate, undoubtedly resulting from crushed mollusc
shells, and low amounts of mineral nutrients, out of which nitrogen and phosphorus are considered
to be of major significance (Grime 1988).

On Jersey, var. vulgaris can be found in man-disturbed inland and coastal localities throughout
the island, such as on waste ground or along roadsides. Since subsp. denticulatus is virtually absent
from ruderal interior sites, this might provide a first clue to the possibility that both taxa are also
ecologically distinct in utilizing ephemeral resources to different degrees. It seems likely that due to
human disturbance (e.g. roads and sand pits at St Ouen’s Bay; building work at Ouaisné Common)
the rudera! var. vulgaris has been recurrently introduced into the coastal habitat of subsp.
denticulatus. Sympatry of the two taxa at St Ouen’s Bay and Ouaisné Common thus might indicate
that their ranges of ecological tolerance are not mutually exclusive. At St Ouen’s Bay, however, it
appears that preference for fertile and highly disturbed soils is a factor limiting the distribution of
var. vulgaris to areas outside Les Quennevais, as larger numbers of var. vulgaris plants were only
found growing in disturbed localities with soils characterized by relatively high amounts of organic
matter and nitrogen, e.g. at Ville des Quennevais, St Brelade and L’Etacq (sites 8, 9 and 11; Table
3). Furthermore, during another visit made to Jersey in August 1992, some tall growing plants of
var. vulgaris were also found on shingle in front of the seawall near L’Etacq, associated with the
drift-laden portion of the beach, an extremely unstable habitat which undoubtedly is very rich in
nitrogen from decaying organic matter (e.g. Chapman 1978).

POLLINATION STRATEGY OF SUBSP. DENTICULATUS AND VAR. VULGARIS

Although both taxa are self-compatible, there is a difference in pollination strategy between subsp.
denticulatus and var. vulgaris. Visits of insects (flies and syrphid flies) to var. vulgaris have been
reported (Abbott & Irwin 1988; Comes & Kadereit 1990), but this taxon normally appears to be an
obligatory selfer with outcrossing rates seldom exceeding 1% under field conditions (Marshall &
Abbott 1982, 1984). From the field observations presented above it seems that subsp. denticulatus is
much more attractive to insects (mostly flies in this case) than either the discoid variant or the short

224 Hees COMES

ligulate hybrid. Thus, in direct analogy to the self-compatible and radiate var. hibernicus that shows
outcrossing frequencies between 3 and 35% (Marshall & Abbott 1982, 1984), it is probable that
subsp. denticulatus has a higher potential for outbreeding than var. vulgaris due to the possession of
female ray florets which may outcross at a higher frequency than the hermaphrodite disc florets and
convey a greater attractiveness of the radiate capitula to pollinators. The difference in pollination
strategy between both taxa probably accounts for the fact that subsp. denticulatus consistently
exhibits a greater amount of variation in the characters contained in Table 6 than var. vulgaris.

GREENHOUSE EXPERIMENTS

The present study has confirmed that, when grown under standard conditions in the greenhouse,
subsp. denticulatus and var. vulgaris differ consistently in a number of morphological and life history
characters, including leaf shape, degree of hairiness, germination behaviour and speed of
development. Moreover, formal genetic (Trow 1912, 1916; Kadereit 1984a) and quantitative
genetic studies (Comes 1994) involving subsp. denticulatus from Jersey have shown that the
differences between both taxa are genetically based, and in general under multigenic control, except
the presence/absence of both seed dormancy and ray florets (and probably also speed of
development). However, at all loci assayed by standard protein electrophoretic techniques, Jersey
subsp. denticulatus has an identical phenotype to British and German var. vulgaris (Ashton &
Abbott 1992; Comes, unpublished results), whereas subsp. denticulatus from Ainsdale has unique
alleles at the 6 EST-2 and 6 EST-3 loci (Ashton 1990; Ashton & Abbott 1992).

As regards the germination behaviour of the cultivated material investigated here, the amount of
difference between both taxa does not conform entirely to experimental results obtained previously.
Kadereit (1984a) found that subsp. denticulatus from Les Quennevais required a minimum time of
111 days after harvest for germination. Contrastingly, no pronounced seed dormancy of subsp.
denticulatus derived from the same locality and two other sites (La Téte du Niér Cote and Ouaisné
Common) was found in this study (Fig. 3). Apart from different experimental conditions after
sowing, these different findings may, in part, be attributable to different conditions experienced by
achenes during formation and ripening on the mother plant, as Kadereit (1984a) used greenhouse-
produced seeds in his experiment. These arguments do not rule out the possibility that phenotypic
variability of germination behaviour in subsp. denticulatus could also be due to multiple allelism at
the seed dormancy locus. However, the findings of the present study still indicate that subsp.
denticulatus shows a markedly slower and less synchronized rate of germination than var. vulgaris
(Fig. 3).

Also, in contrast to what was previously reported by Kadereit (1984a) from a study of Les
Quennevais material of subsp. denticulatus and British var. vulgaris, no differences between both
taxa were recorded for potential reproductive output in the cultivated material (Table 6). Kadereit’s
findings largely resulted from a significantly smaller average number of capitula and buds present at
the time of maturity of the first capitulum in subsp. denticulatus. As plants were grown under
different conditions and there was significant variation among families for this character (Kadereit
1984a), there is the possibility of both a great amount of genotypic variability and a considerable
genotype X environment interaction for this feature.

Perhaps one of the most intriguing results of the progeny tests is that the pattern of germination
behaviour (Fig. 3), the leaf shape characteristics (Fig. 4) and the canonical variate analysis (Fig. 5)
clearly show that all discoid progenies that were raised from the unidentified discoid plants (B3—B6)
from Les Quennevais were very similar to subsp. denticulatus. Although — with the exception of
character AF (time from anthesis to first fruiting) — these variants are consistently associated with
slightly lower values of all phenological characters (Table 6), the phenological differences are slight
compared to those between discoid and radiate subsp. denticulatus on the one hand and Jersey var.
vulgaris on the other. Thus, the present study has demonstrated that the presence of ray florets
alone might be insufficient to identify subsp. denticulatus on Jersey. Evidently, when growing in
drought-prone coastal sites, Jersey var. vulgaris may approach the discoid denticulatus-like variant
in phenotype, and thus may impede identification in the field. Such a possibility has been
demonstrated by the fact that, when collected, discoid seed parents from the fertile site at L’Etacq
~ (B1 and B2) and the dune grassland on Ouaisné Common (B7) could not be assigned to either taxon
using morphological criteria, but progeny analyses proved this material to be typical Jersey var.
vulgaris. It is worth noting that two earlier records exist concerning the occurrence of discoid

SENECIO VULGARIS ON JERSEY (CHANNEL ISLANDS) Pipes

variants of subsp. denticulatus in coastal habitats, namely in the British Isles at Pevensey, Sussex
(Wilmott 1925, 1949), and on Bornholm, Denmark (Lange 1851), but there is no evidence that these
variants are of similar, i.e. hybridogenous, origin as those described here. This will be discussed in
more detail below.

HYBRIDIZATION AT ST OUEN’S BAY

For the following discussion of hybridization at St Ouen’s Bay it is irrelevant whether subsp.
denticulatus represents a local ecotype or a population of a widely distributed monophyletic taxon.
Evidence for the latter comes from the observation of pronounced seed dormancy and strongly
elongated generation time in short ligulate material of S. vulgaris from S. Spain and Sicily (J. W.
Kadereit, pers. comm., 1993).

All plants scored and collected as putative hybrids at St Ouen’s Bay have been interpreted as such
because of their intermediate ray floret length. However, direct evidence of their hybrid status is
lacking because offspring individuals derived from two different short ligulate seed parents collected
on Les Quennevais did not produce the expected intermediate phenotype but resembled subsp.
denticulatus closely (Fig. 5). The fact that both individuals failed to segregate for capitulum type is
not a strong argument against their hybrid status since lack of segregation could have reflected
sampling error due to the very small number of progeny grown (N=3). Nevertheless, it appears
reasonable to assume that the short ligulate plants, together with the discoid denticulatus-like
variants described above, are the products of hybridization between subsp. denticulatus and var.
vulgaris. Alternatively, they could reflect a polymorphism for capitulum type in populations of
subsp. denticulatus, or even represent intermediate stages in the evolution of subsp. vulgaris. The
hypothesis of their hybrid status is preferred, however, because: (i) subsp. denticulatus and var.
vulgaris grow sympatrically at St OQuen’s Bay; (ii) both taxa are visited by a similar set of putative
pollinators; and (iii) fertile crosses can easily be obtained in the greenhouse (e.g. Gilmer & Kadereit
1989).

One morphological marker, absence of ray florets, that is fixed in var. vulgaris but — in general —
absent from subsp. denticulatus, is regulated by an allele of a single gene that is only weakly linked to
the gene(s) controlling speed of development (Comes 1994). If both taxa interact along an extensive
hybrid zone (sensu Harrison & Rand 1989) at St Ouen’s Bay, particularly in the man-disturbed area
north of Les Quennevais, Mendelian segregation of this morphological marker in the resultant
population is to be expected. The observation that the discoid variants tended to complete their life
cycle somewhat earlier than typical subsp. denticulatus (see above) might be attributed to the weak
linkage between the ray floret gene and the gene(s) controlling speed of development. Under the
polymorphism hypothesis, however, this association remains unclear (excluding the highly unlikely
possibility of pleiotropic or developmental effects of the ray floret gene on this phenological
character), and under the intermediate hypothesis it is unclear why no truly intermediate or
vulgaris-like individuals were found on Les Quennevais (Fig. 5).

Although the discoid and short ligulate denticulatus-like phenotypes thus might be considered as
products of hybridization between subsp. denticulatus and var. vulgaris, their mode of origin
remains obscure. It is possible that these phenotypes too are the products of many generations of
introgression with the subsp. denticulatus population on Les Quennevais as the likely recipient
population. Alternatively, they may be early recombinant segregants having arisen following self-
fertilization of an F, hybrid. Evidence in favour of the latter possibility is offered by hybridization
experiments between both taxa (Comes 1994). In these experiments, following spontaneous selfing
of the F, hybrid (57% self seed set), several recombinant individuals occurred in the F, that were
discoid (or short ligulate) but had phenological and morphometric features characteristic of subsp.
denticulatus.

Irrespective of their mode of origin, it remains to be established why all discoid and short ligulate
plants that were found on Les Quennevais showed characteristics typical of subsp. denticulatus. One ~
likely explanation is that the hybridization boundary around this area is less permeable for genes
regulating germination, phenological and morphological characters typical of var. vulgaris that are
more likely to have ecological and adaptive significance, than for those determining capitulum type
that is more likely to be influenced by random drift, especially in the presence of efficient self-
fertilization and in the absence of tight linkage with other life history characters. Moreover,
different mechanisms of mineral uptake and utilization could select for the denticulatus genotype in

226 Hy P? COMES

the Quennevais area. It seems likely that subsp. denticulatus is adapted to harvest and retain scarce
resources under the poor nutrient-supplying power of this area in a similar way to well-known stress
tolerators on calcareous soils, e.g. the associated Saxifraga tridactylites and Myosotis ramosissima
(Grime et al. 1988). Under these circumstances, both a prolonged phase of vegetative growth and a
delayed onset of reproduction in Senecio vulgaris subsp. denticulatus (Table 6) may be indicative of
rather low rates of mineral nutrient capture and utilization (Grime 1988). If this is so, the need for
early, rapid, and heavy resource allocation to reproduction in the ruderal var. vulgaris genotype will
be incompatible with its survival or competitive ability on Les Quennevais. The potential of nutrient
deprivation as a selective factor that might influence the establishment or spread of var. vulgaris has
been demonstrated by Aarssen & Burton (1990). These authors reported evidence that maternal
plants of var. vulgaris grown in nutrient-poor soils produced seeds with lower individual mass, seeds
that germinated later, and seedling offspring that had significantly lower biomass and height than
those individuals with high maternal soil nutrient level (but, unexpectedly, also seedling offspring
that survived longer in the absence of external nutrients). Although it is tempting to attribute
differential survival or competitive ability in the Quennevais area to different metabolic mechan-
isms, other edaphic or biotic factors on Les Quennevais could also select for the denticulatus
genotype. For example, rabbit grazing and poor water supply might well be expected there, but not
(or to a lesser extent) in the man-disturbed localities around this area, and this would also be likely
to influence life history characters like speed of development.

Whatever its selective advantage on Les Quennevais, provided there is in situ selection, and given
both an efficient self-pollination strategy and low levels of hybridization, subsp. denticulatus will be
able to maintain its integrity there. ;

EPILOGUE

There is documentary evidence to suggest that subsp. denticulatus was once more widely distributed
on Jersey. Populations of subsp. denticulatus may have existed on the coastal plains of St Aubin’s
Bay at least until the first half of the 19th century (Babington 1839), but since this locality was not
referred to any more by Lester-Garland’s account in 1903 (Lester-Garland 1903) it seems plausible
that they were extinguished by the stabilizing of the dunes or the building of the sea walls towards
the end of the 19th century. Equally, since Lester-Garland (1903) reported this taxon at St Brelade’s
Bay (including Ouaisné Common), the current population on Ouaisné Common appears to be a
mere remnant of those populations that existed along St Brelade’s Bay at the beginning of this
century. Thus, even if it is not threatened by hybridization with or introgression from var. vulgaris,
subsp. denticulatus might be seriously endangered from progressive destruction of its habitat by
man.

ACKNOWLEDGMENTS

I am very grateful to Mrs F. Le Sueur for her invaluable advice during my field studies on Jersey and
to Dr J. W. Kadereit (Mainz) and Mr C. D. Preston (Monks Wood) for critical discussion of the
manuscript. I also should like to express my thanks to Dr Peter Kirby (Peterborough) for identifying
the insect specimens and for his instructive comments on their biology, and to Anke Berg (Mainz)
for help with the illustrations. The field studies on Jersey were supported by a Landesgraduierten-
Forderungsgesetz (LGFG) grant of the Land Baden-Wirttemberg (Germany).

REFERENCES

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ncn A Ce Na Ne ES SEO sO AL AEA AOA A at COO see

a oe ST ee

a

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London.

(Accepted April 1994)

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Watsonia 20: 229-238 (1995) 229

Rorippa islandica (Oeder ex Murray) Borbas (Brassicaceae) in
Wales

A. O. CHATER
Windover, Penyrangor, Aberystwyth, Dyfed, SY23 1BJ
and
T. C. G. RICH

The Annex, Newgale Farm, Priory Road, Forest Row, East Sussex, RH18 5JD

ABSTRACT

Rorippa islandica (Oeder ex Murray) Borbas sensu stricto is reported from ten localities in Wales, a newly.
discovered stronghold for the species. An updated distribution map of the species in the British Isles is
presented. The habitats of the species on the Afon Teifi are described, including depressions in the flood plain, a
rock outcrop and waste ground. Possible dispersal mechanisms are discussed and the distribution of seeds both
by water and by wildfowl is considered probable.

Keyworps: Distribution, habitat, dispersal, life cycle.

INTRODUCTION

On 17 July 1991, A. O. Chater found a Rorippa growing on rocks by the Afon Teifi at Cenarth,
Cardiganshire (v.c. 46), which agreed with the description of R. islandica, Northern Yellowcress, in
Crucifers of Great Britain and Ireland (Rich 1991). He returned to collect ripe fruits on 25 August
1991, and the material was determined as R. islandica by T. C. G. Rich, and later confirmed by B.
Jonsell. The find was a surprise in two respects. First, this was the first confirmed record for Wales;
old records for ‘R. islandica’ have been treated as the generally commoner R. palustris (L.) Besser
rather than R. islandica (Oeder ex Murray) Borbas sensu stricto (Rich 1991). The nearest known
localities were in the Isle of Man 250 km to the north and western Ireland 250 km to the north west,
so it was a considerable extension of range (subsequently in 1993, another locality was discovered in
Kilkenny 200 km to the west north west). Second, the plant was growing on rocks beside a river
rather than in its more usual habitat in north west Europe of muddy ponds, turloughs and lough
margins.

In 1993, we re-surveyed known Rorippa localities along the Teifi and searched other sites in the
area to determine the status of the plant. We have also examined the excellent regional collection
held at the National Museum of Wales (NMW). In this paper we have documented the occurrence
of R. islandica in Wales and presented an updated distribution map. As there is little ecological
information available, we have also carried out studies on its habitats and vegetation on the Afon
Teifi, and investigated possible dispersal mechanisms. Nomenclature follows Stace (1991).

DISTRIBUTION IN WALES

All Welsh records traced are listed below. As the widespread occurrence of R. islandica in south
west Wales casts some doubt on unconfirmed records of R. palustris in Rich (1991), we have also
included brief details of genuine R. palustris records seen in the field or verified from herbarium
specimens. Fig. 1 shows the distribution of R. islandica in the British Isles as it is currently known,
incorporating the Welsh records and many additional localities in Ireland (Goodwillie 1994).

230 A. O. CHATER AND T. C. G. RICH

aes

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Bisel

i
bu
i
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bE

4

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rf
iF

FicurE 1. Distribution of Rorippa islandica in the British Isles, updated from Rich (1991). @ 1950-1993. O pre-
1950.

Glamorgan, v.c. 41:
1. Garnswllt, near River Loughor (SN/62.10), 13 July 1981, R. G. Ellis (NMW). First v.c. record.

Carmarthen, v.c. 44:

2. 800 m E of Newcastle Emlyn castle (SN/319.408), muddy hollows in silage field south of river and
N. of disused railway, 25 m a.s.l., c. 1000 plants, including many non-flowering rosettes, 23
August 1993, A. O. Chater and T. C. G. Rich (NMW).

3. Afon Bran, near bridge N. of Llandovery (SN/783.362), 11 September 1981, R. D. Pryce (NMW).
Immature plants collected at Cwm Mynys Isaf, N.E. of Llanwrda (SN/726.340), August 1986,
H. J. Killick (NMW), and on a rubbish tip at West Pwll (SN/4.0, tetrad Q), August 1982, I. K.

RORIPPA ISLANDICA IN WALES 231

Morgan (NMW) may also be R. islandica. The former site was searched in 1993 by A. O. Chater
but no suitable habitat for the plant was found.

Other material seen in NMW,, including Pont Stephen, Felinfach (SN/5.4, tetrads T and Y) collected
by A. O. Chater and R. D. Pryce in July 1982, is R. palustris.

Pembroke, v.c. 45:

4. 600 m E. of Llechryd Bridge, in silted-up bed of disused canal south of river and minor road (SN/
223.434), 7 m a.s.l., c. 20 fruiting plants and many non-flowering rosettes (number uncertain
because of presence of Rorippa palustris and consequent doubt about identity of vegetative
rosettes), 23 August 1993, A. O. Chater and T. C. G. Rich (NMW). First v.c. record.

Plants from Castlemartin Corse (SR/9.9) are R. palustris (NMW).

Cardigan, v.c. 46: .

5. 300 m W.S.W. of Llwynduris Farm, in and on banks of overgrown ditch in pasture and alongside
hedge on N. side of river (SN/235.433), 5 maz.s.l., c. 35 plants, mostly fruiting, 23 August 1993,
A. O. Chater and T. C. G. Rich, and 22 September 1993, A. O. Chater (NMW).

6. 50m W. of Cenarth Bridge, on slightly sloping rock platform on north bank of river (SN/246.416),
10 ma.s.1., four plants of R. islandica in crevices in an area of 4m X 5 m, prostrate and level with
the surface of the rocks, 17 July 1991, A. O. Chater (NMW). Three plants with inflorescences and
six non-flowering rosettes, 23 August 1993, A. O. Chater and T. C. G. Rich. No plants were
found on the south bank in v.c. 44 in 1993. First found “‘by the Teifi, Cenarth” in August 1958,
Miss D. E. de Vesian (NMW).

7.200 m S. of Newcastle Emlyn castle, along c. 100 m length of silted-up ox-bow and around ox-bow
pool on north side of river (SN/312.405), 25 ma.s.1., several thousand plants, including many non-
flowering rosettes, 23 August 1993, A. O. Chater and T. C. G. Rich (NMW).

8. The Moat, 1 km E.S.E. of Llandyfriog, muddy depressions and carr in pastures between disused
railway and river (SN/341.408), 30 maz.s.1., c. 1000 plants, including many non-flowering rosettes
(number uncertain because of presence of R. palustris and consequent doubt about identity of
vegetative rosettes), 23 August 1993, A. O. Chater and T. C. G. Rich (NMW).

9. Aberbachnog, 1-7 km E. of Henllan, by pond south of disused railway and in hollows in the
pasture between here and river (SN/374.402), 40 m a.s.l., c. 300 plants, 24 August 1993, A. O.
Chater and T. C. G. Rich. There is a specimen from the pond collected on 1 September 1979,
A. O. Chater in NMW.

10. Lampeter, 500 m N. of the college, marsh by pool in waste ground and on adjacent soil tips (SN/
579.488), 125 m a.s.l., c. 80 plants in marsh and c. 1500 plants on tips including many non-
flowering rosettes, 4 September 1993, A. O. Chater (NMW). Some plants on the tipped soil were
over 1m in diameter, and the ground beneath the prostrate stems was orange with the shed seed.

A number of other sites where it was felt that R. islandica might occur on the Teifi were visited in
1993 without success, and these are listed as grid references in Table 1. Although negative records
are of lesser value, this information may help in any future survey of the species to indicate whether
there has been any change in the distribution on the river.

TABLE 1. AFON TEIFI SITES SEARCHED WHERE RORIPPA ISLANDICA WAS NOT FOUND.
(SITES WHICH SEEMED PARTICULARLY SUITABLE ARE INDICATED BY AN ASTERISK)

SN/154.484* SN/454.401*

SN/182.443 SN/518.435* — 516.439
SN/184.449 SN/552.462* — 581.476*, many suitable sites
SN/225.432* SN/597.484

SN/227.434 SN/606.495 — 616.499*

SN/235.414 SN/617.501

SN/256.416 SN/666.580*

SN/256.422*
SN/296.418*
SN/391.396*

SN/407.403 — 420.411
SN/429.419* — 425.418

SN/673.589*
SN/672.612
SN/683.626
SN/709.664

232 A. O. CHATER AND T. C. G. RICH

Rorippa palustris was also found at three additional sites without R. islandica (voucher specimens
in NMW). These sites were Pont Ceri (SN/296.418), 700 m E. of Dolaugwyrddon-Isaf (SN/564.465)
and 400 m S.S.E. of Dolaugwyrddon-Isaf (SN/559.463).

Anglesey, v.c. 52: |

Jonsell (1968) suggested that ‘‘a constant occurrence (1813-1890s) at a small lake in Anglesey
(Griffith 1895) may be this species” but this was a result of a misunderstanding of Griffith’s text;
Davies and Griffith recorded the plant at different sites. Davies (1813) noted ‘“‘Sisymbrium terrestre
.. . In the clay pits, at Beaumaris, Cors Ddygai, not rare’’, and Griffith (1895) gives ‘‘Nasturtium
palustre DC... . Very rare . . . On the margin of Llyn Frogwy, Bodforth!” R. H. Roberts looked
for R. islandica in the 1970s without success (pers. comm., 1993), and another survey in 1993 by him
of all known sites for R. palustris (including Llyn Frogwy) was equally unsuccessful. R. islandica has
not yet been confirmed for Anglesey.

IDENTIFICATION AND LIFE CYCLE

The field work provided an opportunity to review the identification characters given in Rich (1991).
Plants of R. islandica were often decumbent to ascending at the base, whilst R. palustris was usually
more or less erect, but plants of either species in trampled places could be ascending. Leaf shape was
quite variable, and it was impossible to identify vegetative rosettes with confidence. There was
considerable overlap in flower size between both species, and the floral characters are not very
reliable. The presence of a short-petalled clone of R. sylvestris (L.) Besser on the Teifi also caused
some confusion, though the regularly pinnate leaves, long creeping rhizomes and linear ovaries of
this species are otherwise distinctive.

The fruit and seed characters, however, proved a reliable means of distinguishing the plants,
which could be done from several metres’ distance with some experience. The most striking
morphological feature of the R. islandica plants on the Teifi was that the fruits were secund and
swept downwards so that the whole inflorescence was one-sided, and also quite dense due to the
relatively shorter pedicels. How reliable this character is across the range of the species is unknown,
but it appears diagnostic in Wales. In contrast, the fruits of R. palustris were arranged evenly around
the stem and were not so densely crowded.

With practice, the fruit shapes could be seen to be quite different, with R. islandica having more
rectangular, square-ended fruits. When seeds of both species were examined side by side, those of
R. palustris were darker in colour, slightly larger and the epidermal cells could just be made out with
a X20 lens (the epidermis of R. islandica appearing comparatively featureless). Identification should
always be confirmed from microscopical examination of ripe seeds and comparison with voucher
material. The size of the epidermal cells is diagnostic, although it is difficult to quantify this because
of variation on different parts of the seed.

Rorippa islandica and R. palustris have previously only rarely been found growing together in the
same place (e.g. in Ireland, Goodwillie 1994). One plant was found at The Moat (Site 8) which was
partially sterile (NMW), but this is more likely to be due to environmental factors than hybridisation
since both species are habitually in-breeding. Jonsell (1968) failed to obtain any progeny from
experimental crosses.

Rorippa islandica is usually considered an annual in the wild (e.g. Rich 1991), although Jonsell
(1968) was able to keep some plants in cultivation for up to five years. In August and September
1993 we noticed that many plants (more than half the population at some sites) had not commenced
flowering, and we assumed that these would flower later in the autumn. On revisiting these sites in
early November 1993, there was still no sign of inflorescences on most of these rosettes, and they
were still in the same condition in March 1994. Furthermore, many plants which had fruited in 1993
bore well-developed central rosettes through to March 1994. Several plants were also seen in
November 1993 which had axillary rosettes of leaves on decaying fruiting stems which were about to
break off from the parent plant to act as vegetative propagules (this phenomenon occurs
occasionally in other crucifers, cf. Rich (1984), and Grime et al. (1988) for its occurrence in R.
palustris). In its Teifi sites R. islandica therefore often behaves as a biennial, and possibly at least
sometimes as a perennial, although further field studies with marked plants are needed to confirm

RORIPPA ISLANDICA IN WALES Pa)

the perennial habit. Goodwillie (1994) mentions phenological plasticity in R. islandica as probably
fitting the plant for growth in turloughs where the unpredictability of the water level is the major
environmental variable. The plasticity of life cycles in response to environmental conditions is well
known in other crucifers (Rich 1991).

HABITATS OF RORIPPA ISLANDICA ON THE TEIFI

The Afon Teifi is about 115 km long and rises at 550 m a.s.1. in acidic moorland. When it has fallen to
160 m a.s.1., it meanders slowly for about 8 km through the extensive raised mire complex of Cors
Caron, through beds of silts and clays. For much of the rest of its length, and especially in its lower
half, the river is relatively fast-flowing and rocky. This is a reverse of the usual pattern, in which
rivers have their faster, rocky section above the slower, meandering ones. The Teifi is also unusual
in having no large tributaries. The Cors Caron stretch thus has an unexpectedly dominating effect on
the rest of the river (Holmes 1983; Chater 1994). The underlying rocks of the whole river system are
Ordovician and Silurian shales and mudstones and are largely acidic. Sites 4-9 are situated in the
lower third of the river, and site 10 is three-fifths of the way up the river.

Six of the eight sites on the Teifi, nos. 2, 4,5, 7, 8 and 9, are depressions in the floodplain, within
300 m of the river but separated from it. Four of these depressions originated as meanders or ox-
bows which have become silted up and two still contain areas of more or less permanent open water
adjacent to the Rorippa populations. Site 4 is a long-disused and largely silted-up canal, and site 8 is
a meander anciently modified into a moat, half of which is now silted up. All six sites are regularly
flooded, usually at least once a year and often for some days at a time, when the whole of the
floodplain is inundated. These floods can occur at any time of the year, though rarely in summer. All
six sites are on fine, silty, gley soils, probably mostly of the Conway series (Bradley 1980; Rudeforth
et al. 1984) in situations where they form part of the Teme Association.

Soil pH was measured for each site from one fresh soil sample collected from the top 5 cm of the
soil profile (i.e. the root zone). The samples were mixed with a similar volume of de-ionised water to
form a paste and allowed to stand for about five minutes. pH was measured using a Piccolo ATC pH-
meter (Hanna Instruments Inc.) with a HI 1280 amplified electrode. The soil pHs measured ranged
from 5-2 to 6-6 (Table 2).

Site 6 seems to be unique for R. islandica in that it is a gently sloping platform of rock c. 1 m above
normal water level in a rocky section of the river. The plants grow in crevices in the rock and are very
dwarfed, and some are adjacent to patches of concrete placed to smooth the platform. It is
frequently flooded, and the pH of the soil in the crevices is 6-9, possibly influenced by the concrete.
The site used to be used for sheep-dipping and is now much used by fishermen and picnickers. Our
initial impression that this habitat was unusual has thus been confirmed by further survey.

Site 10 is just west of a tributary, the Afon Dulas, 1-1 km above its confluence with the Teifi. Until
about ten years ago it was a wood yard but is now marshy ground around a shallow pond. The
ground is patchy and partly open and parts of it, including the area with R. islandica, are usually
flooded for much of the winter. Flooding is due to poor drainage and the site, unlike the other seven,
is not part of the Teifi floodplain although it is within the catchment. The pH here averaged 7-6, the
substrate including timber, tarmac, stones and other imported material. Along the south side of the

TABLE 2. SOIL pH VALUES FROM RORIPPA ISLANDICA
SITES IN THE TEIFI VALLEY, WALES

Site 2 pH 6-0
Site 4 pH 6-6
Site 5 pH 6:5
Site 6 pH 6:9
Site 7 pH 5-2
Site 8 pH 5-6
Site 9 pH 5:3
Site 10, marsh pH 7-6

Site 10, tip pH 8-2

234 A. O. CHATER AND T. C. G. RICH

marsh is a large area of soil and rubble of unknown origin, illegally tipped here in early 1992. The pH
of this by the main colony of R. islandica here is 8-2. 100 m to the south-east is another equally recent
soil tip, the material having come from excavation on the site; R. islandica is by far the most
abundant plant on this tip. R. islandica must be a recent arrival at this site, as there was no
vegetation on it until a decade or so ago. It is impossible to guess whether it arrived first in the
marsh, where it could have been brought by birds (Mute Swans nest on the pond), or whether it
arrived with the illegally dumped soil and spread from this to the adjacent tip and to the marsh. R.
islandica has also been reported from rubbish tips in the Isle of Man (L. S. Garrad, pers. comm..,
1989).

VEGETATION

Species associated with R. islandica at the eight Teifi sites are listed in Table 3 (at site 10 only the
marsh part of the site is included). It should be understood that the species lists in Table 3 are not to
be interpreted as floristic tables for the communities present, but are simply lists of species growing
within 1 m of R. islandica plants. The most constant associates were Persicaria hydropiper,
Gnaphalium uliginosum and Callitriche brutia, all also annuals of disturbed open, damp ground.

The vegetation of these sites falls into two main types which can be described in terms of British
plant communities (Rodwell 1991, et seq.). The drier sites support Agrostis stolonifera — Alopecurus
geniculatus grassland (MG13), which is characterised by constant Agrostis stolonifera and Alope-
curus geniculatus, with Ranunculus repens, Holcus lanatus, Poa trivialis, Juncus effusus and Glyceria
fluitans. It occurs in seasonally inundated places, on poorly drained, clayey soils, often in grazed and
trampled conditions, and the sward is usually short and fairly open. It is not uncommon throughout
most of lowland Britain. This was the most characteristic vegetation type for R. islandica which was
very abundant in it at sites 2, 7, 8, and 9.

The British plant communities frequency table for Agrostis stolonifera — Alopecurus geniculatus
grassland appears to underestimate the frequency of the annuals characteristic of these muddy,
trampled sites (e.g. Persicaria hydropiper, Gnaphalium uliginosum), both in western Wales and
elsewhere. An additional sub-community of the Agrostis stolonifera — Alopecurus geniculatus
grassland, characterised by Persicaria hydropiper, might be justified.

The wetter sites support Glyceria fluitans swamp, Alopecurus geniculatus sub-community (S22c),
which occurs in shallow, seasonally-inundated water on nutrient-rich soils, usually with a fine
mineral substrate. It is often grazed and is frequent in ditches in fields, dry ditches and along the
margins of small streams. It is dominated by Glyceria fluitans and Alopecurus geniculatus, with a few
other frequent species such as Rumex crispus and Poa trivialis. It is widespread and common in the
agricultural lowlands of Britain. R. is/andica occurred in this vegetation type at sites 4, 5 and 8, and
in the first two is at a much lower frequency than in the MG13 grasslands.

The Cenarth site (Site 6) supports sparse vegetation which is best treated as MG7 Lolio-
Plantaginion, but can scarcely be considered a typical example of the vegetation.

The vegetation at site 10, disturbed and of recent origin, is probably best considered as M23
Juncus effusus/acutiflorus — Galium palustre rush-pasture, but the R. islandica grows in sparsely
vegetated ground uncharacteristic of the community as a whole.

DISPERSAL

Rorippa islandica is easily overlooked, both because of its inconspicuous nature and because of the
difficulties of identification, and, more than for many other species, absence of records should not
be taken to indicate absence of the species. The earliest record from the Teifi is from Cenarth in 1958
(Site 6 or nearby), and the next is from Aberbachnog in 1979 (Site 9); it is still present at both of
these sites. It was first found at the others in 1993. The only certain comment that can be made about
the history of the populations is that the one at site 8 must be less than about a decade old. Sites 2
and 4~9, all in the lower half of the river, are on the floodplain where seeds could easily have been
transported by regular flooding.

Unlike many other crucifers, the epidermal cells of the seeds of R. palustris and R. islandica do

RORIPPA ISLANDICA IN WALES Mp)

TABLE 3. SPECIES GROWING IN THE SAME COMMUNITIES AS AND WITHIN 1 M OF RORIPPA
ISLANDICA AT EIGHT SITES IN THE TEIFI VALLEY, WALES

Site no.

No}

Associated species 2 - 5 6 7 10 (marsh)
Agrostis capillaris ate
A. stolonifera +
Alisma plantago-aquatica +
Alnus glutinosa

Alopecurus geniculatus =F
Apium nodiflorum

Bidens cernua +
B. tripartita +
Callitriche brutia +
C. stagnalis

Cardamine flexuosa

C. pratensis

Carex remota

C. vesicaria +

Chenopodium rubrum =P

Eleocharis palustris +
Equisetum arvense +

E. palustre +
Festuca rubra +

Filipendula ulmaria
Galium palustre +
Glyceria declinata

G. fluitans ae
Gnaphalium uliginosum +
Juncus articulatus

J. bufonius

J. effusus aa
J. kochii

Lemna minor

Lysimachia nummularia
Lythrum portula

L. salicaria + +
Matricaria discoidea + +

Myosotis laxa +
M. scorpioides +

Oenanthe crocata
Persicaria amphibia
P. hydropiper

P. maculosa
Phalaris arundinacea 4p “F +

Plantago major sF +
Poa annua +
Potentilla anserina

Ranunculus flammula

R. repens

R. sceleratus +
Rorippa islandica +
R. palustris

Rumex obtusifolius

Sagina procumbens

Salix cinerea

Sparganium erectum

Stellaria media

Taraxacum sp.

Trifolium repens

Typha latifolia +
Urtica dioica sP

Veronica beccabunga +

+

+ ++ |
+

++ + +

++++4++ 44+ +

+ +++
+ ++ + +44 +
+
+ +4

++
+
++ + +44
+ +
+
+ ++ +
+ +44 ++

+++ +
+
+
+
+
+

+++ 44
+

236 A. O. CHATER AND T. C. G. RICH

not produce mucilage, but are hollow (Jonsell 1968). This may have the advantage of making the
seeds buoyant, facilitating dispersal in water, and enabling the seeds to be drawn up in water films
as, for instance, by birds as they leave the water. Ridley (1930) reported the presence of R. palustris
in many places by the River Thames at Kew, where its seeds can only have been carried by water, |
and he also found that seeds put in water floated for 24 days. The possibility of dispersal of R. i
islandica in water was tested by a simple experiment. :

FLOTATION EXPERIMENT

Seeds of R. islandica and R. palustris were collected from ripe fruits of plants in cultivation on 7
September 1993. Batches of 20 seeds were floated on 200 ml of tap water in glasses, and the numbers
of seeds floating were counted at intervals for two months. The water was agitated once each day by
shaking the glass enough to cause the seeds to dip below the surface. The influence of surface
tension was tested by adding two drops of household detergent (Fairy Liquid). Seeds in water
floated in a group in the middle of the glass, whilst those in detergent lined up around the margin of
the glass, as would be expected from the shape of the water meniscus. Seeds in detergent initially
sank after agitation, but rose to the surface again after a minute or so. The results are shown in Table
4, and indicate that seeds of both species are buoyant in water for substantial periods of time. In
water with detergent, it is possible that the larger epidermal cells of R. palustris are more easily
infiltrated with water thus reducing their buoyancy.

Transport of seeds of R. islandica down the river by floods (which often occur in autumn when the
seeds are ripe) would thus seem a strong possibility, although ending up in an appropriate habitat is
purely a matter of chance. The dispersal of plants such as Impatiens glandulifera along. river
corridors is well known, and Claytonia sibirica and Rorippa austriaca are currently spreading on the
Afon Tywi and Afon Loughor respectively in v.c. 44 (G. Hutchinson, pers. comm., 1993).

Seeds floating on the surface of the water are easily drawn up in a film of water when an object
(e.g. a finger) is immersed and lifted out. It is possible to imagine the same type of mechanism being
involved with birds’ feet or plumage as they leave water.

BIRD DISPERSAL
Jonsell (1968) suggests that dispersal of seeds by geese is possible for R. islandica and points out the
connections between its distribution and the migration routes of geese. Similar ideas have been put
forward for other species (e.g. Heslop-Harrison 1953; Gornall 1987), and evidence for long-distance
dispersal of seeds in the digestive systems of geese to Britain is given by Welch (1993). He collected
droppings from where skeins of Pink-footed Geese migrating from Iceland first landed, and placed
them in a moderately heated greenhouse on sterile compost, some immediately and some after a few
months storage in a deep freeze. Seedlings which germinated immediately often died, perhaps due
to the toxic conditions of the droppings, but in subsequent years some seedlings were successfully
raised. Seeds may also be dispersed by mud on birds, and Kerner (1903; see also Ridley 1930)
reports finding seeds of R. palustris and many other species, in mud from the feet, beaks and
feathers of Swallows, Snipe, Wagtails and Jackdaws.

Jonsell’s suggestion is especially worth considering in the case of the R. islandica populations on

TABLE 4. NUMBERS OF RORIPPA SEEDS (OUT OF 20) REMAINING AFLOAT DURING A
FLOTATION EXPERIMENT

R. islandica R. palustris

Days from i tel oee eeeee WEEE
start of Water + Water +
experiment Water Detergent Water Detergent

0 20 20 20 20

1 20 20 20 17
13 20 20 20 15
15 20 20 20 14
23 20 20 20 0

61 AN 20 20 0

RORIPPA ISLANDICA IN WALES 237

the Afon Teifi. The Greenland White-fronted Goose (Anser albifrons flavirostris) breeds in west
Greenland, and winters exclusively in Ireland, west Scotland and Wales (Fox & Stroud 1985). A
flock wintered on Cors Caron, the raised mire complex on the upper reaches of the Teifi, from at
least the 1890s (when it probably consisted of c. 30 birds) until the 1960s. Numbers had risen to 250-
300 by 1940, to 400 by 1957, and to 550-600 by 1962. A catastrophic decline followed, and the geese
ceased to winter regularly there after 1967/8 (Fox & Stroud 1985). R. islandica does occur in west
Greenland but is rare there, but of much greater significance is the fact that many of these geese
break their autumn migration to the British Isles on the west and south coasts of Iceland (Cramp
1977). One of the two main areas for R. islandica in Iceland is in Arness Sysla in the south west
(Jonsell 1968) and dispersal from here by migrating geese at a time when seed was ripe could explain
how the seed came to be present in the Teifi river system. R. islandica has not yet, however, been
found anywhere on the Teifi above site 10 at Lampeter, which is some 15 km below the former main
wintering grounds of the geese.

Greenland White-fronted Geese also wintered further north in Cardiganshire throughout the first
half of the present century, including Cors Fochno on the south side of the Dyfi estuary, and since
the 1950s flocks of about 80-250 birds have been wintering regularly on this estuary. Neither R.
islandica or R. palustris have been recorded from this area. Elsewhere in Wales the only regular
wintering flocks that might have been expected to be involved in seed-dispersal direct from Iceland
or Greenland have been in Montgomeryshire and perhaps Anglesey. Small flocks and casual
records have been made in many places, including the lower Teifi, and on the Tywi in
Carmarthenshire (Fox & Stroud 1985).

If Gornall’s (1987) hypothesis that Ranunculus reptans is dispersed to the British Isles mainly by
Pink-footed Geese, Mallard or Wigeon is correct, then one might predict that R. islandica, which
grows in similar habitats and places to Ranunculus reptans at least in Iceland, might be expected to
show some similarities in distribution in Britain. However, the distributions of Rorippa islandica
and Ranunculus reptans in Britain do not overlap, and only Rorippa palustris (which is absent from
Iceland) has been recorded from Loch Leven, the classic site for Ranunculus reptans.

DISCUSSION

The recent discoveries in Ireland (Goodwillie 1994) and the Welsh data presented here indicate that
R. islandica has been widely overlooked, and it should be looked for. elsewhere. It would be
Surprising if the species was not as widespread on the catchments of the Loughor and Bran/Tywi as it
is along the Teifi. We would welcome further records supported by voucher specimens composed of
a few ripe fruits.

The Afon Teifi and the Irish turloughs appear to be the major stronghold for the species in north
west Europe. The Scottish localities are all scattered and isolated and support small populations. It
is somewhat erratic in appearance in the Isle of Man. Whilst there is no definitive evidence, it is
likely that R. islandica is not a new arrival in Wales. Its local abundance would seem to be associated
with dispersal by water, and it is possible that the wider distribution is related to dispersal by geese.

The occurrence along a river system is also unusual. In Ireland it occurs in turloughs and ditches
near the sea. In Scotland it occurs in small ponds, damp ground and loch margins. In Iceland it
occurs around lakes and small bodies of water, moist soil and also around hot springs. In Greenland
it occurs around small lakes and temporary ponds, as weil as on or beside paths. In Norway it occurs
in small coastal pools, ditches and potato fields. In central Europe it occurs on the shores of alpine
lakes (Jonsell 1968).

ACKNOWLEDGMENTS

We are grateful to R. Gwynn Ellis and George Hutchinson, National Museum of Wales, for loan of
specimens, Bengt Jonsell for confirming the Cenarth material, Dick Roberts for help with the
Anglesey records, Richard Pryce and John Killick for help with the Carmarthen records, Roger
Goodwillie for an advance copy of his paper, Lin Gander for telling us of the Lampeter site, and

238 A. O. CHATER AND T. C. G. RICH

Chris Preston and the Biological Records Centre for the updated distribution map and comments on
the draft manuscript. We would also like to thank the local landowners for access.

REFERENCES

BRADLEY, R. I. (1980). Soils in Dyfed. Sheet SN24 (Llechryd). Soil survey record no. 63. Soil Survey of England
and Wales, Harpenden.

Cuater, A. O. (1994). The higher plants and vegetation of Cardiganshire, in Jones, I. G., ed., Cardiganshire
county history 1. University of Wales Press, Cardiff.

Cramp, S., ed. (1977). Handbook of the birds of Europe, the Middle East and North Africa: The birds of the
Western Palearctic 1. Oxford University Press, Oxford.

Davies, H. (1813). Welsh botanology. W. Marchant, London.

Fox, A. D. & Stroup, D. A. (1985). The Greenland White-fronted Goose in Wales. Nature in Wales n.s. 4: 20-
Dik.

GoopwiLuiE, R. (1994, in press). Irish distribution of Rorippa islandica (Oeder ex Murray) Borbas. Irish
Naturalists’ Journal 24.

GorRNALL, R- J. (1987). Notes on a hybrid spearwort, Ranunculus flammula L. X R. reptans L. Watsonia 16: 383—
388.

GriFFITH, J. E. (1895). The flora of Anglesey and Carnarvonshire. Nixon & Jarvis, Bangor.

Grime, J. P., Hopcson, J. G. & Hunt, R. (1988). Comparative plant ecology. Unwin Hyman, London.

HEsLop-Harrison, J. (1953). The North American and Lusitanian elements in the flora of the British Isles, in
Lousley, J. E., ed. The changing flora of Britain, pp. 105-123. Botanical Society of the British Isles,
London.

Homes, N. T. H. (1983). Typing British rivers according to their flora. Focus on nature conservation 4. Nature
Conservancy Council, Shrewsbury.

JONSELL, B. (1968). Studies in north-west European species of Rorippas. str. Symbolae botanicae Upsaliensis 19:
1-222.

KERNER VON MariLaun, A. (1903). The natural history of plants, translated by Oliver, F. W. The Gresham
Publishing Company, London.

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

Ricu, T. C. G. (1991). Crucifers of Great Britain and Ireland. B.S.B.1. Handbook no. 6. Botanical Society of the
British Isles, London.

Riptey, H. N. (1930). The dispersal of plants throughout the world. L: Reeve & Co., Ltd, Ashford, Kent.

RopwELL, J. S., ed. (1991 et seq.). British plant communities. Cambridge University Press, Cambridge.

RuDEFORTH, C. C., HARTNUP, R., LEA, J. W., THompson, T. R. E. & WriGut, P. S. (1984). Soils and their use in
Wales. Soil survey of England and Wales Bulletin No. 11. Soil Survey of England and Wales, Harpenden.

Stace, C. A. (1991). New Flora of the British Isles. Cambridge University Press, Cambridge.

WE LcH, D. (1993). A check on the transmission to Britain of creeping spearwort by pink-footed geese. B.S. BI.
Scottish Newsletter 15: 15-16.

(Accepted April 1994)

Watsonia 20: 239-254 (1995) 239

A biometric survey of Limonium vulgare Miller and
L. humile Miller in the British Isles

H. J. DAWSON and M. J. INGROUILLE

Biology Department, Birkbeck College, Malet Street, London, WC1IE 7HX

ABSTRACT

Limonium vulgare Miller and L. humile Miller (Plumbaginaceae) are closely related species that grow on
saltmarshes around the coasts of northern Europe including the British Isles. They are often found together in
the same marsh. Morphometric analyses were performed on plants collected from sites around the British Isles.
This demonstrated the close relationship of these two species and provided strong evidence for hybridization and
introgression in sites where both species were present.

KeEyworps: hybridization, introgression, self-incompatibility, population, speciation.

INTRODUCTION

Limonium vulgare Miller and the closely related L. humile Miller, of the family Plumbaginaceae,
are found in saltmarshes around the coast of Ireland and Britain northwards to southern Scotland.
They have been recognised as distinct species from the time of Ray (1724) as Limonium majus
vulgatius and Limonium Anglicum minus, caulibus ramosioribus, floribus in spicis rarius sitis.
L. vulgare has short spikes with densely arranged spikelets and L. humile is more laxly branched
with long spikes and spikelets distant from each other (Salmon 1905a & b). L vulgare does not occur
on the coast of Ireland, where it is replaced by L. humile. Both occur on the coasts of north-western
continental Europe. They grow mixed together in some marshes but L. humile is more frequent in
the lower marsh than L. vulgare (Boorman 1966, 1967).

The difficulties of the identification of the two species have been discussed by Dawson (1988). The
confusion in identification between L. humile and L. vulgare has led to misleading reports and
records from some areas and may have obscured the true distribution of both species across the
British Isles. This confusion has arisen because of the high degree of variability in L. vulgare which
has, in the past, led to the description of a wide variety of forms and variants. The variability is not
environmentally determined as was thought previously (Clapham 1987). Boorman (1966) used
comparative cultivation and transplant experiments to demonstrate that some of the variation is
genotypic in origin. Genotypic variation is emphasised in marshes because of the extent of clonal
reproduction by rhizomatous spread in L. vulgare. Large homogeneous patches with distinct
morphologies are found growing next to each other.

The species differ for the most part in chromosome number, L. vulgare is a tetraploid 2n = 36 and
L. humile a hexaploid 2n = 54 (Dawson 1990a). There is aneuploid variation in L. humile and some
plants have 2n = 36, like L. vulgare.

The species possess different breeding systems; L. vulgare is an obligate outbreeder and L.
humile is a facultative inbreeder. Outbreeding in L. vulgare is enforced by a self incompatibility
system accompanied by dimorphism of pollen and stigmas, with either ‘A’ pollen (coarsely
reticulate) and Cob stigma or ‘B’ pollen (finely reticulate) and Papillate stigma. The A/Cob morph is ©
heterozygous (AC.ac) and the B/Papillate morph homozygous (ac.ac). L. vulgare is also slightly
heterostylous, although this character is very variable. L. humile is a self-compatible homostylous
monomorphic species with an ‘A’ Papillate morph with the stigma compatible to B pollen as well as
A pollen (Ac’.Ac'). The monomorphic condition is found rarely elsewhere in the genus, and is also
found in the related genus Armeria (Miller) Willd. and is always derived from the normal dimorphic
system (Vekemans et al. 1990).

240 H. J. DAWSON AND M. J. INGROUILLE

The two species, L. vulgare and L. humile, provide excellent material for the examination of
speciation and the origins and outcomes of different breeding systems.

SAMPLING AND METHODS

Plants were sampled from 52 sites across Britain and Ireland (Fig. 1) some of which had populations
of both species present. Details of sample sites are recorded in Dawson (1990b). Material was
sampled from sites to represent the whole range of variation present. Some populations were either
very small and/or uniform and could be effectively surveyed with a small sample. Different numbers

East Anglia

Brifain

Figure 1. Geographicai distribution of sampled sites of Limonium spp. (Population codes for British Isles as for
O.T.U.s in Fig. 6).

LIMONIUM VULGARE AND L. HUMILE IN THE BRITISH ISLES 241

of plants were collected at different sites. For the main survey of variation in the British Isles 580
plants were sampled and scored.

For the purpose of analysis, where a large number of populations were sampled, a number of
geographical regions were circumscribed; Ireland, North West Britain (Scotland, Cumbria and
Lancashire, North Wales), South Wales, the Solent Region, and East Anglia (Essex, Suffolk,
Norfolk and Lincolnshire).

Plants from each site were first pressed and dried. Flowering spikes were removed from each plant
on collection, labelled correspondingly and preserved separately in 70% ethanol.

The difference in pollen and stigma morph combination has been used to allocate plants to species
in this work: A/Papillate = L. humile, A/Cob and B/Papillate = L. vulgare. This is however
complicated by the mutated morphology of the Papillate stigma in L. humile plants, which is
variable and somewhat intermediate between the Cob and Papillate morphology of the stigmas in L.
vulgare (Dawson 1990b). There is also some variation in pollen morphology. Variation in stigma
and pollen will be reported elsewhere. Pollen and stigma characters and those associated with
heterostyly were not used in the multivariate analyses to investigate similarity.

An initial study was made on plants from two sites, chosen because they provided large, pure
populations of each of the species: L. vulgare from Oxwich, Gower (v.c. 41, SS/514. 877) and L.
humile from Dale, Dyfed (v.c. 45, SM/812.070). After this study an initial set of 33 scored characters
was reduced to exclude those characters which proved difficult to measure accurately.

Following the methods described in Tabachnick & Fidell (1989) a number of characters were
eliminated because of very high correlations with another character. Very high correlations may
arise because of the logical correlation of characters. For example the iength of secondary branches
and the length of primary branches were very highly correlated (r>0.90) so that length of primary
branches only has been included as a measure of branchiness. The length of secondary branches
then becomes a superfluous character. In this situation the inclusion of superfluous characters can
weight particular aspects of the morphology.

A set of characters which were highly correlated are branch length, spike length, number of
spikelets and the distance between the lowest two spikelets. This is a general relationship perhaps
relating to the potential to lengthen internodes. However in previous work it has been clear that
Limonium species vary independently in these characters (Ingrouille 1984; Ingrouille & Stace
1985). For example, some species have an uneven distribution of spikelets on the spike with a
relatively large distance between the first two spikelets. Others with a similar length of spike have an
even distribution of spikelets on a spike.

Other high correlations between characters in the data set were between scape height and other
vegetative measurements. It was clear that there was an overall size effect. Nevertheless scape
height was clearly an important distinct character from overall size, one aspect of the ‘gestalt’ of the
plant. A striking feature of variation on marshes is clones of markedly different ‘gestalt’, growing
adjacent to each other (Dawson 1990b) and others with a similar ‘gestalt’ but differing in overall
size. Scape height is only one aspect of the ‘gestalt’ but should nevertheless be included.

The inclusion of composite variables such as ratios between characters as well as the characters
used to construct them may lead to inflated correlations (Tabachnick & Fidell 1989). In this work
this has been avoided so that outer-bract length and outer bract width were included but not a ratio
of outer-bract length to width. However this general rule has been relaxed in the analysis of ‘leaf
shape’ which is a ratio of leaf length to leaf width. Leaf width, although significantly correlated to
leaf shape, has also been included as a measure of maximum leaf size. In an examination of clones it
was clear that leaf shape varied in two distinct ways: by the maximum size of leaf achieved by any
clone and also by the relative narrowness of the leaf. Leaf length has not been included separately in
the analysis.

Significant correlations between the characters remaining in the data set were the rule but it was
neither possible nor desirable to try to abstract a set of characters which were not significantly
correlated to each other. Significant correlations occur in a complex multidimensional way. In this
large data set with just two species, significant correlations between characters were expected
because particular sets of character measurements were associated with particular taxa or variants.
It would not be desirable, in an analysis designed to measure the extent of relationship between
taxa, a priori to limit the range of characters used on the basis of the presumed distinctiveness of
those taxa.

242 H. J. DAWSON AND M. J. INGROUILLE

It is not the level of statistical significance of correlations which is important but the actual level of
the correlation. A data set with many very high correlations (say r>0-8) lacks information content.

For the major survey 13 metric characters were used in the multivariate analysis. (Table 1, Fig. 2).
There were no missing data. Correlations between these characters are reported in Table 2. The
data were screened for the presence of outliers and non-normality and following the recommen-
dations of Tabachnick & Fidell (1989) those characters which had high levels of skewness or kurtosis
were transformed to their natural logarithmic value (Table 1). A few outliers were recoded as
maximum or minimum values within the normal range. Characters were transformed to Z-scores.

Several different univariate and multivariate analyses were performed. Hierarchical analysis of
variance of variables for plants within populations within regions was carried out to compare the
patterns of variability in species. Multivariate analyses included discriminant function analysis,

Ficure 2. Metric characters of Limonium spp. used in the multivariate analyses (see Table 1).

LIMONIUM VULGARE AND L. HUMILE IN THE BRITISH ISLES 243

TABLE 1. METRIC CHARACTERS OF LIMONIUM SPP. USED IN MULTIVARIATE ANALYSES

Transformed No. of outliers*
Character (natural log) recoded of 580 cases
1. Scape height yes 0
2. Height to first branching node yes 6
3. Primary branch length yes 1
4. Leaf shape yes 0
5. Leaf width yes Z
6. Branch angle no 0
7. Spike length yes 1
8. Spikelet distance yes py
9. Number of spikelets per spike yes 1
10. Outer bract length yes 2
11. Outer bract width yes 1
12. Calyx length no 0
13. Corolla length no 2

* An outlier has a Z score >3.00.

TABLE 2. CORRELATION MATRIX BETWEEN CHARACTERS OF LIMONIUM SPP.
Probability in parentheses (n = 580)

Characters
1 2 3 4 5) 6 7 8 9 10 11 12
2 —0-60
(0-99)
3 0-64 —0-10
(0-00) (0-02)
4 0-07 —0-15 0-12
(0:09) (0-00) (0-01)
5 0-50 0-02 0-19 —0-09
(0:00) (0-55) (0-00) (0-03)
6 0:26 —0-02 0-17 0-08 —0-47
(0:00) (0-69) (0-00) (0-05) (0-00)
7 0-21 —0-30 0-49 0-21 —O0O-11 0-16
(0-00) (0-00) (0-00) (0-00) (0-01) (0-00)
8 O9e 012" 0-08 —0-03. | 0-25 0-03 0-48
(0-00) (0:00) (0:07) (0-53) (0:00) (0-49) (0-00)
9 0-19" —0-24 0-57 0-28 —0:-25 0-18 0-80 0-02
(0-00) (0-00) (0-00) (0-00) (0-00) (0-00) (0-00) (0-58)
10 0-18 —0:04 0-29 0-06 0-08 0-01 0-41 0-22 0-36
(0-00) (0-29) (0-00) (0-17) (0:06) (0-87) (0-00) (0-00) (0-00)
11 0-02 —0O-11 0-24 0-20 —0-12 0-02 0-55 0-15 0-54 0-63
(0-64) (0-01) (0-00) (0:00) (0-01) (0-56) (0-00) (0-00) (0-00) (0-00)
12 0:10 —0:12 0-33 0-17. -—0:04 -0-01 0-37 —0-12 0-47 0-45 0-43
(0:02) (0-01) (0-00) (0-00) (0-31) (0-81) (0-00) (0-00) (0-00) (0-00) (0-00)
13 0:06 -—0:03 0-01 -—0-02 0-27 -—0-17 —-—0-02 0-08 -—0-10 0-20 0-07 0-28

(0:15) (0-44) (0-88) (0-63) (0-00) (0-00) (0-00) (0-55) (0-05) (0-02) (0-08) (0-00)

1=Scape height; 2=Height to first branching node; 3=Primary branch length; 4=Leaf shape; 5=Leaf width;
6=Branch angle; 7=Spike length; 8=Spikelet distance; 9=Number of spikelets per spike; 10=Outer bract
length; 11=Outer bract width; 12=Calyx length; 13=Corolla length.

244 H. J. DAWSON AND M. J. INGROUILLE

principal component analysis (P.C.A.) and cluster analysis on individuals and/or population means
of British and Irish material. Cladistic methods were deemed unsuitable for the analysis of
continuously varying characters.

Each multivariate method simplified the complex data set in different ways. Discriminant analysis
was used to assess the level of intermediacy (hybridization/introgression) between species. P.C.A.
and cluster analysis were used to detect patterns without any a priori allocation of plants to species.
P.C.A. was carried out with VARIMAX rotation of the axes to maximise the interpretation of
components in terms of the original characters. Cluster analysis was carried out on population
means as O.T.U.s using both transformed character scores and also factor scores of factors with an
eigenvalue greater than one. Several different measures of distance and clustering methods were
tried including Ward’s method of minimum variance clustering after the calculation of Squared
Euclidean Distance.

Analyses were carried out using SPSSX (Norusis 1985) and NTSYS-pe (Rohlf 1990) statistical
packages where appropriate.

RESULTS

Hierarchical analysis of variance of individual characters shows a contrasting pattern of variation in
each species. An example is reported for the character corolla length in Table 3 and summarised in
Table 4 for all 13 characters. L. humile is slightly more likely to have populations within regions
more distinct than expected by chance from individual plant variation within populations, and L.
vulgare is slightly more likely to have regions more distinct than expected by chance from variation
between populations within regions. A comparison of amounts of variability which can be allocated
to each source (Table 4) shows that within populations L. vulgare plants are significantly more
variable than L. humile plants in seven characters. No characters are more variable for L. humile
within populations.

No single character or pair of characters is effective for separating species. Character distributions
of three important transformed characters are illustrated in Fig. 3.

Discriminant analysis using a combination of all characters is successful in identifying the two taxa
(Table 5, Fig. 4). An analysis of variance of discriminant scores was highly significant. Characters
highly weighted in the discriminant function include spike length and distance between the first two
spikelets, previously used to identify species and also the outer bract width and calyx length. None
of these characters, either individually or in combination effects a perfect separation of species (Fig.
3). Indeed even using all characters there is an overlap in discriminant scores, with 14 L. humile
plants and 16 L. vulgare plants allocated incorrectly. Of the 16 mis-allocated L. vulgare eleven have
the A/Cob pollen stigma combination. A statistically significant proportion (34) of the 50 L. vulgare

TABLE 3. HIERARCHICAL ANALYSIS OF VARIATION FOR COROLLA LENGTH IN

LIMONIUM SPP.
Degrees
of Sum of Mean
Source of variation freedom squares squares F-ratio
L. humile
Between regions i 966 138-00 ikes)e)
Between populations within regions 34 3075 90-44 4.34*
Between plants within populations 268 5584 20-84
Total 309 9625
L. vulgare
Between regions 8 2683 447-17 7-38*
Between populations within regions ral 1636 60-59 1-43
Between plants within populations 236 10005 42-39

Total ‘ TiN 14324

*F-ratios significant at p<0-05.

LIMONIUM VULGARE AND L. HUMILE IN THE BRITISH ISLES 245

TABLE 4. VARIABILITY OF LIMONIUM VULGARE AND L. HUMILE COMPARED;
FREQUENCY OF SIGNIFICANT F-RATIOS (AT PS0-05 LEVEL) IN 13 CHARACTERS

No. of significant
F-ratios (13 maximum)

L. humile L. vulgare
F-ratio within species
Between regions vs 6 8
between populations within regions
Between populations within regions vs s) 7
plants within populations
L. humile/ L. vulgare/
L. vulgare L. humile
F-ratio between species
Regions 0 1
Populations within regions 3 1
Plants within populations 0 7

plants with the least discriminating scores (i.e. the 50 most similar to L. humile) are A/Cob (Table
6). Mis-allocated plants and plants with small discriminant scores are found almost entirely in mixed
populations and concentrated in particular regions (Fig. 5).

Species distinctiveness differs markedly between regions (Table 7). Within species there are
significant differences between regions for discriminant score (Table 8). The most distinct L. humile
is found in the extreme north and west, in Ireland and Scotland. The most distinct L. vulgare is
found in South Wales and East Anglia. In part this is related to the relative abundance of plants of
each species in each region. It is most difficult to identify species in mixed populations in the Solent
region, where in four mixed populations there are plants which are mis-allocated and in three of
these marshes mean discriminant scores for different species are not or only just significantly
different: probabilities of a significant difference are East Head p<0-089, Itchenor p<0-050,
Chidham pS0-134. Elsewhere discriminant scores are significantly different even in mixed
populations. The Solent Region has 66% of all mis-allocated plants in the discriminant analysis.

Within regions some populations are significantly different for discriminant score. Commonly this
is related to whether the populations come from mixed sites or not. For example Cumbria/
Lancashire Ravenglass L. humile is distinct from the other populations of L. humile which are from
mixed sites. Similarly the populations of L. vulgare found growing with L. humile in the Solent
Region and Norfolk are statistically distinct from pure L. vulgare ones in the same regions.
Elsewhere, occasional individual populations can be significantly distinct. Exceptionally L. humile
plants from the far west of Ireland, where only pure L. humile populations are found have low
discriminant scores. One L. humile plant from Poulnasherry Bay in County Clare was mis-allocated
by discriminant analysis.

Cluster analysis of Squared Euclidean Distance by Ward’s method proved most effective at
clustering species separately. A small number of populations were clustered with populations of the
other species, but often close to those from the same region. There were only minor differences
between clustering using characters or factor scores. Clustering using character scores was slightly
more successful than clustering factor scores; only five populations are mis-clustered compared to
six using factor scores (Fig. 6). Only clustering using character scores is described in more detail
here.
Mis-clustered L. vulgare populations are all from mixed sites. Itchenor and Needs Ore Point L.
vulgare populations from the Solent are clustered in the major L. humile cluster in a small sub-
cluster with the populations of L. humile also from the Solent. The Holme Island L. vulgare
population clusters with the L. humile populations from its region of Lancashire and Cumbria. Two
pure L. humile populations are mis-clustered: Treaddur Bay in North Wales and Rine Point in the
far west of Ireland.

246 H. J. DAWSON AND M. J. INGROUILLE

No. of plants

Both species

Std. Dev = .45
Mean = 3.15

Spike length (transformed)

wy 120
ic
&
Q
S 100
fe)
Z
80
60
40
Both species
20
Std. Dev = .59
Mean = 3.60
0 N = 580.00
4 ad 2 e e vy v. Ss
‘ 2) i) io) 2) 2) i) io)
Spikelet distance (transformed)
no 1005
Tt
AS}
& 1404
eo) nike
fe)
Z 120
ic
80
60
40 Both species
20 Std. Dev = .21
Mean = 3.23
0 N = 580.00
= <=) e e e eo
% > C7) ‘SS i.) “~S

Outer bract width (transformed)

Ficure 3. Histograms of transformed character measures of three characters: Spike length, Spikelet distance and
Outer bract width of Limonium spp. e

LIMONIUM VULGARE AND L. HUMILE IN THE BRITISH ISLES 247

w 80;
i
&
Qa
—
{eo}
fe)
Z 604
40+
204 Both species
Std. Dev = 2.12
Mean = 0.00
04 N = 580.00
. ° 7 s S 7 <7 2
&s kes LS <y Bb 0 D DH >
Discriminant function score
w 805
i=
&
Q
—
le)
fo) 4
S 60
a
20-4 L.humile
Std. Dev = .88
Mean = 1.74
0 N = 310.00
ane ol! ps 1 —= Tv = = po
. . . . ‘ SD 7 2 fea 4
RSG) SS OR ee ea
Discriminant function score
w 60;
iS
&
Q.
So 304
fo)
ZZ

BS
oO
1

305
204
L.vulgare
os Std. Dev = 1.12
Mean = -2.00
: maa rel ah cnt N = 270.00
Ss Y Vv 7 2 S B 5) rr =

Discriminant function score

Ficure 4. Histogram of discriminant scores for: all plants of both species, Limonium humile plants only and L.
vulgare plants only.

248

KEY: sampled plants

h L.humile only
V L.vulgare only

Mm mixed sample

H. J. DAWSON AND M. J. INGROUILLE

e misallocated
plants in

FiGuRE 5. Summary of geographical origin of plants of Limonium spp. mis-allocated by the discriminant analysis.

TABLE 5. CHARACTERS OF LIMONIUM SPP. ORDERED BY
CORRELATION WITHIN THE DISCRIMINANT ANALYSIS FUNCTION

Character

nl

. Spikelet distance

. Spike length

. Outer bract width

. Calyx length

. Leaf width

. Primary branch length

. Number of spikelets per spike
. Branch angle

. Corolla length

. Outer bract length

. Leaf shape

. Height to first branching node
. Scape height

Pooled within groups correlations
between characters and canonical
discriminant functions

0-63
0-34
0-27
0-26
—0-24
0-22
Oli
0-17
—0-14
0-11
0-09
0-09
0°02

|
|

LIMONIUM VULGARE AND L. HUMILE IN THE BRITISH ISLES 249

TABLE 6. ANALYSIS OF THE DISTRIBUTION OF STIGMA/POLLEN MORPHS IN THE MOST
INTERMEDIATE L. VULGARE PLANTS

Stigma/pollen morph Low Discriminant Scores High Discriminant Scores Totals
A/Cob 34 99 133
B/Papillate 16 121 B57)
Totals 50 220 270

Chi square = 8-62; pX0-01.

TABLE 7. DISCRIMINANT SCORES FOR L. HUMILE AND L. VULGARE IN DIFFERENT REGIONS

Mean Mean

score score Difference Standard error
Region L. humile L. vulgare between means _ of difference
1. Norfolk/Lincolnshire 1-49 (n=7) —2-39 (n=61) 3-88 0-366 (df=66)
2. Suffolk/Essex 1-51 (n=7) —2-22 (n=60) 3-73 0-356 (df=65)
3. Solent 0-89 (n=21) —1-44 (n=62) 2-33 0-311 (df=81)
4. South Wales 1-53 (n=26) —2-60 (n=38) 4-13 0-226 (df=62)
5. North Wales 1-68 (n=55) —1-50 (n=23) 3-18 0-218 (df=76)
6. Cumbria/Lancashire 1-24 (n=24) —1-74 (n=17) 2:98 0-307 (df=39)
7. Scotland 2-24 (n=40) —0-90 (n=9) 3-14 0-317 (df=47)
8. Ireland 1-91 (n=130)
Total 1-74 (n=310) —2-00 (n=270)

n = number of plants; df = degrees of freedom.
All differences significant at p<0-01.

TABLE 8. DIFFERENCES BETWEEN REGIONS FOR DISCRIMINANT SCORES OF LIMONIUM
HUMILE AND L. VULGARE, REGIONS ORDERED BY SIZE OF DIFFERENCE

Region
L. humile 3 6 1 2 4 5) 8 7

. Solent

. Cumbria/Lancashire
. Norfolk/Lincolnshire
. Essex/Suffolk

. South Wales

. North Wales

. Ireland

. Scotland

NON RNRP AW
+
>

Region
L. vulgare + 1 2 6 5 3 i

. South Wales

. Norfolk/Lincolnshire
. Essex/Suffolk

. Cumbria/Lancashire
. North Wales

. Solent

7. Scotland

WOonNnr
+ + +
+ + xt

x

Duncan test, * = p<0-05.

250 H. J. DAWSON AND M. J. INGROUILLE

Rescaled Distance Cluster Combine

: 0 5 10 15 20 25
Population pan ee gee des is patty gecccsee qonosensee a
HstBchVUL iL
NortonvuL 16
ThnhamVvUL 14
BeauliVUL 4

BlkneyVUL 19
WarhamVUL ALT)
BrnctrVUL aS)

LnstneVUL 5
ChidhmVUL atal
OxwichVUL 61

LongRnVUL 36
Wht frdvVUL 62
RcekffevuL 32
PreeslVUL 40
EastHdVUL 7
TrdBayHUM 43
TthDulVUL 41
ShelIdvUL 47
RinePtHUM 60
StnbryVUL aIEZ
HolbrkVUL ZS,
GldhgrVUL 26
AldbrhvVuL Zak
Iken VUL 20
LtOklyVUL 24
TolbryVUL 25
GibltrVUL 13
BshptnHUM 30
TrMoreHUM 52
HolbrkHUM 22.
RvnG1lsHUM 34
ShlIsdHUM 46
WtesdeHUM 50
YoughlHUM 51
BrwBayHUM 56
AbysdeHUM 49
CromneHUM 54
Dale HUM 63
PwfootHUM 33
ClghneHUM 55
KldsrtHUM 58
DrmrWdHUM 53
LongRnHUM 35
PlnsryHUM 59
HlmeIsHUM 37
BlyLfdHUM Si
BlkneyHUM 18
PreeslHUM 39
HlmeIsVUL 38

NdsOreVUL 3
ItchnrVUL 9
NdsOreHUM 2
EastHdHUM 6
ItchnrHUM 8

ChidhmHUM 10
RckffeHUM shal
TramreHUM 48
AlwEstHUM 42
RhsngrHUM 45
FrM1LBrHUM 44
WigtwnHUM PAU
RssBayHUM 29
CretwnHUM 28

FicurE 6. Phenogram of Limonium vulgare and L. humile populations from the British Isles. O.T.U. code
numbers as in Fig. 1. (HUM = L. humile, VUL = L. vulgare.)

There is some clustering together of populations from the same region. For example, all six
populations of L. vulgare from Essex and Suffolk cluster together with a population from
Lincolnshire. All five L. vulgare populations from North Norfolk and four (of eight) from the Solent
cluster together along with one from South Wales. Four (of six) L. humile populations from
Scotland cluster with three others from North Wales and one from Ireland.

LIMONIUM VULGARE AND L. HUMILE IN THE BRITISH ISLES 251

P.C.A. produces axes of variation which partially separate plants of different species. The first
component was strongly correlated with the same characters discovered by discriminant analysis to
be important discriminators. However multi-dimensional plots of factor scores do not effectively
separate the two species but serve only to emphasise the overlap in species morphology. The second
principal component was strongly correlated to overall size.

DISCUSSION

It is clear that L. vulgare and L. humile are morphologically very close. The ability to transform
characters in a very simple way to produce near normal distributions of characters from the
combined data set of both species is remarkable evidence for the continuity of variation between the
two species. Of the best species discriminators only Spikelet Distance shows any trace of bimodality
after transformation (Fig. 3b). No simple combination of easily measured characters can be used to
identify species precisely. The best discriminating character is Spike Length but it is a character of
debatable value; in eastern England and the Solent, Spike Length is plainly longer in L. humile than
in L. vulgare but taxometric analyses of L. humile from several pure populations in Ireland show
that here it is not necessarily significantly different from L. vulgare (Dawson 1990b).

The shape of the outer bract character given by Clapham (1987) is, in practice, unworkable.
Although outer bract width is generally greater in L. humile there is very considerable overlap.
Another character which has been used in keys is the degree of branching above or below the middle
of the stem. This is one of the weakest discriminants.

Patterns of variation may be complicated by hybridization and introgression. Putative hybrid
plants, intermediate plants as determined by discriminant analysis, are concentrated where both
parental species are present in a marsh. This is strong evidence for hybridization, and it could have
arisen because of the presence of intermediate habitats in these marshes. Nevertheless a
concentration of intermediacy is the Solent region where a high degree of aneuploidy has also been
detected. This is in plants with the L. humile stigma/pollen morph combination. L. humile is
normally 2n=54. Aneuploid plants are generally intermediate in gross morphology (Dawson
1990b). Intermediate L. vulgare plants have 2n=36.

Intermediate/hybrid plants do have reduced pollen stainability though there is a broad range of
values and some hybrids may be partly fertile. Those with the L. humile pollen/stigma morph have
mean stainability of 68% (standard deviation = 18-9, n = 10) and those with the L. vulgare morphs
have stainability of 84% (standard deviation = 14-4, n=15). This compares to stainabilities of 95%
and 96% for the pure species L. humile and L. vulgare with little variation (Dawson 1990b).

Artificial hybrids are vigorous and at least partly fertile. The results of a small number of crossing
' experiments (Dawson 1990b) have shown that hybridization between the species is unidirectional
and is only successful when L. vulgare is used as the male parent and L. humile as the female. This
unidirectional inter-specific incompatibility is normal in other groups in crosses between a self-
incompatible and self-compatible species; pollen from the parent with the intact self-incompatibility
mechanism germinates successfully on the stigma of the self-compatible species but not the other
way around.

However there is little evidence in nature that introgression is unilateral. Approximately equal
numbers of intermediate plants are found in each species. Of the 40 plants mis-clustered in a cluster
analysis of plants (Dawson 1990b) 17 were L. humile and 23 L. vulgare. Of the 30 plants
misclassified by the discriminant analysis 14 were L. humile and 16 L. vulgare using their pollen/
stigma combination.

There is a significant difference in the pollen/stigma morph of intermediate L. vulgare (Table 4).
_ They more commonly have the A/Cob morph than the B/Papillate morph. This is expected because
hybrids of a cross between an A/Cob L. vulgare plant (genotype AC/ac) and a L. humile plant
_ (genotype Ac'/Ac') produce A/Cob plants (genotype AC/Ac') and A/Papillate plants (genotype
_ Ac’/ac) in equal proportions.

It is unlikely that A/Cob intermediate plants can self-pollinate because they have a functioning
Cob stigma gene. Nevertheless they may be able to act as males in backcrosses to either parental
species: to L. humile because it has a mutated stigma gene and to L. vulgare because of the presence
of B/Papillate plants. The consequences of having an unmutated stigma allele in intermediate A/

Jas H. J. DAWSON AND M. J. INGROUILLE

Papillate plants is not known. The A pollen could act in the same way as A pollen from the A/Cob
intermediates. If selfing is possible it would give rise to a range of genotypes identical for pollen/
stigma morph to pure B/Papillate L. vulgare or A/Papillate L. humile as well as hybrids, with
genotypes in proportion 1 ac/ac(B/Cob):2 Ac'/ac(hybrid A/Papillate):1 Ac'/Ac! (A/Papillate L.
humile type.

Another possibility is shown by Armeria maritima (Miller) Willd., which has an identical
incompatibility system including some similar A/Papillate monomorphic populations. It also has
some dimorphic populations, found on polluted soils, in which both A/Cob and B/Papillate morphs
are partly self-compatible (Vekemans et al. 1990). In these populations it is the papillate morph
which is the better selfer but the cob morph may also self effectively.

In L. vulgare/L. humile there is some evidence that introgression occurs in different directions in
different regions. In North Wales and Scotland L. vulgare is the more intermediate in mixed
populations. In East Anglia neither species has a high level of intermediacy in mixed populations. In
Cumbria and Lancashire it is L. humile which is more intermediate in mixed populations. In the
Solent Region both species have a high level of intermediacy in mixed populations.

The selective pressures that cause the breakdown of self-incompatibility systems are difficult to
identify (Jain 1976) but are frequently associated with the colonization of marginal habitats
(Stebbins 1950; Moore & Lewis 1965; Barrett 1988). In these circumstances the more assured
production of seed shown by L. humile is an advantage. A generally lower position down the marsh |
may be for Limonium a more marginal position. L. humile is found in more open communities,
often lower down the marsh than L. vulgare, where competition between species is lower. Its
distribution is positively correlated with patches of bare mud (Boorman 1967, 1968, 1971)..

Regional patterns of variation within each species are complicated by the different abundance of
each species. The presence of both species within a marsh indicates a greater range of habitats,
including intermediate habitats where plants with an intermediate morphology can grow success-
fully. There may have been selection within the gene pool of either species, enlarged or not by
hybridization, for characteristics which allow them to grow within the ecological range of the other
species. Different relative abundance also alters the potential for hybridization and introgression in
different regions. L. vulgare is absent from Ireland and in Scotland and North Wales L. humile is the
much commoner species. In East Anglia it is L. vulgare which is much more abundant.

Clear patterns of geographical variation in salt-marsh vegetation have been noted in Britain both
on a broad geographical scale and more locally (Adam 1978). Part of this is clearly related to
climate. For example, Scottish marshes are distinct because they are beyond the climatic limits of
many important southern salt-marsh species. They are also more frequently subject to fresh-water
run-off from the dry-land. Another important factor is the varying importance of grazing in
marshes. Those around the Irish Sea are often very grassy as a result of grazing. They are also
sometimes narrow or have a small altitudinal range over a large part of them. All these factors may
influence the patterns of variation within and between regions. However apart from the continuum
of variation between L. vulgare and L. humile there is no broad geographical pattern of variation,
say from south-east to north-west. Differences in variation are more haphazard, but here too may
reflect local variation between marshes. This kind of variation has been noted in Salicornia even in
some of the same marshes sampled here (Ingrouille & Pearson 1987; Ingrouille et al. 1990).

It has been frequently suggested that inbreeding species and obligate outbreeders have
contrasting patterns of variation (Loveless & Hamrick 1984). Inbreeders are homogeneous within
populations but have distinct populations. Outbreeders are more variable within populations with
less distinct populations. This pattern has been found in the Limonium species pair described here.
Neighbouring individual plants of L. vulgare are more likely to be distinct than neighbouring plants
of L. humile (Table 3). This observable difference is exaggerated because L. vulgare grows as large
homogeneous clonal patches producing tens or hundreds of genetically identical flowering scapes.
L. humile plants are smaller and more discrete with fewer scapes. This contrast is part of the
differing strategies exhibited by the species. It will be described in detail elsewhere.

There are clearly two distinct species if one resorts to stigma and pollen morph to identify them.
Since these are characters of considerable biological importance, relating to reproductive isolation,
it is taxonomically worth maintaining two distinct species. However this kind of difference in
Armeria maritima is not normally considered significant enough to merit specific recognition.
Northern populations of A. maritima are monomorphic like L. humile with the A/Papillate morph.

LIMONIUM VULGARE AND L. HUMILE IN THE BRITISH ISLES 239

There is considerable gross morphological variation within A. maritima which might be used to
distinguish species in characters such as hairiness, calyx size and leaf shape for example (Lefebvre
1971; Philipp 1974). The monomorphic variant of Armeria is sometimes given specific rank as A.
sibirica but is only one of several subspecies of A. maritima recorded in Flora Europaea (Gorentflot
& Roux 1972; Pignatti 1972).

The correct identification of L. vulgare and L. humile can be fraught with difficulties for the non-
expert even if pollen/stigma morph is used. This was graphically illustrated to M.J.I. when a class of
final year B.Sc. students asked to identify the species from stigma and pollen morph routinely made
an incorrect identification even with the aid of microscopes. Their difficulty was not a result of poor
observation, but because stigma and pollen morphology is more variable than has been recognised
previously. The stigma of L. humile is somewhat intermediate between the clearly contrasting Cob
and Papillate morphs of L. vulgare (Dawson 1990b) and the ‘A’ pollen is variable.

Nevertheless, despite extensive hybridization where they grow together, L. humile and L. vulgare
are distinct species, with different chromosome numbers and, perhaps related to this, some
restriction on their ability to produce fully fertile hybrids.

ACKNOWLEDGMENT

H. J. Dawson acknowledges the financial support of the S.E.R.C. during the course of this work.

REFERENCES

ApDaAMS, P. (1978). Geographical variation in British saltmarsh vegetation. Journal of ecology 66: 339-366.

Barrett, S. C. H. (1988). The evolution, maintenance and loss of self incompatibility systems, in Lovett-
Doust, J. & Lovett-Doust, L., eds. Plant reproductive ecology, pp. 98-124. Oxford University Press,
Oxford.

Boorman, L. A. (1966). Experimental studies in the genus Limonium. D. Phil. thesis, University of Oxford.

Boorman, L. A. (1967). Limonium vulgare Miller and L. humile Miller, in the Biological Flora of the British
Isles. Journal of ecology 55: 221-232.

Boorman, L. A. (1968). Some aspects of the reproductive biology of Limonium vulgare and L. humile. Annals
of botany 32: 803-807.

Boorman, L. A. (1971). Studies in salt marsh ecology with special reference to the genus Limonium. Journal of
ecology 59: 103-120.

CLapuHaM, A. R. (1987). Limonium Miller, in CLAPHAM, A. R., Tutin, T. G. & Moore, D. M., eds. Flora of the
British Isles, 3rd ed. Cambridge University Press, Cambridge.

Dawson, H. J. (1988). Limonium vulgare Miller/L. humile Miller, in Ricu, T. C. G. & Ricu, M. D. B., eds.
Plant crib, pp. 71-72. Botanical Society of the British Isles, London.

Dawson, H. J. (1990a). Chromosome numbers in two Limonium species. Watsonia 18: 82-84.

Dawson, H. J. (1990b). Variation and evolution in Limonium vulgare Miller and L. humile Mill. (Plumbagina-
ceae) in the British Isles. Ph.D. thesis, University of London.

GORENTELOT, R. & Roux, M. (1972). Traitement numérique de populations d’Armeria maritima (Mill.) Willd.
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254 H. J. DAWSON AND M. J. INGROUILLE

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(Accepted May 1994)

Watsonia 20: 255-262 (1995) ZI)

Potamogeton X schreberi G. Fisch. (P. natans L. X P. nodosus
Poir.) in Dorset, new to the British Isles

C. D. PRESTON

Institute of Terrestrial Ecology, Monks Wood, Abbots Ripton, Huntingdon, PEI7 2LS

ABSTRACT

Potamogeton X schreberi G. Fisch. (P. natans L. X P. nodosus Poir., Potamogetonaceae) was discovered in
1992 in the River Stour, Dorset, the first record from the British Isles. Both P. natans and P. nodosus are
recorded from the R. Stour, although neither has been found growing with P. x schreberi. The Dorset plant is
described and compared with material from Germany and Switzerland. P. x schreberi differs from P. natans in
having laminar submersed leaves and from P. nodosus in having phyllodes near the base of the stem and
narrower submersed leaves which are entire rather than minutely denticulate. The differences between this
hybrid and P. X fluitans Roth (P. natans x P. lucens L.) and P. X sparganiifolius Laest. ex Fr. (P. natans X
P. gramineus L.) are discussed. P. X schreberi appears to be sterile, but is apparently able to persist and spread
by vegetative reproduction.

Keyworbs: hybrid, Potamogeton X fluitans, P. X sparganiifolius, vegetative reproduction, England.

INTRODUCTION

Potamogeton nodosus Poir. (Loddon Pondweed) has been known from the River Stour in Dorset
since 1928, when it was first recorded (as P. drucei Fryer) by Druce (1929). In 1988-89 its
distribution in the river was surveyed for the Nature Conservancy Council by Lady Rosemary
FitzGerald, D. A. Pearman and other local botanists (FitzGerald 1990). The plant proved to be
locally abundant in the river for about 9 km from immediately above Hayward Bridge at Child
Okeford, grid reference ST/825.122, downstream to Blandford Bridge at Blandford Forum, ST/
884.060 (see Fig. 1). All known herbarium specimens collected prior to this survey come from the
same stretch of river. Most of them were collected at Hayward Bridge, the best known site for the
species and the one where Druce was photographed examining it in 1931 (Allen 1986).

Although all the authenticated records of P. nodosus have been made at Hayward Bridge or
downstream of it, there are a number of unconfirmed reports of the species at sites some distance
upstream of Child Okeford. Good (1969) reported that P. nodosus “‘was found in considerable
quantity, and flowering and fruiting well’? during a Dorset Natural History and Archaeological
Society excursion to West Mill, Stalbridge, on 27 July 1968. In 1981 the Rev. A. J. C. Beddow
reported P. nodosus from Marnhull, and the distribution of the species was summarised by Good
(1984) as “plentiful in the R. Stour between Marnhull Ham and Child Okeford’’. Both Marnhull
and Stalbridge are more than 10 km upstream of Child Okeford. FitzGerald (1990) and her
colleagues made determined searches for P. nodosus at West Mill and other localities upstream of
Child Okeford, but failed to find it.

In 1992 Dr H. J. M. Bowen and Dr C. Turner reported that they had recently seen P. nodosus in
quantity at Marnhull, in a stretch of river which had not been surveyed by FitzGerald (1990). This
appeared to provide an opportunity of confirming the presence of the species upstream of Child
Okeford. I visited the area on 14 October 1992 with D. A. Pearman, and we found the population
Bowen and Turner had discovered. Examination of these plants showed that they were not
P. nodosus but a hybrid between P. natans L. (Broad-leaved Pondweed) and a species with broad
submersed leaves. At first I tentatively concluded that the other parent must be P. Jucens. More
detailed examination of the material, following a second visit to the site in July 1993, revealed
characters which were inconsistent with this parentage, but could be explained if the broad-leaved
parent was not P. lucens but P. nodosus. The putative hybrid between P. natans and P. nodosus

256 C. D. PRESTON

Marnhull Ham

West Mill © @ Marnhull

Ds

e
e Stalbridge Mounters
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R. Lydden R. Stour

Sturminster
Newton

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Blandford
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8 9

FicureE 1. Map of the River Stour between Fifehead Mill and Blandford Forum, Dorset, showing the 10-km
squares of the national grid and the place-names mentioned in the text. The stretch of river in which P. X
schreberi is found is shown by a solid arrow. P. nodosus grows in the river between the open arrows. Details of
the records of P. natans are given in the text.

(P. X schreberi G. Fisch.) has not hitherto been recorded in the British Isles, but is known from
Germany and Switzerland. Comparison of the Dorset plant with continental material has shown
that it is similar, and I have therefore concluded that it is indeed P. X schreberi.

A preliminary analysis of the isozymes of P. nodosus and the putative hybrid from the R. Stour
and P. natans from the Moors River (v.c. 11, S. Hants) has added support to the hypothesis that the
Stour hybrid is P. x schreberi (P. M. Hollingsworth 1993, unpublished data). P. natans and P.
nodosus differed consistently in five enzyme systems, for all of which the hybrid showed a banding
pattern which combined the bands of both putative parents. The remaining three enzyme systems
produced banding patterns which were invariant across all three taxa.

More fieldwork is needed to establish the distribution of the hybrid and its putative parents in the
R. Stour, but this paper has been written to document the occurrence of the plant in the British Isles
prior to its inclusion in a forthcoming B.S.B.I. handbook.

DESCRIPTION

The following description is based on fresh material and pressed specimens collected from the R.
Stour in Dorset. f

POTAMOGETON X SCHREBERI, NEW TO THE BRITISH ISLES Pasi

Rhizome 2:5—4-2 mm in diameter. Stem 0-5—2-2 m long, 1-75—4-0 mm in diameter near the base,
terete, unbranched. Phyllodes present at base of stem, 95-133 x 2:3-2-6 mm, 41-52 times as lon gas
wide, opaque, green, slightly canaliculate, acute at the apex. Submersed leaves with lamina 56-182
x 2-5-14-5 mm, (6—)10—20(-31) times as long as wide, translucent, pale brown when young, green
when mature, linear-elliptical, very gradually tapering into the petiole, acute at the apex, entire, the
midrib bordered on each side by a band of lacunae up to 1 mm wide, the lateral veins 1-4 on each
side of the midrib, the secondary veins transverse or ascending, rather irregular; petioles (60—)100—
355 mm. Transitional leaves present between the submersed and the floating leaves. Floating leaves
with laminae all pointing in the same direction on the surface of the water, 68-140 x 16-45 mm, 2-3-
5-8 times as long as wide, opaque, subcoriaceous or coriaceous, rather flaccid, brown or brownish
green when young, rather dark olive green when mature, elliptical to oblong-elliptical, tapering into
the petiole, acute at the apex, entire, the lateral veins 5—10 on each side of the midrib, paler than the
lamina when the living leaf is held up to the light, the secondary veins inconspicuous; petioles 89-
180(—330) mm, without a distinct, discoloured section between the petiole and the lamina. Stipules
60-141 mm, rigid, translucent, brownish green, green, colourless with a greenish tinge or pale pink,
with two slight or strong ridges along their length, obtuse to rounded and slightly hooded at the apex
but appearing acute when rolled inwards. Inflorescences 22-28 x c. 6 mm; peduncles 46-79 mm,
slightly compressed towards the base, similar in diameter from base to apex, spongy. Flowers 33-48,
crowded together, with 4 carpels which protrude through the closed perianth segments; pollen
apparently sterile. Fruits not developing.

IDENTIFICATION OF P. X SCHREBERI

Potamogeton X schreberi can be distinguished without difficulty from its putative parents. It has
linear phyllodes at the base of the stem, but above these there are submersed leaves with a distinct
lamina. These submersed leaves vary from leaves which resemble phyllodes but are expanded into a
narrow, rather opaque lamina at the distal end, to leaves with a much more distinct, translucent
lamina. P. natans has phyllodes but lacks laminar submersed leaves; P. nodosus lacks phyllodes but
has laminar submersed leaves. The submersed leaves of P. nodosus are broader than those of P. x
schreberi and are minutely denticulate (the teeth on fresh leaves can be seen in the field with a x 20
lens) whereas those of P. X schreberi are absolutely entire, even when examined at high power
under the microscope. All three taxa produce floating leaves. The floating leaves of P. natans
usually (but not invariably) have a discoloured, flexible section between the petiole and the lamina.
This is not found in either the hybrid or P. nodosus. However, in some living leaves of P. X schreberi
the petiole adjacent to the lamina is slightly browner than the green lamina or greenish brown
petiole, and this may represent a vestige of the flexible section of P. natans. Because they lack this
flexible section, the floating leaves of P. X schreberi tend to point to the same direction whereas
those of P. natans do not.

At anthesis the flowers of the putative parents open, whereas the carpels of P. X schreberi
protrude through closed perianth segments (as in some other sterile hybrids). P. natans fruits
regularly. I have not seen any fruiting material of P. nodosus collected in Britain in the wild,
although plants from the River Loddon cultivated by Fryer (1899) did develop some well-formed
fruit (specimens in BM). Druce (1929) reported that the species was fruiting freely in the R. Stour in
September 1928, but although he distributed specimens collected on this visit I have not yet seen any
with fruits. The inflorescences of P. < schreberi in Dorset show no sign of developing fruit, and the
pollen appears to be sterile (under the microscope the grains appear to be collapsed like a burst
football).

Potamogeton X schreberi is more similar to two other hybrids than it is to its putative parents: P.
x fluitans Roth (P. natans X lucens) and P. X sparganiifolius Laest. ex Fr. (P. natans X gramineus —
L.). Both of these (like P. X schreberi) are hybrids between P. natans and species with broad
submersed leaves, and both have phyllodes at the base of the stem, laminar submersed leaves and
possess the capacity to produce fioating leaves. P. X fluitans is a variable hybrid, but all the
populations I have examined have had stipules with two narrow wings at the base, extending for
most of the length of the stipule. This character is derived from P. Jucens, which has stipules winged
for most of their length. The midrib of the submersed leaves of P. X fluitans lacks a border of

258 C. D. PRESTON

lacunae and can be excurrent for up to 12 mm. Both the submersed and floating leaves of P. x
fluitans have shorter petioles than those of P. x schreberi. P. X sparganiifolius is more difficult to
distinguish from P. X schreberi but its submersed leaves can be much longer both in absolute terms
and in relation to their width (especially in running water), and the fully developed submersed
leaves have a shorter petiole than those of P. X schreberi. In both P. X fluitans and P. x
sparganiifolius the lamina is usually longer than the petiole, whereas in P. X schreberi it is shorter
than or only slightly exceeds the petiole. The shorter petioles of the submersed leaves of P. x
fluitans and P. X sparganiifolius reflect the fact that the submersed leaves of P. lucens have a short
petiole, and those of P. gramineus are sessile, whereas those of P. nodosus have a long petiole.
When comparing the submersed leaves of these taxa the true submersed leaves should be
considered, not leaves which are transitional between submersed and floating leaves.

The differences between P. natans, P. nodosus, P. X fluitans, P. X sparganiifolius and P. X
schreberi are summarised in Table 1.

COMPARISON OF THE DORSET PLANT WITH CONTINENTAL MATERIAL

Potamogeton X schreberi is recorded from Germany and Switzerland (Dandy 1975). Fischer (1907) —

described the Bavarian plant in detail. I have seen specimens of P. xX schreberi in BM, CGE, E, M,
MANCH, RNG, Z and ZT; those in M (German plants) and ZT (Swiss) are most useful in showing
the range of variation of continental plants. Specimens circulated by Koch (1933, 1934) to members
of the Watson Botanical Exchange Club are typical of much Swiss material. Some characters of the
German and Swiss plants are compared to those of the Dorset P. X schreberi and to related taxa in
Table 1. Details of some of the specimens I have seen are listed at the end of this paper.

Fischer (1907) described the lowest 1-3 leaves on his material as ‘““+ aphyllis (phyllodis vel
pseudo-phyllodis)”’. He described the floating leaves as variable in shape, some subcordate or
rounded at the base, others tapering into the petiole. Examination of plants collected by Fischer
(and of other Bavarian material) confirms that those submersed leaves which are present are of the
type which resemble phyllodes with an expanded, distal lamina, and that floating leaves
predominate. None of the floating leaves has a well-developed discoloured section between the
petiole and the lamina, although some of them appear to have a slight discolouration in this area.
Some of the floating leaves are broader in relation to their length than those of the Dorset plant, and
have more lateral veins.

Swiss specimens of P. X schreberi show a full range of-submersed leaves, from expanded
phyllodes to leaves with a long petiole and a distinct, translucent lamina with a band of lacunae on
each side of the midrib. The latter closely resemble the well-developed submersed leaves of the
Dorset plant. On some Swiss plants the slight discolouration between the petiole and the lamina of
the floating leaves is more apparent than on any material I have seen from Dorset or Bavaria, and is
clearly derived from the strongly discoloured and (in herbarium specimens) shrunken section typical
of P. natans. (One specimen in ZT determined as P. cf. natans X nodosus by W. Koch has such a
section, but it has no submersed leaves and in my opinion it is probably P. natans.)

Fischer (1907) described the stipules of P. X schreberi as ‘‘nervosis 1—2 carinatis acutis’’; both
German and Swiss specimens usually have stipules with two distinct ridges along their length but
these ridges are not winged.

Plants of P. x schreberi from Dorset, Bavaria and Switzerland are essentially similar. J. E. Dandy
annotated the only Fischer specimen in BM, ‘“‘apparently is P. natans X nodosus as G. Fischer
thought”’. P. x schreberi shows some variation, with the German plants rather closer to P. natans
than those from elsewhere.

DISTRIBUTION AND HABITAT

Ihave seen P. X schreberi at intervals in a stretch of the R. Stour some 1-5 km long. It is particularly
frequent for about 0:5 km between ST/764.183 and ST/768.184, where there are extensive stands of

259

POTAMOGETON X SCHREBERI, NEW TO THE BRITISH ISLES

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the hybrid. Isolated patches occur downstream of this stretch as far as the footbridge at ST/764.176,
which is the only point along this stretch of the river beside a public right-of-way. I have not searched
other stretches of the river systematically, and P. x schreberi could well be more widespread.

The R. Stour in the vicinity of the P. x schreberi site flows through an area of Oxford Clay. The
hybrid grows over a soft clay substrate in water from 0-2 m to at least 1 m deep. In those areas where
it occurs it tends to be the most frequent macrophyte, and in places stands span the narrow river
from bank to bank. Associated species include Callitriche sp., Nuphar lutea, Potamogeton
pectinatus, Sagittaria sagittifolia and Sparganium emersum. There are well-developed stands of
emergents (Phalaris arundinacea, Phragmites australis, Schoenoplectus lacustris and Sparganium
erectum) along the sides of the river. According to Mr H. Cluett, whose farm lies along the stretch of
the river where P. X schreberi is particularly frequent, the river was last dredged some 10-15 years
ago.

I have not seen either of the putative parents of P. x schreberi growing with it. Both are, however,
recorded in the R. Stour. P. nodosus is, as discussed above, locally abundant downstream of
Hayward Bridge, some 10 km from the nearest recorded P. X schreberi. The stretch of the river with
P. nodosus flows through a variety of geological formations including Upper Chalk, Lower Chalk
and Upper Greensand. FitzGerald (1990) noted that P. nodosus grows over a gravelly substrate,
and is never found where the bottom of the river is muddy. This suggests that P. x schreberi either
has different ecological requirements from P. nodosus, or tolerates a wider range of substrate, as the
hybrid grows over muddy clay at Marnhull. P. natans is apparently rarer than P. nodosus in the
river. J. C. Mansel-Pleydell collected P. natans in the R. Stour at Durweston (May 1865, DOR) and
in the R. Lydden between Stalbridge and Sturminster Newton (29 August 1883, DOR). Durweston
lies on the stretch of the R. Stour where P. nodosus occurs; FitzGerald (1990) recorded P. nodosus
both above and below Durweston Bridge. The R. Lydden is a tributary of the R. Stour which enters
the R. Stour S.S.W. of Marnhull. There are no recent records of P. natans in the R. Stour supported
by herbarium specimens, and the species was only recorded once during the 1988-89 survey of P.
nodosus. FitzGerald and A. Horsfall recorded P. natans in the R. Stour at ST/764.202, downstream
of Fifehead Mill, but they noted that the floating leaves were uncharacteristically elongated. It is
possible that they actually saw P. xX schreberi, which has been recorded a short distance
downstream.

A detailed survey of the R. Stour would be required to establish the distribution of P. X schreberi
and its relationship to that of its parents. The absence of both parents at Marnhull is not particularly
surprising. The ability of Potamogeton hybrids to persist and spread vegetatively often leads to the
presence of a hybrid in sites from which one or both parents are absent.

REPRODUCTIVE BIOLOGY

The one stand of P. X schreberi which I visited in July 1993 was flowering sparingly. There was no
sign of developing fruit on these plants, and their pollen appeared to be sterile. The hybrid has,
however, a robust rhizome and it is clearly able to persist and spread vegetatively. In addition, a
specimen collected in October 1992 had a short shoot, terminated by a bud, arising from a node
towards the base of a stem which had lost its lower leaves. This resembled similar shoots found in P.
nodosus (Dandy & Taylor 1939) and other species, which can act as a means of dispersal
downstream. It is possible that the population of P. x schreberiin the R. Stour represents one clone,
which has spread vegetatively following the initial establishment of a single hybrid plant.

I have seen some inflorescences on both Swiss and German specimens. The young inflorescences
are very compact, with closed perianth segments; some (perhaps more mature) inflorescences are
more elongated with the flowers less closely spaced, but there is no sign of developing fruit. Koch
(1934) noted, of Swiss material which he circulated, “plantu statu florendi rarissima’”’. Fischer
(1907) reported that Bavarian plants failed to fruit and material from Glattbrugg cultivated by Koch
(ZT) also appears to be sterile. Nevertheless, the hybrid can persist in one site for a long period, as
shown by the fact that it was first collected in the Seebach by J. C. D. von Schreber in 1775, and
refound there by Fischer in 1904 (Fischer 1905, -1907).

POTAMOGETON X SCHREBERI, NEW TO THE BRITISH ISLES 261

NOMENCLATURE

G. Fischer first mentioned the possibility of a hybrid between P. natans and P. nodosus in 1903,
when he suggested that a specimen collected by Schreber in the Seebach at Dechsendorf in 1775 was
probably referable to this hybrid (Fischer 1903). In 1904 he rediscovered Schreber’s plant in the
Seebach, and decided that it was definitely the hybrid P. natans X nodosus (Fischer 1905). He
applied the name P. X schreberi to the hybrid, and although he did not provide a formal description
of the plant he included just enough morphological detail in his discussion to validate the name.
Shortly afterwards he provided a full account of the hybrid, based on specimens from the Seebach
and two other localities in Bavaria (Fischer 1907). In his publications Fischer used the name
P. fluitans Roth for the fertile species which is called P. nodosus in this paper and the name P. noltei
G. Fisch. for the hybrid P. lucens X natans.
The nomenclature of the hybrid is therefore:

Potamogeton X schreberi G. Fisch., Mitteilungen der Bayerischen Botanischen Gesellschaft 1: 471
(1905) (P. natans L. X P. nodosus Poir.).

LIST OF SPECIMENS EXAMINED

The British specimens of P. X schreberi which I have collected, and some of the German and Swiss
specimens which I have examined, are listed below. The characters of other taxa mentioned in the
text are (unless stated) based on descriptions which I have prepared for the forthcoming B.S.B.I.
handbook. These descriptions are based on living plants and herbarium specimens from Britain and
Ireland.

BRITISH ISLES

England: Dorset, v.c. 9: River Stour by footbridge S. W. of Mounters, near Marnhull, ST/764.176,
14 October 1992, D. A. Pearman & C. D. Preston (Preston 92/149) (BM, CGE); —, 22 September
1993, J. M. Croft, D. A. Pearman & C. D. Preston (Preston 93/84) (CGE). River Stour W. of
Mounters near Marnhull, ST/765.183, 14 October 1992, D. A. Pearman & C. D. Preston (Preston
92/150) (CGE). River Stour W. of Mounters near Marnhull, ST/767.182, 10 July 1993, D._A.
Pearman & C. D. Preston (Preston 93/30, 93/31) (BM, CGE, NMW); —, 22 September 1993, J. M.
Croft, D. A. Pearman & C. D. Preston (Preston 93/85) (CGE).

GERMANY

Bavaria: Im Seebach nahe dem Ausfluss, alt. 270 m, Oberfranken, July 1904-1905, G. Fischer
(Flora exsiccata Bavarica 997c) (E, M). Seebach bei Erlangen, alt. 270-275 m, Oberfranken, June—
September 1904-1905, G. Fischer (Flora exsiccata Bavarica 997d) (E, M). Im Seebach bei
Mohrendorf (Erlangen), alt. 272 m, Oberfranken, July 1904, G. Fischer (Flora exsiccata Bavaria
997a) (E). Im Seebach a) bei Mohrendorf, b) bei dem altdeutschen Grabhiigel, alt. 274 m,
Oberfranken, July & September 1904 & 1905, G. Fischer (Flora exsiccata Bavarica 997b) (E, M).
Mohrendorf, September 1916, G. Fischer (BM: conf. J. E. Dandy & G. Taylor, 1938). In der Vils
ober- und unterhalb Hahnbach, alt. 385 m, August 1905, W. Niebler (det. G. Fischer) (Flora
exsiccata Bavaria 996) (E, M). In der Vils bei Hahnbach, alt. 375 m, August 1906, W. Niebler
(MANCH). B[ei] Hahnbach, Oberpfalz, 15 July 1946, L. Oberneder 8230D (BM: det. J. E. Dandy,
1962). Kanalisierter Vilslauf zwischen Neumthle und Amberg, Oberpfalz, 27 July 1946, L.
Oberneder 424 (BM).

SWITZERLAND ;
Schwyz: Im Rieselgiessen, Tuggen, 13 October 1918, W. Koch (Z, ZT). Im untern Rieselgiessen
nahe der Miindung in die Spettlinth, Tuggen, 27 August 1919, W. Koch (ZT). Unter dem
Sandsteinbriicklein iiber den Rieselgiessen im Stafelreit, 23 August 1920, W. Koch (ZT). In der
Spettlinth unterhalb des Zusammenflusses mit dem Rieselgiessen, Tuggen, alt. 410 m, 31 August
1922, W. Koch (ZT).

St Gallen: Graben nahe Bahnhof, Rebstein, alt. 413 m, 28 September 1919, W. Koch & E. S. Biiel

262 C. D. PRESTON

(ZT). Rheineck, Bodenseegebiet, August 1904, C. S. Biiel (ZT). Neumihlekanal Rheineck,
Bodenseegebiet, 2 November 1909, C. S. Biiel (ZT: teste G. Fischer).

Zurich: Zurich, August 1847, E. Miiller (Z). Dibendorf, July 1849, C. Cramer (ZT). In einem rasch
fliessenden Bach an der Strasse zwischen Wallisellen (Station) und Diiberndorf, June 1892, F. v.
Tavel (Z). In der Glatt unterhalb Neugut, Wallisellen, 24 June 1917, W. Koch (ZT); —, 3 August
1931, W. Koch (ZT). In der Glatt beim [i.e. bei dem] Neugut, Wallisellen, 17 September 1917, E.
Baumann (ZT: teste G. Fischer). In der Glatt bei Glattbrugg, alt. 425 m, 11 June 1932, W. Koch
(BM, CGE, E, RNG (conf. J. E. Dandy & G. Taylor, 1938), ZT); —, 27 June 1933, W. Koch (CGE,
E, RNG (conf. J. E. Dandy & G. Taylor, 1938), ZT).

ACKNOWLEDGMENTS

I am very grateful to D. A. Pearman for bringing the original record of the plant which proved to be
P. X schreberi to my attention, for joining me for all my fieldwork and for much additional help. I
also thank Mrs J. M. Croft for help with fieldwork and Mrs A. Pearman for hospitality in Dorset.
P. M. Hollingsworth kindly provided preliminary details of his isozyme studies and Lady Rosemary
FitzGerald, Dr T. Franke, Prof. J. W. Kadereit and Dr M. F. Watson helped in other ways. I am
grateful to the herbaria cited for the loan of specimens, to English Nature for permission to cite
FitzGerald (1990) and to A. O. Chater, P. T. Harding and P. D. Sell for commenting on a draft of
this paper.

REFERENCES

ALLEN, D. E. (1986). The botanists, p. 135. St Paul’s Bibliographies, Winchester.

Danpy, J. E. (1975). Potamogeton L., in Stace, C. A., ed. Hybridization and the flora of the British Isles, pp.
444-459. Academic Press, London.

Danpy, J. E. & Taytor, G. (1939). Studies of British Potamogetons — IV. The identity of Potamogeton Drucei.
Journal of botany, London 77: 56-62.

Druce, G. C. (1929). Potamogeton Drucei Fryer. Report of the Botanical Society and Exchange Club of the
British Isles 8: 928. ;

FiscHER, G. (1903). Beitrag zur Kenntnis der bayerischen Potamogetoneen. III. Mitteilungen der Bayerischen
Botanischen Gesellschaft zur Erforschung der heimischen Flora 1: 301-306.

FIscHER, G. (1905). Beitrag zur Kenntnis der bayerischen Potamogetoneen. V. Mitteilungen der Bayerischen
Botanischen Gesellschaft zur Erforschung der heimischen Flora 1: 471-476.

FiscHER, G. (1907). Die bayerischen Potamogetoneen und Zannichellien. Berichte der Bayerischen Botanischen
Gesellschaft zur Erforschung der heimischen Flora 11: 20-162.

FirzGERALD, R. (1990). Rare plant survey of south-west England. 4, Dorset. Unpublished report (Nature
Conservancy Council C.S.D. confidential report no. 1061).

Fryer, A. (1899). Potamogeton Drucei Fryer. Journal of botany, London 37: 524.

Goon, R. D’O. (1969). Botany. Proceedings of the Dorset Natural History and Archaeological Society 90: 45—46.

Goop, R. D’O. (1984). A concise flora of Dorset, p. 170. Dorset Natural History and Archaeological Society,
Dorchester.

Kocu, W. (1933). Potamogeton natans L. X nodosus Poir. (= P. schreberi Fischer). Report of the Watson
Botanical Exchange Club 4: 195.

Kocu, W. (1934). Potamogeton natans L. X nodosus Poir. Report of the Watson Botanical Exchange Club 4: 243.

(Accepted July 1994)

eee ee

Watsonia 20: 263-273 (1995) 263

A new variety of Narrow-leaved Marsh-orchid in
South Hampshire (v.c.11)

M. N. JENKINSON

Merriefield House, 21 St Clair Road, Canford Cliffs, Poole, Dorset, BH13 7JP

ABSTRACT

Morphometric study of a population of Dactylorhiza majalis (Rchb. fil.) P. F. Hunt & Summerh. subsp.
traunsteineri (Saut. ex Rchb. fil.) H. Sund. (Orchidaceae) near Exbury, South Hants. (v.c. 11) reveals some
distinct morphological differences from most other populations of the subspecies. These differences are
considered sufficient to justify the proposal of a new variety, var. bowmanii M. N. Jenk., var. nov., which is
described. It is now known from three sites in v.c. 11 and one in Dorset (v.c. 9). Another study population, in
North Hants. (v.c. 12), is determined as subsp. traunsteineri var. traunsteineri.

Keyworps: Orchidaceae, Dactylorhiza, morphometrics, multivariate analysis, Yorkshire, Berkshire, Anglesey,
Hampshire.

INTRODUCTION

In ‘Plant Records’, Watsonia 19: 152 (1992), the discovery in 1984 by R. P. Bowman of a new
population of Dactylorhiza traunsteineri (Saut. ex Rchb. fil.) Sod (Orchidaceae) in S. Hants. (v.c.
11) was reported. The find was determined by Dr Francis Rose, and is a first record for the vice-
county, this taxon having been previously recorded in Hampshire only from N. Hants. (v.c. 12).

For reasons outlined in detail elsewhere (Jenkinson 1992), related to morphological overlap
between taxa (cf. Bateman & Denholm 1983), and the presence of morphological intermediates,
particularly in Hampshire and Dorset populations, I regard this taxon as a subspecies of D. majalis
(Rchb. fil.) P. F. Hunt & Summerh. Although strictly objective scientific data (in the form of
compatible morphometric data) to justify merging this taxon with the continental D. traunsteineri
(Saut. ex Rchb. fil.) So6 are actually still lacking, I am persuaded by the weight of informed opinion
of experts who have seen both taxa in the field that they should be regarded as conspecific. The
correct nomenclature for this taxon is therefore Dactylorhiza majalis (Rchb. fil.) P. F. Hunt &
Summerh. subsp. traunsteineri (Saut. ex Rchb. fil.) H. Sund. (Sundermann 1980; R. M. Bateman,
pers. comm., 1987; Bateman & Denholm 1989; Jenkinson 1992).

R. P. Bowman (pers. comm., 1985) drew my attention to the Exbury population as there
appeared to be some particularly interesting features of some of the plants at the site. Growing in a
flushed marshy clearing in the 10-km square SU/4.0, the population of Narrow-leaved Marsh-orchid
(c. 200 plants) formed a small but distinct morphological population within a much larger mixed
colony of Dactylorhiza spp., consisting predominantly of D. majalis subsp. praetermissa (Druce) D.
M. Moore & So6, D. fuchsii (Druce) So6 and hybrids between the two. There was also a small
number of putative hybrids between D. fuchsii and D. majalis subsp. traunsteineri. The whole
colony numbered c. 2000 plants at that time.

The site itself is a very wet flush in a clearing between a belt of old oak woodland and a coniferous
plantation. The clearing was cut as a strip through the woodland in the early 1960s to enable a gas
pipeline to be installed. The site was subjected to soil-test samples by R. M. Bateman in 1986, and
the pH was found to be c. 6-6, slightly lower than that of most British and Irish sites for the taxon,
which is widely regarded as being restricted to calcareous fens and basic flushes. Working on the
assumption that the orchids were not present in the wood before the strip was cut, it would appear
that the orchid colony could not have been more than 20 years old when it was first discovered. It
may be felt that this is a rather short time for a site to acquire such a large and heterogeneous colony

264 M. N. JENKINSON

of Dactylorhiza. The oil pipeline from Wytch Farm, Dorset, to Fawley Refinery was also installed
through the cut strip recently, to one side of the main concentration of orchids, without any
apparent damage to the population of D. majalis subsp. traunsteineri, although on a recent visit
(June 1993) it was found that much of the site had become overgrown with an umbellifer (? Oenanthe
crocata L.) and Bracken (Pteridium aquilinum (L.) Kuhn), as a result of which the full extent of the
mixed colony had declined somewhat.

A number of authors (e.g. Bateman & Denholm 1983; Foley 1990) have given recent detailed
morphological descriptions of D. majalis subsp. traunsteineri (see Table 1). The Exbury plants are
clearly distinct from D. majalis subsp. praetermissa at the site in many characters regarded as
diagnostic of subsp. traunsteineri, such as small stature, thin stems, few narrow leaves, short lax few-
flowered inflorescences, strongly reflexed lateral lobes to the labellum, and the presence (sometimes
intense) of diffuse anthocyanins in upper stem and bracts. Certain other morphological features of
these plants, however, are not typical of the subspecies. The most obvious of these in the field is the
presence in the vast majority of plants of exceptionally deep sinuses and an unusually elongated
central lobe to the labellum. In addition the labella are proportionately narrower in relation to their
length than in most plants of this taxon, a feature that is exaggerated in the field by the strong
reflexion of the lateral lobes. The flowers are also rather darker in base colour than those in most
populations I have seen (with the exception of one Yorkshire population). The vegetative
characteristics are generally more robust than most populations of the taxon (closer to some Irish
populations — see Foley 1990), in some respects intermediate between this taxon and subsp.
praetermissa. There are usually two non-sheathing leaves and three (but sometimes four) sheathing
leaves, which are generally longer and broader than those of most populations of the taxon.

R. M. Bateman, having carried out a detailed morphometric comparison, expressed the opinion
(pers. comm., 1986) that these plants were intermediate in overall morphology between subsp.
traunsteineri (as epitomised by populations in Ireland and Anglesey) and subsp. praetermissa, but
closer to, and therefore referable to, subsp. traunsteineri, thus concurring in essence with Rose.

It was clear, however, that whilst some vegetative characteristics could be described as
‘intermediate’, certain floral characters were even more extreme than most subsp. traunsteineri. I
felt that the unusual morphology of these distinctive plants merited more detailed investigation.
Accordingly in 1988 and 1989 morphometric data were obtained from recognised populations of
subsp. traunsteineri in Oxfordshire (Berks., v.c. 22), Yorkshire (Mid-W. Yorks., v.c. 64) and
Anglesey (v.c. 52), in order to incorporate the broadest possible spectrum of natural and normal
variation within the taxon.

In addition, another taxonomically ambiguous colony of marsh-orchids, at Mapledurwell Fen
near Basingstoke in N. Hants. (v.c. 12) in SU/6.5, was examined, as it has long been believed to
contain a number of plants referable to subsp. traunsteineri (Doherty & Pilkington 1983), although
Dr Francis Rose, in correspondence with R. P. Bowman (R. P. Bowman, pers. comm., 1984),
expressed the view that it was ‘“‘not quite pure”’ at the site, implying some introgression, presumably
with D. fuchsti and/or D. majalis subsp. praetermissa, both of which are also present. In an attempt
to resolve the status of these plants, they were incorporated into the current study.

I have since found three further smaller populations of plants similar in morphology to those at
Exbury, all in the administrative county of Dorset, but two in v.c. 11 (S. Hants.) and one in v.c. 9
(Dorset). The largest population (c. 50 plants), discovered in 1988 at Avon Forest Park near
Ringwood, in SU/1.0, was measured for this research. The other populations, six plants in 1992 at St
Leonards near Ringwood in SU/1.0, and a single plant in 1993 on Studland Heath in SZ/0.8, were
too small for significant statistical analysis. All three new populations differed from the original
Exbury population in minor morphological features (Avon Forest Park, slightly more robust with
slightly less deeply three-lobed lips; St Leonards, slightly paler in base colour; and Studland, with a
slightly less deeply three lobed lip), but the overall similarity to the Exbury plants was such that
there was no doubt that they were referable to the same taxon.

Details of the eight study populations from which data were obtained for the purpose of this
research are listed on Table 2.

An interesting feature of all three recently discovered sites is that they are all in dry acid
grassland. The plants are growing amongst Bracken (Pteridium aquilinum) and acid-soil indicators
Galium saxatile L. and Potentilla erecta (L.) Raeusch. are present at all three sites. Pteridium
aquilinum and Potentilla erecta are also present at Exbury. The dry grassland orchids Orchis morio

Nn WN

10.

11.

122

i

14.

113y.

16.

ils

A NEW VARIETY OF NARROW-LEAVED MARSH-ORCHID 265

Character

Stem height

. Stem width

. Sheathing leaves

. Non-sheathing leaves
. Length of longest leaf
. Width of widest leaf

. Leaf markings

. Inflorescence

. Peripheral bract cells mean size

Labellum size

Labellum shape

Labellum markings

Labellum colour

Lateral outer perianth segments
Spur

Anthocyanins

subsp. traunsteineri
12-30 cm, only occasionally >30 cm

<5 mm, flexuous

Up to 3

0-1

Up to 12 cm, rarely more

Usually 8-12 mm, only occasionally
>15 mm

Occasionally present, solid spots
(except var. eborensis), 1-2 mm in
diameter, round or transversely
elongated, usually concentrated
towards leaf-tips

Rarely >7 cm, 7-12 flowers

60-115 um, usually >80 um in mean
length, subacute to serrate

>7-°5 X 9-5 mm, usually broader than
long, often broadest below middle,
usually c.10 mm wide

Usually three-lobed, central lobe often
elongated (>1 mm) well beyond
laterals, lateral lobes usually markedly
reflexed

Dashes or flecks with loop markings,
usually covering most of lip

Base colour variable from pale rose-
pink to reddish-purple. Markings much
darker, sometimes intensely so

Usually nearly vertical, rarely with
annular markings

Occasionally <8-5 mm, usually
tapering

Stem/bract anthocyanins usually
present, often intense

TABLE 1. RANGE OF VARIATION AND DESCRIPTION OF MAIN DISTINGUISHING CHARAC-
TERS OF DACTYLORHIZA MAJALIS SUBSP. TRAUNSTEINERI AND SUBSP. PRAETERMISSA

subsp. praetermissa

20-50 cm, often
exceeding 30 cm

>5 mm, stout, erect
3 or more

2 or more

Usually >12 cm
Usually >15 mm

Unmarked, except var.
Junialis

Usually >7 cm, usually
>20 flowers

50-100 um, but rarely
>80 um, rounded
obtuse

>7:5 X 9-5 mm,
usually broadest + at
middle, usually >10
mm wide

Usually slightly three-
lobed, rarely entire,
central lobe only
occasionally elongated
well beyond laterals,
laterals usually + flat
Dots or dashes, more
or less concentrated
towards centre of lip
Base colour usually
pale rose-pink, rarely
deeper rose-red.
Markings slightly
darker than base colour
Usually more or less at
45°, unmarked
Usually <8-5 mm,
tapering

Stem/bract
anthocyanins
occasionally present,
rarely intense

L. and Spiranthes spiralis (L.) Chevall. are present in quantity at the St Leonards site, as are D.
fuchsii and D. maculata (L.) So6; At Avon Forest Park, D. majalis subsp. praetermissa occurs in a
damp hollow nearby, Listera ovata (L.) R. Br. and Spiranthes spiralis both occur occasionally with
D. majalis subsp. traunsteineri, and Orchis morio occurs in some numbers in the vicinity. At the |
Studland site, the single plant occurred on a dry track verge amongst Pteridium aquilinum with no
other orchids, although D. majalis subsp. praetermissa occurs in a damp patch of the same verge
some 100 m away, and D. incarnata (L.) So6 subsp. pulchella (Druce) So6 occurs in the adjacent
acid bog.

Long regarded as typically an inhabitant of wet calcareous fen or basic flushes (Heslop-Harrison

1953; Lacey 1955; Lacey & Roberts 1958; Tennant 1979; Kenneth & Tennant 1983; Bateman &

266 M. N. JENKINSON
TABLE 2. STUDY POPULATIONS OF DACTYLORHIZA TAXA

Population Grid Ref. Habitat No. plants © Accompanying Dactylorhiza

1. Wharfedale, SD/97.67 Basic flush >200 Dii, Df, Dm, hybrids
Mid-W. Yorks., v.c. 64

2. Cothill, Berks., v.c. 22 SU/46.99 Fen <10 Df

3. Parsonage Moor, Berks., SU/46.99 Fen <50 Dmpr, Df, hybrids
VG 22

4. Rhos-y-Gad, Anglesey, SH/51.79 Fen >200 Dmp, Dip, Dic, Df, Dm,
Vc: D2 hybrids

5. Cors Erddreiniog, SH/47.82 Fen >1000 Dmp, Dip, Dic, Df, Dm,
Anglesey, v.c. 52 hybrids

6. Exbury, S. Hants., v.c. 11 SU/4-.0- Neutral flush >200 Dmpr, Df, hybrids

7. Avon Forest Park, SU/1-.0- Acid grassland <S50 Dmpr, Df, Dm
S)- Hants vecHd

8. Mapledurwell Fen, SU/67.52 Fen <100 Dmpr, Df, hybrids

N. Hants., v.c. 12

Key to accompanying Dactylorhiza: Df = D. fuchsit, Dm = D. maculata, Dii = D. incarnata subsp. incarnata,
Dip = subsp. pulchella, Dic = subsp. coccinea, Dmp = D. majalis subsp. purpurella, Dmpr = subsp.
praetermissa.

Note: Populations 2 and 3, as a result of county boundary changes, are now in the administrative county of
Oxfordshire, and similarly, population 7 is now in the county of Dorset.

Denholm 1983; Kenneth, Lowe & Tennant 1988; Roberts 1988; Foley 1990), D. majalis subsp.
traunsteineri has not hitherto been recorded, as far as is known, in any dry or markedly acid habitats
in Britain, although some authorities (e.g. Davies, Davies & Huxley 1983) have cited Sphagnum
bog as an occasional habitat for the taxon in continental Europe. It should be noted however that in
the case of Davies, Davies & Huxley (1983) the authors express the cautionary opinion that this may
be due to confusion between morphologically similar marsh-orchids.

Since the Narrow-leaved, Wicklow or Pugsley’s Marsh-orchid was first described from Co.
Wicklow in Ireland by H. W. Pugsley (Pugsley 1936), various authorities have sought to delimit the
taxon more precisely. Arguably the best descriptions are those published by Bateman & Denholm
(1983) and Foley (1990). These have been amalgamated and summarised in Table 1. In addition, for
the purpose of comparison, a description of D. majalis subsp. praetermissa based on Bateman &
Denholm (1983) and the present writer’s own data has also been incorporated into Table 1.

MATERIALS AND METHODS

It was considered that in order to quantify accurately the similarities and dissimilarities of the
proposed new variety with D. majalis subsp. traunsteineri as it is currently recognised in Britain, it
was necessary to obtain morphometric data from a widely separated selection of well-known
populations of the taxon, in order to incorporate as much as possible of the range of variation to be
found, in what is a very variable taxon. ‘Control samples’ were therefore obtained from five
populations (numbers 1—5 on Table 2) of the subspecies which have been the subject of previous
morphometric research (Heslop-Harrison et seq. above), and which are widely recognised, well-
determined sites for this taxon.

Morphometric data were obtained on a total of 59 characters for each of ten randomly selected
plants of the target taxon in each population, according to criteria outlined by me elsewhere
(Jenkinson 1992). These data were then subjected to the multivariate analysis described in that
paper.

Two specimen plants (minus roots) were taken from the Exbury population in June 1989, and
pressed and mounted. These are now conserved at BM. They were drawn prior to pressing, and that
original drawing is reproduced as Fig. 1. The rt -hand of the two plants is the holotype, the left-
hand a paratype.

A NEW VARIETY OF NARROW-LEAVED MARSH-ORCHID 267

10

aj iff |
fl |, Af,

i y/ oll
AN 0

Figure 1. Dactylorhiza majalis subsp. traunsteineri var. bowmanii M. N. Jenkinson. Holotype (right hand
plant): near Beaulieu, S. Hants. (v.c. 11), 13 June 1989. Paratype (left hand plant): as holotype. Drawn from
specimens deposited in BM.

A separate sample of ten excised flowers was also obtained, mounted and forwarded to BM with
the holotype. Drawings of the excised labella of that sample, and of the survey samples from Exbury
and Avon Forest Park, are reproduced as Fig. 2.

RESULTS

Table 3 is a summary of population means, for the eight study populations, for those characters
(listed in Table 1) regarded as ‘diagnostic’ for subsp. traunsteineri.

M. N. JENKINSON

268

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270 M. N. JENKINSON

Samples from Exbury 13 June 1989 (conserved at BM with holotype)

FiGurE 2. Drawings of 30 excised and mounted labella of Dactylorhiza majalis subsp. traunsteineri from Exbury
(20) and Avon Forest Park (10).

It will be seen that for the vast majority of character-states, the three Hampshire populations from
which data have been obtained fall within the parameters of subsp. traunsteineri according to
accepted morphological criteria. The only difference from typical subsp. traunsteineri noted in the
Mapledurwell Fen population was a relatively low mean density of stem anthocyanin. This is not
considered significant. At Exbury and Avon Forest Park, however, certain clear distinctions in
morphology become apparent. These are listed in Table 4, in which characters are numbered as in
Table 1. j

A NEW VARIETY OF NARROW-LEAVED MARSH-ORCHID 7a) Mh

TABLE 4. MEAN VALUES (WITH RANGE IN PARENTHESES) FOR MAIN DISTINGUISHING
CHARACTERS OF D. MAJALIS SUBSP. TRAUNSTEINERI VAR. BOWMANII

Avon Forest

Character Exbury Park Remarks
3. Sheathing leaves 3-0 3°5 Avon Forest Park more
(3-0) (3-0-4-0) than most
4. Non-sheathing leaves 1-4 9 More than most
(1-0-2-0) (1-0-3-0)
5. Length of longest leaf (cm) 12-81 10-51 Longer than most
(8-5-16-7) (7-0-12:5)
6. Width of widest leaf (cm) 2-03 2-25 Broader than most
(1:6-2-45) (1-6-3-4)
8. Number of flowers 16-8 19-1 More than most
(13-28) (8-40)
11. Length of central lobe (mm) Sal 2:18 Longer than most —
(2:25-4-5) (1-75-3-0) to max. 7 mm at
(survey Exbury— very marked
sample) in the field
14. Lateral outer perianth Both populations between Lower than most
45° and vertical.
15. Length of spur (mm)* 7-60 6-65 Shorter than
(5-5—9-0) (5-5—8-25) most — especially

Avon Forest Park

*See Bateman & Denholm (1989) for their remarks re compatibility of measurements for length of spur. All
measurements for this study, including those for the ‘control sample’ populations, are lower than those given by
Bateman & Denholm (1983) for this character. It depends on the point of excision of the spur. Those at Avon
Forest Park, however, were shorter than those of most plants measured in all populations.

DISCUSSION AND CONCLUSIONS

The absence of significant differences between the morphology of the plants measured at
Mapledurwell Fen and that of the ‘control’ populations measured I consider to be sufficient
evidence to support a firm determination of D. majalis subsp. traunsteineri (i.e. of the type variety,
cf. Bateman & Denholm 1983) at this site. Furthermore, Dr Rose now takes the view (pers. comm.,
1993) that there are some early-flowering plants at the site which are referable to D. majalis subsp.
traunsteineri var. traunsteineri, although the situation is somewhat complicated later in the flowering
season by the presence of hybrids and intermediates. There would therefore be no justification in
separating these plants from var. traunsteineri merely on the basis of a few very minor morphological
differences.

The situation at Exbury and Avon Forest Park, however, is somewhat different: the differences
from typical subsp. traunsteineri var. traunsteineri at these sites (and in the few plants found at St
Leonards and Studland) I consider to be of some importance. Whilst in the context of their overall
morphology it is clear that they are sufficiently close to subsp. traunsteineri to warrant inclusion in
that taxon, and indeed have been determined as such by both Rose and Bateman, it is my view that
the differences identified and described above are comparable in magnitude to those that delimit the
other two British named varieties of the subspecies (Roberts & Gilbert 1963; Bateman & Denholm
1983; Lowe, Tennant & Kenneth 1986), viz.:
var. eborensis (Godfery) R. M. Bateman & Denholm: short stem, leaves often marked with annular

markings, small labellum, short spur;
var. francis-drucei (Wilmott) R. M. Bateman & Denholm: short stem, short unmarked leaves, few-

flowered inflorescence, small labellum often longer than broad, elongated central lobe, short
spur.

A new variety is therefore proposed to take account of the distinct morphological characteristics
of plants at the four locations so far identified. It is named in honour of the finder of the Exbury
population, R. Paul Bowman, B.S.B.I. recorder for many years for v.c. 11 (S. Hants.).

272 M. N. JENKINSON
DESCRIPTION

Dactylorhiza majalis (Rchb. fil.) P. F. Hunt & Summerh. subsp. traunsteineri (Saut. ex Rchb. fil.)
H. Sund. var. bowmanii M. N. Jenk., var. nov.
Caulis plerumque 15 cm (interdum 30 cm) altus superans, raro 5 mm diametro superans. Folia
vaginantia 3-4, plerumque secus caulem + aequaliter dispersa, folium longissimum plerumque
secundum infimum, 8-16 cm longum, latissimum interdum secundum infimum, plerumque
infimum, 1-6—3-4 cm latum; folia invaginantia plerumque 2, interdum 1; folia omnia immaculata.
Inflorescentia plerumque minus quam 7 cm longa, plerumque 20% longitudinis caulis excedens,
plerumque laxa, interdum plus quam 18-flora. Cellulae peripherales bractearum plerumque
majores quam 70 wm longae. Labellum plerumque 7-5-9 x 8-5-10-5 mm, plerumque + circa
mediam partem latissimum; color basicus plerumque atrorubens, cum maculis lineatisque fusciori-
bus, labello incomplete obtecto; sinus semper praesentes atque profundi, lobus medianus
plerumque longus (>2-5 mm), lobi laterales raro indentati, plerumque valde reflexi. Sepala lateralia
plerumque plus verticalia quam horizontalia (>45°), modice sed non annulatim maculata. Calcar
raro minus quam 6-5 mm longum, plerumque 7-5—8-5 mm, plerumque decrescens, raro cylindricum
vel sacciforme.

Aestate florens (Junio, primam ad tertiam hebdomadem). Habitat in pascuis asperis udis vel
aridis atque + acidis; quattuor populationes in Anglia australi, tota distributio ignota; rarissima.

Ho otypus: Exbury, near Beaulieu, S. Hants. (v.c. 11), England, very wet neutral flush, 13 June
1989, M. N. Jenkinson (holo. BM) (illustrated in Jenkinson (1991), Plates 62 & 63, p. 112).

Stem usually exceeding 15 cm, only occasionally exceeding 30 cm, rarely exceeding 5 mm in
diameter. Sheathing leaves 3-4, usually + evenly distributed along stem, longest leaf usually second
lowest on stem, 8-16 cm long, widest leaf occasionally second lowest, but usually lowest leaf, /-6-3-4
cm in width, non-sheathing leaves usually 2, occasionally 1; leaf markings absent. Inflorescence
usually less than 7 cm long, usually more than 20% of stem length, usually lax, occasionally more
than 18 flowers. Mean length of peripheral bract cells usually exceeding 70 um. Labellum usually
more than 7:5 X 8-5 mm, but Jess than 9 xX 10-5 mm, usually broadest + at middle; base colour
usually dark magenta, markings usually darker dashes and loops, /ess than complete coverage of
labellum; sinuses always present, well-developed; central lobe usually long (>2:5 mm); lateral lobes
rarely indented, usually strongly reflexed. Lateral outer perianth segments usually nearer vertical
than horizontal (>45°), usually lightly marked with solid, not annular, markings; spur rarely less
than 6-5 mm long, usually 7-5—8-5 mm, usually tapering, rarely cylindrical or sac-like.

Flowering June (best weeks 1-3). Habitat: damp or dry, neutral to slightly acidic rough grassland;
only four known populations in southern England, full distribution unknown. Very rare. (N.B.
Italics in the English description follow the practice of Bateman & Denholm (1983) in order to
emphasise those characters considered to be most diagnostic, but concentrate in this case on those
which best distinguish the newly-described variety from the typical subspecies and the other named
varieties. )

ACKNOWLEDGMENTS

I gratefully acknowledge the assistance of the following: Drs R. M. Bateman and I. Denholm, for
advice and helpful discussion, permission to use the results (published and unpublished) of their
own extensive research, and critical appraisal of this paper; R. P. Bowman, for much useful
fieldwork, advice and discussion, on Dactylorhiza in particular, culminating in the discovery and
investigation of the Exbury colony; F. Horsman, for guidance and assistance in the field in
Yorkshire; Nicholas Jenkinson, for assistance with the fieldwork in North Hants., Oxfordshire and
Yorkshire; Dr F. Rose, for helpful discussion with regard to the Mapledurwell Fen population; Dr
R. J. White, for helpful advice and discussion, and invaluable assistance with the multivariate
analysis on the computers of Southampton University; and F. A. Woodhead, for assistance with the
Avon Forest Park colony, much helpful fieldwork, and the loan of her microscope for the peripheral
bract cell measurements. In addition, my thanks go to the following individuals and organisations,
who have variously assisted with site information, accompanied visits to sites, and the provision of

A NEW VARIETY OF NARROW-LEAVED MARSH-ORCHID 273

permits: Berkshire, Buckinghamshire and Oxfordshire Naturalists’ Trust (F. La Forte); Hampshire
Wildlife Trust; the former Nature Conservancy Council (Berkshire, Dyfed-Powys and North Wales
Regions); and the North Wales Wildlife Trust.

REFERENCES

BATEMAN, R. M. & DENHOLM, I. (1983). A reappraisal of the British and Irish dactylorchids, 1. The tetraploid
marsh-orchids. Watsonia 14: 347-376.

BATEMAN, R. M. & DENHOLM, I. (1989). Morphometric procedure, taxonomic objectivity, and marsh-orchid
systematics. Watsonia 17: 449-455.

Davies, P., Davigs, J. & Hux ey, A. (1983). Wild orchids of Britain and Europe. Chatto & Windus, London.

Douerty, J. & PILKINGTON, J. (1983). Hampshire’s countryside heritage: 3. Rivers and wetlands. Hampshire
County Council, Winchester.

Fotey, M. J. Y. (1990). An assessment of populations of Dactylorhiza traunsteineri (Sauter) So6 in the British
Isles and a comparison with others from Continental Europe. Watsonia 18: 153-172.

HesLop-Harrison, J. (1953). Studies in Orchis L. Il. Orchis traunsteineri Saut. in the British Isles. Watsonia 2:
371-391.

JENKINSON, M. N. (1991). Wild orchids of Dorset. Orchid Sundries Ltd, Stour Provost, Gillingham, Dorset.

JENKINSON, M. N. (1992). The marsh orchids in Hampshire and Dorset: some recent research. Proceedings of the
Hampshire Field Club and Archaeological Society 47: 225-252.

KENNETH, A. G., Lowe, M. R. & TENNANT, D. J. (1988). Dactylorhiza lapponica (Laest. ex Hartman) Soo in
Scotland. Watsonia 17: 37-41.

KENNETH, A. G. & TENNANT, D. J. (1983). The Scottish records of Dactylorhiza traunsteineri (Sauter) Sod.
Watsonia 14: 415-417.

Lacey, W. S. (1955). Orchis traunsteineri (Saut.) in Wales. Proceedings of the Botanical Society of the British
Isles 1: 297-300.

Lacey, W.S. & Roberts, R. H. (1958). Further notes on Dactylorhiza traunsteineri (Saut.) Vermeul. in Wales.
Proceedings of the Botanical Society of the British Isles 3: 22-27.

LowE, M. R., TENNANT, D. J. & KENNETH, A. G. (1986). The status of Orchis francis-drucei Wilmott. Watsonia
16: 178-180.

PuGsLey, H. W. (1936). New British marsh orchids. Proceedings of the Linnean Society of London 148: 121-125.

Roserts, R. H. (1988). The occurrence of Dactylorhiza traunsteineri (Sauter) So6 in Britain and Ireland.
Watsonia 17: 43-47.

Roserts, R. H. & GiLBert, O. L. (1963). The status of Orchis latifolia var. eborensis Godfery in Yorkshire.
Watsonia 5: 287-293.

SUNDERMANN, H. (1980). Europdischer und mediterrane Orchideen — eine Bestimmungsflora, 3rd ed. Brucke-
Verlag, Hildesheim.

TENNANT, D. J. (1979). Dactylorhiza traunsteineri in Yorkshire. Naturalist (Hull) 104: 9-13.

(Accepted August 1994)

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Watsonia 20: 275-278 (1995) 275

Erica X stuart: the authorship reconsidered

ESC. NELSON

National Botanic Gardens, Glasnevin, Dublin 9, Ireland

ABSTRACT

The author of the epithet stuartii for the naturally-occurring hybrid Erica x stuartii (E. mackaiana X tetralix)
(Ericaceae) was J. Muirhead Macfarlane, and the combination at specific level was made by the editor of The
Gardeners’ Chronicle, Maxwell T. Masters; the binomial’s correct form is Erica X stuartii (Macfarl.) Mast.

Keyworps: Ericaceae, Ireland, nomenclature.

INTRODUCTION

This paper does not propose a radical name-change for a plant that is relatively common in western
Ireland (W. Galway (Connemara), W. Mayo and W. Donegal; v.cc. H16, H27, H35), and (as
named cultivars) in gardens. Its purpose is to establish the authority for the established name Erica
x stuartil.

An essential principle of the /nternational code of botanical nomenclature (I.C.B.N.) is that a
name is validly published when it is accompanied by a diagnosis that “‘in the opinion of its author
distinguishes the taxon from others” (/.C.B.N. 1988; Art. 32.2). For nomenclatural precision,
therefore, it is crucial that the first place of valid publication of a Latin name and the author’s name
are exactly established and clearly stated. After 1 January 1935, the diagnosis must be in Latin
U.C.B.N. 1988; Art. 36); before 1935 a diagnosis in any language is sufficient to validate a Latin
name.

Even today plants grown in gardens may be dubbed with names which are not validly published —
or not published at all. Sobriquets of this kind can seep into print through horticultural periodicals
and nurserymen’s catalogues. Such seepage has no consequences nowadays — after 1 January 1953
publication of names in trade catalogues and non-scientific newspapers does not constitute valid
publication (/.C.B.N. 1988; Art. 29.4), and the inadvertent or unintentional printing of a name is
not effective.

STUART'S HEATHER: DISCOVERY AND NAMING

Erica X stuartii, a naturally-occurring hybrid between FE. mackaiana Bab. and E. tetralix L., is an
example of a wild plant taken into cultivation, informally named by gardeners, and the garden
sobriquet subsequently was effectively published in the horticultural press.

The principal modern accounts (e.g. McClintock 1979, 1980) report that this binomial was
published by the Rev. E. F. Linton in The annals of Scottish natural history in July 1902. But Linton
(1902a) was not the first author to publish a valid Latin name for this heather.

On 11 August 1890, Dr Charles Stuart, a Scottish naturalist of some renown who was visiting
Connemara (Co. Galway, western Ireland, v.c. H16) with other members of the Scottish Alpine
Botanical Club, collected an unusual heather with a strikingly coloured and constricted corolla
(Nelson 1979) in the peatlands near Craiggamore. Other club members also gathered specimens,
and some brought cuttings to Edinburgh, where, because several were skilful gardeners, this
peculiar plant was soon established in cultivation including at the Royal Botanic Garden.

The heather’s discovery was reported at a meeting of the Botanical Society of Edinburgh, chaired
by Robert Lindsay, held on 11 December 1890. At that time Dr J. Muirhead Macfarlane agreed “‘to

276 E. C. NELSON

examine its microscopic structure with a view to assist in its identification” (Craig 1891), but already
was inclined to the view that it was ‘“‘no hybrid but only a form, like E. Mackayi [i.e. E. mackaiana},
of E: Tetralix.4

Macfarlane (1893), having examined Stuart’s heather, decided that it was a subspecies of E.
tetralix and that it deserved a name. Thus, in Transactions and proceedings of the Botanical Society
of Edinburgh, he published the name E. tetralix subsp. stuarti accompanied by a good formal
description to diagnose it from three other subspecies recognized by him. Macfarlane’s name is
valid, even though it is now considered that Stuart’s heather represents a unique clone — in
cultivation it may be deemed a cultivar.

SUBSEQUENT PUBLICATIONS

Eight years later, in The gardeners’ chronicle of 13 July 1901, the editor noted that a Mr Lindsay —
evidently Robert Lindsay, one of the Scottish Alpine Botanical Club members who had been on the
Connemara tour — had sent specimens of E. stuarti (sic); the editorial ({[Masters] 1901) included a
precise description of the specimen. Under the /.C.B.N. this paragraph constitutes a valid
publication of the binomial FE. stuarti, even though the orthography is not in accord with modern
practice. Who was the author; Robert Lindsay, or the editor of The gardeners’ chronicle (Maxwell
T. Masters)? And, was Masters publishing a new name or merely repeating Macfarlane’s name at
species level? |

A few weeks later, on 7 September, a second notice of the plant appeared in The gardeners’
chronicle, this one signed by R. Lindsay (Kaimes Lodge, Murrayfield, Midlothian) and retrospec-
tively dated 4 July (Lindsay 1901).

In April 1902 an obituary of Dr Charles Stuart, who died on 12 February 1902, was published in
The annals of Scottish natural history: Evans (1902) wrote that:

‘“Aquilegia stuarti [sic] commemorates his work in this direction [i.e., crossing species], as does
Erica tetralix stuarti [sic] his discovery of a very distinct subspecies of heath in Connemara.”’
Other obituaries (e.g. in The garden (R. D. 1902)) also included explicit references to the heather.

Several months after Stuart’s death, Linton (1902a) published his article, entitled ‘Erica stuarti,
nov. hybr.’ There is no sense in stretching the interpretation of the /.C.B.N. to confirm Linton’s
(1902a) paper as the original publication of a valid name for the plant found by Dr Stuart. Indeed,
any attempt to promote such an argument would be in error because Linton’s name then becomes
an illegitimate homonym. Thus Linton’s claim to authorship is spurious.

ERICA X STUARTII: AUTHORSHIP

To whom should the name be credited? I have no doubt that Scottish gardeners affectionately
dubbed Dr Stuart’s heath Erica stuarti soon after its discovery and propagation. Furthermore, I
conclude that Macfarlane should be credited with publication of the epithet ‘stuarti’ — the fact that
he identified the plants as a subspecies of E. tetralix reflects only the general state of knowledge of
indigenous heathers during the 1890s. The subsequent use of the epithet at species level — as E.
stuarti— by M. T. Masters in The gardeners’ chronicle does not indicate anything other than that the
name was currently in use. To suggest, as one interpretation of the /.C.B.N. might require, that
Masters’s binomial is a distinct new name is unhelpful, but he did raise the epithet to specific level.

As noted above, Stuart’s Connemara heather is now identified as a mutant clone of the hybrid
between E. tetralix and E. mackaiana, also formerly called E. X praegeri Ostenf. ‘Stuartii’
(McClintock 1979) but correctly (following a change in the /.C.B.N.) E. X stuartii ‘Stuartii’ (and
not, as recently proposed, E. X stuartii ‘Charles Stuart’ (Clarke 1988, [Coombes] 1991). Following
is an expanded bibliographic summary:

- Erica X stuartii (Macfarl.) Mast., The gardeners’ chronicle 30 (ser. 3): 34 (13 July 1901).
Basionym: Erica tetralix L. subsp. stuartii (‘stuarti’?) Macfarl., Transactions and proceedings of the
Botanical Society of Edinburgh 19: 63-64 (1893). -

THE AUTHORSHIP OF ERICA x STUARTII BIT

FicureE 1. Type specimen of Erica tetralix L. subsp. stuartii Macfarl.; designated here as neotype of Erica x
stuartii (Macfarl.) Mast. (Royal Botanic Garden, Edinburgh).

NEortyPus (here designated): “‘Erica sp. - Near Roundstone Connemara [Co. Galway v.c. H16]. . .
E. Tetralix sub-sp. Stuarti, Macf... .”’, 9 August 1890, anon. (E).
Synonym: Erica X praegeri Ostenf., New phytologist 11: 114-127 (1912).

ACKNOWLEDGMENTS

My thanks are due to the Curator of the Herbarium, Royal Botanic Garden Edinburgh, for a loan of
specimens; to Gina Douglas, Librarian at the Linnean Society of London, for help in tracing
bibliographic materials; to Dr R. K. Brummitt, Royal Botanic Gardens, Kew, for his guidance; and
to David McClintock for his comments on an early draft of this paper.

REFERENCES

CiarKE, D. (1988). W. J. Bean’s Trees and shrubs hardy in the British Isles. Supplement. J. Murray, London.
[Coomses, A. J.] (1991). The Hillier manual of trees and shrubs, 6th ed. Hillier Nurseries (Winchester) Ltd,
Romsey.

278 E. C. NELSON

Craic, W. (1891). Excursion of the Scottish Alpine Botanic Club to Connemara, in August 1890. Transactions
and proceedings of the Botanical Society of Edinburgh 19: 15-25.

R.D. (1902). Dr Charles Stuart. The garden 61: 132.

Evans, A. H. (1902). The late Dr. Charles Stuart of Chirnside. The annals of Scottish natural history 11: 65-66.

Linpsay, R. (1901). Erica stuarti. The gardeners’ chronicle 30 (ser. 3): 190.

Linton, E. F. (1902a). Erica stuarti, nov. hybr. The annals of Scottish natural history 11: 176-177.

Linton, E. F. (1902b). A new Erica hybrid. The Irish naturalist 11: 177-178.

McCuintock, D. (1979). The status of, and correct name for, Erica ‘Stuartii’. Watsonia 12: 249-252.

McCuintock, D. (1980). Erica X stuartii E. F. Linton — a correction. Watsonia 13: 59.

MaAcrFaRLANE, J. M. (1893). An examination of some Ericas collected by the Scottish Alpine Botanical Club in
Connemara during 1890. Transactions and proceedings of the Botanical Society of Edinburgh 19: 58-64,
plate 1.

[Masters, M. T.] (1901). Erica stuarti. The gardeners’ chronicle 30 (ser. 3): 34.

NELson, E. C. (1979). Historical records of the Irish Ericaceae, with particular reference to the discovery and
naming of Erica mackaiana Bab. Journal of the Society for the Bibliography of Natural History 9: 289-299.

(Accepted August 1994)

Watsonia 20: 279-284 (1995) 279

Notes

GENTIANELLA CILIATA (L.) BORKH. IN WILTSHIRE (V.C. 8)

While working on Gentianaceae at The Natural History Museum, London (BM) in 1992, T.N.H.
found a previously unrecorded specimen of Gentianella ciliata (L.) Borkh. from South Wilts. (v.c.
8).

This species has only been recorded three times before in the British Isles, two records being from
the same locality in Buckinghamshire (v.c. 24), where it was first collected in 1875 and rediscovered
in 1982, with specimens of both collections being in BM (Knipe 1982; Stace 1991). The third record,
from Surrey (v.c. 17), is supported by a specimen at K, but is considered to be of an alien, rather
than a native, plant (Knipe 1982).

The recently found specimen was collected at Pitton, S. Wilts, ““on Down at junction of chalk and
Tertiary beds”, by E. J. Tatum in September 1892, and was included in the herbarium of A. W.
Bennett.

Edward John Tatum (1851-1929) was a solicitor who was active botanically from c.1880—c.1890.
He is known to have been a diligent and meticulous worker who was responsible for several first
records to the Wiltshire flora. It would appear that he was puzzled by his Gentianella and sent it to
Bennett for comment, for it is one of the few Tatum specimens known to be in BM, the bulk of his
herbarium being untraced (Kent & Allen 1984).

T.N.H. and her Chinese colleagues prefer to treat Gentianella ciliata as a member of the genus
Gentianopsis. If they are followed the species is known as Gentianopsis ciliata (L.) Ma.

This interesting record of a very rare plant provides evidence to support Knipe’s hypothesis that
G. ciliata is native to southern England. It also shows that valuable ‘new’ records can be found even
in comparatively well-known herbaria.

ACKNOWLEDGMENT

We wish to thank Miss A. Hutchinson, B.S.B.I. Recorder for v.c. 8, for information on E. J.
Tatum.

REFERENCES

Kent, D. H. & ALLEN, D. E. (1984). British and Irish herbaria. Botanical Society of the British Isles, London.
KnireE, P. R. (1982). Gentianella ciliata (L.) Borkh. in Buckinghamshire. Watsonia 17: 94.
Stace, C. A. (1991). New Flora of the British Isles. Cambridge University Press, Cambridge.

C. M. DOWLEN

Department of Botany, The Natural History Museum, London, SW7 5BD
T. N. Ho

Northwest Plateau Institute of Biology, Academia Sinica,

78 Xiquan Street, Xining, Qinghai, People’s Republic of China —

A FORGOTTEN SHETLAND DANDELION

In 1907 W. H. Beeby discovered an unusual Taraxacum now classified in section Spectabilia
(Dahist.) Dahlst. near Lang Clodie Loch, North Mainland (GR HU/31.87), in the remote country

280 NOTES

north of Ronas Hill, Shetland (v.c. 112). Late-grown as the material was, it must have struck Beeby
as different from any form of T. faeroense (Dahlst.) Dahlst. (which he knew as T. spectabile
Dahlst.), the common dandelion of rocks, marshes, hillsides, etc., in Shetland. He not only
preserved a specimen of this new dandelion but brought back a root to grow, and was so impressed
by its differences that he described it as a new subspecies of T. spectabile, subsp. geirhildae Beeby
(Beeby 1909).

Beeby was handicapped by making his Shetland visits at a time when most dandelions were past
flowering and by having to base his knowledge of them largely on cultivated plants. His description
was clearly drawn up from material, wild and cultivated, of the Lang Clodie Loch plant; but when he
goes on to say that subsp. geirhildae grows not only near Lang Clodie Loch, but also in parts of
Central Mainland, many miles to the south and with a quite different terrain, he was apparently
confusing forms of T. faeroense with his taxon. At all events no recent botanist has seen the Lang
Clodie Loch plant in Central Mainland, and Beeby’s single specimen from this area (Burn of Quoys,
5 August 1908, SLBI), which is in leaf only, is merely 7. faeroense. Beeby’s epithet is romantic but,
unfortunately, not very appropriate for a Taraxacum of North Mainland: it commemorates Geirhild
(daughter of the Viking pioneering seafarer Floki Vilgerdarson), who is said to have been drowned
in the Loch of Girlsta (Palsson & Edwards 1972), a large loch in Central Mainland which Beeby
believed (wrongly) to be a site of his new taxon. Druce (1922, p. 501) misinterpreted subsp. |
geirhildae, and the records of his own which he cites can only have referred to forms of T. faeroense.
The confusion surrounding subsp. geirhildae may have contributed to the subsequent neglect of this
taxon by British botanists. In 1980, however, its rediscovery by one of us (W.S.) in the type locality
showed clearly that it was something quite distinct from any form of T. faeroense. Richards &
Haworth (1984) at first referred it to the Scandinavian T. ornatum G. Hagl., and it appeared under
that name in our Flora (Scott & Palmer 1987); but this identification, it now appears, was mistaken.
It seems best to treat the Lang Clodie Loch plant as a new species of the section Spectabilia (Dahlst. )
Dahlst., raising Beeby’s subspecies to specific rank; a new combination is therefore called for.

Taraxacum geirhildae (Beeby) R. C. Palmer & W. Scott, comb. et stat. nov. (Fig. 1).

Basionym: Taraxacum spectabile Dahlst. subsp. geirhildae Beeby, Annals of Scottish Natural
History 18: 105 (1909). .

Lectotypus: “‘on the Kattar6énis, e. of Lang Kl6di Loch, Northmaven, Shetland’’, 28 August 1907,
W. H. Beeby 1669 (SLBI). (The specimen on the lower half of the sheet; the upper half contains
material cultivated by Beeby, 1908.)

As Beeby characterised his taxon only very briefly, a fuller description might be helpful.

Plant medium-sized, robust. Leaves prostrate to widely spreading, typically broadly obovate,
(60—)90-100(-140) mm xX (22—)25-28(-30) mm, thick, rough above with very short stout hairs,
yellowish-green to dark apple green, very sparingly and lightly spotted brownish-red, unlobed, with
up to five teeth or small denticulations on either side of the proximal margin, midrib conspicuously
reddish, typically very shortly petiolate, occasionally + wanting. Scape to 18 cm, dull brownish-red.
Exterior bracts adpressed, dark green, ovate-triangular, typically 7-8 x 3 mm, with narrow paler
border. Capitulum deep yellow, 45-50 mm in diameter; ligules striped dark purplish-grey; styles
discoloured (in fresh material); pollen present. Achene 4-3-4-6 mm long, straw-brown, shortly
spinulose in apical quarter, otherwise + smooth.

T. geirhildae resembles T. faeroense but, as Beeby noted, has markedly larger capitula of a darker
yellow, and unlike 7. faeroense is polliniferous. The very lightly spotted leaves are yellowish-green
to dark apple green, firmer in texture than in T. faeroense and distinctly glossy (rather than dull);
they also have shorter petioles and are somewhat different in shape, being obovate (sometimes
broadly so) and never lobed in the wild. (Contrary to Beeby’s statement, we find that in cultivation
T. geirhildae sometimes produces weakly lobed leaves.) The ligule stripe may be best described as a
dark purplish-grey (not pinkish-red as is usual in 7. faeroense). In fresh material of T. geirhildae the
styles are obviously discoloured; in T. faeroense the discoloration is less pronounced. |

Flowering in early June, T. geirhildae is quite frequent in the type locality, on grassy ledges among
rocky outcrops to the south-east of Lang. Clodie Loch and towards the south end of Birka Water.
This attractive dandelion appears to be an endemic Shetland species.

NOTES 281

PAY D- TYPE SPECIMEN
lam ryt 4.

Figure 1. Lectotype of Taraxacum spectabile Dahlst. subsp. geirhildae Beeby (SLBI), basionym of T. geirhildae
(Beeby) R. C. Palmer & W. Scott. As with most of the specimens in Beeby’s North Isles Collection, the label
giving all the details appears on the back of the sheet.

ACKNOWLEDGMENT

Thanks are due to Dr A. J. Richards for his help and encouragement.

REFERENCES

Beesy, W. H. (1909). On the flora of Shetland. Annals of Scottish natural history 18: 103-107.

Druce, G. C. (1922). Flora Zetlandica. Report of the Botanical Society and Exchange Club of the British Isles 6
(supplement): 457-546.

PAtsson, H. & Epwarps, P., eds. (1972). The book of settlements. Landnamabok, p. 17. University of
Manitoba Icelandic Studies, part 1.

RicHarps, A. J. & HAaworty, C. C. (1984). Further new species of Taraxacum from the British Isles. Watsonia
15: 85-94.

Scotr, W. & PALMER, R. [C.] (1987). The flowering plants and ferns of the Shetland Islands, p. 302. The Shetland |
Times Ltd, Lerwick.

O08 6 LS ee ee,

R. C. PALMER
11 Fleet Way, Didcot, Oxon., OX11 8BZ
W. Scotr

New Easterhoull, Castle Street, Scalloway, Shetland Islands, ZE1 0TP

282 NOTES

A NEW SHETLAND HIERACIUM OF THE
SECTION ALPESTRIA [FRIES] F. N. WILLIAMS

The Hieracium described below was first collected in 1952 by the late Professor D. H. N. Spence at
West Burrafirth, West Mainland, Shetland (v.c. 112) (Spence 417, STA, two sheets), and was
subsequently found by W. S. in various places in the West Burrafirth area and in the Norby and
Bousta districts of Sandness, also in West Mainland. When describing H. attenuatifolium Sell & C.
West from the mouth of the Laxo Burn, on the east coast of Mainland and some 20 km E.N.E. of
West Burrafirth, Sell & West (1965) referred W.S.’s material from West Burrafirth and Bousta to
their new species while recognising certain differences. In our Flora (Scott & Palmer 1987) we
included the West Mainland plant under H. attenuatifolium without comment. Doubts as to whether
it was really H. attenuatifolium were raised by R.C.P. after a visit to West Burrafirth in 1986, and
these misgivings were endorsed by J. Bevan when he visited Shetland in 1987. Bevan (1988) later
referred to the West Mainland plant as ‘Taxon A’, an undescribed species. Further careful study of
the plant both in the wild and in cultivation has confirmed us in our view that the West Mainland
plant is fully distinct from H. attenuatifolium; it appears to match no known species, and is therefore
described as a new species.

Hieracium spenceanum W. Scott & R. C. Palmer, sp. nov.

Ho.otypus: common on steep brackeny slopes, head of West Burra Firth, West Mainland, Shetland
Islands, v.c. 112, GR HU/257.567, 5 August 1986, R. C. Palmer S1986/80 (OXF, sheet 1, left-hand
specimen).

Caulis robustus, (10—-)20-40(—80) cm altus, plerumque ubique rubellus sed interdum parte basali
rubellus superne viridis, raro ubique viridis, modice ubique (praesertim inferne) pilosus, parcissime
inferne densius superne floccosus, eglandulosus. Folia saturate viridia, subtus atque interdum supra
purpureo-tincta, marginibus modice pilosis, rubellis. Folia basalia (0Q—)3—S(—7), sailtem nonnulla
florendi et fructificandi tempore paene semper praesentia, eis temporibus bene rosulata; folia
primigenia parva, subrotunda, late elliptica vel ovato-elliptica, + abrupte in petiolum alatum
contracta, apice + late rotundato, saepe minute apiculato, folia posteriora grandiora, alioquin
similia; omnia subintegra vel minute denticulata, subtus parcissime floccosa parce pilosa (pilis
simplicibus in costa numerosioribus), supra modice pilosa, efloccosa. Folia caulina 3-7, plerumque
elliptica, 2-5-plo-—3-5-plo longiora quam latiora, saepe in caule inferiore coacta, infima in petiolum
alatum angustata, mediana et superiora ovato-elliptica, sessilia, semiamplexicaulia; omnia minute
usque distincte et regulariter denticulata, interdum paucis vadosis dentibus utrinque praedita, apice
minuto apiculo acuta, utraque pagina (praesertim subtus) pilis stellatis dispersis vestita, modice
subtus (praesertim in costa) et parce supra pilis simplicibus induta, superiora glabrescentia. Anthela
compacte cymosa, acladio brevi et capitulis (1—-)2—-6(-24), interdum (in plantis luxuriantibus
locorum umbrosorum) ramos longos ex axillis foliorum superiorum emittens. Pedunculi modice
floccosi, necnon pilis aliis simplicibus sparsis pallidis nigribasibus, aliis glanduliferis paucis obscuris
inaequalibus induti. Capitula c. 35 mm diametro, basi subtruncata. Involucri squamae 10-11 mm
longae, 2 mm latae, lineari-lanceolatae, ante anthesin incumbentes, apice obtuso, obscurae, parce
floccosae, parce pilosae pilis pallidis nigribasibus, uberrime pilis glanduliferis obscuris valde
inaequalibus vestitae. Ligulae flavae, apice glabro. Styli mediocriter lividi, id est mediocriter
olivacei. Cypsela 4-5 mm longa, saturate fulva. Receptaculi alveoli margine dentati, dentibus
projecturis brevibus filamentosis armatis.

Stem robust, (10—)20—40(—80) cm high, normally reddish throughout but sometimes reddish at base
and green above, rarely green throughout, moderately pilose throughout (especially below), very
sparingly floccose below but more densely floccose above, eglandular. Leaves deep green, suffused
with purple beneath and sometimes also above, with moderately pilose reddish margins. Basal
leaves (0—)3-5(-7), at least some nearly always present at flowering and fruiting and often forming a
well-developed rosette at those times, the primordial small, subrotund, broadly elliptic or ovate-
elliptic, + rapidly contracted below to a winged petiole, apex + broadly rounded, often minutely
apiculate, the later basal leaves larger but otherwise similar. All basal leaves subentire or finely
denticulate, very sparingly floccose beneath, sparingly pilose beneath (but pilose hairs more
numerous on the midrib), moderately pilose and efloccose above. Stem-leaves 3-7, typically elliptic,
2-5—-3-5 times longer than broad, often concentrated in the lower part of the stem, the lowest

NOTES 283

TABLE 1. A COMPARISON OF THE CHIEF CHARACTERS OF HIERACIUM SPENCEANUM,
H. AUSTRALIUS AND H. ATTENUATIFOLIUM

Character H. spenceanum H. australius H. attenuatifolium
Presence or absence of Present Absent Absent
lower leaves at flowering time
Number of stem-leaves 3-7, 5-10, + 8-10, +
concentrated evenly spaced evenly spaced
towards base
Stem-leaf length/width ratio ISB) 2-5-4 3-25-3-75
Shape of stem-leaves elliptic elliptic widest above
middle
Diameter of capitula 35 mm c. 35 mm 50 mm
Glandular hairs on phyllaries markedly of + uniform markedly
unequal in length unequal in
length length
Style colour medium livid dark livid yellow
Cypsela length 4-5 mm 3-5 mm c.4mm

narrowed to a winged petiole, the median and upper becoming ovate-elliptic, sessile and
semiamplexicaul. All stem-leaves finely to distinctly and + regularly denticulate, sometimes with a
few shallow teeth on each side, apex acute with a minute apiculus, with scattered floccose hairs on
both surfaces (especially beneath), moderately pilose beneath (especially on the midrib), sparingly
pilose above, upper stem-leaves becoming glabrescent above. Inflorescence compactly cymose with
short acladium and (1—)2-6(—24) capitula, sometimes with long branches from the upper leaf-axils in
well-grown plants from sheltered sites. Peduncles moderately floccose with scattered, pale, dark-
based pilose hairs and a few unequal dark glandular hairs. Capitula c. 35 mm diameter, subtruncate-
based. Phyllaries incumbent in bud, 10-11 xX 2 mm, linear-lanceolate, tip obtuse, dark, sparingly
floccose, sparingly pilose (the hairs pale with dark bases), and with abundant very unequal dark
glandular hairs. Ligules yellow, glabrous-tipped. Styles medium livid (medium olive-green).
Cypsela 4:5 mm long, deep reddish-brown. Receptacle pits dentate, the teeth with short
filamentous projections.

H. spenceanum is quite different from H. attenuatifolium; indeed, its affinities lie more with H.
australius (Beeby) Pugsley. The chief characters of the three species are set out in Table 1.

H. attenuatifolium, as here delimited, seems restricted to the mouth of the Laxo Burn, where it
has become almost extinct as a consequence of grazing pressure. All the other stations cited for H.
attenuatifolium by us (Scott & Palmer 1987) in fact belong to H. spenceanum, which occurs in a
handful of stations, usually in small numbers, in the West Burrafirth and Sandness areas, both on
the northern coastal fringe of West Mainland. There is, fortunately, a particularly splendid colony
of H. spenceanum in the type locality, on low rocky sea-banks and brackeny slopes near the head of
West Burra Firth, which may well have been Spence’s station. H. spenceanum, like H. attenuatifo-
lium, is probably endemic to Shetland.

We have named H. spenceanum in honour of the late Professor David Spence of the University of
St Andrews, in recognition of his pioneering work on the ecology of Shetland (in particular his
studies of the serpentine and relict scrub sites), and also, on a more personal note, to record our
appreciation for the help he gave us in many ways in our own studies of the Shetland flora.

ACKNOWLEDGMENTS

Grateful thanks are due to J. Bevan, whose stimulating observations encouraged us to look more
closely at the West Mainland plant we had been calling H. attenuatifolium. Our thanks must also go
to Professor M. B. Usher, who spurred us on to describe the present taxon as a new species.

284 NOTES
REFERENCES

Bevan, J. (1988). A survey of Hieracia (Hawkweeds) in Shetland 1987. Unpublished report to the Nature
Conservancy Council, Peterborough.

Scott, W. & PALMER, R. C. (1987). The flowering plants and ferns of the Shetland Islands, p. 320. The Shetland
Times Ltd, Lerwick.

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

W. ScoTrT
New Easterhoull, Castle Street, Scalloway, Shetland Islands, ZE1 OTP
R. C. PALMER

II Fleet Way, Didcot, Oxon., OX11 8BZ

1g

Watsonia 20: 285-301 (1995) 285

Plant Records

Records for publication must be submitted to the appropriate Vice-county Recorder (see B.S.B.I. Year Book
for 1995), and not the Editors. The records must normally be of species, hybrids or subspecies of native or
naturalized plants belonging to one or more of the following categories: 1st or 2nd v.c. record; 1st 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. 102-104 and 110. Only 1st records can
normally be accepted for Rubus, Hieracium and hybrids. Records for subdivisions of vice-counties will not be
treated separately; they must therefore be records for the vice-county as a whole. Records of Taraxacum are now
being dealt with separately, by Dr A. J. Richards, and will be published at a later date.

Records are arranged in the order given in the List of vascular plants of the British Isles by D. H. Kent (1992),
from which the species’ numbers, taxonomy and nomenclature are taken. The B.S.B.I. has set up a procedure to
register changes to this list, and a number of records of additional species have been held over until the new
arrangements can take effect. 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: 1-6, 9, 11, 13, 14, 17, 22, 24-26, 28,
29, 35, 36, 38, 41, 43-51, 58, 59, 61, 64, 68-70, 72-75, 77-81, 83, 86, 89, 91-93, 96, 98, 99, 101. 104, 108.

The following signs are used:

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

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

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

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

1/4.1. DIPHASIASTRUM ALPINUM 83, Midlothian: Moorland, Mauldslie Hill, NT/316.518.
C. Dixon, 1992, E, det. D. R. McKean. Ist record since 1863.

4/1.1. EQUISETUM HYEMALE *78, Peebless.: Base-rich flushes, Medwyn Mains, NT/12.49.
B. D. Wheeler, 1993, E, det. D. J. McCosh.

4/1.4 X 5. EQUISETUM FLUVIATILE X E. ARVENSE (E. X LITORALE) *24, Bucks.: Edge of lake,
Great Linford Gravel Pits, SP/836.430. R. Maycock, 1989, det. C. N. Page.

4/1.4 X 8. EQUISETUM FLUVIATILE X E. PALUSTRE (E. X DYCEI) *104, N. Ebudes: Abhainn
a’Ghlinne, Dalavil, Skye, NG/60.05. M. Veit, 1991, herb. M.V.

4/1.5 X 8. EQUISETUM ARVENSE X E. PALUSTRE (E. X ROTHMALERI) *101, Kintyre: Shore just
above water level, Aucha Lochy, NR/726.228. C. D. Preston & N. F. Stewart, 1988, E, det. C. N.
Page.

5/1.1. OPHIOGLOSSUM VULGATUM 73, Kirkcudbrights.: Grassland, I.C.I. factory, Cargen-
bridge, NX/946.747. O. M. Stewart, 1993. 2nd extant locality.

+8/1.1. PTERIS CRETICA *5, S. Somerset: Porlock Wood, Porlock, SS/877.467. P. Green,
1983: *29, Cambs.: Hole in brick wall, Market Passage, Cambridge, TL/449.585. C. D. Preston,
1993, CGE, conf. J. R. Akeroyd.

11/1.1 x 2. PoLYPODIUM VULGARE X P. INTERJECTUM (P. X MANTONIAE) *28, W. Norfolk:
Ditch-side, East Winch, TF/704.159. H. Williamson, 1993, BM, det. A. C. Jermy.

14/1.1. THELYPTERIS PALUSTRIS 41, Glam.: Fen meadow near Cefn Cribwr, SS/8.8.
J. Woodman, 1993, NMW, det. G. Hutchinson. Only extant locality.

15/2.4. ASPLENIUM MARINUM 81, Berwicks.: Sandstone rocks near Ladykirk, NT/8.4. M. E.
Braithwaite, 1993. Confirmation of an 1838 record at an inland site 14 km from sea.

286 PLANT RECORDS

15/2.5a. ASPLENIUM TRICHOMANES subsp. TRICHOMANES *50, Denbs.: River gorge, Ysbytty
Ifan, SH/817.458. J. A. Green, 1993.

16/3.3. ATHYRIUM FLEXILE 92, S. Aberdeen: Near Derry Cairngorm, NO/0.9. D. J. McCosh,
C. A. Stace & D. J. Tennant, 1993, BM, conf. A. C. Jermy & A. Paul. 2nd record.

16/4.2. GYMNOCARPIUM ROBERTIANUM *46, Cards.: Overgrown stoney mounds, Mynach Vale
lead mine, SN/772.775. D. K. Reed, 1993, NMW.

+17/2.1. CYRTOMIUM FALCATUM *11, S. Hants.: Wall of dock, Portsmouth Naval Base, SU/
628.003. D. E. Bulloch, 1992, herb. R. P. Bowman, det. E. J. Clement.

17/3.5. DRYOPTERIS AEMULA 41, Glam.: Sandstone cliff in oak woodland near Blackmill,
Bridgend, SS/9.8. P.S. Jones, N. Ward & A. J. Williams, 1993, NMW, det. G. Hutchinson & Q. O.
N. Kay. Only extant locality. *73, Kirkcudbrights.: Shaded rocks, Castramont Burn, NX/
592.617. R. Tydewell, 1989, refound by P. Ripley & O. M. Stewart, 1993.

17/3.8 X 9. DRYOPTERIS CARTHUSIANA X D. DILATATA (D. X DEWEVERI) *50, Denbs.: Shore of
Alwen Reservoir, SH/927.554. G. Battershall, 1993, det. A. C. Jermy.

719/1.1. AZOLLA FILICULOIDES *46, Cards.: Ditch E. of Llangorwen church, SN/603.838. S. P.
Chambers, 1993. Covering surface of fish pond, Llain-wen, SN/392.550. R. J. Williams, 1993,
NMW. Ist and 2nd records.

27/1.1. CERATOPHYLLUM DEMERSUM *99, Dunbarton: Forth & Clyde Canal between Temple
and Kelvindale, NS/55.69. K. J. Watson & K. J. Murphy, 1988.

+28/6.1 X var. ACONITUM NAPELLUS X A. VARIEGATUM (A. X CAMMARUM) "1, Wallankor
Trackside, Craigend, NS/82.33. P. Macpherson, 1993, herb. P.M.

28/13.4. RANUNCULUS SARDOUS 38, Warks.: Edge of cereal crop, Wellesbourne, SP/286.542.
M. J. Senior, 1992, WAR, det. J. M. Mullin. 1st post-1930 record. 50, Denbs.: Gravel near the
sea, Foryd, Rhyl, SH/991.808. J. A. Green, 1993. 2nd extant locality. 75, Ayrs.: By muddy farm
track, Standard Farm, Barrhill, NX/264.818. J. McCleary, 1993, GLAM, conf. A. McG. Stirling.
2nd, and Ist recent, record.

28/13.10. RANUNCULUS AURICOMUS 108, W. Sutherland: Deciduous woodland, Strath Beag,
Loch Eriboll, NC/390.514. M. Barron, 1993. 2nd record.

28/15.1. MYOSURUS MINIMUS 3, S. Devon: Hundreds of plants on damp edge of green lane
near Colyton, SY/23.95. L. J. Margetts, 1991. 1st post-1930 record.

28/17.3. THALICTRUM FLAVUM *45, Pembs.: Phragmites australis-dominated fen, Castlemartin
Corse, SR/900.997. J. O. Mountford, 1984.

30/1.5b. PAPAVER DUBIUM subsp. LECOQII 46, Cards.: Vegetated waste ground, Penrhyn-coch,
SN/638.839. S. P. Chambers, 1993. 2nd record.

30/1.6. PAPAVER HYBRIDUM 5, S. Somerset: Arable field, Kilton, Nether Stowey, ST/160.451.
I. Green, 1993. 1st record since 1897.

30/5.1. CHELIDONIUM MAJUS 781, Berwicks.: Woodland edge by road near Lennel, NT/
849.406. M. E. & P. F. Braithwaite, 1993. Only extant locality.

31/4.1. CERATOCAPNOS CLAVICULATA 104, N. Ebudes: Boulder scree N. of Eyre, Raasay, NG/
577.351. S. J. Bungard, 1993, conf. C. W. Murray. Only extant locality on Raasay.

31/5.3. FUMARIA BASTARDII 79, Selkirks.: Disturbed ground, Boleside, Galashiels, NT/
500.335. M. E. Braithwaite, 1992, herb. R. W. M. Corner, det. M. G. Baker. 2nd record, Ist since
1930.

31/5.8. FUMARIA DENSIFLORA 61, S.E. Yorks.: Arable field on chalk, Etton Wood, SE/
945.433. J. Dews, 1993, det. M. G. Daker. Only extant locality and 1st record of a native
population. :

PLANT RECORDS 287

40/2.2. ALNUS INCANA *77, Lanarks.: Marshy ground, Hamilton, NS/72.55. P. Macpherson,
1993, herb. P.M.

+43/1.3. CHENOPODIUM CAPITATUM *13, W. Sussex: Maize field S. of Coates Common, SU/
998.172. H. W. Matcham & N. J. H. Sturt, 1993, det. J. M. Mullin.

43/1.6. CHENOPODIUM RUBRUM 46, Cards.: Abundant on manure heap, Tygwyn, SH/523.647.
A. O. Chater, 1993. 2nd post-1930 record.

43/3.2. ATRIPLEX PROSTRATA 81, Berwicks.: Verge of A697 near Crook’s Lodge, NT/815.405.
M. E. Braithwaite, 1993, herb. M.E.B. Ist inland record.

43/3.2 X 4. ATRIPLEX PROSTRATA X A. LONGIPES (A. X GUSTAFSSONIANA) *46, Cards.: Bare
disturbed mud in saltmarsh, W. bank of Afon Leri, Ynys-las, SN/616.934. A. O. Chater, 1993,
NMW.

43/3.5. ATRIPLEX PRAECOX 104, N. Ebudes: Seashore, Eilean Tigh, Raasay, NG/609.533. S. J.
Bungard, 1990, E, det. D. R. McKean. 1st Raasay record.

43/3.6. ATRIPLEX LITTORALIS *35, Mons.: Margin of old sludge pond, Alpha Steel Works,
Newport, ST/338.848. T. G. Evans & M. Jones, 1993, NMW. *46, Cards.: Bare disturbed mud
in saltmarsh, W. bank of Afon Leri, Ynys-las, SN/616.934. A. O. Chater, 1993, NMW. Stoney
ground at top of saltmarsh, E. side of Aberystwyth harbour, SN/582.811. A. O. Chater, 1993,
NMW. Ist and 2nd records.

43/3.10. ATRIPLEX PORTULACOIDES +*46, Cards.: Edges of creeks in saltmarsh, Dyfi estuary,
SN/641.941. A. O. Chater, 1993, NMW. Planted in 1950s by E. H. Chater, now naturalized over
several hectares.

43/8.1. SALSOLA KALI 104, N. Ebudes: Sandy shore, Laig, Eigg, NM/46.88. J. Bevan, P. F.
Braithwaite & C. W. Murray, 1992. Only extant locality on Egg.

46/1.1c. ARENARIA SERPYLLIFOLIA subsp. LEPTOCLADOS *81, Berwicks.: Ha-ha, The Hirsel,
NT/828.409. M. E. Braithwaite, 1993, herb. M.E.B., det. G. Halliday.

46/4.2. MINUARTIA VERNA +*41, Glam.: Site of abandoned experimental plots on smelter
waste tips, Pentre-chwyth, Swansea, SS/669.953. S. J. Wainwright, 1993, conf. C. R. Hipkin.
Probably a deliberate but unrecorded introduction.

46/7.7b. CERASTIUM FONTANUM Subsp. HOLOSTEOIDES *73, Kirkcudbrights.: Damp rock ledge
near Door of Cairnsmore of Fleet, NX/512.643. O. M. Stewart, 1993.

46/7.12. CERASTIUM SEMIDECANDRUM 79, Selkirks.: Top of old wall, Newark Castle, Yarrow,
NT/422.294. R. W. M. Corner, 1993, herb. R.W.M.C. 2nd record.

46/9.1. MOENCHIA ERECTA 44, Carms.: Dry S.-facing rocky bank, Cwm Crychan, SN/820.390.
I. K. Morgan, 1992. 2nd extant locality.

746/18.1. LYCHNIS CORONARIA *46, Cards.: Vegetated ballast of long-disused railway, site of
Llanfarian station, SN/591.778. S. P. Chambers, 1993. Well established and persistent.

147/1.7. PERSICARIA AMPLEXICAULIS *77, Lanarks.: Rough grassy bank, Cessnock, Glasgow,
NS/56.64. E. K. Lindsay & P. Macpherson, 1993, herb. P.M.

47/1.8. PERSICARIA VIVIPARA 73, Kirkcudbrights.: In two flushes, Cairnsmore of Fleet, NX/
512.653 & 517.644. D. Hawker, 1992. 2nd record.

747/5.1 X 2. FALLOPIA JAPONICA X F. SACHALINENSIS (F. X BOHEMICA) *3, S. Devon: Edge of
carpark, Trenchford Reservoir, SX/804.824. L. J. Margetts & W. H. Tucker, 1992, herb.
L.J.M. *69, Westmoriand: Beck Head, Witherslack, SD/446.846. T. C. G. Rich & M. Baecker,
1991, LANC, det. A. P. Conolly & J. P. Bailey.

47/4.6. POLYGONUM RURIVAGUM *68, Cheviot: Gravel by road near roundabout on Al,
Berwick, NT/990.547. M. E. Braithwaite, 1992, herb. G. A. Swan, det. J. R. Akeroyd.

288 PLANT RECORDS

47/8.8 X 13. RUMEX LONGIFOLIUS X R. cRISPUS (R. X PROPINQUUS) *79, Selkirks.: With
parents on waste ground, A7 3 km S. of Selkirk, NT/468.250. R. W. M. Corner, 1993, herb.
R.W.M.C., det. J. R. Akeroyd.

47/8.15 X 19. RUMEX SANGUINEUS X R. OBTUSIFOLIUS (R. X DUFFTII) *46, Cards.: With
parents on waste ground, Glanyrafon Industrial Estate, Llanbadarn Fawr, SN/611.803. A. O.
Chater, 1992, NMW, det. J. R. Akeroyd.

47/8.21. RUMEX MARITIMUS 81, Berwicks.: Muddy edge of pool in fen, Lithtillum Loch, NT/
80.40. M. E. Braithwaite, 1993. Only extant locality.

50/1.1. ELATINE HEXANDRA *96, Easterness: Loch Dochfour, NH/607.387. N.C.C. Loch
Survey Team, 1988.

51/1.6 X 7. HYPERICUM PERFORATUM X H. MACULATUM subsp. OBTUSIUSCULUM (H. X DESETANGSII
nothosubsp. DESETANGSII) *24, Bucks.: Calcareous bank of disused railway, Buckingham, SP/
698.330. R. Maycock & A. Woods, 1989, det. N. K. B. Robson. *44, Carms.: Hay meadow,
Pencarreg, SN/567.460. H. J. Killick, 1985, NMW. Roadside, farm above Goitre, SN/373.234.
J. Bevan, H. J. Killick & M. E. Smith, 1992, NMW. Ist and 2nd records, both det. N. K. B. Robson.

52/1.1. TILIA PLATYPHYLLOS 50, Denbs.: Hedgerow, Colwyn Bay, SH/835.809. G. Battershall,
1993. 2nd record.

+53/1.6. MALVA PUSILLA 4, N. Devon: Recently disturbed area, Kenwith Nature Reserve,
Bideford, SS/448.272. W. H. Tucker, 1993. 1st post-1930 record.

53/2.1. LAVATERA ARBOREA 50, Denbs.: Hedgerow, Llysfaen, SH/899.784. G. Battershall,
1993. 2nd record. 51, Flints.: Disturbed ground near dunes, Ffrith Beach, Prestatyn, SJ/
047.833. J. Phillips, 1992. 1st post-1930 record.

53/2.2. LAVATERA CRETICA +*3, S. Devon: Waste ground, Topsham Quay, SX/966.877. L. J.
Margetts, 1991, BM, det. N. K. B. Robson.

754/1.1. SARRACENIA PURPUREA 98, Main Argyll: In Sphagnum at margin of bog pools,
Rannoch Moor, NN/3.4. J. Rees, 1992. 31 plants counted by G. P. Rothero & B. H. Thompson, 13
July 1993.

59/1.1. FRANKENIA LAEVIS +*3, S. Devon: Face of low cliff, Lympstone, SX/989.838.
R. Lawrence, 1986.

61/2.5 X 9 X 11. SALIX PURPUREA X S. VIMINALIS X S. CINEREA-(S. X FORBYANA) *77, Lanarks.:
Shingle, Glengavel Water, NS/64.36. P. Macpherson, 1993, herb. P.M., det. R. D. Meikle.

761/2.6. SALIX DAPHNOIDES *77, Lanarks.: Grassy river-bank, Meadowside, NS/55.66.
P. Macpherson, 1991, herb. P.M., det. R. D. Meikle.

61/2.9 X 10. SALIX VIMINALIS X S. CAPREA (S. X SERICANS) *75, Ayrs.: Gailes Marsh, Irvine,
NS/32.36. K. J. Watson, 1992.
61/2.9 X 16. SALIX VIMINALIS X S. REPENS (S. X FRIESIANA) *69, Westmorland: Sandy car-

park, Sandscale, SD/200.757. P. Burton, 1993, LANC, det. R. D. Meikle. 3rd British record.

61/2.10b. SALIX CAPREA subsp. SPHACELATA *78, Peebless.: Steep hill slope, Talla Linns, NT/
138.202. N. P. Ashmole, 1993, E, det. D. R. McKean & R. D. Meikle.

61/2.10 xX 14. SALIX CAPREA X S. MYRSINIFOLIA (S. X LATIFOLIA) *104, N. Ebudes: Rocky
outcrop below bridge over R. Lealt, Skye, NG/516.604. C. Westall, 1989, E, det. R. D. Meikle.

61/2.11 X 16. SALIX CINEREA X S. REPENS (S. X SUBSERICEA) *11, S. Hants.: Heathland,
Whitten Bottom, SU/204.006. G. H. Forster, 1993, herb. R. P. Bowman, det. R. D. Meikle.

61/2.14 xX 15. SALIX MYRSINIFOLIA X S. PHYLICIFOLIA (S. X TETRAPLA) *93, N. Aberdeen:
Roadside by moorland, Glass, NJ/445.382. D. Welch, 1993, ABD, det. R. D. Meikle.

+62/1.6. SISYMBRIUM ORIENTALE ~~ *99, Dunbarton: Waste ground near river bank, Dumbarton
Common, NS/39.76. A. McG. Stirling, 1993, GLAM.

PLANT RECORDS 289

+62/9.1. MALCOLMIA MARITIMA *49, Caerns.: Marine embankment below Mor Hel Hotel,
Criccieth, SH/498.377. R. E. Hughes, 1987, NMW.

62/10.1. MATTHIOLA INCANA 746, Cards.: Well naturalized on vegetated shingle, Borth, SN/
608.891. A. O. Chater, 1993, NMW. 2nd record, Ist since 1930.

62/12.3. RORIPPA ISLANDICA *44, Carms.: River shingle, River Bran N. of Llandovery, SN/
783.362. R. D. Pryce, 1981, NMW. Muddy hollow in silage field near Newcastle Emlyn Castle, SN/
319.408. A. O. Chater & T. C. G. Rich, 1993, NMW. Ist and 2nd records, both det. T. C. G.
Rich. 46, Cards.: Muddy hollow in old ox-bow near Newcastle Emlyn Castle, SN/312.405.
A. O. Chater & T. C. G. Rich, 1993, NMW, det. T. C. G. Rich. 2nd record.

62/12.5 X 6. RORIPPA SYLVESTRIS X R. AMPHIBIA (R. X ANCEPS) *61, S.E. Yorks.: Winter-
fiooded marshy grassland, Wheldrake Ings, SE/704.438. C. D. Preston, 1993, CGE.

+62/14.4. CARDAMINE RAPHANIFOLIA 79, Selkirks.: Edge of R. Yarrow below Newark Castle,
NT/422.293. R. W. M. Corner, 1993, herb. R.W.M.C. 2nd record.

62/21.4. DRABA MURALIS +79, Selkirks.: Old walls, Newark Castle, Yarrow, NT/422.294.
R. W. M. Corner, 1993, herb. R.W.M.C. Introduced as seed 10 years ago. Ist record since 1875.

62/22.3. EROPHILA GLABRESCENS 46, Cards.: Rabbit-grazed sandy grassland, Penyrergyd
dunes, SN/162.485. A. O. Chater, 1991, NMW, det. T. T. Elkington. 2nd record. 13:
Kirkcudbrights.: Crevices in road and pavement, Black Water Bridge, NX/616.883. O. M. Stewart,
1993:

apa Goen BAKA OFFICINALIS 81, Berwicks.: Bank of Al, Bowshiel Wood, NT/794.670.
M. E. & P. F. Braithwaite, 1993. Ist inland record.

62/23.5. COCHLEARIA DANICA 81, Berwicks.: Verge of Al, Brockholes, NT/823.648. M. E. &
P. F. Braithwaite, 1993. 1st inland record.

62/27.1. TEESDALIA NUDICAULIS *81, Berwicks.: Near Gaitheugh, NT/5.3. A. Brotherston,
1877, BM.

62/31.1. CORONOPUS SQUAMATUS *77, Lanarks.: Rough bank, Coatbridge, NS/71.67.
P. Macpherson, 1993, herb. P.M.

+62/34.4. BRASSICA JUNCEA 41, Glam.: Flower-bed, Hill’s Street, The Hayes, Cardiff, ST/
186.764. K. L. Davies, 1986, NMW, det. T. C. G. Rich. Still present in 1993. Only extant locality.

+62/37.1b. COINCYA MONENSIS subsp. RECURVATA *58, Cheshire: Waste ground, Bromborough
Dock, SJ/346.854. V. Gordon & K. Watson, 1992, herb. G. M. Kay, det. T. C. G. Rich.

62/41.1. CRAMBE MARITIMA 75, Ayrs.: Shingle N. of Girvan Mains, NS/19.00. A. McG. Stirling
et al., 1993. 2nd extant locality.

65/10.2. ARCTOSTAPHYLOS ALPINUS 104, N. Ebudes: Windswept ridge, Beinn Bhuidhe, Skye,
NG/77.21. C. W. Murray, 1990, herb. C.W.M. Ist record since 1772 record from same area.

69/6.3. ANAGALLIS MINIMA’ 104, N. Ebudes: Rubha na Cloiche, NG/565.337, & near Eyre,
NG/572.339, Raasay. S. J. Bungard, 1993. Ist Raasay records.

+72/1.1. ESCALLONIA MACRANTHA *46, Cards.: Self-sown plants on cliff, E. side of Ystwyth
estuary, Aberystwyth, SN/580.806. A. O. Chater, 1993, NMW.

73/1.1. CRASSULA TILLAEA *13, W. Sussex: Sandy arable field near Bignor Park Cottage, SU/
990.169. M. Edwards, 1993, conf. M. Briggs.

+73/1.3. CRASSULA HELMSII *35, Mons.: Edge of stream, Trethomas, ST/187.886. J. Wood-
man, 1992, NMW. Margin of borrow pit, Monmouth, SO/503.127. J. F. Harper, 1992, NMW. ist
and 2nd records, both conf. T. G. Evans. *49, Caerns.: Pond, Port Meirion village, SH/58.37.
R. G. Ellis, 1993. *91, Kincardines.: Pool in rocks above high water-mark, Cove Bay, NJ/
957.007. D. Welch, 1993, ABD. *104, N. Ebudes: Pond in grounds of Skeabost House Hotel,
Skye, NG/413:.487. C. W. Murray, C. D. Preston & N. F. Stewart, 1989, herb. C.W.M.

290 PLANT RECORDS

+73/5.8. SEDUM STOLONIFERUM *4, N. Devon: Roadside verge under trees, Ayshford, ST/
045.160. L. J. Margetts & W. H. Tucker, 1990, det. H. J. M. Bowen.

+73/5.17. SEDUM DASYPHYLLUM 50, Denbs.: Rhos-on-sea, SH/843.808. G. Battershall, 1993.
2nd extant locality.

+74/1.2. ASTILBE RIVULARIS “98, Main Argyll: Clearing in estate woodland N. of Inveraray
Castle, NN/096.099. B. H. Thompson, 1993, BM, det. J. M. Mullin.

+74/5.8 X 9. SAXIFRAGA UMBROSA X S. SPATHULARIS (S. X URBIUM) 50, Denbs.: Damp wood,
Nant y Garth Pass, Ruthin, $J/150.518. J. A. Green, 1993. Only extant locality.

+75/1.2. SORBARIA TOMENTOSA *29, Cambs.: Old wall, Humberstone Road, Cambridge, TL/
458.595. G. M. S. Easy, 1993, herb. G.M.S.E.

+75/4.1. ARUNCUS DIOICUS *47, Monts.: Meadow below Powis Castle, Welshpool, SJ/
216.063. G. Hutchinson, 1989. 1st Welsh record.

75/8.14. RUBUS ARRHENIIFORMIS *9, Dorset: Common land, Lambert’s Castle, SY/36.98. L. J.
Margetts, 1991, herb. L.J.M., det. A. Newton. *28, W. Norfolk: Wet heath and scrub, East
Winch, TF/70.14. A. L. Bull, 1993, conf. A. Newton.

75/8.27. RUBUS PLICATUS *104, N. Ebudes: Skeabost, Skye, NG/41.48. A. Newton, 1991,
herb. A.N.

75/8.50. RUBUS EBUDENSIS *104, N. Ebudes: Side of B884, Colbost, Skye, NG/212.492.
A. Newton, 1991, herb. A.N.

75/8.51. RUBUS ERRABUNDUS *2, E. Cornwall: Scrub on heathy hill, Kit Hill, SX/376.715.
R. W. Gould, 1993, det. A. Newton.

*75/8.55. RUBUS LACINIATUS *75, Ayrs.: Daljarrock, Colmonell, NX/1.8. R. C. L. Howitt,
1970, GLAM.

75/8.68. RUBUS PERDIGITATUS *3, S. Devon: Ride in forestry plantation, Trinity Hill,
Axminster, SY/305.954. L. J. Margetts, 1992, herb. L.J.M., det. A. Newton. .

75/8.76. RUBUS PYRAMIDALIS *104, N. Ebudes: Kyle Farm road, Skye, NG/74.25. A. Newton,
1991, herb. A.N.

75/8.82. RUBUS SILURUM *69, Westmorland: Single large bush by side of farm road to
Beckside Hall, Middleton, SD/628.882. G. Halliday, 1993, LANC, conf. A. Newton. Northernmost
known locality.

75/8.90. RUBUS AMPLIFICATUS *80, Roxburghs.: Railway S. of Newcastletown, NY/4.8. C. W.
Muirhead, 1969, GLAM, det. A. McG. Stirling.

75/8.92. RUBUS BOUDICCAE *9, Dorset: Woodbury Hill near Bere Regis, SY/85.94. M. Porter,
1991

75/8.93. RUBUS CARDIOPHYLLUS *2, E. Cornwall: Scrub, Red Lake Reserve between Lostwit-
hiel and Lerryn, SX/12.58. A. Newton, 1988.

+75/8.102. RUBUS ELEGANTISPINOSUS *69, Westmorland: Hedge, Arrad Foot, Greenodd, SD/
306.813. G. Halliday, 1993, LANC, det. A. Newton.

75/8.111. RUBUS MILFORDENSIS *11, S. Hants.: Single bush by gateway by Dunwood near
Romsey, SU/306.230. D. E. Allen, 1985, BM, conf. A. Newton.

75/8.116. RUBUS PISTORIS *9, Dorset: Lightly wooded heath, Bewley Down, ST/28.06. L. J.
Margetts, 1988, herb. A. Newton, det. A.N.

75/8.125. RUBUS SUBINERMOIDES . *104, N. Ebudes: Portree, Skye, NG/47.43. A. Newton,
199], herb. A.N.

PLANT RECORDS 291

+75/8.134. RUBUS ARMENIACUS *70, Cumberland: Verge of lane W. of Renwick, NY/575.440.
G. Halliday, 1993, LANC, conf. A. Newton. *75, Ayrs.: Embankment of disused railway,
Parkhouse Community Nature Reserve, Ardrossan, NS/23.42. A. McG. Stirling, 1993.

75/8.140. RUBUS ROSSENSIS *2, E. Cornwall: Path by Lower Tamar Lake, SS/294.110. W. H.
Tucker, 1993, conf. A. Newton.

75/8.143. RUBUS WINTERI *3, S. Devon: Under roadside trees near Exmouth, SY/02.83. R. D.
Randall, 1991, conf. A. Newton.

75/8.144. RUBUS ADSCITUS *9, Dorset: Hedgerow, Hawkchurch, SY/34.99. L. J. Margetts,
1988, herb. A. Newton, det. A.N.

75/8.159. RUBUS ORDOVICUM *50, Denbs.: Hedge, Capel Garmon, SH/812.582. G. Batter-
shall, 1993, conf. A. Newton.

75/8.169. RUBUS FUSCICORTEX *50, Denbs.: Hedge, Capel Garmon, SH/812.582. G. Batter-
shall, 1993, conf. A. Newton.

75/8.176. RUBUS AEQUALIDENS *3, S. Devon: W. of Dawlish, SX/93.76. R. D. Randall, 1991,
herb. R.D.R., conf. A. Newton.

75/8.188. RUBUS HETEROBELUS *3,S. Devon: Lympstone Common, SY/03.85. R. D. Randall,
1991, herb. R.D.R., conf. A. Newton.

75/8.199. RUBUS RADULOIDES *70, Cumberland: Hedge by road W. of Rose Castle, Nether
Welton, NY/363.464. G. Halliday, 1993, LANC, det. A. Newton.

75/8.239. RUBUS INSECTIFOLIUS *2, E. Cornwall: Scrub, Red Lake Reserve between Lostwit-
hiel and Lerryn, SX/12.58. A. Newton, 1988.

75/8.242. RUBUS LONGITHYRSIGER *9, Dorset: Woodland, Bewley Down, ST/28.06. L. J.
Margetts, 1988, herb. A. Newton, det. A.N.

75/8.243. RUBUS MALVERNICUS *3, S. Devon: Shady roadside, Venn Ottery, SY/064.915.
R. Gould, L. J. Margetts & R. D. Randall, 1992, herb. L.J.M., det. A. Newton.

75/8.253. RUBUS RUDIS *28, W. Norfolk: Conifer plantation, Shouldham Warren, TF/66.10.
A. L. Bull, 1993, conf. A. Newton.

75/8.280. RUBUS PHAEOCARPUS *9, Dorset: Footpath across Warmwell Heath, SY/758.872.
D. E. Allen, 1993, BM, conf. A. Newton.

75/8.286. RUBUS SCABRIPES *17, Surrey: One large bush on margin of woodland, Holmbury
Hill, TQ/098.432. Surrey Fiora Committee meeting, 1993, det. D. E. Allen & A. Newton.

75/8.312. RUBUS LATIFOLIUS *104, N. Ebudes: Roadside verge, Elgol, Skye, NG/528.146.
M. Gregory, 1989, herb. C. W. Murray, det. A. Newton.

75/8.319. RUBUS TUBERCULATUS *77, Lanarks.: Grassy waste ground, Kenmuir, Glasgow, NS/
66.62. P. Macpherson & A. McG. Stirling, 1993, herb. P.M.

+75/9.9. POTENTILLA NORVEGICA 80, Roxburghs.: Waste ground by the Slitrig, Whitlaw,
Hawick, NT/499.134. R.W. M. Corner, 1992, herb. R.W.M.C. 2nd record.

75/9.13b. POTENTILLA ERECTA subsp. STRICTISSIMA *3, S. Devon: Gorse scrub, Hackpen Hill,
Culmstock, ST/111.119. Gorse scrub, North Hill near Broadhembury, ST/096.065. Both L. J.
Margetts, 1993, NCE, det. A. J. Richards. 1st and 2nd records. *4, N. Devon: Rough grassland, .
Marshall Farm estate, Woolfardisworthy, SS/310.185. M. Bristow & W. H. Tucker, 1993, herb.
L. J. Margetts, conf. L.J.M. *5,S. Somerset: Foches Corner, West Buckland, ST/183.172. L. J.
Margetts, 1993, det. A. J. Richards. *46, Cards.: With subsp. erecta in dense Calluna vulgaris/
Empetrum nigrum heath, Draws Drum E. of Ponterwyd, SN/797.805. Bank in conifer forest,
Myherin valley E. of Ysbyty Cynfyn, SN/772.790. Both A. O. Chater, 1991, NMW, conf.
B. Harold. ist and 2nd records.

292 PLANT RECORDS

75/9.14. POTENTILLA ANGLICA *93, N. Aberdeen: Grassland on abandoned railway, Ellon, NJ/
947.305. D. Welch, 1993, ABD.

75/9.14 X 15. POTENTILLA ANGLICA X P. REPTANS (P. X MIXTA) *46, Cards.: Laneside grassy
bank, Felin Ganol, Penparc, SN/199.487. A. O. Chater, 1991, NMW. Mown grassland, R.A.E.
site, Aberporth, SN/241.522. A. O. Chater, 1992, NMW. Ist and 2nd records, both conf. B. Harold.

75/13.3. GEUM MACROPHYLLUM *46, Cards.: Abundant in wooded dingle and on roadside
banks and verges, Cwmyrolchfa N. of Bronnant, SN/640.688. G. Jones, 1993, NMW, det. A. O.
Chater. Known here to site owner for 40 years.

75/15.2. AGRIMONIA PROCERA 28, W. Norfolk: Pasture, Larling, TL/963.903. J. Humphris,
1993. Only extant locality. +104, N. Ebudes: Edge of woodland path, Cuillin Hills estate,
Portree, Skye, NG/48.43. J. E. Duncan & H. Lefevre, 1982, herb. C. W. Murray. Ist Skye record.

+75/18.1. ACAENA NOVAE-ZELANDIAE *46, Cards.: Shaley slopes under pines, U.C.W.
Campus, Aberystwyth, SN/597.817. S. P. Chambers, 1993.

$75/19.5. ALCHEMILLA TYTTHANTHA *77, Lanarks.: Rough grassland, Cuningar Loop, Glas-
gow, NS/62.63. K. J. Watson, 1990, GL, conf. S. M. Walters.

+75/21.8. ROSA GLAUCA *17, Surrey: Self-sown in rough grass, Bagshot Heath, SU/903.632.
S. Berrett & K. Page, 1993. :

75/21.11 X 12. ROSA STYLOSA X R. CANINA (R. X ANDEGAVENSIS) *17, Surrey: Scrub around
old farm pond, Dormansland, TQ/428.404. G. Fookes, 1993, det. A. L. Primavesi.

75/21.12 X 14. ROSA CANINA X R. OBTUSIFOLIA (R. X DUMETORUM) *25, E. Suffolk: Sea-wall,
Buss Creek Marshes, Southwold, TM/495.761. C. D. Preston & S. E. Yates, 1992, det. A. L.
Primavesi. :

75/21.17. ROSA MOLLIS *75, Ayrs.: Garnock Floods, Irvine, NS/30.41. K. J. Watson, 1992.
Sandy waste ground, I.C.I. Nobel Works site, Ardeer, NS/28.40. A. McG. Stirling, 1993. Ist and
2nd records.

75/21.19. ROSA MICRANTHA *26, W. Suffolk: Roadside verge near Mayday Farm, Brandon,
TL/796.833. C. D. Preston, 1992, det. A. L. Primavesi. 1st post-1930 record.

+75/22.7. PRUNUS CERASUS 50, Denbs.: Hedge, Cefn Berain, SH/979.687. J. A. Green, 1993.
Only extant locality. ;

+75/22.13. PRUNUS LUSITANICA *49, Caerns.: Bird-sown plants in grassland, Brynian Farm
near Llanbedr-y-Cennin, SH/785.719. R. Lewis, 1993, NMW.

75/28.24. SORBUS TORMINALIS 41, Glam.: Carboniferous limestone cliff, Castell Coch wood-
lands, ST/128.828. G. M. Barter, 1992. 2nd extant locality.

+75/32.5. COTONEASTER AFFINIS *3, S. Devon: Edge of East Budleigh Common, SY/040.847.
L. J. Margetts, 1991, herb. L.J.M., det. J. Fryer & J. R. Palmer. *46, Cards.: Ty-rhos near
Plwmp, SN/371.513. M. Evans, 1978, NMW, det. J. Fryer. Still present 1993, when dominant in
roadside hedges and with abundant self-sown seedlings.

+75/32.10. COTONEASTER SALICIFOLIUS 41, Glam.: Railway cutting, Gabalfa, ST/166.791.
G. Hutchinson, 1986, NMW, det. J. Fryer. 2nd record. *47, Monts.: Scrub on limestone,
Llanymynech Rocks, SJ/260.217. J. Pedlow & M. Wainwright, 1989, det. J. Fryer.

+75/32.11 xX 18. COTONEASTER DAMMERI X C. CONSPICUUS (C. X SUECICUS) *41, Glam.: Waste
ground, Bridgend Industrial Estate, SS/92.79. G. Hutchinson, 1990, NMW, det. J. Fryer.

+75/32.16. COTONEASTER INTEGRIFOLIUS *77, Lanarks.: Carrick Quay, Glasgow, NS/59.64.
P. Macpherson, 1992, herb. P.M., conf. J. Fryer.

+75/32.18. COTONEASTER CONSPICUUS . *41, Glam.: Railway embankment, Gabalfa, ST/
168.791. G. Hutchinson, 1986, NMW, det. J. Fryer. *49, Caerns.: Steep roadside slope between

PLANT RECORDS | 293

Tyn-y-Groes and Talycafn Bridge, SH/785.719. R. Lewis, 1992, NMW, conf. J. Fryer. lst Welsh
records.

+75/32.21. COTONEASTER HORIZONTALIS *47, Monts.: Disused limestone quarry, Llanymynech
Rocks, SJ/264.217. H. Webster, 1986. *73, Kirkcudbrights.: Amongst willows in quarry E. of
Creetown, NX/500.585. O. M. Stewart, 1992. *75, Ayrs.: Embankment of disused railway,
Parkhouse Community Nature Reserve, Ardrossan, NS/23.42. A. McG. Stirling, 1993.

+75/32.22. COTONEASTER HJELMOQVISTII *41, Glam.: Railway embankment, Gabalfa, ST/
168.791. G. Hutchinson, 1986, NMW, det. J. Fryer.

+75/32.34. COTONEASTER BULLATUS *25, E. Suffolk: Open woodland, The Dales, Ipswich,
TM/155.462. M. A. Hyde, 1981. Ditch, Piper’s Vale, Ipswich, TM/175.416. T. Hutton, 1993, det.
E. M. Hyde. Both herb. E. & M. Hyde. 1st and 2nd records. *26, W. Suffolk: Roadside hedge,
Harper’s Hill, Nayland, TL/967.334. B. H. Harley, 1993, IPS, det. G. M. S. Easy. *47, Monts.:
Limestone scree, Llanymynech Rocks, SJ/260.218. R. Dawes & I. C. Trueman, 1989, det.
J. Fryer. *75, Ayrs.: Embankment of disused railway, Parkhouse Community Nature Reserve,
Ardrossan, NS/23.42. A. McG. Stirling, 1993.

+75/32.35. COTONEASTER REHDERI *46, Cards.: Scrub below A487 S.W. of Ffos-y-ffin, SN/
444.601. A. O. Chater, 1993, NMW. *49, Caerns.: Woodland, Bryn Maelgwyn S. E. of
Llandudno, SH/797.805. R. Lewis, 1991, NMW, det. J. Fryer. Edge of woodland between
Pontwgan and Roewen, SH/7.7. R. Lewis, 1993. Ist and 2nd records.

+75/32.37. COTONEASTER DIELSIANUS *47, Monts.: Woodland on limestone, Llanymynech
Hill, SJ/266.217. M. Wainwright, 1989, det. J. Fryer.

+75/32.40. COTONEASTER STERNIANUS *4, N. Devon: Wall above river-bank, Hillsford Bridge,
SS/742.478. W. H. Tucker, 1988, det. J. R. Palmer. Still present in 1993. *29, Cambs.: Bramble
thickets on waste ground, Wisbech, TF/462.103. G. M. S. Easy, 1993, herb. G.M.S.E. "41,
Glam.: Waste ground, Birchgrove, Cardiff, ST/165.804. A. D. Tipper, 1988, NMW, det.
J. Fryer. *44, Carms.: Lane to Waungron Farm, S. of Llanybydder, SN/52.42. G. Hutchinson,
1991, NMW, det. J. Fryer. Stradey Woods, Llanelli, SN/495.014. I. K. Morgan, c.1969, NMW, det.
G. Hutchinson.

77/4.2. ASTRAGALUS DANICUS *108, W. Sutherland: Sandy Bua Bay of Clachtoll, NC/
040.273. A. McG. Stirling & A. A. P. Slack, 1993.

177/7.1d. ANTHYLLIS VULNERARIA subsp. CARPATICA *25, E. Suffolk: yarns of bridle-path and
farm-track, Wherstead, TM/147.413. Ipswich District Natural History Society, 1993, IPS, det. J. R.
Akeroyd.

77/14.12. VIcIA LATHYROIDES 79, Selkirks.: Old wall top, Newark Castle, Yarrow, NT/
422.294. D. J. Methven, 1993. Ist record since 1926 record in this locality.

77/14.13. VICIA LUTEA +58, Cheshire: Road bank, Tarvin, SJ/496.661. M. Stead, 1991, herb.
G. M. Kay. Ist record since 1862.

77/15.2. LATHYRUS JAPONICUS 11, S. Hants.: Bare shingle, Hurst Castle Spit, SZ/319.898.
G. D. Field, 1993, conf. R. P. Bowman. Only extant locality. *91, Kincardines.: Stoney beach
just above high-water mark, Nigg, NJ/965.047. D. Bale, 1993, herb. D. Welch, conf. D.W.

{77/15.7. LATHYRUS GRANDIFLORUS *73, Kirkcudbrights: Ravenshall, NX/525.523. O. M.
Stewart, 1993. Well naturalized on roadside and river-bank, Palnure, NX/455.631. O. M. Stewart,
1993. Ist and 2nd records.

77/15.12. LATHYRUS NISSOLIA +*44, Carms.: Roadside verge near Derwen Farm, Cae’r-bryn,
SN/597.143. R. D. Pryce & K. Sanderson, 1993, NMW. Side of B4310 near Porthyrhyd, SN/
521.166. G. Hutchinson, 1993, NMW. Ist and 2nd records.

77/18.5. MEDICAGO ARABICA +83, Midlothian: Waste ground, Granton, Edinburgh, NT/23.77.
O. M. Stewart, 1990, E. Grassy bank, Warriston Road, Edinburgh, NT/253.754. S. Maxwell, 1993.
1st and 2nd post-1930 records.

794 PLANT RECORDS

77/19.5. TRIFOLIUM GLOMERATUM 5, S. Somerset: Gadd’s Bottom, Kingston St Mary, ST/
230.297. I. Green, 1993. 1st record since 1916.

+77/25.1. GENISTA MONSPESSULANA *5, S. Somerset: Hedge bank, Luccombe, SS/913.443.
J. Clarke, 1993.

78/1.1. HIPPOPHAE RHAMNOIDES +*77, Lanarks.: Grassy waste ground, Cessnock, Glasgow,
NS/56.64. P. Macpherson, 1991, herb. P.M. Still present in 1993.

+79/2.2. MyYRIOPHYLLUM AQUATICUM *46, Cards.: In Azolla filiculoides-dominated pond,
Ljain-wen near Synod Inn, SN/392.550. A. O. Chater, 1993, NMW.

+80/1.1. GUNNERA TINCTORIA *75, Ayrs.: Waste ground near sea, Auchengarth, NS/190.643.
A. McG. Stirling, 1993. *99, Dunbarton: Bank of A83 near Cameron House, Balloch, NS/
37.83. A. McG. Stirling, 1993.

84/1.3 X 8. EPILOBIUM MONTANUM X E. CILIATUM *104, N. Ebudes: With both parents on
waste ground, Park Lane, Portree, Skye, NG/48.43. G. Kitchener, 1991, E.

84/1.7. EPILOBIUM ROSEUM 73, Kirkcudbrights.: Old Creetown station, NX/475.599. O. M.
Stewart & A. White, 1993. Only extant locality.

+84/1.14. EPILOBIUM KOMAROVIANUM *2, E. Cornwall: China clay waste, Carvear Moor, SX/
051.547. G. C. Matthews & R. J. Murphy, 1993. *98, Main Argyll: Roadside bank, grounds of
County Hospital, Oban, NM/86.30. E. L. S. Macpherson, 1961, herb. P. Macpherson, conf. E. J.
Clement.

+91/2.14 X wal. EUPHORBIA ESULA X E. WALDSTEINII (E. X PSEUDOVIRGATA) *3,S. Devon: By
Exeter Canal, SX/92.91. L. M. Spalton, 1991, det. A. Radcliffe-Smith.

+100/1.1. RHUS HIRTA *50, Denbs.: Waste ground, Llandulas, SH/918.782. G. Battershall,
1993.

+102/1.6. OXALIS STRICTA *50, Denbs.: Weed in flowerbeds, Loggerheads, SJ/199.626.
R. Maycock & A. Woods, 1993.

7102/1.10. OXALIS DEBILIS *49, Caerns.: Waste ground: near railway halt, Dolgarrog, SH/
784.669. R. Lewis, 1991, NMW.

+103/1.1 xX 2. GERANIUM ENDRESSII X G. VERSICOLOR (G. X OXONIANUM) *77, Lanarks.:
Wood, Thorntonhall, NS/58.54. P. Macpherson, 1989, herb. P.M. Still present and spreading, 1993.

+104/1.1. TROPAEOLUM SPECIOSUM *75, Ayrs.: Roadside hedge, Pinmore, Girvan, NX/20.91.
A. E. White, 1993, conf. A. McG. Stirling.

105/1.1. IMPATIENS NOLI-TANGERE +*46, Cards.: Gulley, Capel Bangor, SN/6.7. C. Morley,
1939, ms notebook in IPS.

+105/1.3. IMPATIENS PARVIFLORA *93, N. Aberdeen: Railway siding, Inverurie, NJ/776.219.
D. Welch, 1992, ABD.

+107/32.1. AMMI MAJUS *38, Warks.: Churchyard, Birmingham, SP/070.871. A. D. Grenfell,
1983. Roadside, Coventry, SP/356.786. M. J. Senior, 1992, det. J. C. Bowra. Ist and 2nd records.

108/3.2. CENTAURIUM ERYTHRAEA 80, Roxburghs.: Steep bank, Bowden Burn, Newtown St
Boswells, NT/56.30. J. Blane, 1992. 2nd extant locality.

108/4.1. BLACKSTONIA PERFOLIATA *28, W. Norfolk: Bank of New Cut by Whitedyke Farm,
Feltwell, TL/69.89. J. Secker, 1992.

108/5.2. GENTIANELLA CAMPESTRIS 50, Denbs.: Limestone turf, Maeshafn, SJ/200.615.
R. Maycock & R. Woods, 1993. 2nd post-1930 record.

108/5.4a. GENTIANELLA AMARELLA subsp. AMARELLA 1, W. Cornwall: Dune grassland, Rushy
Green near Crantock, SW/787.608. H. Meredith, 1993, conf. L. J. Margetts. 1st record since 1912.

PLANT RECORDS 295

109/1.herb. VINCA HERBACEA {81, Berwicks.: Delete record published in The botanist in
Berwickshire by M. E. Braithwaite & D. G. Long (1990), which is based on a form of V. minor, fide
M. E. Braithwaite. |

110/3.1. ATROPA BELLADONNA 75, Ayrs.: Sandy waste ground, I.C.I. Nobel Works site,
Ardeer, NS/28.40. A. McG. Stirling & E. Millar, 1993. 1st record this century.

111/3.2b. CALYSTEGIA SEPIUM subsp. ROSEATA *45, Pembs.: Behind shingle ridge, Newgale,
SM/849.218. S. B. Evans, 1987, NMW.

111/3.2 x 4. CALYSTEGIA SEPIUM X C. SYLVATICA (C. X LUCANA) *73, Kirkcudbrights.: With
both parents on hedgebank, Eldingham industrial estate near Dalbeattie, NX/838.626. O. M.
Stewart, 1993.

113/2.1. NYMPHOIDES PELTATA +*68, Cheviot: Edge of R. Tweed N.E. of Union Bridge, NT/
934.512. J. M. Croft & C. D. Preston, 1991, CGE. Still present in 1994.

116/1.2. LITHOSPERMUM OFFICINALE 46, Cards.: Weedy vegetation on sandy slope between
road and beach N.W. of Nantyferwig, Gwbert, SN/165.484. A. O. Chater, 1993, NMW. ist record
since 1929, when recorded in this area by J. H. Salter.

116/4.3. SYMPHYTUM TUBEROSUM [47, Monts.: Delete record published in Watsonia 19: 288
(1993); specimen in NMW is S. grandiflorum, det. F. H. Perring.]

4116/4.4. SYMPHYTUM GRANDIFLORUM *47, Monts.: Churchyard, Penstrowed, SO/070.915.
C. A. Small, 1992, NMW, det. F. H. Perring. *73, Kirkcudbrights.: Wood edge E. of
Kirkcudbright, NX/691.507. O. M. Stewart & A. White, 1992.

+116/4.6. SYMPHYTUM ORIENTALE *50, Denbs.: Roadside, Eglwysbach, SH/804.716.
W. McCarthy, 1993, NMW. ist Welsh record.

+116/4.8. SYMPHYTUM BULBOSUM *6, N. Somerset: Edge of path, Butleigh, Street, ST/520.338.
PeGreen-) 1993" det. F. HH. Perring.

+116/5.1. BRUNNERA MACROPHYLLA *64, Mid-W. Yorks.: Roadside, Marston Moor, SE/
499.525. P. P. Abbott, 1993.

116/15.1 X 4. MyosotIs scorPIOIDES X M. LAXA (M. X SUZAE) *46, Cards.: Poached, muddy
margin of pond, The Moat, Llandyfriog, SN/342.408. A. O. Chater, 1993, NMW, det. P. M. Benoit.

116/15.9. MyosoTIs RAMOSISSIMA 73, Kirkcudbrights.: Short turf near shore, Fauldbog Bay,
NX/643.441. O. M. Stewart, 1992. 2nd extant locality.

+118/4.1a. LAMIASTRUM GALEOBDOLON subsp. ARGENTATUM *25, E. Suffolk: By wall alongside
watercourse, Walberswick, TM/499.746. G. Kitchener, 1992. Off Dunwich road, Westleton, TM/
445.693. G. Kitchener, 1992. 1st and 2nd records.

7118/13. rac. X nep. NEPETA RACEMOSA X N. NEPETELLA (N. X FAASSENII) *46, Cards.: Dune
by caravan site, The Patch, Penyrergyd, SN/162.485. A. O. Chater, 1993, NMW.

118/22.1. LycopUS EUROPAEUS 104, N. Ebudes: In Sphagnum and running water under Salix
aurita, N. of Lochan Gun Ghrunnd, Raasay, NG/613.519. S. J. Bungard, 1991. 1st Raasay record.

118/23.43 X 4. MENTHA SPICATA X M. SUAVEOLENS (M. X VILLOSA) *24, Bucks.: Disused
orchard, Ravenstone, SP/849.507. R. Maycock & A. Woods, 1989, det. R. M. Harley as var.
alopecuroides.

118/23.4. MENTHA SUAVEOLENS *83, Midlothian: Waste ground by sea, Granton, Edinburgh,
NT/23.77. O. M. Stewart, 1988, E, det. R. M. Harley. 1st confirmed record.

118/25.3. SALVIA PRATENSIS *13, W. Sussex: Grazed chalk grassland, Anchor Bottom S. of
Upper Beeding, TQ/205.095. G. Hart, 1992, herb. P. Harmes, det. A. C. Leslie. 1st confirmed
record.

121/1.3b. PLANTAGO MAJOR subsp. INTERMEDIA *25, E. Suffolk: Side of ditch, Shipmeadow,
TM/389.905. C.D. Preston & N. F. Stewart, 1989, CGE. *46, Cards.: Margin of potato field N.

296 PLANT RECORDS

of Ty-gwyn, Mwnt, SN/197.521. J. R. Akeroyd, A. O. Chater & C. D. Preston, 1987. Recently
reconstructed bank of A44, Lovesgrow, SN/636.810. S. P. Chambers, 1992. 1st and 2nd records.

+123/1.sus. X vir. FORSYTHIA SUSPENSA X F. VIRIDISSIMA (F. X INTERMEDIA) *35, Mons.: Bank
of R. Ebbw, Risca, ST/2.9. J. Harper, 1992.

124/1.2. VERBASCUM VIRGATUM +*35, Mons.: Rail ballast, old shunting lines, Undy, ST/
447.875. T. G. Evans, 1993, herb. T.G.E.

124/2.2. SCROPHULARIA AURICULATA 104, N. Ebudes: S. of Eaglais Breige, Raasay, NG/58.43.
S. J. Bungard, 1990, herb. C. W. Murray, det. D. R. McKean. Ist Raasay and 2nd v.c. record.

124/8.2. CHAENORHINUM MINUS *93, N. Aberdeen: Cess by railway, Gartly, NJ/527.314.
D. Welch, 1993, ABD.

+124/16.23. VERONICA FILIFORMIS 104, N. Ebudes: Inverarish, Raasay, NG/555.357. S. J.
Bungard, 1993. Ist Raasay record.

+124/17.1. HEBE SALICIFOLIA *50, Denbs.: Woods, Pwllycrochan, SH/842.785. G. Battershall,
1993.

124/20.1 X 9. EUPHRASIA ROSTKOVIANA subsp. ROSTKOVIANA X E. CONFUSA *44, Carms.:
Acidic, sheep-grazed turf above Pont Clydach, Mynydd Du, SN/740.195. R. D. Pryce, 1982, NMW,
det. A. J. Silverside. 2nd Welsh record.

124/20.3 X 7. EUPHRASIA ANGLICA X E. NEMOROSA *44, Carms.: Rough pasture, Cefn Parcin,
Llanybydder, SN/54.42. I. M. Vaughan, 1972, NMW, det. A. J. Silverside. 3rd Welsh record.

124/20.5 X 9. EUPHRASIA. ARCTICA subsp. BOREALIS X E. CONFUSA *44, Carms.: Disturbed
ground on recolonised ash-tip, Machynys, Llanelli, SS/51.98. R. D. Pryce, 1986, NMW, det. A. J.
Silverside. 2nd Welsh record. *46, Cards.: Leadmine, Llwynmalus, Tyn-y-graig, SN/690.679.
S. P. Chambers, 1992, det A. J. Silverside.

124/20.5 X 19. EUPHRASIA ARCTICA subsp. BOREALIS X E. SCOTTICA *44, Carms.: Heathy
roadside E.N.E. of Llidiadnenog, SN/553.367. R. D. Pryce, 1986, NMW, det. A. J. Silverside. 1st
Welsh record.

124/20.6 X 9. EUPHRASIA CONFUSA X E. TETRAQUETRA *2, E. Cornwall: Short turf by the sea,
Tintagel, SX/05.89. G. G. & P. S. Graham, 1983, SUN, det. A. J. Silverside.

124/20.20. EUPHRASIA HESLOP-HARRISONII 104, N. Ebudes: Saltmarsh turf, Ardnish, Skye,
NG/679.242, and Kensaleyre, Skye, NG/418.518. A. J. Silverside, 1982, herb. A.J.S. 1st and 2nd
Skye records.

124/23.1. PARENTUCELLIA VISCOSA +*5, S. Somerset: Edge of newly made pond, Fitzhead,
Wiveliscombe, ST/109.299. P. Green, 1993. +64, Mid-W. Yorks.: Old quarry, West Bradford,
SD/731.471. K. A. Crowther, 1992. 2nd record. Limestone grassland below slag heap, Micklefield,
SE/433.324. Leeds Naturalists’ Club excursion, 1993. 2nd extant locality.

124/25.2b. PEDICULARIS SYLVATICA subsp. HIBERNICA *41, Glam.: Molinia caerulea bog, Broad
Pool, Gower, SS/51.91. E. Nic Lughadha et al., 1993, NMW. Coastal heath, Overton Cliff, SS/
454.849. T. C. G. Rich et al., 1993. 1st and 2nd records. *46, Cards.: With subsp. sylvatica in
Molinia caerulea-dominated heathy pasture, N. of Nanthen-foel, Pont Creuddyn, SN/542.521.
A. O. Chater & D. Glyn Jones, 1993, NMW. With subsp. sy/vatica in flush, Cefnyresgair-fawr,
E.S.E. of Tregaron, SN/707.581. A. O. Chater, 1993. 1st and 2nd records.

*4125/1.2. LATHRAEA CLANDESTINA *48, Merioneth: Woodland, W. bank of Afon Dulas near
Esgair-geiliog, N. of Machynlleth, SH/7.0. S. Grasse & A. Morton, 1990, NMW. Open Salix scrub,
Dwyryd estuary, SH/5.3. R. E. Hughes, 1993, NMW. Introduced c.30 years ago and now well
naturalized. Ist and 2nd Welsh records.

125/2.2. OROBANCHE RAPUM-GENISTAE 4, N. Devon: On Cytisus scoparius subsp. maritimus on
steep rock face above landing beach,. Lundy, SS/142.439. L. Farrell & R. Key, 1993. 1st post-1930
record for Lundy.

PLANT RECORDS 297

+129/1.4. CAMPANULA PERSICIFOLIA *46, Cards.: Overgrown bank of Afon Einion, Eglwys
Fach, SN/684.958. A. O. Chater, 1993, NMW.

130/6.7. GALIUM MOLLUGO 79, Selkirks.: Field on S.E. side of Selkirk, NT/476.284.
J. Muscott, 1992. 2nd record.

130/6.7b. GALIUM MOLLUGO subsp. ERECTUM *50, Denbs.: Edge of lane, Eryris, SJ/209.571.
J. A. Green, 1993.

130/6.9. GALIUM STERNERI 81, Berwicks.: Rock outcrop, Gaitheugh, Gledswood, NT/59.34.
M. E. Braithwaite, 1993, herb. M.E.B. 1st record since J. H. Balfour’s 1852 record in this locality.

*4131/1.1. SAMBUCUS RACEMOSA *6, N. Somerset: Postlebury Wood, Trudoxhill, ST/739.427.
G. Read, 1993.

*131/2.3. VIBURNUM TINUS *50, Denbs.: Woodland, N. side of Bryn Euryn, Colwyn Bay, SH/
83.80. and bank of A470 N.E. of Talycafn Bridge, SH/79.72. R. Lewis, 1993. 1st and 2nd records.

131/6.4. LONICERA XYLOSTEUM 7*75, Ayrs.: Sourlie Wood, Irvine, NS/34.41. K. J. Watson,
1993.

133/1.1. WALERIANELLA LOCUSTA *96, Easterness: Dunes, Nairn, NH/896.574. M. Barron,
1993.

133/1.2. VALERIANELLA CARINATA 50, Denbs.: Colwyn Bay, SH/833.802. G. Battershall, 1993.
2nd post-1930 record.

¥133/1.5. WALERIANELLA ERIOCARPA *11, S. Hants.: Base of cliffs below Royal Bath Hotel,
Bournemouth, SZ/091.908. R. M. Walls, 1993, herb. R.M.W., conf. R. P. Bowman.

135/11.1. CENTAUREA SCABIOSA 81, Berwicks.: Railway cutting, Lamberton, NT/96.59. J. O.
Mountford & C. Sargent, 1980. 2nd record.

4135/11.2. CENTAUREA MONTANA 50, Denbs.: Mynydd Llanelian, SH/850.744. G. Battershall,
1993. 2nd record.

135/17.1. PicRIS ECHIOIDES +*98, Main Argyll: Edge of estate track near Loch Craignish,
Lunga, NM/797.070. B. H. Thompson, 1993, herb. B.H.T., conf. C. D. Preston.

135/22.1. LACTUCA SERRIOLA *43, Rads.: Recently excavated ground, Presteigne by-pass, SO/
321.634. D. R. Humphries, 1993. *51, Flints.: Roadside bank in industrial estate, Sandycroft,
SJ/328.667. T. Edmondson, 1992.

135/22.2. LACTUCA VIROSA *5, S. Somerset: Waste ground, Minehead, SS/974.462. I. Green,
1993.

7135/27.5a. PILOSELLA CAESPITOSA subsp. COLLINIFORMIS *77, Lanarks.: Rough grassland near
Stevenston, NS/76.59. P. Macpherson, 1993, herb. P.M.

135/28.4. HIERACIUM SALTICOLA *99, Dumbarton: Waste ground, Dumbarton Common, NS/
39.76. A. McG. Stirling, 1993.

135/28.5. HIERACIUM VAGUM *75, Ayrs.: R. Stinchar, Ballantrae, NX/0.8. A. Somerville,
1902, LCN, det. P. D. Sell & C. West.

135/28.10. HIERACIUM SUBUMBELLATIFORME *91, Kincardines.: Rocky riverside, Banchory,
NO/687.957. D. Welch, 1993, ABD, det. D. J. McCosh.

135/28.94. HIERACIUM SUBHIRTUM *75, Ayrs.: Rocky streamside, Cross Burn, Kirriereoch, -
NX/40.87. O. M. Stewart & A. McG. Stirling, 1988, E, det. D. J. McCosh.

135/28.116. HI1ERACIUM DURICEPS *80, Roxburghs.: Calcareous streamside rocks, Green
Needles, Kielderstone, NT/64.01. B.S.B.I. field meeting, 1993, det. A. McG. Stirling.

135/28.140. HIERACIUM EXOTERICUM *91, Kincardines.: Old Red Sandstone outcrop in river
gorge, The Burn, Edzell, NO/599.710. D. Welch, 1993, herb. D.W., det. D. J. McCosh.

298 PLANT RECORDS

135/28.147. HIERACIUM CALEDONICUM *91, Kincardines.: Rock outcrop, Slack of Birnie, NO/
672.807. D. Welch, 1993, herb. D.W., det. D. J. McCosh.

135/28.176. HIERACIUM DICELLA *75, Ayrs.: Sea cliffs S. of the castle, Dunure, NS/24.15.
R. Mackechnie, 1936, GLAM, det. D. J. McCosh. Still there in 1993.

135/28.209. HIERACIUM VENNICONTIUM *104, N. Ebudes: Cliff above Old Man of Storr, Skye,
NG/49.54. O. M. Stewart, 1987, E, det. P. D. Sell.

135/28.236. HIERACIUM GLOBOSIFLORUM *104, N. Ebudes: Rocky outcrops, Beinn na Greine,
Skye, NG/74.22. C. W. Murray, 1989, herb. C.W.M., det. J. Bevan as ‘dark-styled form’.

135/28.248. HIERACIUM HOLOSERICEUM *104, N. Ebudes: Rough gravelly ground, Beinn na
Greine, Skye, NG/74.22. R. M. & C. W. Murray & A. A. P. Slack, 1986, herb. C.W.M.., det.
J. Bevan.

135/30.4. FILAGO MINIMA 46, Cards.: Disturbed sandy soil, Penparc sand quarries, SN/
199.484. A. O. Chater & C. D. Preston, 1993, NMW. Only extant locality.

+135/41.3 X 4. ASTER LAEVIS X A. NOVI-BELGI (A. X VERSICOLOR) *46, Cards.: Long grass at
tidal limit of Afon Cletwr, Craigypenrhyn, SN/653.929. A. O. Chater, 1992, CGE, det. P. F. Yeo.

7135/43.4. ERIGERON KARVINSKIANUS 50, Denbs.: Crags above woodland, W. side of Bryn
Euryn, Colwyn Bay, SH/83.79. R. Lewis, 1993, det. Mrs R. Lewis.

135/62.41 X 10. SENECIO CINEREA X S. JACOBAEA (S. X ALBESCENS) *59, S. Lancs.: Disused
cinder track, Fazakerley, SJ/380.978. B.S.B.I. meeting, 1993. *75, Ayrs.: Waste ground in
industrial estate, Cambuslea Road, Newton-upon-Ayr, NS/35.23. A. McG. Stirling, 1993.

+135/71.3. PETASITES ALBUS 46, Cards.: Extensive colony in scrub and woodland, Falcondale
grounds, SN/565.491. A. O. Chater, 1993. 2nd record.

+138/4.2. ELODEA NUTTALLII *61, S.E. Yorks.: Barmston Drain, Beverley, TA/05.40.
J. Dews, 1993. Slowly flowing ditch, Wheldrake Ings, SE/703.433. C. D. Preston, 1993. 1st and 2nd
records. 73, Kirkcudbrights.: Carsfad Loch, NX/605.858. O. M. Stewart, 1993. 2nd record.

142/1.8. POTAMOGETON PRAELONGUS 29, Cambs.: Wicken Lode, Wicken Fen, TL/56.70. N. F.
Stewart, 1992. 2nd extant locality. |

142/1.14. POTAMOGETON OBTUSIFOLIUS 46, Cards.: Mesotrophic lake, Llyn Maesllyn, SN/
693.627. A. O. Chater & C. Moscrop, 1993, NMW. 2nd record.

142/1.16. POTAMOGETON TRICHOIDES *3, S. Devon: Drainage ditch, Exminster Marshes, SX/
9.8. M. C. F. Proctor, 1969, BM, det. J. E. Dandy. *61, S.E. Yorks.: Pond and slowly flowing
ditch, Wheldrake Ings, SE/704.434. & 703.433. C. D. Preston, 1993, CGE.

142/2.1. GROENLANDIA DENSA +*104, N. Ebudes: Pond in hotel grounds, Skeabost House
Hotel, Skye, NG/413.487. C. W. Murray, C. D. Preston & N. F. Stewart, 1989, herb C.W.M.
Perhaps introduced unintentionally with plants from Derbyshire.

148/2.2 LEMNA MINOR *108, W. Sutherland: Recently re-excavated peaty ditch near Elphin
school, NC/215.119. I. M. Evans, 1993.

+148/2.4. LEMNA MINUTA *38, Warks.: Pond, Solihull, SP/164.817. J. W. Partridge, 1989,
WAR, det. A. C. Leslie. R. Leam near Leamington Spa, SP/333.655. J. W. Partridge, 1991, det.
J. C. Bowra. 1st and 2nd records.

+151/1.2. JUNCUS TENUIS *79, Selkirks.: Track, Thirlestane Tower, Ettrick, NT/281.154.
R. W. M. Corner, 1993, herb. R.W.M.C.

151/1.6. JUNCUS FOLIOSUS *73, Kirkcudbrights.: Edge of damp, shingley burn, glen N. of
Corse of Slakes road, NX/54.58. O. Mi. Stewart, 1993. *86, Stirlings.: Sandy shore of Loch
Lomond, Balmaha, NS/42.90. A. & D. A. Pearman & C. D. Preston, 1993, CGE.

152/3.2. ELEOCHARIS AUSTRIACA ~° *59, S. Lancs.: Silty mound in bed of River Ribble 2:5 km
N.N.E. of Clitheroe Castle, SD/752.441. P. Jepson, 1993, det. S. M. Walters.

PLANT RECORDS 299

152/7.2. SCHOENOPLECTUS TABERNAEMONTANI *81, Berwicks.: Pond, Morningbank, Swinton,
NT/833.477. M. E. Braithwaite, 1993, herb. M.E.B.

152/12.1. SCHOENUS NIGRICANS *48, Merioneth: With Juncus maritimus in upper saltmarsh
near Pennal, SN/6.9. R. Squires, 1993, NMW. Ist confirmed record.

152/16.1. CAREX PANICULATA *86, Stirlings.: Mugdock Wood, Milngavie, NS/545.763.
A. McG. Stirling, 1950, BM, det. R. W. David.

152/16.1 < 3. CAREX PANICULATA X C. DIANDRA (C. X BECKMANNII) *98, Main Argyll: Loch
Ederline, Ford, NM/865.021. A. McG. Stirling & B. H. Thompson, 1992, BM, det. R. W. David.

152/16.3. CAREX DIANDRA *78, Peebless.: Bridgehouse, NT/1.5. No collector, 1865, BM.

152/16.5. CAREX OTRUBAE - 104, N. Ebudes: Shore rocks, Rubha na Leac, Raasay, NG/
598.381. S. J. Bungard, 1993. 1st confirmed Raasay record.

152/16.7. CAREX SPICATA *74, Wigtowns.: Port Castle Bay, Whithorn, NX/425.357.
A. Rutherford & A. McG. Stirling, 1985, BM, det. R. W. David.

152/16.8b. CAREX MURICATA subsp. LAMPROCARPA *89, E. Perth: Stenton, Dunkeld, NO/
07.40. J. H. Penson, 1961, BM, det. R. W. David.

152/16.11. CAREX DISTICHA *93, N. Aberdeen: Marsh by flooded brick-pit, Cruden Bay, NK/
08.36. M. Innes, 1992.

152/16.27. CAREX PSEUDOCYPERUS 46, Cards.: Large pond dug out from swampy, wooded
hollow in 1992, Cwmsaeson, Oakford, SN/453.585. A. O. Chater & D. Glyn Jones, 1993. 2nd
record.

152/16.47. CAREX PALLESCENS 83, Midlothian: Upland grassland, Moorfoot Hills S.W. of
Hirendean Castle, NT/294.501. C. Dixon, 1993, E, conf. D. R. McKean. Upland heath, Linhouse
Water E. of Murieston, NT/075.649. S. Maxwell, 1993. 1st and 2nd post-1930 records.

152/16.49. CAREX ORNITHOPODA 64, Mid-W. Yorks.: Grassy areas amongst limestone pave-
ment at 380 m, Twistleton Scars, SD/715.764. B.S.B.I. meeting, 1992, det. G. M. Kay. 2nd record,
1st this century.

4

152/16.59. CAREX MAGELLANICA *46, Cards.: Fringing pool and in nearby wet mire, E. of
Drybedd, Dyffryn Castle, SN/779.832. A. O. Chater, 1993, NMW.

152/16.68. CAREX ELATA 36, Herefs.: Pondside, Wallbrook Wood, Allensmore, SO/440.360.
J. Port, 1993, det. A. O. Chater. 1st confirmed record this century.

153/11.1. MILIUM EFFUSUM 81, Berwicks.: Policy woodland, Swinton House, NT/813.473.
M. E. Braithwaite, 1993, herb. M.E.B. 2nd extant locality.

7153/12.7g. FESTUCA RUBRA subsp. MEGASTACHYS 25, E. Suffolk: Edges of garden lawn,
Shottisham, TM/31.44. A. Morgan, 1992, det. P. J. O. Trist. 2nd record.

153/12.9. FESTUCA VIVIPARA 83, Midlothian: Disturbed ground, Emly Bank, Moorfoot Hills,
NT/293.474. C. Dixon, 1993, E, det. D. R. McKean. 2nd post-1930 record.

153/12.12. FESTUCA HUONII *3, S. Devon: Rock outcrop near Prawle Point, SX/767.361.
R. Takagi-Arigho, 1992, herb. R.T.-A., det. T. A. Cope. 1st record for British Isles outside
Channel Islands.

153/18.1. POA INFIRMA *9, Dorset: Cliffs, Bournemouth, $Z/12.91. R. M. Walls, 1993.

153/18.6. POA ANGUSTIFOLIA *81, Berwicks.: Sandstone wall top, side of R. Tweed below
Ladykirk House, NT/886.455. M. E. Braithwaite, 1993, herb. M.E.B., det. J. R. Edmondson.

153/20.1. CATABROSA AQUATICA 78, Peebless.: peaty ditch, Kippit Farm, Dolphinton, NT/
11.47. D. J. McCosh, 1993, herb. D.J.M. Ist post-1930 record.

153/24.1. GLYCERIA MAXIMA 81, Berwicks.: Side of R. Tweed, Ladykirk, NT/896.464. M. E.
Braithwaite, 1993, herb. M.E.B. 2nd record.

300 PLANT RECORDS

4153/28.3. AVENA FATUA 79, Selkirks.: Waste ground, Clovenfords, NT/451.367. R. W. M.
Corner, 1991, herb. R.W.M.C. 1st post-1930 record.

+153/29.1. GAUDINIA FRAGILIS *11, S. Hants.: Old grassland in meadow, Curdridge, SU/
52.13. J. Rowe, 1993, HCMS, det. F. Rose.

153/32.1b. DESCHAMPSIA CESPITOSA subsp. PARVIFLORA *35, Mons.: Shady cliff top near
Lover’s Leap, Piercefield Park, ST/523.969. T. G. Evans, 1988, herb. T.G.E. Shady bank of River
Honddu N.N.W. of Cwmyoy, SO/291.249. T. G. Evans, 1993, herb. T.G.E. 1st and 2nd records.

153/31.2. KOELERIA MACRANTHA 79, Selkirks.: Basic. grassland, Selkirk Hill, NT/481.284.
J. Muscott, 1992, herb. R. W. M. Corner. 2nd extant locality.

+153/38.5. PHALARIS PARADOXA *14, E. Sussex: Arable field near Wilmington East, TQ/
539.053. P. A. Harmes, 1989, herb. P.A.H., det. E. J. Clement. Established in quantity; still present
in 1993.

+153/39.3. AGROSTIS CASTELLANA *3, S. Devon: Waste ground, The Maer, Exmouth, SY/
011.800. R. Takagi-Arigho, 1990, RAMM. *58, Cheshire: New road verge, Ringway, SJ/
807.858. D. Shaw, 1992, herb. G. M. Kay, det. R. M. Payne.

7153/39.9. AGROSTIS SCABRA 83, Midlothian: Boggy area of disused railway, Edinburgh, NT/
763:251. D. R. McKean & U. L. Moorhead, 1993, E. Ist record since 1915.

153/40.2. CALAMAGROSTIS CANESCENS *24, Bucks.: Edge of woodland ride by hazel coppice,
Leckhampstead Wood, SP/727.403. R. J. Hornby, L. M. Jones-Walters & J. Spencer, 1989.

153/44.1. APERA SPICA-VENTI 13, W. Sussex: Arable set-aside on chalk downland, Thunders-
barrow Hill, TQ/22.09. H. Matcham & A. Spiers, 1993, herb. P. A. Harmes. Sandy set-aside field E.
of Coates Castle, SU/996.177. A. Spiers, 1993. 1st and 2nd records since 1909.

153/45.1. MIBORA MINIMA +*9, Dorset: Dune-slack, Studland Heath, SZ/033.838. F. Wood-
head, 1993, RNG.

+153/51.4. BROMOPSIS INERMIS *69, Westmorland: S. side of Kaber-Barras road, NY/815.116.
. Atkins, 1993, LANC, det. G. Halliday.

+153/52.1. ANISANDRA DIANDRA 38, Warks.: Side of bridle path, Pillerton Priors, SP/292.486.
J. C. Bowra, 1993, WAR, det. P. J. Copson. 2nd record, Ist this century.

+153/53.1. CERATOCHLOA CATHARTICA 11, S. Hants.: Side of farm track near Sandy Lane,
Shedfield, SU/556.147. E. A. Pratt, 1992, det. E. J. Clement. By footpath along Danes Stream,
Milford on Sea, SZ/292.917. V. Scott, 1992, det. P. J. O. Trist. 1st records since 1925.

+153/53.2. CERATOCHLOA MARGINATA *22, Berks.: Field track, Pusey House, Southover, SU/
3.9. T. C. G. Rich, 1993, RNG.

153/56.1 X 2. ELYTRIGIA REPENS X E. ATHERICA (E. X OLIVERI) *3, S. Devon: Waste ground,
The Maer, Exmouth, SY/009.800. R. Takagi-Arigho, 1989.

153/57.1. LEYMUS ARENARIUS 104, N. Ebudes: Rocky skerry S. of Oskaig Point, Raasay, NG/
544.377. J. Bungard, 1992, conf. S. J. Bungard. Ist Raasay record.

153/63.1b. MOLINIA CAERULEA subsp. ARUNDINACEA *44, Carms.: Damp grassy area N.E. of
Witchett Pool, Laugharne Burrows, SN/283.077. G. Hutchinson & I. K. Morgan, 1992, NMW.

7153/68.1. ECHINOCHLOA CRUS-GALLI *46, Cards.: Abundantly naturalized in marsh by pond
N. of Lampeter College, SN/579.488. A. O. Chater, 1993, NMW.

*153/71.1. DiIGITARIA ISCHAEMUM *3, S. Devon: Derelict sidings, Exeter, SX7O17-9me:
D. Bolton, 1989, RAMM, det. R. Takagi-Arigho. Waste ground, Exmouth, $X/992.807. R. Takagi-
Arigho, 1989, RAMM. Ist and 2nd records.

154/1.4. SPARGANIUM NATANS *5, S. Somerset: Pinkworthy Pond, SS/723.423. C. Giddens &
P) Green, 1993 det: Cs DAe Cook :

Quy

;

PLANT RECORDS 301

155/1.2. TYPHA ANGUSTIFOLIA 35, Mons.: Galvanised tank, sludge bed pond, Alpha Steel
Works, Newport, ST/33.84. T. G. Evans & M. Jones, 1993. 2nd record.

4157/1.1. PONTEDERIA CORDATA *59, S. Lancs.: Disused canal, White Gate, Oldham, SD/
895.034. G. M. Kay & P. D. Stanley, 1993.

158/7.1. COLCHICUM AUTUMNALE 41, Glam.: Damp grassland near Lakston, Bridgend, SS/8.7.
C. Lindley, 1993. Only extant locality.

158/11.1. FRITILLARIA MELEAGRIS +*58, Cheshire: Sloping wet pasture, Hassal Green, SJ/
W927 2.5. Jones, 1993.

4158/23.1. Muscari NEGLECTUM *83, Midlothian: Woecd, Bowshank near Stow, NT/45.41.
M. Little, 1984.

+158/23.2. MUSCARI ARMENIACUM *83, Midlothian: Under birch trees on old railway line,
Warriston, Edinburgh, NT/25.75. D. R. McKean, 1986. Still present in 1994.

4158/24.5. ALLIUM SUBHIRSUTUM *17, Surrey: Roadside, Purley, TQ/326.538. G. Fookes,
1993.

158/24.14. ALLIUM SCORODOPRASUM *98, Main Argyll: Grassland between beach and fields,
Toward Point, NS/130.675. C. Gibson, 1991, conf. G. P. Rothero.

158/24.16. ALLIUM VINEALE *104, N. Ebudes: Rock ledges S. of Sithean Mor, Raasay, NG/
599.475. S. J. Bungard, 1991, herb. C. W. Murray.

7158/33.3a. NARCISSUS POETICUS subsp. POETICUS *49, Caerns.: Grassy bank at top of rock
outcrop, Tyn-y-Groes, SH/775.718. R. Lewis, 1993, NMW.

7159/4.1. HERMODACTYLUS TUBEROSUS 6, N. Somerset: Sandy bank, Brean, ST/297.557.
P. Green, 1993. 2nd record.

162/3.1. EPIPACTIS PALUSTRIS 2, E. Cornwall: Marshy area in dunes, Rock Dunes, SW/
934.766. T. J. Dingle & M. G. C. Atkinson, 1993. 2nd record.

162/3.4. EpIpACTIS HELLEBORINE 72, Dumfriess.: Above Liddel Water, Penton Bridge, NY/
433.774. M. S. Porter, 1991. 2nd record this century.

162/8.1. GOODYERA REPENS 81, Berwicks.: Pinewood near Hirsel Law, NT/83.42. M. E.
Braithwaite, 1993. Only extant locality.

162/13.2. PLATANTHERA BIFOLIA *79, Selkirks.: Hill pasture, Selkirk Hill, NT/4.2. G. French,
1993.

162/16.1 X 18.1. GYMNADENIA CONOPSEA X DACTYLORHIZA FUCHSII (X DACTYLODENIA ST-
QUINTINII) *104, N. Ebudes: Grassland, Harris area, Rum, NM/3.9. C. W. Murray, 1984, det.
M. Bates & S. J. Clarke.

162/17.1. COELOGLOSSUM VIRIDE 46, Cards.: Unimproved pasture, Tynygwndwn, S.S.E. of
Llanfair Clydogau, SN/6.4. A. O. Chater & L. Gander, 1993. 2nd extant locality.

162/18.3 X 5. DACTYLORHIZA INCARNATA X D. PURPURELLA (D. X LATIRELLA) *81, Berwicks:.:
Old quarry, Grantshouse, NT/827.645. Lady Margaret Elliot, 1993, E, det. S. Clarke.

162/18.8. DACTYLORHIZA LAPPONICA *104,N. Ebudes: Damp fiush, Dibidil, Rum, NM/4.9.
B. & J. Allan, 1989, E (photographs), det. D. J. Tennant.

162/20.4. ORCHIS USTULATA *41, Glam.: Limestone grassland, Llanmadog, SS/43.93.
N. Benson, 1993, conf. D. Painter. Ist certain Welsh record.

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Watsonia 20: 303-316 (1995) 303

Book Reviews

Exkursionsflora von Osterreich. W. Adler, K. Oswald & R. Fischer; edited by M. A. Fischer. Pp.
1180, 510 figures. Eugen Ulmer, Stuttgart & Vienna. 1994. Price DM 78 (ISBN 3-8001-3461-6).

Although Austria has long been a favourite holiday destination for those in search of Alpine plants,
the only pocket-sized Flora in the language of that country which was available until now was the
1973 reprint of K. Fritsch’s Exkursionsflora fiir Osterreich, first published in 1897. French-speaking
visitors are well served by books centred on Switzerland: see, for example, the review of
Aeschimann & Burdet’s Flore de la Suisse (1989) in Watsonia 18: 229, 1990. For the monoglot
English visitor, the Swiss Alpine Club’s Our Alpine Flora (1989) by E. Landolt & K. M. Urbanska is
available in English, and there are many other popular guides to Alpine plants available in English,
French, German and Italian. i

The appearance of Exkursionsflora von Osterreich is doubly welcome, as it effectively replaces as
weil as updates Janchen’s rather inadequate account of Austria’s flowering plants in Catalogus
florae Austriae, edited by K. Hafler & F. Knoll. This was published in four parts from 1956-60 with
four supplements from 1963-67. Within a concise format, the new Excursion Flora provides (to
quote its subtitle) an identification book for all vascular plants growing in the wild in Austria as well
as the most important cultivated plants, together with information on their ecology and distribution.
Compiled by three main authors with major contributions from eleven others and smaller
treatments and corrections from a further 26, the book was edited by Manfred Fischer. Illustrations
of critical parts are mainly by Gabriele Hofer-Sabek and Anton Igersheim, together with seven
other artists; in contrast to Aeschimann & Burdet, the figures are included within the keys.

As is usual with key-based Floras, most of the descriptive matter is presented within the terminal
couplet of the keys, including notes on the economic uses of the plant. Diagnostic features are
underlined. Introduced taxa are treated in a smaller font than native taxa, as are keys to infraspecific
taxa, a distinction which is unlikely to help users in the field. It also contains a 16-page summary of
the literature, and a very detailed 50-page glossary which includes an analysis of the Latin
geographical epithets of the scientific names of plants.

A considerable degree of detail is provided in the taxonomic treatments of some of the more
critical genera. The account of Alchemilla, for instance, occupies 22 pages; the more cursory
treatment of Rubus, by contrast, still covers 13 pages. Even under familiar taxa there are some
critical observations, such as the possibility that Potentilla crantzii subsp. serpentini is more
deserving of varietal status.

A useful aspect of the book is its attention to synonymy. The second edition of Flora Europaea
vol. 1 appeared (in 1993) too late for detailed consultation; thankfully, the Green Spleenwort here
appears with its more familiar scientific name, Asplenium viride. Aithough the great majority of
accepted names follow those given in W. Gutermann et al.’s Liste der GefapBpflanzen Mitteleuropas
(1973), the exceptions are shown by placing the synonym in bold italic font; other synonyms,
including deviations from Flora Europaea, are given in medium italic font. In a similar way,
standard vernacular names of plants are given in bold font, followed in some cases by alternative
names in medium font, both being abbreviated wherever feasible. These conventions are easier to
read than to explain.

Most botanists visiting Austria will wish to know the distribution of a particular plant within the -
country, and this is shown partly by means of a summary of occurrences within each of the eleven
Lander (provinces) or, for commoner species, a list of provinces from which it is absent. Where
infraspecies are included, this information is given at subspecific level. A summary of the
distribution of each species outside Austria is also given, except for endemics; for these, a more
detailed statement of distribution within Austria is provided. Geographical information is given in
highly compressed form; the front end papers include not only a map of Austria and its provinces,

304 BOOK REVIEWS

principal rivers, lakes and mountain ranges, but also a Key to the abbreviations of places shown on
the map which greatly aids legibility. This map also solves the problem, for German speakers at
least, of what to call the new Czech and Slovak republics: “Ischechien’ and ‘Slowakei’. These,
together with another recently established country, ‘Slowenien’ (Slovenia), not to be confused with
Slovakia, form three of the eight countries now sharing a border with Austria.

It is evident from a glance at the bibliography that local floristic studies are being actively pursued
in Austria, and I would expect that the appearance of this new Flora will be welcomed
enthusiastically by Austrian botanists as well as by visitors. Although its format is more complex, its
practicality and comprehensiveness is on a par with Stace’s New Flora of the British Isles (1991).
Thanks to the extensive use of abbreviations, even users whose knowledge of German is patchy
would find it eminently usable if a translation into English of the abbreviated terms listed on pp. 21—
24 were to be provided in a future edition.

Students of botany will find the detailed chapter on ‘““Morphology and Phytography” (pp. 38-91)
interesting, and the summary data on endemics and near-endemics on pp. 114-116 will be
convenient both for Flora-writers and conservationists. One must also mention the short chapter on
the history of botanical studies of the flora of Austria (pp. 157-163) which ranges from Clusius to
Janchen and ends with a sketch of a bulging plant press. At a price of around £33 at mid-1994 prices,
this closely printed and meticulously edited book represents good value for money and may soon
become the preferred German-language field Flora for much of central Europe as well as for Austria
itself.

J. R. EDMONDSON

Illustrated Flora of Mallorca. E. Beckett. Pp. 223, with 96 colour plates and an illustrated glossary.
Editorial Moll, Palma de Mallorca. 1993. Price Pta. 5500 (ISBN 84-273-0714-4).

This excellent illustrated guide to the flora of Mallorca, in a carefully and well edited format, follows
the model of The concise British Flora in Colour of W. Keble Martin (1965). The most important
part of the book is the inclusion of 96 plates, each one containing between five and 19 plants (the
majority 10-14) drawn in pen and ink and watercolours. They represent both the commonest plants
of the Mallorcan flora, those which the visitor sees most frequently, and also some of the rarer ones.
The drawings were made by the author during repeated visits to Mallorca since 1976, the majority
being from living specimens or photographs. In some cases, clearly indicated in the text, the
drawings were made from non-Mallorcan plants. In a few cases the latter include some plants whose
presence in Mallorca needs to be confirmed, such as Carduncellus pinnatus, or which have been
recently deleted from the Balearic flora, like Ranunculus monspeliacus. The drawings include
diagnostic details and, with rare exceptions (we could mention Helianthemum salicifolium and
Cistus clusii as examples of those which have an unusual appearance) are very accurate
reproductions of the form, appearance and true colour of the plants. Many of the drawings are really
excellent; however some of the white-flowered Umbelliferae are on plates which also have a white
background, and while they are not themselves of poor quality they have not printed well. The
monocotyledons, with the exception of Urginea maritima (= Drimia maritima), have been drawn
without the basal parts, doubtless to avoid having to uproot the plants.

To compile the list of Mallorcan plants, the author has used not only the first edition of Flora
Europaea and an unpublished catalogue by Professor Llorens, a Mallorcan botanist, but also the
general Floras and catalogues currently in use for the Balearic islands, as well as some specialised
works about particular genera which are grouped within the bibliography. It is surprising that the
most recent Floras of Spain which included the Balearic islands in their circumscription have not
been included: Bolos & Vigo, Flora dels Paisos Catalans (vol. 1, 1984; vol. 2, 1990), and Castroviejo
et al. (eds), Flora Iberica (four vols, 1986—). To have consulted them would have resolved some
doubts expressed in the book such as the presence or absence in Mallorca of plants like Asarum
europaeum or the previously mentioned Ranunculus monspeliacus.

The plants are ordered systematically, from the ferns to the monocotyledons, following the order
of Flora Europaea. The information on each species includes the Latin name, the English name
(where it exists), a short description based on Flora Europaea, the flowering period and indications

BOOK REVIEWS 305

of ecological preference and degree of rarity. Flora Europaea has also been followed for the
nomenclature, except in some cases where she uses names given in Med-Checklist by Greuter,
Burdet & Long or in the previously mentioned Llorens catalogue. In these cases the name used in
Flora Europaea is cited as a synonym. On occasion, this has led to the unnecessary inclusion of
synonymous names older than the accepted ones, such as Quercus humilis Mill. (1768) vs Q.
lusitanica Lam. (1785), Gagea iberica Terracc. (1804) vs G. nevadensis (1838) or Asparagus
stipularis Forsk. (1775) vs A. horridus L. in Murray, 1774; L. f., 1781; this error has already been
noted in Flora Europaea. This is a slightly embarrassing feature for a professional botanist, but it is
probably unimportant for the majority of potential users of the book. The nomenclatural or
taxonomic deviations from Flora Europaea are not always accurate; this is the case with the
previously mentioned Gagea iberica, which actually has the correct name G. durieui Parl. ex Batt. &
Trab. (1895) or in the case of Teucrium balearicum (Pau) Castrov. & Bayon, which scarcely differs
from T. subspinosum and according to a recent publication (Candollea 46: 47-51 (1991)) must be
included within 7. marum L.

The work includes a preface, notes on the flora and vegetation of Mallorca, an illustrated glossary
and an index of plates, and concludes with an index of Latin familial and generic names. The format
of the book is not ideally suited to being taken into the field, but because of its very solid covers it can
be handled without reducing its ease of use.

The book fills a large gap, in spectacular fashion, in the existing coverage of Mallorcan and
Balearic Floras. I expect that it will be welcomed enthusiastically not only by the many nature lovers
who visit Mallorca and the other Balearic islands each year, but also by the many Mallorcans with an
interest in botany.

G. LOPEZ

The art of botanical illustration. W. Blunt & W. T. Stearn. New edition, revised and enlarged. Pp.
368, illustrated. Antique Collectors’ Club, Woodbridge in association with The Royal Botanic
Gardens, Kew. 1994. Price £29.95 (ISBN 1-85149-177-S).

A new edition of such a familiar and treasured book is cause for excitement. There have been
several recent histories of botanical illustration but ‘Blunt’ has earned a respect and devotion that
deserves to be maintained. In the introduction to the new version we learn that, when the book was
first mooted in 1946, two of the original editors of Collins’ New Naturalist series separately asked
Blunt and Stearn to write it! That situation surely highlights one of the difficulties of writing a history
of this subject — whether to treat it as a branch of the history of art or of botany. Plants occupied a
significant place in the whole canvas of subjects chosen for artistic depiction long before the needs of
descriptive botany tailored that specially disciplined representation that we understand as ‘botanical
illustration’. So perhaps it was more fitting that Blunt, a student and teacher of the art and history of
drawing, should have been the eventual choice as author of this history. A measure of his success is
the classic status according to this history in the four decades since its publication and the fact that
two other publishers have brazenly ‘borrowed’ his title for more recent histories of their own —
sincere flattery one hopes!

Stearn’s contribution to the original work, which Blunt graciously acknowledged, obviously
amounted to an essential improvement in the specialised botanical view of the subject. Blunt had
more feeling for the poetry of drawing, and the aesthetics of media, syntax, rhythm, texture, light
and shade, perspective, etc., — but always with sensitivity for the ‘personality’ of the plant. Stearn’s
interest was more concerned with the importance of botanical illustration as one of the tools
developed by descriptive botanists of the classical period and its significance as a visual information
resource. Long before his death in 1987, Blunt had declined to undertake a full revision of the book,
his view being that his history should stand as a product of its day. However, now that revision is
possible, we are indeed fortunate to have the authority of knowledge of Blunt’s original collaborator
to perform that task.

In fact, although the outward appearance is radically altered, the internal structure of Blunt’s
work is not, and the text is mostly the same. Stearn has lightly revised it — altering outdated facts,
Inserting significant comments to strengthen botanical emphasis, adding biographical dates, and

306 BOOK REVIEWS

supplying or enlarging footnotes that direct the reader’s attention to more recent literature. Blunt’s
text has been slightly modernized here and there but, despite the title-page statement of dual
authorship of this edition, those who knew him will still hear Blunt’s unmistakable deep and
cultured voice coming off the page and will appreciate his personal expressions of unashamed
pleasure at the loveliness of some illustrations or the repellence of others!

It is not until Chapter 23 ““The Twentieth Century” that we really hear Stearn’s voice. He has
extended the densely packed account of contemporary artists, giving emphasis to British, Greek,
South African and Australian work, and has included thematic passages on the illustration of
comparative plant morphology and orchids. It must be said that Stearn’s selection of contemporary
artists for mention is a very personal one that only hints at the number of talented artists that have
emerged since 1950. A notable trend of botanical art during the post-war period has been a
broadening of the range of styles and the introduction of entirely new media, some of which have
achieved general acceptance. Unfortunately for the reader, Stearn barely mentions the 32 year-old
Hunt Institute for Botanical Documentation and its collection of over 30,000 examples of botanical
art that includes a fully representative share of post-war artwork, merely alluding to its exhibitions
on pages 12 & 13 of the Introduction, and does not touch on the prestigious 24 year-old Guild of
Natural Science Illustrators and its informative Handbook of scientific illustration (1989). Also
omitted is any mention of the famous Broughton Collection of some 119 flower paintings, 38 albums
and almost 900 mounted drawings, a collection distinguished for the excellence of its examples from
the classical period of European flower painting and botanical art. Given to the Fitzwilliam
Museum, Cambridge between 1966 and 1973, it is probably Britain’s most important collection of
fine botanical art. 2

The authors had expressed disappointment at the distinctly poor quality of the illustrations in the
original edition. The present publishers clearly decided that the visual element of the original should
be radically enlarged and improved using much more colour. To achieve this they replaced the
modest octavo format of the New naturalist series with a 27 X 21 cm quarto. In keeping with
contemporary fashion in book-design, illustrations are variously placed about the pages within the
discipline of the designer’s invisible ‘grid’; one, two or three illustrations may occupy the full width
of a page, some are full-page. The column of text is fitted into this grid system leaving outer margins
of 4.5 cm throughout the book, most of which are blank.

Most of the original choice of 55 colour images have been retained, not always in the same size,
but some are much improved by re-photography with a wider field of view. Omissions from the
original selection are usually filled with alternative illustrations from the same sources. The number
of examples is increased making a new total of 125 colour illustrations and, of these, just over a
dozen are devoted to post-1950 artwork. Of the original 47 monochrome plates, most are retained
and ten more are added. Unfortunately, some plates that Blunt intentionally juxtaposed are now
separated (e.g. Plates 8 & 13, now six pages apart), and several are reduced in size. The 61 figures,
which were generally poor in the 1950 printing, are all repeated but with virtually no improvement
in quality, and five more have been added. Three early intaglio illustrations are so badly reproduced
as to obscure all fine detail (Figs 41, 45 & 46). Two others (Plates 32 & 33) are reproduced in half-
tone but in such reduced size that they look like tonal drawings, the linear quality of the engraving
being virtually obscured. This is doubly unfortunate because Blunt describes them as a “landmark in
botanical illustration’, a special feature of which was that plants were engraved life size.

Many illustrations are chosen from rare, sometimes unique, originals that most readers may never
chance to see. Those who are fortunate enough to know and handle these source works may not
need to be told, for example, that the 9 X 6 cm Colour Plate 44 illustrates an original painting of
some 53 X 36-5 cm in the Victoria & Albert Museum. But what size is the drawing reproduced in
Plate 24 at 8-5 x 4-5 cm, one of a thousand paintings in what Blunt calls “‘five noble volumes” in far-
away St Mark’s Library, Venice? Three colour reproductions from Thornton’s famous Temple of
Flora are presented in different sizes (Colour Plates 81-83). In short, the reader, however
knowledgable, seriously needs statements of the sizes of the original images.

A few textual errors deserve mention: on p. 171, the three-colour printing process is that of J. C.
Le Blon (1667-1741), not to be confused with A. Le Blond (1819-1894), a colour-printer by the
Baxter process; it was that same Baxter process that Wm. Dickes (1815-1892), a Baxter licensee,
employed to print Anne Pratt’s illustrations, and not chromolithography as stated on p. 276; the co-
author of the Nature-printed British seaweeds, mentioned on p. 158, is not A. Crumb but actually

BOOK REVIEWS 307

Alexander Croall (1809-1885); the ““Hunt Botanical Collection”, on p. 347, should read Hunt
Botanical Library.

The accurate colour reproduction of artwork is still difficult even today. Films, lighting and
colour-printing techniques have all improved but many modern colour reproductions of artwork on
paper still show disturbingly odd background tints: the present work is no exception. Generally,
though, the colour work is good and quite acceptable for the price. What one gets for one’s money is
a much nicer version of Blunt but without a great deal of new information. It remains a selective
outline of the development of a rich segment of specialised art, elegantly chronicled in a personal
style, and providing a very reliable platform from which to launch into a broad range of specialised
studies.

G. D. R. Bripson

Dictionary of British & Irish botanists and horticulturists, including plant collectors, flower painters
and garden designers, revised edition. R. Desmond, assisted by C. Ellwood. Pp. xi + 825. Taylor &
Francis and the Natural History Museum, London. 1994. Price £120.00 (ISBN 0-85066-843-3).

Ray Desmond’s Dictionary, which was published in 1977, has since that time been the standard
reference work of biographical data of British and Irish botanists and horticulturists. The author has
now produced a new edition which is even bigger and better than the original volume, xl + 825
against xxvi + 747 pages, with an enlarged format which permits more entries per page, and with
many more references in the bibliography. As well as providing data on a further 3000+ botanists,
gardeners and nurserymen, the scope of the book has been extended to include flower painters,
botanical artists and garden designers.

The book follows the arrangement of the earlier edition with dates and places of birth and death
of each of the individuals, followed by a concise biography. A further paragraph provides data on
books and papers written by the individual, and a final paragraph gives details of sources of further
information. The rearrangement of the different categories into separate paragraphs adds greatly to
the clarity of the work.

Eight years’ scanning of periodicals and books was undertaken in the production of the new
edition, but it is a pity that the termination date for periodicals was as early as December 1990 as this
excludes entries for such worthies as John Codrington (1898-1991), John George Dony (1899-
1991), Ronald d@’Oyley Good (1896-1992), Mary Patricia Happer Kertland (1902-1991), William
Arthur Sledge (1904-1991) and Bryan Thomas Styles (1934-1992). There appear to be few
omissions, though Richard Pearse Libbey (1911-1987), a Norfolk botanist who studied Oxalis and
Poaceae and whose herbarium is now at the University of Leicester (LTR) is not mentioned, nor are
Clara Winsome Muirhead (1915-1985), an authority on the flora of Cumberland who worked at the
Royal Botanic Garden, Edinburgh, and whose plants are at Carlisle Museum (CLE) and Plymouth
(PLH), or Effie Moira Rosser (1923-1987), author of Senecio cambrensis, whose types and other
plants are at Manchester Museum (MANCH).

Errors, as are to be expected in a work of this high standard, appear to be remarkably few, but
John Blackstone’s plants (p. 77) were donated to the British Museum (Natural History), London
(BM) by Ripon Museum as long ago as 1947, Frederick J. Hutchison died in 1891, not 1981, and
Barbara Welch’s maiden name (p. 728) was Gullick not Gullicky.

Ray Desmond is again to be congratulated on a fine revision of his most useful reference work,
which despite its high price should be in the hands of all those who are seriously interested in the
history of Irish- and British-born botanists and gardeners.

D. H. KENT

Field guide to wild flowers of Britain and Northern Europe. R. Gibbons & P. Davies. Pp. 330,
illustrated. The Crowood Press, Marlborough. 1994. Price £9.99 (ISBN 1—85223-784-8).

Field guide to trees of Britain, Europe and North America. A. Cleave. Pp. 320, illustrated. The
Crowood Press, Marlborough. 1994. Price £10.99 (ISBN 1—85223-801-1).

308 BOOK REVIEWS

The field guide market is a crowded one and some people feel it is in danger of being overwhelmed
by a flood of clonal works offering little more than recycled information packaged in slightly
different ways. As an author of field guides myself I have been told that no new works of this genre
should be written. Obviously I disagree with this extreme sentiment. Nomenclature changes over
the years if nothing else, and even favourite references eventually become outdated or too worn to
be risked in the field, so it is worth perusing new works with an eye to finding suitable replacements,
preferably ones offering a little extra—a greater coverage (in terms of species or area) than is already
available, new information or easier access, better illustrations or some other significant improve-
ment.

How, then, do these latest offerings from the Crowood Press measure up? They are genuinely
pocket-sized, well-bound with attractive covers and modestly priced for books illustrated with over
720 colour photographs. Descriptions are brief and avoid jargon, those for wild flowers being very
succinct while the guide to trees uses a more discursive style. The two books are clearly meant to
form part of a series but beyond the points mentioned, the similarities diminish. The Field guide to
wildflowers embraces Britain and Northern Europe (i.e. the British Isles and northern France
eastwards to Germany, Denmark and the southern tips of Norway and Sweden). The usefulness of
the concept of Northern Europe is at best questionable and in my view usually denotes a publisher’s
ruse to encourage take up of foreign editions while maintaining a British market, rather than an
attempt to add real value to the book. Another sure sign that the Northern European factor has
been added simply to increase sales potential is the giving of separate distributions of species for
Britain and for the continent, as is done here. Like almost all modern flower guides, this one
contains only a selection of the flora of the area and its claim to be comprehensive is somewhat
tongue-in-cheek. However, with a species list exceeding 1400, it compares favourably with other
works. While there are many good photographs the overall quality is patchy. I found this surprising
since the authors are experts who normally produce a consistently high level of work.

The Field guide to trees fares better. The much smaller total number of tree species allows a better
coverage in the book and, while its usefulness as a reference for North America in doubtful, it does
include most of the common introductions from that continent encountered in Europe. The overall
quality of the photographs is also better and many species are represented by several shots to
illustrate the different features of the tree. The descriptions are followed by comments giving
additional characters and snippets of interesting information.

My biggest single complaint about these books (and many similar ones) is how to get into them.
There are no keys in either book, with Gibbons & Davies claiming these ‘“‘demand a level of detail
. . . finer than most people will want’’. Fair enough, though I find the reluctance of authors to tempt
inexperienced users with simple, usable keys regrettable. In photoguides of this kind, especially for
wild flowers where lack of space restricts species to a single photograph, many species will be
inadequately portrayed for rapid identification and some bear little resemblance to the plants as
they would be seen in the field. Can less experienced botanists (presumably the main target
audience) expect to ‘get a result’ simply by skimming the pages and ‘picture spotting’ before
checking descriptions, as Gibbons & Davies suggest? I think the answer frequently must be no.
Without any alternative means of entry, potentially useful and attractively packaged books of this
kind remain less accessible than they need be. Experienced botanists, on the other hand, will find
the books little different from many already available. On balance then, Field guide to wildflowers
falls into the ‘clone’ category, while Field guide to trees, by virtue of the extra information, escapes —
just.

J. R. PRESS

Index of garden plants. M. Griffiths. Pp. lxi + 1234; 16 figs. Macmillan, London. 1994. Price £35.00
(ISBN 0-333-59149-6).

This is not so much an index to the Royal Horticultural Society Dictionary of Gardening, rather than
a condensed version of it compiled by Mark Griffiths with unfortunately only a cursory
acknowledgment to some 250 contributors to the original book whose work it mainly is. In order to

BOOK REVIEWS 309

reduce the four volumes to one, the general entries, biographies and the many pages of plates
(except those which illustrate the glossary) have been lost. Family entries have been reduced to a list
of included genera, which is quite useful unless one is looking at Fumariaceae. This family is now
often included in Papaveraceae and this seems to have been the fate of Corydalis while related
genera such as Dicentra remain in Fumariaceae. Descriptions have been cut to the bare essentials,
with no information on cultivation, but room is found for number of species, hardiness and
distribution details. One particularly important piece of information which seems to have been lost
is reference to the poisonous nature of any of the plants listed.

In several ways I prefer the layout to that used in the original. Common names are incorporated
into the alphabetical list, and synonyms, instead of being relegated to the end of each genus, are
incorporated into the list of accepted names. One problem I am sure some will have, is the size of the
text which is very small and may test the eyes of those with less than perfect sight.

Although the introduction has been much reduced, the comprehensive illustrated glossary is
retained and incorporates a glossary of plant taxonomy. A useful section is included on the naming
of plants. Examples show how names are applied to the various ranks of cultivated plants, and how
different terminations are used. These are often a cause of considerable confusion, for while the
example Crinodendron hookeranum is given, the epithet is correctly cited as hookerianum in the
main body of the text.

Confusion also seems to have arisen over the designation of hybrids, with x used in the examples
but X used in the text. While both are acceptable, usage should be consistent within one work.
When a multiplication sign is used it should always be lower case; however, in the main text (but not
in the examples, where the letter x is used) upper case multiplication signs are used for intergeneric
hybrids, except for x Pyracomeles where a combination can be found. Another confusing and
annoying typographical error is the consistent placing of the multiplication sign immediately before
the name of the second parent in a hybrid formula as if it were an intergeneric hybrid.

The R.H.S. Dictionary itself has been heavily criticised for its numerous spelling mistakes and
other errors. Although some of these have been corrected (for example the cultivars incorrectly
assigned to Buxus harlandii are now correctly placed under B. microphylla) others have not. Ilex x
altaclerensis is still in as altaclarensis although its correct spelling has long been settled. Acer
palmatum ‘Senkakv is still incorrect, A. platanoides ‘Crimson King’ now appears twice, once as
‘Crimson king’, and Helichrysum italicum still grows to 50 m. I could go on.

Although brief, the individual descriptions in most cases give enough information to be useful but
it could be argued that describing the leaves of Pieris japonica ‘Pygmaea’ as “‘blade-like”’ says little
about this very distinct form. Works such as this often fall short on their inclusion of cultivars but we
are told that out of more than 60,000 names, some 30,000 are cultivars. Some inconsistencies do
arise, however, when hybrid cultivars are treated. These are occasionally (as in Ceanothus) given
individual entries, or (as in Pieris) listed under one of the parents, or (as in Diascia) listed at the end
of the entry where (as in the original Dictionary) only very scanty information is given.

The list of entries appears to contain most of those in the Dictionary itself except for “‘the vast
array of cultivars of Rosa or Rhododendron or those of economic plants’’. Indeed I have come
across several additions the inclusion of which seems to have been prompted by works published too
late to influence the Dictionary itself. These include Colutea buhsei (as “‘bushei’’) and C. multiflora,
both unfortunately with incorrect author citations. :

While this work will undoubtedly appeal to gardeners and students of cultivated plants, its use to
those interested in the British flora is less obvious. It is certainly a valuable guide to plants in
cultivation but it is unlikely that many of these will escape and naturalise in this country. Those that
do will not be easy to identify from this work with no illustrations and no keys. I find it hard to
believe, for example, that many of the species of Cotoneaster naturalised here will be easy to name
from the descriptions (apart from the fact that not all naturalised species are included), and one
species prominent by its absence, because it is rarely cultivated, is the only native member of the
genus, C. cambricus.

In spite of its far from minor defects, this will be a useful book and it has, in addition to its
extensive coverage, a major factor in its favour, the price. At £35 it is a bargain, and one that I have
no hesitation in recommending.

A. J. COOMBES

310 BOOK REVIEWS

Index Kewensis on CD-ROM. Compiled by Royal Botanic Gardens, Kew. One Compact Disk and
spiral-bound manual. Oxford University Press, Oxford. 1993. Price £995.00 + VAT (ISBN 0-19-
268003—-X); additional manuals £7.50 (ISBN 0—19-268004-8).

Index Kewensis, that wonderful work that is treasured, cursed, and used daily in almost any
herbarium, is now available on Compact Disk. The books have been like old friends to us — over the
years they have become tattered and sometimes slightly unglued, the pages darkened from the
touch of thousands of fingers over many decades, the edges rounded from wear. Many of us have
copies that have been cut up and pasted back together, and the publication of each Supplement has
required another round of this. Those of us who don’t cut and paste are familiar with the long slow
crawl through each Supplement. /ndex Kewensis on CD-ROM brings this enormous amount of data
into one easily searchable place.

Charles Darwin believed it would be useful to have an updated version of Steudel’s Nomenclature
(1840-1841). Index Kewensis was begun in January 1882 and funded at first by a provision in
Darwin’s will. Index Kewensis today is a list of plant names with bibliographic references to the
place of first publication and the geographic distribution reported in the original publication. It is
compiled by a team of people at the Royal Botanic Gardens, Kew, and is one of the many useful
references produced by that institution.

The Index Kewensis on CD-ROM provides the information from the original publication plus all
19 Supplements, cleaned up and standardized somewhat, in a form that allows a wide variety of
searches and report generation. This was done by electronically scanning the text using an optical
character reader. The project was started some years ago, when scanners were not as smart as they
are now, and a great deal of time and effort has had to be invested in cleaning up errors in
misreading text due to changes in and peculiarities of type-faces over the years. The addenda to the
original two volumes and to Supplements | and IJ were not read by the optical character reader, but
those addenda that could be interpreted have been added to the CD-ROM files.

When one purchases Index Kewensis on CD-ROM, one receives a floppy diskette, the CD-ROM
itself, and a spiral-bound manual which also describes how the software can be installed on a
network. The manual is extremely useful not only because it leads one step-by-step in the use of the
CD-ROM but also because it gives a very clear history of Index Kewensis, including an explanation
of conventions that have been used. To use Index Kewensis on CD-ROM one requires, at a
minimum, an IBM AT or PS/2 or compatible equipped with a hard-disk drive; 2 Mb of free hard-
disk space; 640 Kbd of RAM with 420 Kb free; and DOS version 3-0 or higher. One also needs a CD-
ROM drive and MS-DOS CD-ROM extensions version 2-0 or higher.

It is possible to search seven lists or fields using one of two search methods. The lists include
family, infrafamily, genus, infragenus, species, infraspecies and full author names (data for the
‘infra’ categories only after 1971). Three additional files can be searched using a text search:
publication, notes and author. A few notes on specific files may be useful: the manual warns that
familial and generic concepts have changed over the years. Considerable effort has been made to
standardize author names on Brummitt and Powell’s Authors of plant names (1992), and a search on
the full name will pull up records regardless of the abbreviation used in the original Index Kewensis
entry.

Searches can be done using wild cards and Boolean operators, and one can move from field to
field using a mouse or a combination of keys or arrow keys. At any point one can click on a field and
see a display of the item. One can display or download full records for any taxon or author name, or
one can list names of taxa with publication information or notes. Distribution information is
contained in notes and searches on distribution need to be carefully constructed. It is really a treat to
be able to list all names published in a certain journal, or described by a given individual, or
described over a specified period of years.

Inevitably there are some problems, but considering the magnitude of this project, they are few.
Firstly, the quality of the data of the CD-ROM is not much better than it was in the original book
(except for the author files), and many errors have crept in as a result of the optical scanning
exercise. The folks at Kew welcome corrections. To prevent users from simply absorbing all the data
into their own databases, there is an upper limit on records that can be downloaded at one time.
Names of families and genera are in upper case letters. The infraspecific rank is not displayed in lists.
Searches using many wild cards, or wild cards at the beginning rather than at the end of a complete

eae

=

age + pairs 8 nce abe ME pe INN, AP es cart aids A Pp nal SS

BOOK REVIEWS 311

search with many wild cards may seem to take forever, and I know that some searches have been
terminated after failing to be completed overnight.

These problems are minor, however, compared with the overall utility of Index Kewensis on CD-
ROM, and I think every herbarium that uses the books regularly, and most libraries that might be
reluctant to buy the books but that have patrons who need to look up names of plants, will want to
invest in it. Updates will be issued regularly on subscription; the first is due in 1995 and thereafter
the publishers plan to issue them on a yearly basis.

N. R. Morin

Trees of Ireland, native and naturalized. E. C. Nelson & W. F. Walsh. Pp. 247, with 30 coloured
plates and 30 black and white vignettes. The Lilliput Press, Dublin. 1993. Price IR£17.99
(paperbound) and IR£35.00 (cloth bound) (ISBN 1—874675-24-4).

Trish trees and shrubs. P. Wyse Jackson. Pp. 72, with 32 coloured plates. Appletree Press, Belfast.
1994. Price £3.99 (ISBN 0-86281-420-0).

The nearly simultaneous appearance of two illustrated books on Irish trees suggests, at first sight,
either bad planning or bad luck. This is not so, however, as the books are of very different type, and
aimed at different markets. Nelson & Walsh is an archetypal coffee-table book, and will be bought
by most purchasers mainly with the illustrations in mind. Wyse Jackson’s book is a cheap pocket
manual designed for day-to-day reference.

Nelson & Walsh deal only with trees; their delimitation from shrubs is fully discussed. It is all the
more surprising to find Rhamnus catharticus and Frangula alnus included. I will not deny that they
can attain tree size, but in sixty years’ botanizing in Ireland I have never seen a specimen
approximating to tree dimensions. It makes me wonder whether the text may have been written to
fit a pre-existing set of illustrations. Mrs Walsh’s illustrations, amounting to 30 full-page plates, each
with three or four detailed paintings in colour, and a pencil sketch to show the general form of the
tree, have the attractive elegance and botanical accuracy characteristic of her work. In particular she
must be congratulated on the remarkable accuracy of the colour throughout. It is a pity,
accordingly, that a few plates show one of her characteristic defects — a poor choice of specimen. It is
impossible, for example, to deduce from these pictures what is the shape of a horse-chestnut leaf, or
how many leaflets there are normally on the leaf of an elder.

Nelson’s text contains the necessary minimum of botanical description, but the greater part
consists of a mixture of history, folk-lore and poetry, the last-named often rather thinly linked to the
plant in question. Folk-lore is a legitimate field of serious study, but I think it a mistake to mix in odd
snippets of it with botanical facts; the borderline between fact and fancy is too easily blurred. Nelson
has done some very good work in Irish botanical history, but his enthusiasm sometimes runs away
with him, and there are rather too many quotations from Dr Rutty.

The scope of the book is restricted to native and naturalized trees, and excludes those that are
merely planted. It is rather surprising that Pyrus pyraster should be included; the tree is extremely
rare in Ireland, and I believe that most of the trees alleged to be naturalized have been, in fact,
planted. It could, with advantage, have yielded place to Sorbus intermedia, which is abundantly
naturalized in some districts. Equally eccentric is his treatment of the genus Betula. After a needless
mockery of the methods of distinguishing the two species (fairly difficult, but by no means
impossible), he decides that the only method is to count the chromosomes, and flies in the face of the
virtually unanimous opinion of European botanists by proclaiming that there is only one species, B.
alba L. There are quite a few interesting and reliable pieces of information to be picked out of
Nelson’s causeries, but on the whole one is forced to the conclusion that Mrs Walsh’s illustrations
deserve a less self-indulgent text.

Peter Wyse Jackson’s book is very different. It slips easily into the hip-pocket; it includes shrubs
as well as trees; and it covers a few planted conifers, which are all too often treated as pariahs. In fact
it includes all the woody plants one is apt to see when wandering round the country. The only serious
omission is Populus x canadensis, which is by far the commonest poplar in most of Ireland, and is
recognizable from afar. Pinus radiata could have been sacrificed to allow for its inclusion. In other
respects, however, the choice is sensible. Each species is given about 200 words of text and a half-

312 BOOK REVIEWS

page (75 cm’) of illustration. The text is practical and down-to-earth, with a minimum of folk-lore,
though even Jackson cannot resist telling us that the carrying of a hazel-nut in the pocket “‘was said”’
(more honest than the usual ‘is said’) to keep away rheumatism. The illustrations are variable in
quality; many are entirely satisfactory, but some (Rhododendron and Escallonia, for example) are
misleading in their colour, and a few others have faults in design. In most of the willows and a few
others the leaves appear too glossy. Nevertheless, with only a few exceptions they will serve well
enough as aids to identification.

If Jackson has a tendency to political correctness it is to be found in the ‘Food for free’ syndrome.
His assessment of the gastronomic value of the various ‘wines’ and other products which he cites as
derivable from the trees and shrubs of the countryside is often too enthusiastic. He even tries to
make us eat ash-fruits: “‘its fruits can be eaten if picked very young, boiled and pickled in vinegar’.
‘Can’, I think, is the operative word.

D. A. WEBB

Sedges and their allies in Dorset. An atlas of the distribution of the family Cyperaceae in Dorset. D. A.

Pearman. Pp. 108. Dorset Environmental Records Centre, Dorchester. 1994. Price £5.00 (SBN 0- | !

951139444).

Having been brought up to believe that Scotland was the place to visit to see sedges, I was surprised
to learn that Dorset contained 69 species of Cyperaceae, probably more than in any other county
and almost 70% of all the British species. For this reason among many others the Atlas under review
is of much more than local interest. It covers modern administrative Dorset, chiefly v.c. 9 and a part
of v.c. 11, and includes maps on a 1-km square grid for all species and subspecies, post-1980 records
being distinguished from pre-1980 ones (what has happened to the 1980 records is not made clear).
The text for each taxon is full of interesting information on the Dorset habitats for the plants,
associations with other sedges, hints on separation from other similar species, the distribution in
Dorset with lists of sites for the rarer species and a note on the British distribution.

There is a great emphasis on historical records and on changes in abundance or distribution.
Several of the larger waterside species seem to have disappeared from most of their former sites,
and changes in grazing regimes, lowering of water-tables and spread of Phragmites australis
(because of increase in nutrients) are suggested as possible causes..Lack of grazing and consequent
overgrowth of many of the sites for other species seems another major reason for losses. Much use
has been made of the uniquely detailed survey of Dorset by Professor R. d’O. Good in the 1930s,
when he made species lists for over 7,000 vegetation stands. I was sorry to learn from the Ad/as that
the forthcoming new Flora of Dorset will not have space for similar discursiveness on historical
changes. It is ironic that whilst one of the most frequently cited justifications for producing local
Floras is to document changes, this aspect is so often given short shrift when it comes to the crunch.
The present Aflas is a most welcome attempt to draw conclusions and not just to add yet more
undigested information.

There is a most interesting introductory chapter on habitats, including discussions of the historical
changes, species to be found in each type and a selection of accessible representative sites. The few
drawings by Robin Walls scattered through the book are so expressive of the characters of the
species depicted that I wish there had been room for more. The Latin nomenclature follows Stace
(1991) and Kent (1992). Among the English names used are several that are new to me. Whether
they are of local or personal origin is not made clear, but I hope that one of them at least,
Gingerbread Sedge for Carex elongata, will become popular. Errors seem few, but a couple could
seriously mislead: /solepis cernua is said to be “easily told from J. setacea by the wrinkled seed’’, but
it is the latter that has the longitudinally ribbed nut; and the male glumes of Carex acutiformis
cannot be described as ‘“‘ovate or truncate” for they are oblong-lanceolate and subacute or obtuse.
The subspecific epithet of what used to be Carex demissa is several times mis-spelled, perhaps
reflecting the author’s stated difficulty in accepting the taxonomy. An odd omission is any mention
of the subspecies of Eleocharis palustris. Subsp, palustris is shown in the Critical supplement to the

BOOK REVIEWS 313

atlas of the British flora for two 10-km squares (SZ/1.9 and SU/1.0) that are partly covered by the
Dorset Atlas, so it would have been helpful to have made clear whether any of the Dorset plants are
this or whether they are all subsp. vulgaris.

Anyone interested in sedges should learn a lot from this work. As Clive Jermy says in his preface,
it illustrates how plant recording on a local level can, when properly written up, contribute to
conservation and academic understanding in a wider context. The author and the Dorset
Environmental Records Centre (which only last year produced its County red data book) are to be
congratulated on producing this work and on making it available in such an attractive format and at
such a reasonable price. The rest of us must look to our laurels.

A. O. CHATER

Wild and garden plants. S. M. Walters. Pp. 200 with 16 pp. of colour plates. HarperCollins
Publishers, London. 1993. Price £27.50 (ISBN 0-00219376-0).

How far are the English passions for gardening and ‘wild nature’ linked and how far are they
opposed? Max Walters poses these questions in his foreword and explicitly makes the answering of
them the underlying theme of this volume, the 80th, in “The New Naturalist Library’.

Wild and garden plants is divided into three parts, each comprising four chapters. The first quartet
explores general topics under these headings: “The sources of wild and garden plants’, “The kinds of
plants’, “The variation of plants’ and “Hybridisation and sterility’. The author’s emphasis is on basic
botanical matters including, for example, the definition of a species and a genus, the application of
Latin names, plant ecology and the variability of plant populations, the history of the native flora,
and the recognition of a ‘native’ plant. Dr Walters also pursues briefly such topics as hardiness, and
the impoverished flora of ‘‘what we now call the British Isles’’. The second part, headed ‘Life forms
and adaptations’, explores another four subjects, broadly defined groupings of plants: native and
exotic trees, shrubs and shrubberies, some ‘special’ life forms including parasites, carnivores,
succulents and epiphytes, and in a chapter on ‘Herbs, ‘Flowers’ and ‘Grasses’’ Dr Walters
expatiates on the different meanings that the word ‘herb’ has for pedantic botanists and common-or-
garden folk. The last set of four chapters is headed “Botany and horticulture as modern hobbies’: the
individual chapters are entitled “The science of genetics and horticultural practice’, ‘Botanists,
gardeners and social change’, “The rise of ecology’ and ‘Late twentieth century attitudes’. The final
chapter includes a diversity of subjects from the work of the N.C.C.P.G. (National Council for the
Conservation of Plants and Gardens), to the European garden Flora project, and the spread of alien
plants of garden origin.

The book is illustrated with black and white photographs, a variety of line drawings and 39 colour
plates.

‘The New Naturalist Library’ is an estimable and collectable series of natural history books. The
aim of these monographs remains, as the editors affirm in their preface, to make up-to-date
information and ideas accessible to all with an interest in the countryside and its wildlife. To this
end, previous volumes have had excellent, comprehensive bibliographies. The bibliography in Wild
and garden plants is restricted to three pages, albeit set in double columns, which is parsimonious
compared with as many as 30 pages in former volumes.

The flora of gardens within our lands is immeasurably rich in exotic species and those plants of
various origins, called cultivars, that have been deliberately selected and are deliberately
maintained merely to beautify and exhilarate gardeners. Max Walters’ book provides some insights
into the biological background to the garden flora, and interweaves fascinating details about the less
bountiful native flora and the naturalized plants that inhabit the envelope of countryside beyond
garden hedges.

With such phrases as “‘the green English countryside”’ or “‘the two ancient Universities” sprinkled
liberally throughout, indeed from the first sentence, this is very much an England-centred book,
which is a pity for surely the other nations that share the Celtic Archipelago possess equally
interesting wild floras, and their people have as much of a passion for gardening and wilderness and
have contributed as significantly to the creation, maintenance and embellishment of fascinating

314 BOOK REVIEWS

gardens. Indeed, the repeated use of the adjective ‘English’ becomes irritating when it was inexactly
juxtaposed with statements or illustrations referring obviously to a wider context (for example, a
discussion of epiphytes in ‘English’ woodlands illustrated by a photograph of ivy (Hedera helix) and
polypody (Polypodium vulgare) on a tree in Anglesey). The author and his editors might have taken
more care to ensure that this book served a wider context than eastern England.

Gardening is an intensely personal activity and many gardeners are eccentrics with contagious
enthusiasm, yet there were very few people in this book. Plants also have characters — colours,
scents, textures, extraordinary habits — but they are rarely described with enough enthusiasm to
ignite the reader’s mind to paint enticing pictures in the mind’s eye. I tried hard to discover that
passion for gardening and wilderness presaged in the foreword. The English attitude towards
gardens and wild nature is strangely impersonal and muted when envisioned through Wild and
garden plants.

Wild and garden plants is an erudite book, informed by the author’s life as a botanist and director
of the University of Cambridge Botanic Garden, but its pages failed to enthrall me.

E. C. NELSON

Flora of the Christchurch area. F. Woodhead. Pp. 120, ten figures, four colour plates. Privately
published, Bournemouth. 1994. Price £7.95 (ISBN 0-9522857-0-3). Available from 28 Hungerford
Road, Bournemouth, BH8 0EH, price £9.00 including postage and packing.

This new Flora, on a 1-km square basis, covers much of Britain’s fastest growing urban area, most of
which is on the acid Tertiary soils which extend west towards Dorchester and east to the New Forest
and beyond. E. F. Linton, in his Flora of Bournemouth, covered this area too, but that was 80 years
ago. There is tremendous pressure here from roads, housing, airports, and from leisure activities,
but there are some superb remaining sites sandwiched in between, such as Christchurch harbour,
the lower Avon valley, the heaths around Hurn, and the Bournemouth cliffs themselves, where
many ‘Red Data’ and ‘Scarce’ species have been found.

This work covers an area of v.c. 11, South Hampshire, which was absorbed into Dorset in 1974,
with the addition of a small part of modern Bournemouth from v.c. 9, Dorset. Thus although it has
no natural boundary, it is a whole in itself. The same area will be covered by the long-awaited new
Flora of Hampshire, but that work will be on a tetrad basis and will, I understand, cover records
from the 1960s onwards, whereas this volume incorporates records from 1981 to 1993. The proposed
new Flora of Dorset might await developments from the boundary commissioners before covering
the same area again! Incidentally the part of Bournemouth that is most densely urban is not actually
covered in this work. Perhaps it should have been for completeness.

The introductory chapters are concise but comprehensive, with clear maps and explanations.
Figure 4 shows the extent of the conurbation, and reinforces one’s surprise that eight S.S.S.I-s are
found within the survey area. The records are all made by the author, and incorporate a few pointed
out by others and subsequently verified. It is this aspect that would be the basis of the only
reservation I feel about the book. By all means check anything rare, or critical or unlikely, but to
eschew all records of others seems slightly surprising, and is certainly at variance with almost every
other recent county Flora. Perhaps the very smallness of the survey area — 122 1-km squares — is an
excuse for this approach.

Overall the 1-km square grid coverage works well indeed. I often feel most dubious about what
tetrad maps actually show, other than a generalised demonstration of surface geology. Site surveys
seem far more preferable, and their incorporation into maps at least serves two useful purposes. The
1-km square grid is intermediate between the two methods and is fine enough to convey some of the
benefit of a site approach. The maps are clear, and of course, are far better interwoven with the text
than in a separate section at the end. I feel it is carrying secrecy too far to omit maps of Osmunda
regalis, Orchis morio and all dactylorchids; there are enough real threats without inventing more.
The commentary is to the point, and places the records within the context of the historical position
and that of the rest of Dorset (but not Hampshire).

The book is spaciously and pleasingly laid out. As with many recent county Floras there are the
obligatory colour photographs, presumably for the benefit of a ‘wider public’. I like very much the

BOOK REVIEWS 315

idea of a ‘local’ Flora, especially of a definite area. Whilst this particular area might not have the
immediate appeal of the Lizard or the Avon Gorge, perusal of the contents reveals a wealth of rare
plants and it can only be a timely contribution to conservation by its presentation of hard facts on a
threatened area.

D. A. PEARMAN

Flora of Flintshire. G. Wynne. Pp. 400, with 25 colour and three black and white plates. Gee & Son,
Denbigh. 1993. Price £25.00 (ISBN 0-7074-0224-7).

Flintshire can now be crossed off the decreasing list of counties that has never had a county Flora
thanks to this work by Goronwy Wynne. Although no longer existing as an administrative unit,
being amalgamated with Denbighshire to form Clywd in 1974, Flintshire covers the Watsonian v.c.
51. This makes a compact recording unit some 38 km X 16 km forming the north-east corner of
Wales, every part of which is within half an hour’s drive of the author’s home. It is 30 years since the
death of Arthur Dallman who, although he never published his Flora of Flintshire, left extensive
botanical notes covering the first half of this century and indeed a draft manuscript for his projected
Flora. These are housed in the Botany Department at the National Museum and Galleries on
Merseyside. This current Flora, which draws heavily on Dallman’s work and for which fieldwork
started in the early 1970s, therefore spans virtually the entire 20th Century. It is apparent that
Goronwy Wynne’s surveys are no less meticulous than those of Arthur Dallman before him.

The Flora is divided into three sections. Part I deals with the description of the county and
includes sections on botanical exploration and a guide to some of the best botanical localities
(omitting some sensitive sites). The climatic data are well presented and demonstrate that even in
such a relatively small area as Flintshire, climatic variations do occur. The effect of climatic variation
on species distribution within the County is dealt with in Part II.

Part II explains the survey techniques and analyses the results. Species recording is at the tetrad
level, in common with the majority of recent county Floras. There are 172 tetrads in Flintshire. In
order to attempt to explain patterns of plant distribution in Flintshire, a total of 119 environmental
attributes were identified and assigned to each tetrad. These environmental attributes range from
soil type to altitude and include land use such as golf courses, railways and built up areas. By using
the TWINSPAN ordination technique on these environmental attributes, eleven different land
classes were identified in Flintshire. It could be argued that the 2-km square is rather large for such a
technique, but it appears that a series of clearly defined units has been produced.

The species records were also subjected to TWINSPAN ordination and produced 13 different
floristic classes. Clearly one test of whether such techniques is valid is to see if there is any
correlation between the land and floristic classes. Although statistically tested (using DECOR-
ANA) it is also visually apparent that there is such a correlation. Geographical elements, floristic
gains and losses as well as individual habitat studies are also included in Part II.

Part III is concerned with the details of individual species occurring in Flintshire. This is
comprehensively and clearly presented. Of great value are the tetrad distribution maps for every
species with more than one record. Many of these additionally have a small graph attached showing
the species’ position on the first two axes of the DECORANA ordination.

The tri-lingual index is rather lengthy. It might have been more manageable to index Latin names
separately from Welsh and English ones. A similar problem occurs with the gazetteer. All the places
named in the Flora are listed in the gazetteer with a 4-figure grid reference. Their inclusion on a map
' is therefore superfluous, especially as this necessitates the use of very small print.

___ Presentation of the Flora is most attractive and the text is well laid-out, although not without the
- occasional typographical error (such as the addition of a ‘t’ in rainfall on page 81). The excellent
_ colour plates are a welcome addition. Both colour photographs and plant portraits by Margaret
Gillison Todd and Jean Hughes are included. Their presence in such a competitively-priced book is
due to sponsorship of individual plates by organisations such as the Countryside Council for Wales
and the Liverpool Botanical Society. Whilst admittedly the Wood Horsetail (Equisetum sylvaticum)
is perhaps no great botanical art subject, its black and white photograph along with three other

316 BOOK REVIEWS

uncommon Flintshire plants (Pyrola rotundifolia, Epipactis phyllanthes and E. helleborine), is
unfortunately lifeless in comparison to the colour plates in the book.

The Flora of Flintshire goes a long way to satisfying a wide readership. It has a high scientific
content, but at the same time is very readable and will be of value to land managers and
conservationists as well as botanists.

B. D. GREENWOOD

Watsonia 20: 317-326 (1995) Si

Obituaries

ERIC SMOOTHEY EDEES
(1907—1993)

Eric Edees was born at Runcorn, Cheshire, on 5 June 1907 and died on 14 October 1993. He was the
son of Isaac Edees, a Methodist minister, and Eric himself was for many years a lay preacher in the
Methodist church. The family moved to Manchester, where he was educated at Manchester
Grammar School, and went on to read Classics at Manchester University. In 1930 he moved to
Stoke-on-Trent as a history teacher, and from that time Staffordshire was his adopted county.

He joined the North Staffordshire Field Club in 1938 and was elected Chairman of their Botany
Section in 1941 on the retirement of W. T. Boydon Ridge. He held this post until his death and was a
most active and distinguished member of the Field Club. He joined our Society, then the Botanical
Exchange Club, in 1943. Eric’s long-term study of Staffordshire plants resulted in his Flora of
Staffordshire, published in 1972. He was a founder member of the Staffordshire Wildlife Trust.

Eric Edees’ important contributions to British field botany were recognised by the award of the
North Staffordshire Field Club’s Spanton Medal (1948) and Robert Garner Silver Medal (1955), the
Linnean Society of London’s H. H. Bloomer Silver Medal (1975) and a Master of Science degree by
Keele University. He was elected an Honorary member of the B.S.B.I.

He was a quiet and courteous man, but held strong opinions. He was a natural teacher with a wide
knowledge of plants, who would patiently explain in the field the diagnostic characters of his
beloved Rubus to everyone interested. His wife Margaret, who gave him much support and help
with the Flora, predeceased him, but he is survived by two brothers and a sister.

B. R. FOWLER

ERIC EDEES’ CONTRIBUTION TO BRITISH BATOLOGY
I first met Eric at a Breckland field meeting in 1965, but it was not until 1967, when I needed to make
progress with Rubus for the Cheshire Flora, that a closer relationship developed. I had sent a few
specimens for his determination, but had omitted to ensure that the pressings retained the essential
characters of the plants. They were cursorily returned unnamed, but he invited me to meet him on
our county march to inspect the local brambles on both sides of the line. Thus began a 20-year period
of fruitful cooperation in bramble study.

Eric first came to grips with Rubus in 1936 when he submitted local gatherings to W. C. Barton.
Later he turned to Francis Rilstone for enlightenment and assistance in naming specimens.
Unfortunately, Rilstone’s detailed knowledge was mainly of the Cornish bramble flora, but
nevertheless he was able to name the more widespread Staffordshire plants satisfactorily. This early
collaboration, conducted energetically on both sides, culminated in the joint publication of a local
Staffordshire bramble as Rubus daltrii in 1945.

Thereafter, specimens were submitted to William Watson, then the Botanical Exchange Club’s
referee for Rubus. Eventually, in 1950, Watson visited Staffordshire to examine growing bushes _
which he could not name satisfactorily in the dried state. Eric was not impressed by the initial
hesitation on encountering the plants, followed later by pronouncement of an obscure name
exhumed from the recesses of Sudre’s Rubi Europae. Watson’s view at that time apparently was that
inability to ascribe a name amounted to a stain on his reputation, an attitude completely at variance
with Eric’s careful, methodical, and above all cautious approach to the determination of Rubus
specimens.

318 OBITUARIES

Following Watson’s death in 1954, Eric became the B.S.B.I.’s referee for Rubus and published
the first of his ‘county’ articles, on the brambles of Staffordshire, in 1955. These consisted of a list of
species, with notes on the features, taxonomic history and distribution of each. As referee, he was
approached for help by several county Flora writers, which resulted in accounts for Derbyshire in
1959 and Lincolnshire in 1966. While these, together with visits to Cheshire in support of my own
1971 Flora, were a natural development of his Staffordshire experience, receipt of gatherings from
Scottish and Welsh botanists, in particular Miss U.K. Duncan, Miss M. McCallum Webster and T.
A. W. Davis, reinforced his developing conviction that a number of taxa lacked descriptions in the
literature and could not be satisfactorily named from either Rogers’ or Watson’s handbooks. His
outlook on methodology is well set out in “The difficulties of a Rubus referee’ (Proceedings of the
Botanical Society of the British Isles 3: 281-282, 1959). Visits were made further afield during
summer to Scotland, Mid-Wales, Cumbria, Norfolk (at regular intervals), Suffolk, Hampshire and
Cornwall.

B. A. Miles had, from 1961 onwards, set himself the task of refinding Watson’s and J. E.
Woodhead’s collecting sites, amassing much definitive material of taxa described in Watson’s
Handbook. Based in south-eastern England and applying himself to the task with vigour, he built up
a detailed knowledge of the brambles of the Home Counties, both in field and herbarium, as well as
a burgeoning distrust of Watson’s taxonomy. Eric was pleased to find a batologist familiar with the
South-East to complement his own mainly Midland and Northern experience. Specimens were
exchanged, problems ventilated and provisional conclusions reached about many of the con-
undrums discussed in a lively correspondence. A few tentative proposals were made, e.g. to embark
on regional reviews of bramble taxonomy and nomenclature. Eric visited Miles in 1968, when
specimens were examined and cooperation within spheres of influence mooted. No definitive plan
of action had, however, been agreed before Miles’ untimely death in January 1970.

I had already decided to publish some of the unnamed Cheshire brambles for inclusion in my
Flora (virtually my batological apprenticeship). Fortunately our contiguous counties allowed
frequent Saturday visits to Eric’s house at Newcastle-under-Lyme for reviews at first of MANCH
specimens, then of our own collections, as well as parcels from enthusiasts, who now included
Messrs Kenneth and Stirling from Scotland and Messrs Ironside-Wood and Bull from eastern
England. Eric had been given a number of Rilstone’s specimens (including continental sheets) by
Woodhead and also a selection of Watson’s herbarium. This material, together with his own
expanding gatherings, became the reference point for comparison and discussion.

When I broached the subject of possible publications, including a revision of Rubus in Britain, he
was lukewarm on anything other than staged regional accounts, insisting that our knowledge was
inadequate. It was clear that, if the British Rubi were to be better understood, syntype material of
British and Continental authors must be critically examined. ~

In 1972 I determined to collect together the relevant material for study, no easy task as it is
scattered across the herbaria of Europe. I was fortunate to enlist the support of the late David
Valentine, then Professor of Botany at Manchester University, in gaining access to the biggest and
best British collection, the Barton and Riddelsdell herbarium (BM). Barton’s photographs and
notes, the fruit of 25 years’ study, including discussion of Muller’s and Focke’s taxa, as well as the
whole of Sudre’s Batotheca Europaea, were sent on loan to Manchester Museum, where Charles
Bailey’s Rubi were also made available. These, together with sheets loaned from Kew, the South
London Botanical Institute and Lausanne, with Rogers’ BM syntypes, enabled me to assemble
groups of associated specimens. Referring to the literature passim, I began to review the Rubus list:
as assessment was completed, I took the groups of specimens to Eric for his independent opinion.
We then compared notes and found that in most cases our views coincided. It became clear that it
was sensible for each of us to concentrate on our adopted fields of interest. I began to compile
articles for publication, and after some hesitation, Eric followed suit, not however until I had faced
him with the thought that a chance encounter with a bus could render his hard-won expertise
abortive: his published contribution was vital if progress was to be made.

Two developments in the mid-1970s served to advance British batological aspirations. First, our
growing contacts with the Continent, particularly H. Weber in Germany and H. Vannerom in
Belgium, enabled concepts to be challenged or confirmed and elucidation of the N. W. European
bramble flora to reach an advanced state. Second, there was the launch of a projected ‘Flora of
Great Britain and Ireland’, for which Eric and I committed ourselves to write Rubus for Volume 2.

OBITUARIES 319

This task filled almost four years and required a disciplined, systematic approach. Eric
concentrated on descriptions and keys, while my contribution embraced the geography, ecology and
nomenclature. There was, of course, regular discussion on problems in both spheres. Joint meetings
were held in the field and once in the Cambridge University Herbarium, where the profusion and
diversity of material readily available, along with library facilities, provided a major stimulus for
research.

Unfortunately the Flora project was becalmed by the ill-health of two of the three editors and our
Rubus account languished until, in 1985, I approached the Ray Society with a view to publishing our
work as a monograph. By this time Eric had retired from active batology and compilation of the
introductory material fell to my lot, as well as the assembly of supporting features and liaison with
our stalwart editor, D. H. Kent. Eric’s eagle eye, was, however, cast fruitfully over the final draft
and he was quietly pleased and proud to look through the finished product in July 1988.

Through the years of our collaboration, his steadfast adherence to defined objectives, sound
judgement based on meticulous attention to detail, with discussion lightened by bluff, occasionally
quirky humour, made for a congenial relationship which I shall always cherish.

A. NEWTON

ERIC EDEES, NORFOLK AND NORFOLK BRAMBLES
Eric Edees’ wife, Margaret, was born in the village of Necton, near Swaffham, Norfolk, and her
family were still living there a few years ago, and probably still do. As a result, many holidays were
spent at Necton, in the course of which Eric collected sufficient material to produce a short MS
entitled ‘The Flora of the Swaffham district’. It was never published, but his efforts were not wasted,
as the MS was passed on to Eric Swann, co-author of The Flora of Norfolk, in which the use of
Edees’ work is acknowledged. Eric (Edees) told me on one occasion that, when the family went to
the sea for the day, either to Blakeney or to the Wells/Holkham area of the North Norfolk coast, he
would go with them, then turn round and walk the 30 or so miles back to Necton, botanising all the
way.

I first met him at the end of the 1960s. In the previous two seasons I had been struggling
unsuccessfully to come to terms with Rubus, using W. C. R. Watson’s Handbook, and sending a
small package of wildly inaccurately identified specimens to Eric for ‘confirmation’. The only one I
ever got right at that stage was R. ulmifolius! In 1970 I wrote and asked if I might impose on his
goodwill again and send another parcel, to which he replied that he would be visiting his wife’s
relatives during October, and if I cared to wait until then, he would call and name them. Then, if I
had a site nearby with a number of species in that we could walk round, he would name those that he
could so late in the season. I could mark them, and examine them during the summer of the
following year, and I would then have a basis on which to build.

We lived at Foxley at the time, with the large area of ancient woodland, Foxley Wood, just 1 km
away, which I had permission to visit. The day duly arrived, and so did Eric and Margaret Edees, my
gatherings were named, and he expressed himself surprised at one or two of my discoveries. We
spent an hour walking around the wood and I had 14 named species to collect next season — indeed
the basis for all my subsequent work on Rubus. Within a year or two, it became obvious that in
Norfolk, as in most other counties, there were brambles that did not have names, and two stood out
as being widespread and locally abundant.

In 1977, Eric came to stay with us for a week, and we visited many sites in Norfolk and Suffolk.
From gatherings made on this visit, the names R. boudiccae and R. norvicensis were both
subsequently published. Since then, both have been discovered to have a much wider distribution
than East Anglia, with the former here and there across central and southern England, and a
number of sites in southern Ireland, and the latter as far south and west as Hampshire and
Guernsey.

I felt sometimes that Eric’s fascination for brambles was something of a ‘love-hate’ relationship,
which was borne home to me whilst investigating R. norvicensis in depth. We stood in a large
woodland clearing with a seemingly endless mix of plants all around, some of which were nameable,
several others not. Turning slowly through 180°, glaring silently all around as he did so, he suddenly

320 OBITUARIES

exclaimed: “I hate blackberries!””. We all feel like that from time-to-time, but 25 years of study of
them has given me a deal of satisfaction — a satisfaction which I should not have had, had Eric not
gone some distance out of his way to help a beginner.

A. BULL

RICHARD WILLIAM (‘DICK’) DAVID
(1912—1993)

To his large circle of friends and colleagues, Richard David, who died on 25 April 1993, will always
be known and remembered as ‘Dick’. It is some indication of the breadth of his interests and the
attractive qualities of his personality that very many botanical friends had little or no knowledge of
his considerable academic achievements as a Shakespearean scholar or his professional career in
learned publishing as University Publisher and Secretary to the Syndics of Cambridge University
Press.

Dick David was born on 28 January 1912 in Winchester, where his father, the Rev. F. P. David,
was a Housemaster at the College, and Dick went there as a Scholar. His academic career was
smoothly brilliant, and he entered Corpus Christi College to read first Classics and then English in
1931. After graduating, followed by a year’s research in Paris, he wrote a Prize Essay on
Shakespeare’s dramatic poetry, which was published by Cambridge University Press in 1935 when
the author was only 23. In the following year he joined the Press as assistant secretary. This
publishing career was interrupted by wartime service in the Royal Navy, in which he rose to be a
Lieutenant-Commander, and after the war Dick returned to the Press and rose steadily to his
leading position among academic publishing, a service recognised by his C:B.E. in 1967.

Whilst English literature and Shakespearean studies were Dick’s main academic interest, and
publishing his professional expertise, he was keenly interested in the countryside and in field botany
in particular, having acquired at an early age his mother’s love of flowers. Dick’s own delightful
account of how he was given the “Bentham & Hooker Illustrations on his fourth birthday and
encouraged by his mother to paint in the daisy and the dandelion” can be found in his 1980
Presidential Address to the B.S.B.I., ““Gentlemen and Players” (Watsonia 13: 173-179, 1981).
Throughout his career, the family holiday home at Polzeath on the beautiful Camel estuary in
Cornwall drew Dick most summers, and immersed him in a very different botanical world from that
provided by Cambridge; and his friendship with John and Faith Raven, together with his enthusiasm
for fly-fishing, also made him a regular holiday visitor to Ardtornish on the marvellous Morvern
peninsula in Argyll, again a totally contrasting botanical scene.

Dick’s Presidential Address reminds us that his own contribution to botany can be taken as a
model of the value of that close link between amateur and professional which is undoubtedly one of
the main reasons why the Botanical Society of the British Isles continues to serve the science of
taxonomy so well. With long service on the Society’s Publications and Records Committees, and
following an established tradition of alternation between amateur and professional in the post, Dick
became President of the B.S.B.I. for the customary two-year stint from 1979 to 1981, and in that
position made his unique contribution. It was a time of continuing change in the B.S.B.I., which
needed to address important questions of orientation and policy, especially in the field of the
conservation of the natural environment. The Society’s pioneer role in the Distribution Maps
Scheme in the 1950s and 1960s had firmly established the value of systematic recording of the British
flora in determining policies for nature conservation, and Dick put much effort into liaison with both
official and voluntary conservation bodies to shape the way ahead. He had, as early as 1955, become
the B.S.B.I.’s Recorder for East Cornwall (v.c. 2), and later in 1961 took on the post for West
Cornwall (v.c. 1) as well. From these interests developed his friendship with Len Margetts, with
whom he collaborated to produce the Review of the Cornish Flora, 1980 in 1981. Len recalls (in
Davies (1994)) how “‘his quiet humour, modesty and expertise made him an ideal companion, and
our shared fascination for the lesser critical groups — Euphrasia (eyebrights) and Fumaria
(fumitories) in particular — ensured a succession of Cornish venues.”

OBITUARIES 321

What of the sedges, the botanical passion of Dick’s later years? In his account of his own
friendship with Dick, Clive Jermy (in Davies 1994) reminds us how it all began: “In 1968 the
Botanical Society of the British Isles published a little book aimed at helping botanists identify
British and Irish sedges. It had been written by myself, together with my former tutor, Tom Tutin,
under whom I had researched the group at Leicester University. As with all books, we as authors
had a few copies to give to those friends who had helped, and when those were satisfied I had one
left. Tom said “I think you should send that to Dick David at the Cambridge University Press. I
think he likes sedges!”

Tom Tutin’s pithy sentence was proved to be a monumental understatement, for, in his
retirement, Dick became devoted to the British sedges and obviously found, in his careful, planned
survey of the surviving field populations in the British Isles of many of the rarer Carex species, a
perfectly satisfying way of combining his zeal and expertise in field botany with his deep enjoyment
of the ‘wild places’ of the British and Irish countryside. In this he was following an honourable
tradition of gifted amateurs. He did much miore than mere recording, however; stimulated by his
growing friendship with two outstanding professional botanists, Arthur Chater and Clive Jermy, he
tackled the notorious ‘critical’ sedge groups of Carex muricata and C. flava, and the fruits of their
joint labours are available to all to appreciate in the second edition of the Society’s Handbook,
entitled Sedges of the British Isles (Jermy, A. C., Chater, A. O. & David, R. W., 1982) — a book,
incidentally, that is much appreciated and used by many Continental European botanists as
providing an up-to-date and definitive survey of an important European genus.

In his attitude to that other great traditional British hobby, gardening, Dick differed significantly
from many of his fellow-botanists in having a foot in both camps. He could search for hours for an
insignificant rare alpine sedge of no conceivable interest to gardeners, but he also developed a
parallel passion for that very attractive horticultural genus, Crocus, which provided him with
occasions for field trips abroad in the winter and early spring when the sedges were ‘lying low’. In
part, the stimulus to take up the study of Crocus was provided by his association with the University
Botanic Garden in Cambridge, an institution for which he developed a deep affection, especially
after he and his wife Nora moved to a house nearby. A frequent visitor to the Botanic Garden,
especially to the rock garden and alpine house in early spring, Dick served as a Manager of the Cory
Fund, which administers the important Reginald Cory bequest to the Garden, from 1979 to 1982.
During my time as Director (1973-83) Dick began to use the facilities of the Experimental and
Research Area of the Garden to grow wild-origin stocks of his critical sedges under comparable
conditions, and it is pleasant to record that we still have plants derived from these stocks in
cultivation in the Garden on the Systematic Beds.

No picture of Dick would be remotely adequate without some reference to his happy marriage
and family life. In 1935 he married Nora Blakesley, a graduate of Newnham College who shared
Dick’s interests in the countryside, travel and the theatre in particular, but whose own career took
her into local and national Labour Party politics, culminating in a life peerage in 1978. It was a
constant source of admiration to their many friends that Dick and Nora happily pursued their
separate careers from a firm and affectionate family basis with shared pleasure in the careers of their
four children, Dick coping with amused tolerance with the unusual social problem of being plain ‘Mr
David’ to Nora’s ‘Lady David’.

Dick was active to the end. He died in Corsica on a field excursion to see the lovely Crocus. We all
feel a great loss, not least because the modern world seems to produce fewer selfless ‘gentlemen’ of
Dick’s calibre, but we realise that increasing incapacity in old age would have been difficult for Dick
to tolerate, and the end was appropriate. Our sympathies go out to Nora and the family.

I am personally indebted to Nora and others concerned for permission to use and to quote from
draft texts of some contributions to a volume of essays, Dick David remembered by his friends,
covering the many facets of Dick’s life. This volume, edited by Dick’s son-in-law Tony Davies
(1994), contains inter alia a bibliography of Dick’s published works, including the botanical ones
(available from L. A. Davies, 50 Clarence Road, Birmingham, B13 9UH).

S. M. WALTERS

322 OBITUARIES

KATHLEEN MARGARET HOLLICK
(19131993)

Kathleen was a keen plantswoman and an able artist. Her garden at the Old House at Ashbourne, a
shadow of its former self, is sadly neglected since she tragically lost all her mental powers — she was
in a nursing home for some years before death released her on 6 December 1993. But her drawings
will not die.

She was born on 26 October 1913, the only daughter of a much respected G.P. of the town, where
she lived all her life. She was educated by a governess, Miss Orme, who kindled her life-long
absorption in plants.

She was B.S.B.I. Recorder for Derbyshire (v.c. 57) for 36 years up to 1985. Professor A. R.
Clapham wrote in his Flora of Derbyshire, which came out in 1961, that he was ‘“‘very deeply
indebted” to her. She was the mainspring of the two Supplements to the Flora in 1974 and 1979,
which she produced with Susan Patrick of the Derby Museum. She knew her county very well, its
landscapes, people, history and all, her travelling done by bicycle, bus or train. She had close
connections with the lovely parish church next door to her really old house, and she wrote its
history. She helped her father plant the daffodils in the local churchyard, and it is due to her
insistence that the grass should not be mown until the flowering was over and the foliage had died
down, that they are still the magnificent show they are today.

We are told that when the steeple was being repaired, she went up the steeple-jack’s ladder to the |

top — she was then over 60. This was typical of her ‘go’ and dauntless spirit, in all weathers too.

Her garden was open to the public on occasions for charity, with good plants which she would talk
interestingly about. Several of them had been sent her for drawing, to ensure she had fresh material.

She made scores and scores of drawings of wild flowers. The first were for Dr R. W. Butcher’s
two-volume New Illustrated British Flora of 1961, no fewer than 359 of them. She also did many for
the sadly never-finished British Alien Flora, originally conceived as Volume 3 of Butcher’s work.

Her interests included local history (her notes on early tiles are in the Derbyshire County Records
Office), birds, regular duck counts, and being a member of the Conservation Committee of the
Derby Wildlife Trust; and she was no sleeping Fellow of the Linnean Society of London.

Our happy memories of her go back over 50 (D.McC.) and over 20 (A.B.) years, as always a
welcoming, lively and inspiriting companion and friend. Her herbarium and drawings are in the
Museum at Derby. Her private collection of paintings has been kept within the family.

We are grateful to her niece Mrs Joanna Neal for help with details of her life.

D. McCuintock & A. BURNS

KENNETH GUY MESSENGER
(1920-1993)

All who knew him were shocked and saddened at the news of the sudden death of Guy Messenger in
November 1993, especially as some were expecting to meet him at the B.S.B.I. Annual Exhibition
Meeting to be held on the following day, and at which Guy had been intending to mount an exhibit.

Kenneth Guy Messenger was born on 26 February 1920 at Hampstead, London. His father was a
dentist, and Lecturer and Consultant at Guy’s Hospital; his mother had been a nurse at Guy’s. He
had one brother and one sister. His childhood was spent at Hendon. He was educated at Grove Park
Preparatory School and Felsted School, Essex, at the latter as a scholar. In 1937 he began the course
of Medical Studies at Guy’s Hospital, but he abandoned this in 1939 when he went up to Emmanuel
College, Cambridge to read for the Natural Sciences Tripos in Chemistry, Zoology and Botany.
This course was interrupted by military service, most of which he spent in India and Ceylon in the
Royal Corps of Signals. After the Second World War he returned to Cambridge, and graduated with
Second Class Honours in 1949.

At first he thought of specializing in Marine Biology, but decided on a career as a teacher. He was
appointed Senior Biology Master at Uppingham School in September 1949, and taught there for 31
years, retiring in 1980. He had read Zoology for the second part of his Tripos, and claimed to have

OBITUARIES 323

been somewhat shaky on the Botany at the beginning of his teaching career. However, he
acknowledged his debt to Humphrey Gilbert Carter at Cambridge and the stimulus provided by
acquaintance with other botanists such as Max Walters and Paul Richards. In 1957, at the instigation
of John H. Chandler of Stamford and Franklyn Perring, he joined the B.S.B.I. and began recording
for the Atlas Distribution Maps Scheme. In the Preface to his Flora of Rutland Guy expresses
amazement at his own temerity in undertaking a 10-km square for this survey in the state he
considered his own botanical knowledge to be, but he had the cooperation of a very competent
botanist in John Chandler. His efforts for the Distribution Maps Scheme provided some of the
groundwork for a complete survey of the county of Rutland on a tetrad basis, with the assistance of
John Chandler and an enthusiastic team of Uppingham schoolboys. The outcome of this survey was
Guy’s Flora of Rutland, published in 1972.

I first became acquainted with Guy Messenger when he joined the Leicestershire Flora
Committee at its inauguration in 1967. His experience on the Rutland Flora was invaluable, and he
was also a very useful committee member with a shrewd capacity for cutting Gordian knots and
breaking vicious circles. He and I made several ‘square-bashing’ excursions together, sometimes in
my area and at others in the large area of Leicestershire adjacent to Rutland for which he had
assumed the responsibility. My friendship with him really matured after 1974 when he undertook
the specialist work on the critical groups Ulmus, Rubus, Hieracium and Taraxacum, and I took on
the work for Rosa. Together we covered the greater part of Leicestershire, Guy recording elms and
I roses. Our conversations during these forays would have amused an outside observer, consisting as
they did of monologues on Rosa and Ulmus, delivered almost simultaneously with little mutual
comprehension. Later I spent some days with him at Uppingham, helping him to make a complete
survey of the Rutland roses for the supplement to the Flora of Rutland which, alas, he did not live to
complete.

Guy’s interest in the genus Ulmus began with his Rutland survey, and was enhanced by the survey
of Leicestershire, during which he was working against time, because the ravages of Dutch Elm
Disease were rapidly destroying nearly all the adult elm trees in the county. He had two long
sessions with R. H. Richens at Cambridge on U/mus, and the study of the genus became one of the
two absorbing interests during his retirement. His object was to devise a practicable scheme which
would reconcile the two violently opposed current taxonomic views of the treatment of the genus.
He accumulated a vast quantity of data, but had not arrived at a workable solution to the problem.
He was not, I think, a very systematic worker, and was also a perfectionist who would not dream of
publishing anything until he was absolutely sure of his facts. He was probably, too, diffident of
setting himself up as an authority against the eminent people who have studied the genus. It is to be
hoped that the notes he has left can be effectively used by others in the future, and that all his work
will not have been wasted.

Guy never married, and after the death of his mother lived a solitary life with only his Siamese cat
for company. Nevertheless he was a sociable person and was good company, with a wry and
sometimes slightly bawdy sense of humour. He derived great pleasure from classical music, and
played the double bass in the Uppingham School orchestra. His second main absorbing interest
besides Ulmus, and of longer standing, was in cartobibliography. He amassed a collection of more
than 5000 Ordnance Survey maps, and published a monograph on the 1-inch maps. He was also a
leader in the Brathay Exploration Group, and was one of the authors of the Flora of Foula. He
contributed several articles to the B.S.B.I.’s journals, and his interest in maps and mapping no
doubt inspired him to contribute the topographical and botanical gazeteers for his own Flora of
Rutland and the Flora of Leicestershire. He was elected a fellow of the Linnean Society of London in
1974.

Allin all, Guy was a warm-hearted man with a keen mind, wide interests, and sound views on all
subjects. He will no doubt be remembered with affection by many of his former Uppingham Schooi
pupils. He certainly made a name for himself in botanical circles, and will be sadly missed by his -
many friends.

A. L. PRIMAVESI

324 OBITUARIES

SIR GEORGE TAYLOR
(1904—1993)

Sir George Taylor, Keeper of Botany at the British Museum (Natural History) 1950-1956, Director
of the Royal Botanic Gardens, Kew, 1956-1971, and a Life Member of the Botanical Society of the
British Isles, which he joined in 1933, died at Dunbar, Lothian, on 12 November 1993, in his
ninetieth year. He was born in Edinburgh on 15 February 1904, the only son (though he had three
sisters) of William Taylor and Jane Sloan, and was educated at George Heriot’s School, where, it is
said, encouraged by a housemaster, George Scott, he developed a keen interest in field botany,
cycling extensively around the countryside in search of rare plants. Although remembered chiefly as
a botanical and horticultural administrator, he had a sound knowledge of British plants, with a
particular fondness for, and interest in, the Scottish flora. Indeed his attachment to Edinburgh,
Scotland and things Scottish, persisted throughout a lifetime spent largely in the south of England.
He was proud of his ‘Scottishness’, and to the end of his life retained, almost unmodified, his
Scottish accent.

On leaving school he went to Edinburgh University, where he had a distinguished academic
career, graduating in 1926 with first class honours in Botany, and being awarded the Vans Dunlop
Scholarship. After a collecting trip to South Africa, in company with Reginald Cory, he joined the
staff of the Botany Department of the British Museum (Natural History) in 1928, where — apart
from wartime secondment to the Air Ministry — he remained until 1956, becoming Deputy to the
easy-going John Ramsbottom in 1946, and succeeding him as Keeper of Botany in 1950. Those of us
who first became acquainted with the B.M. Botany Department in the immediate post-war years,
may remember the occasion as a somewhat intimidating one, with the choleric A. J. Wilmott ruling
like one of the less predictable Roman Emperors over the European section of the herbarium, while
those who ventured into exotic botany risked the sharp-tongued derision of the disputatious J. E.
Dandy. It was not a place for the weak-nerved; in such company, George Taylor, formidable though
he could be, seemed the very model of normality.

George Taylor had developed a special interest in the botany of the Himalayan region during his
Edinburgh days, and this enthusiasm remained with him throughout his life. In 1934 he published an
excellent monograph of the genus Meconopsis, for which he was awarded a doctorate, and in 1938,
just before the outbreak of the Second World War, he accompanied Frank Ludlow and George
Sherriff on what was to prove for him a memorable collecting trip to Bhutan and S. E. Tibet.

Early in his career at the B.M. he had begun to take an interest in Potamogeton (pondweeds), and
working with J. E. Dandy, published a series of critical revisions of the species and hybrids in
Journal of Botany and Watsonia. Along with Dandy he was for many years a B.S.B.I. referee for
this difficult genus. For most of his career he was a leading authority on the family Podostemaceae, a
remarkable group, many of the species superficially resembling bryophytes rather than flowering
plants.

In 1956, on the retirement of Sir Edward Salisbury, George Taylor was appointed Director of the
Royal Botanic Gardens, Kew, a post for which he was eminently well qualified. He already knew
most of the Kew botanical staff; he had served as Botanical Secretary of the Linnean Society from
1951 to 1956, and as a member of the Royal Horticultural Society’s Council from 1951. Furthermore
he was a keen practical gardener, very proud of the rarities he had successfully established in his
garden at Rickmansworth.

Kew in 1956 was still recovering from the effects of the war years and the austerities of the early
post-war period; considerable areas of the Gardens had changed little since the beginning of the
century, and there were uninviting expanses of dull, commonplace evergreens, by this time aged
and often overgrown. These the new Director replaced with rhododendrons, camellias and other
flowering trees and shrubs. A Heath Garden provided colour at a season when there was little else to
catch the eye. Waste ground at the back of Kew Palace (which the Duke of Edinburgh had
accurately described as ‘‘a bit of a dump’’) was transformed into the popular Queen’s Garden. A
new Jodrell Laboratory was built, and a new wing added to the Herbarium; Kew took over control
of the fine gardens at Wakehurst Place, Ardingly, Sussex. Looking back, one wonders how so much
was accomplished in a relatively short time. But George Taylor was a vigorous and determined man,
always ready to push ahead where a project met with his approval, and no less ready to reject any
proposal which he reckoned unsound. Naturally such a decisive personality made enemies, but all

OBITUARIES 325

except the most irreconcilable of these would have had to admit that he was a most effective
administrator. Certainly, under his Directorship, botanical research at Kew rapidly extended
beyond the traditional taxonomy and anatomy into cytology, physiology and phytochemistry, and
he would, I am sure, be very gratified to note the many recent advances and developments in these
and other disciplines. In one respect he might not see eye-to-eye with the current programme: he
once told me emphatically that he did not want Kew to be actively involved in educating the general
public. He wanted it to be a fine garden and research centre, where the intellectually curious could
add to their knowledge, and the others simply enjoy themselves.

He received many honours, amongst them the Victoria Medal of the R.H.S. (1956) and a
knighthood (1962), and was elected a Fellow of the Royal Society in 1968. He is commemorated in
the genus Tayloriophyton M. P. Nayar, and in no fewer than 14 species, including one British plant,
the very rare willow hybrid, Salix x taylorii Rechinger f.

On retirement from Kew in 1971, Sir George became the first director of the Stanley Smith
Horticultural Trust, with its headquarters at Belhaven House near Dunbar. He was back in
Scotland, not far from Edinburgh, and settled in a fine house in a good garden, where he was to
enjoy many happy years, still in touch with many of his friends and former colleagues, and still able,
through the Trust, to play an active role in assisting a wide range of horticultural and botanical
projects.

Until his final illness he remained the same Sir George, restless, forthright and pungently
humorous — an original character and, to me, an engaging one.

R. D. MEIKLE

SIR GEORGE TAYLOR’S STUDIES OF THE GENUS POTAMOGETON
Although George Taylor was best known as a botanical and horticultural administrator, it is for his
studies of the genus Potamogeton that he will be remembered by field botanists in Britain and
Ireland. This work was carried out with J. E. Dandy, in a remarkable and perhaps unique
partnership which lasted until Taylor’s promotion to administrative posts meant that the work had
to be continued by Dandy alone.

When asked towards the end of his life when he first became interested in aquatic plants, Taylor
replied that he had ‘paddled in water’ for as long as he could remember. The first specimens of
Potamogeton collected by Taylor which are recorded in Dandy’s card index were gathered in
Scotland in 1926, and the fact that he found at least eleven taxa that year (including the rare hybrid
P. X olivaceus) suggests that he already had some knowledge of the genus, or immediately
discovered that he had an affinity with it. If Taylor collected much material in the next few years it
has not survived, but he collected extensively in both England and Scotland in 1932. These
collections included the very rare P. X griffithii from Loch na Creige Duibhe, Argyll; this
population had been described as P. macvicarii by Arthur Bennett in 1907 but Taylor was the first to
attempt to refind the plant. The specimen was submitted to the acknowledged expert in the genus,
W. H. Pearsall, who misidentified it as P. alpinus; Dandy and Taylor themselves were later to sort
out the taxonomy of this hybrid.

J. E. Dandy became interested in Potamogeton when he worked up some specimens collected by
Taylor on an expedition to Africa in 1934-35. These included some species which also occurred in
Britain, and while working on them Dandy and Taylor resolved to write a monograph of the genus
in Britain and Ireland. Eighteen preliminary papers were published in the series “‘Studies of British
Potamogetons” between 1938 and 1942. These established the correct application of the name
Potamogeton pusillus and clarified the taxonomy and British distribution of numerous species and
hybrids. A 19th paper in the series, on P. X sudermanicus, was at an advanced stage of preparation
when the Journal of Botany ceased publication in 1942, a calamity which Taylor attributed to the
laziness of the editor, J. Ramsbottom. After he moved to Kew, in 1956, Taylor was unable to
continue his work on Potamogeton. The projected monograph had by then been drafted but it was
never submitted for publication: the demise of the Journal of Botany and Taylor’s move to Kew may
have been partly responsible, but the main reason was probably the reluctance of Dandy, a
notorious perfectionist, to commit himself to print. Taylor did not undertake any serious work on
the genus after his retirement, but he collected one or two specimens: it gave him particular pleasure

326 OBITUARIES

to discover P. polygonifolius in East Lothian in 1982, as he had searched for the species in the county
for many years.

Taylor’s contribution to the partnership with Dandy included extensive fieldwork (see below).
During his collecting trips Taylor spent many hours in the evenings floating out specimens, usually in
a bath in the hotel in which he stayed. His specimens are lodged in BM, with duplicates in numerous
other collections. They are of outstanding quality, usually displaying entire plants from root to apex,
but are not accompanied by ecological notes. During his fieldwork Taylor made the first (and, so far,
the only) British collections of P. x lanceolatifolius, and also discovered the rare P. X cognatus in
Scotland. Some of Taylor’s wartime Yorkshire collections which appeared in the field to be P.
pectinatus turned out on detailed examination to be P. X suecicus (P. filiformis X pectinatus), and
thus established the presence of this hybrid south of the current limit of the rarer parent, P.
filiformis. In addition to these discoveries, Taylor made a point of visiting sites where other rare
hybrids had been discovered, and collecting representative material for the herbarium. He visited
the Outer Hebrides in 1951 and Colonsay in 1953, for example, to follow up records made by J. W.
Heslop-Harrison and his team. In addition to the fieldwork, Taylor was responsible for dissecting
the stipules of many of the linear-leaved specimens determined by Dandy and Taylor, including the
type specimen of P. pusillus in the Linnaean herbarium. The publications were, in Taylor’s words,
‘polished’ by Dandy and therefore bear the imprint of Dandy’s personality.

In his old age Taylor was justly proud of his work on Potamogeton. He was delighted by a
chronological printout of his collections which I gave to him, and asked repeatedly whether it
qualified him for an appearance in the Guinness Book of Records! His only regret was that the
monograph which he and Dandy had planned had never materialised.

Dandy once remarked to W. T. Stearn that ‘““George does the collecting and I do the thinking”, a
caustic comment which doubtless contains an element of truth. However, it is perhaps fairer to
conclude with Taylor’s view that theirs was a true partnership in which it was difficult and perhaps
undesirable to assess their individual contributions. Certainly, Taylor’s own hard work and his
determination to publish must have been partly responsible for the stream of papers on the genus
Potamogeton which appeared in the period when they were working together, a series of
publications which must constitute one of the most critical and scholarly contributions made this
century to the taxonomic study of the British flora.

The years in which Taylor is known to have collected Groenlandia and Potamogeton specimens
are listed below, with the vice-counties in which he gathered them. I hope to publish a full
bibliography of the work of Dandy and Taylor on the British and Irish Potamogeton taxa in a
forthcoming B.S.B.I. Handbook.

1926283988, Ole

1928: 88.

1932 Shvi7 2 OM 2iD4 O22283h- S85 Vie

1933: 97S ASOnSileS3 4.879095, S706:
1934526251655 6696094 wid: 19s SI Son O27:
LOZ 5478280) SSiy SSMS GMCS ooo Ss:
1936575894 1058

193724205 230 S95 55—97/2209% TON S80h 88-86,

88-90, 97, 98, 105.
1938322:

1939247 Sal 7920-225 7A S05; Sil. 6S5 1045168"

72-74, 76, 78, 81-86, 92, 93, 95, 96.
1940: 720-62 1le 58a 9> (G65?
1941: 32, 53-57, 61-65, 67, 78, 80, 82-84.
1942: 32, 53, 54, 56, 62-64, 68, 77, 81, 83, 84.
1943: 23, 59, 64, 77, 85, 86, 90.
1944: 54, 56, 61-67, 86, 90.
1945: 30, 56, 59; 61-67,,77, 78, 83, 85, 86.

1946: 7-9, 11, 16, 20-22, 24, 30, 31, 56, 62,64,
68577, 78;,80=83 857-88) 91925 Ose 9Gn Ost

1947: 21, 23,025; 27528) 32)383/945) Sona eaGUs
62=655:08; 7307/5 pT 16282, 8690 O

1948: 14, 16, 20-23, 32-34, 36, 38, 43, 80, 86,
87, 90, 108.

1949: 6, 17, 31, 32, 34, 38, 47, 50, 54, 64, 65, 67,
OSH Swit:

Se:

1952: Zon SEB SH itso:

1953218, 19,31 485:555 565 72=74A ORSON ae
805 JOR 925) 96502

1954: 73, 74.

19552 Bie Ole

1956: 84.

IO FORSO:

1OSZeO2:

1984: 78.

C. D. PRESTON

Watsonia 20: 327-328 (1995) 327

Report

ANNUAL GENERAL MEETING, 14 MAY 1994

The Annual General Meeting of the Society was held at the University of Oxford, in the
Department of Plant Sciences by kind permission of Professor C. J. Leaver, Sibthorpian Professor
of Botany, at 11.45 a.m. Dr F. Perring, President, took the Chair and 161 members were present.

Apologies for absence were read and Minutes of the 1993 Annual General Meeting, published in
Watsonia 20: 85-87 (1994), were approved.

REPORT OF COUNCIL

The Chairman took members through the Report of Council, commenting first on the number of
members who had died in 1993, not only the quantity but the quality of those members who would
be hard to replace. At the request of the Chairman all stood in silence for a minute to their memory.
The Report had been circulated to members, and with the 1993 A.G.M. attendance figure corrected
to 80, the adoption of the Report was proposed by Miss E. Young, seconded by Mrs M. Lindop and
accepted unanimously.

HON. TREASURER’S REPORT AND ACCOUNTS

Mr M. Walpole, Hon. Treasurer, taking members through the Accounts, invited queries. There
being none, the adoption of his Report and the Accounts was proposed by Mr D. A. Pearman,
seconded by Mr R. G. Ellis and accepted unanimously.

ELECTION OF PRESIDENT-ELECT

The President stated that it was a pleasant duty to propose Council’s nomination of Mr. D. A.
Pearman, who had worked with dedication on the Society’s projects during the past years. The
nomination was endorsed by the meeting, and Mr Pearman was elected with applause.

RE-ELECTION OF HON. GENERAL SECRETARY AND HON. TREASURER

Proposing the re-election of these Officers from the Chair, the President thanked Mrs M. Briggs for
ensuring that the administration of the Society synchronised as clockwork, and Mr M. Walpole for
competent management of our finances which had ensured our peace of mind. Dr Perring
commented that he could not have managed this year without these officers, and they were re-
elected with applause and a presentation.

ELECTION OF COUNCIL MEMBERS

In accordance with Rule 11 nominations had been received for Mrs M. Lindop, Professor C. A.
Stace and Mr R. M. Walls. Profiles had been published, and these members were elected
unanimously by the meeting.

328 REPORT
ELECTION OF HONORARY MEMBERS

Council had nominated three members: proposing Dr D. E. Allen, Professor W. T. Stearn highly
commended him for his scholarly publications, including The Botanists — A history of the Botanical
Society of the British Isles through 150 years.

Proposing Dr R. J. Gornall, Dr J. R. Akeroyd, in writing, referred to him as a past Honorary
Receiving Editor of Watsonia, who had enhanced the quality and prestige of the Society’s journal,
and now, as a teacher and researcher at the University of Leicester, was furthering our traditional
links with the wider taxonomic community.

Proposing Mr D. McClintock, Mr R. S. R. Fitter described him as a major player on the botanical
scene since the 1950s when the Pocket guide to wild flowers, McClintock & Fitter, 1956, was the first
in the series of modern field guides to wild flowers in Britain.

The elections of Dr D. E. Allen, Dr R. J. Gornall and Mr D. McClintock were unanimous, and
with applause.

RE-ELECTION OF HONORARY AUDITORS

The Hon. Treasurer, proposing the re-election of Grant Thornton, West Walk, Leicester, reported —

that they were willing to continue as Hon. Auditors. Their re-election was seconded by Mrs J.
Robertson and passed unanimously with appreciation and applause.

ANY OTHER BUSINESS

The President thanked all the Secretaries and Editors who had contributed to the projects and
activities of the Society during the year, particularly mentioning Mrs E. Wiltshire, Hon. Field
Secretary and all the Regional Secretaries who had together produced an excellent programme of
field meetings in all parts of the British Isles. Also he thanked Mr C. D. Preston, retiring member of
Council not eligible for immediate re-election, for his considerable and constructive help with
recording projects and liaison for these.

Finally, on behalf of all present, the President thanked Dr Q. C. B. Cronk, local organiser for this
Annual General Meeting in Oxford and his team of helpers, for the excellent arrangements and
interesting programme provided for the weekend.

There being no other business, the meeting closed at 12.22 p.m.

Mary BrIGGSs

INSTRUCTIONS TO CONTRIBUTORS

Scope. Authors are invited to submit Papers and Notes concerning British and Irish vascular plants,
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Submission of manuscripts

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SM cmt ecu — . u
Se SB ote at I I i are tie SP ;
v = SS LL ST a a TT A eat a Ae

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 1995-96

President, Mr D. A. Pearman

Vice-Presidents, Mr A. O. Chater, Mr. R. G. Ellis, Mr C. D. Preston
Honorary General Secretary, Mrs M. Briggs

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Editors of Watsonia

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© 1995 Botanical Society of the British Isles

Watsonia 20: 329-343 (1995) 329

DNA fingerprinting and biometry can solve some taxonomic
problems in apomictic blackberries (Rubus subgen. Rubus)

T. KRAFT

Department of Systematic Botany, University of Lund, Ostra Vallgatan 18, S-223 61 Lund,
Sweden

and
H. NYBOM*

Balsgard-Department of Horticultural Plant Breeding, Swedish University of Agricultural
Sciences, Fjalkestadsvagen 123-1, S-291 94 Kristianstad, Sweden

ABSTRACT

The feasibility of using DNA fingerprinting and biometry in apomictic blackberries (Rubus subgen. Rubus) was
investigated on five sets of material, chosen to represent various taxonomic problems: 1. Four similar but
according to biometrical results distinct species were compared: the Swedish R. pseudopallidus and R.
hartmanii, and the German R. fuscus and R. pallidus. Surprisingly, R. hartmanii and R. fuscus exhibited
identical DNA fingerprints, whereas the other taxa in this group could be clearly distinguished; 2. R. insularis
and R. septentrionalis, both from Sweden, were rather similar but still well separated by each of the two
methods; 3. Samples of R. infestus from Sweden and Germany were almost identical with DNA fingerprinting
and did not differ appreciably with biometry; 4. Samples of R. polyanthemus from Sweden and Germany were
completely identical with DNA fingerprinting and deviated only slightly with biometry; and 5. R. scheutzii from
Sweden and R. muenteri from Germany could not be separated with either method.

KeEyworps: Taxonomy, microspecies, apomixis, DNA analysis, M13 probe, (AC)/(TG) probe, Rosaceae.

INTRODUCTION

Taxonomists have for a long time struggled with the notorious blackberries, Rubus subgen. Rubus
(Rosaceae). These perennial, bushy plants are characterized by vigorous vegetative reproduction,
either through root suckering or through tip rooting. Thus a genetically homogeneous clone,
consisting of several ramets, may spread over a substantial area. The taxonomic problems are,
however, caused primarily by the system for seed reproduction. Most species in this group are
polyploid and pseudogamous, 1.e. the egg cell is not fertilized but pollination is still necessary to
trigger embryo development. The few existing diploid species instead appear to be sexual
(Gustafsson 1930). Crossing experiments between polyploid species have demonstrated that
apomixis is not complete since at least a minor proportion of the resulting offspring carry some
paternal traits and thus emanate from sexual seed set (Lidforss 1905; Nybom 1988). Moreover,
many of these interspecific hybrids in their turn appear to have reverted to sexuality (Lidforss 1905;
Nybom 1995).

In Europe, Sudre (1908-1913) recognized over 100 blackberry species, many of which were split
into units of lower rank. The fact that his index contains more than 3000 names is, however, also
caused by a vast number of synonyms. At the beginning of this century, a considerable number of
botanists — often non-professionals — were occupied with Rubus taxonomy but activity eventually.
dwindled. A new boost was delivered by the work of Weber (1972) on the Rubus-flora in northern
Germany and the Nordic countries, and a comprehensive treatise of the British brambles published

GWNitt SUN IAy Se

[ sep 98 1995 )
—UeanieS

* Corresponding author.

330 T. KRAFT AND H. NYBOM

by Edees & Newton (1988). In a later study, Weber (1985) arranged the Rubus taxa into four
different taxonomic categories: 1. widespread species with a distribution of more than 500 km in
diameter; 2. regional species with a distribution of 20-250 km in diameter; 3. local species with a
distribution of less than 20 km in diameter; and 4. individual species consisting of a few vegetatively
derived plants. Only the first two categories were thought to merit taxonomic interest.

Phenotypic variation in Rubus is to a large extent associated with environmental factors, thus
rendering identification of isolated populations very difficult. At the end of the last century, several
Swedish Rubus species with a restricted distribution were described as new species. Later, many of
these were lumped with previously described species, often of British or German origin. In some
cases, this led to a rather intense debate among Rubus taxonomists.

Rubus taxonomy has always emphasized morphological characters, and the different species have
been defined as morphologically uniform groups of plants. Often a major part of the taxonomic
work has been based on subjective evaluation of dried herbarium specimens. In other apomictic
genera, morphological data have sometimes been subjected to more sophisticated biometrical
treatments, e.g. in Crataegus (Dickinson & Phipps 1985; Dickinson 1986). Also allozyme
electrophoresis has yielded valuable results in apomicts, e.g. in Antennaria (Bayer 1989) and
Taraxacum (Van Oostrum et al. 1985). Recently, the very sensitive method of DNA fingerprinting
has provided an efficient means to identify genotypes and to separate clones in, for example,
Taraxacum (Van Heusden ef al. 1991) and Rubus (Nybom & Schaal 1990; Antonius & Nybom
1994). To what extent DNA fingerprinting could be considered a generally useful tool for taxonomy
in apomicts is, however, not yet investigated. In this study we therefore attempt to solve some
taxonomic problems in Rubus subgen. Rubus using two different approaches: DNA fingerprinting
and biometry. |

1. R. pseudopallidus (C. E. Gustafsson) A. Gustafsson is a species endemic to a restricted area on
the Swedish east coast. When first described, suggestions were made that this could be a subspecies
of R. hartmanii Gandoger (Lund 1877), which also occurs on the Swedish east coast and, very
rarely, in Great Britain (Edees & Newton 1988). Areschoug (1886), however, treated it as identical
to R. pallidus Weihe, whereas several other botanists regarded it as R. fuscus Weihe & Nees
(Hylander 1941; Oredsson 1970; Weber 1972). Both R. pallidus and R. fuscus are widespread in
N.W. Europe but do not otherwise occur in Sweden (Weber 1972). Thus, does R. pseudopallidus
merit species rank, or should it be synonymized with any one of R. fuscus, R. pallidus or R.
hartmanit?

2. R. septentrionalis Watson occurs in a very restricted area onthe Swedish west coast, in southern
Norway, in northern Denmark and in Great Britain (Weber 1984; Edees & Newton 1988).
Sometimes this taxon has been considered a subspecies of R. insularis F. Aresch. (Areschoug 1886;
Weber 1972). This latter species occurs in the southernmost part of Sweden, in Denmark and in the
eastern parts of Germany (Pederson & Schou 1989). Therefore, should R. septentrionalis and R.
insularis be treated as separate species?

3. The endemic species R. taeniarum Lindeberg has been described from a restricted area on the
Swedish west coast (Lindeberg 1858). However, it was soon afterwards synonymized with R.
infestus Weihe & Nees (Focke 1877), a species which occurs in Great Britain, Germany and
Denmark (Weber 1972; Edees & Newton 1988). Should R. taeniarum and R. infestus be treated as
separate species?

4. R. polyanthemus Lindeberg was first described from Sweden where it occurs in a very restricted
area on the southwestern coast (Lindeberg 1883). Since then, it has also been reported from Ireland,
Great Britain, Denmark and Germany (Weber 1985; Pedersen & Schou 1989). Can the Swedish and
German populations of R. polyanthemus be treated as conspecific?

5. It has been suggested that R. scheutzii Lindeberg, endemic to the Swedish east coast, is
identical to R. muenteri Marsson, a species that only occurs in Germany (Areschoug 1886; Weber
1985). Are R. scheutzii and R. muenteri separate species?

MATERIALS AND METHODS

Nomenclature is according to Weber (1972), except in two cases: the Swedish Rubus population
which Weber regarded as R. fuscus in 1972, is here called R. pseudopallidus (as suggested by Weber,

DNA FINGERPRINTING, BIOMETRY AND RUBUS Soil

pers. comm., 1992); and R. insularis subsp. confinis in Weber (1972) is here called R. septentrionalis
in accordance with Weber (1984).

DNA FINGERPRINTING

Recently hypervariable regions of genomic DNA have been detected, which can be used to yield
genotype-specific ‘DNA fingerprints’ in various organisms (Jeffreys et al. 1985; Nybom 1991, 1993).
This so called minisatellite-DNA consists of a number of tandem repeats of a core sequence with c.
10-60 base pairs. Restriction enzymes will usually cut the DNA at either end of a series of tandem
repeats. The length of the resulting DNA-fragment is therefore proportional to the number of
repeats. Hybridization of sample DNA to a minisatellite DNA probe allows us to analyze a
considerable number of such DNA fragments simultaneously.

Fresh Rubus leaves were collected in Sweden and Germany (Table 1, Fig. 1). Plants were con-
sidered from different localities if collected at least 1 km apart. Voucher specimens are deposited at
LD. Determinations of the German plants have been verified by Prof. Dr Dr H. E. Weber.

DNA was extracted from fresh or frozen leaf material and subsequently digested with the
restriction enzyme HaellII. The resulting DNA fragments were size-separated by electrophoresis in
an agarose gel, denatured and transferred to a nylon filter by Southern blotting. These nylon filters
were hybridized to a **P-labelled minisatellite DNA probe derived from the M13 bacteriophage
(Vassart et al. 1987). Autoradiography was carried out for 2-12 days at —80°C with intensifying
screens. Residual probe was stripped off and filters rehybridized, this time to a synthetic (AC)/(TG)

Bye,

\ | \

AG \
j »Bremen ;
gy p (
- pa « Hannover di 200.4
qo~r? MM f : Ny
Y if (
f |
/ xX

Figure 1. Map of southern Sweden, Denmark and northern Germany showing where material of Rubus spp. was
collected for the DNA fingerprinting. f = R. fuscus, h = R. hartmanii, i = R. insularis, if = R. infestus, m = R.
muenteri, p = R. polyanthemus, pa = R. pallidus, ps = R. pseudopallidus, s = R. septentrionalis, sc = R.
scheutzit. Some letters represent several closely situated localities.

332 T. KRAFT AND H. NYBOM

TABLE 1. ORIGINS OF THE RUBUS MATERIAL USED FOR DNA FINGERPRINTING AND THE
DISTANCE BETWEEN THE TWO MOST DISTANT LOCALITIES

Sweden Germany
Number of Number of Maximum Number of Numberof Maximum

Species localities shrubs distance (km) _ localities shrubs __ distance (km)
R. fuscus 0 0 - 1 1 -

R. hartmanii d. 8 7 0 0 -

R. infestus 4 11 72) 1 2 -

R. insularis 5 6 180 0 0 -

R. muenteri 0 0 - 2 3 7

R. pallidus 0 0 - 1 1

R. polyanthemus 2 2 2 1 1 -

R. pseudopallidus 4 6 5 0 0 -

R. scheutzii 6 10 20 0 0 -

R. septentrionalis a 4 i] 0 0 -

polydinucleotide (Lonn et al. 1992). For a more detailed description of DNA isolation and
hybridization with the M13 probe, see Nybom & Schaal (1990). Labelling and hybridization with the
(AC)/(TG) probe was performed in the same way as with the M13 probe except that the final washes
were in 0.2 SSC instead of 2x. :
DNA fragment patterns were scored manually for presence or absence of individual bands. A
bandsharing value was calculated for each pairwise comparison between two different plants: D = 2
x number of shared fragments/(number of fragments in plant A + number of fragments in plant B).
For a more detailed description of DNA fingerprint evaluation, see Nybom & Rogstad (1990).

BIOMETRY

We have chosen morphological characters for the biometrical investigation according to two
criteria: they should be easy to measure on dried material as well as considered generally useful for
species identification within Rubus. Distribution of prickles on the stem and in the inflorescence is
an important character. Some species have prickles of more or less even size, whereas others have a
characteristic mixture of both large and small prickles. Thus we noted the frequency of prickles
occurring in three different length-classes. Lengths of the petiolules and the shape of the terminal
leaflet of the three-, five-, or seven-foliated leaves are often employed to separate species. Number
of glands and hairs on the stem and in the inflorescence are also commonly used characters. For the
present investigation we ultimately decided on the following measurements: length/width ratio of
the terminal leaflet (A/B, Fig. 2); lengths of leaf petiolules (C,D,E, Fig. 2); distance from petiolule
base of the middle leaflets (leaf centre) to the point of attachment for the lower leaflet petiolule (F,
Fig. 2); length of petiole (G, Fig. 2); number and size distribution of prickles on 5 cm of the stem;
number of hairs on 1 cm of the stem; distance from inflorescence apex down to the base of the
uppermost inflorescence leaf; number of prickles and glands on the pedicel; number of hairs on the
pistils; and number of hairs on the anthers.

Measurements were taken on herbarium specimens in LD (Table 2, Fig. 3). The herbarium sheets
have been marked with ‘TK’ and a number from 1 to 212. For each species—country combination we
selected 20 specimens that were as complete as possible. We-also used plants from our own
collections as well as some herbarium specimens in C of R. muenteri and R. polyanthemus from
Germany. Still, only five specimens of each of these two species—country combinations were
available.

All comparisons were performed between plants of similar stem diameter to correct for variation
caused by differences in plant vigour. F-tests were calculated, for each character separately, to
determine the level of variation between taxa as compared to within. Multivariate F-tests, including
all of the above-mentioned characters, were also performed utilizing the Hotelling-Lawley trace
statistic (Morrison 1976). All calculations were performed with SYSTAT for Windows (1992).
When necessary, logarithmic transformations were used to adjust for inequality of variances.

DNA FINGERPRINTING, BIOMETRY AND RUBUS 333

FicureE 2. Leaf of blackberry (Rubus sp.) and enlarged portion of the centre of the leaf with some of the
characters used for the biometrical analysis: length/width-ratio of the terminal leaflet (A/B); length cf the
petiolule of the terminal leaflet (C), middle leaflet (D) and lower leaflet (E); distance from petiolule base of the
middle leaflets (leaf centre) to the point of attachment for the lower leaflet petiolules (F), and length of the
petiole (G).

TABLE 2. ORIGINS OF THE RUBUS MATERIAL USED FOR BIOMETRY AND THE DISTANCE
BETWEEN THE TWO MOST DISTANT LOCALITIES

Sweden Germany
Number of Number of Maximum Number of Numberof Maximum

Species localities shrubs distance (km) _ localities shrubs distance (km)
R. fuscus 0 0 - 5 17 190

R. hartmanii 10 20 20 0 0 -

R. infestus 13 30 30 5 20 160

R. insularis 12 21 180 0 0 -

R. muenteri 0 0 - + 5 380

R. pallidus Ona, ¢ 0 - 12 15 150

R. polyanthemus 2 20 5 3 5 300

R. pseudopallidus 5 19 10 0 0 -

R. scheutzii 6 20 5 0 0 —

R. septentrionalis 6 20 5 0 0 =

334 T. KRAFT AND H. NYBOM

4

BS: ;

3 (9

}
. . ‘
a8 Na ‘
»Bremen }
r ue C
| DDal Boannoves Ne
eee pa 200 km
elseeWorl ment If | |

¢ £ \
Ficure 3. Map of southern Sweden, Denmark and northern Germany showing where material of Rubus spp. was
collected for the biometrical analysis, f = R. fuscus, h = R. hartmanii, i = R. insularis, if = R. infestus, m = R.
muenteri, p = R. polyanthemus, pa = R. pallidus, ps = R. pseudopallidus, s = R. septentrionalis, sc = R.
scheutzii. Some letters represent several closely situated localities.

\
y pa \
te

RESULTS

R. PSEUDOPALLIDUS
Does R. pseudopallidus merit species rank, or should it be synonymized with any one of R. fuscus,
R. pallidus or R. hartmanii? Quite surprisingly, the German R. fuscus and the Swedish R. hartmanii
exhibited identical DNA fingerprints with both DNA probes (Table 3, Fig. 4). Barring minor
mutations, which do not manifest themselves in DNA fingerprinting, these two species appear to
have essentially the same genotype. By contrast, the DNA fingerprint of R. pallidus differed
considerably from those of R. pseudopallidus and R. fuscus/R. hartmanii. R. pseudopallidus was
more similar to R. fuscus/R. hartmanii than to R. pallidus (Table 3). Nine fragments could be scored
with the M13 probe in R. fuscus/R. hartmanii, eight in R. pseudopallidus and seven in R. pallidus.
With the (AC)/(TG)-probe 13 fragments could be scored in R. fuscus/R. hartmanii, 15 in R.
pseudopallidus and eleven in R. pallidus. Intraspecific variation was not encountered.

A multivariate F-test on the biometrical data showed a significant difference between R.

DNA FINGERPRINTING, BIOMETRY AND RUBUS 335

TABLE 3. AMOUNT OF BANDSHARING (D-VALUES) AS OBTAINED IN DNA FINGERPRINTING.
FOR PAIRWISE COMPARISONS AMONG RUBUS SPP.
Results from hybridization with the (AC)/(TG) probe in the upper right half, and from hybridization with the
M13 probe in the lower left half.

Species R. pseudopallidus R. fuscus R. hartmanii R. pallidus
R. pseudopallidus = 0:79 0-79 0-54
R. fuscus 0-82 - 1-00 0-67
R. hartmanii 0-82 1-00 - 0-67
R. pallidus 0-53 0-62 0-62 ~
A 8 C D Ec G - ; A J
9.4 — sin

6.6 —

4.4 — '

2.3 —|

Figure 4. DNA samples from leaves of Rubus fuscus (A), R. pseudopallidus (B—F), R. hartmanii (G-L) and R.
pallidus (M) digested with HaellI and hybridized with the M13 probe. Size markers (kb) were obtained by
digestion of A DNA with HindIII.

pseudopallidus and R. fuscus (F=3-25, p=0-007), between R. pseudopallidus and R. hartmanii
(F=6-98, p<0-001), and between R. pseudopallidus and R. pallidus (F=9-46, p<0-001). Single-
variate F-tests for the same interspecific comparisons showed significant variation (p<0-05) for five,
six and seven characters, respectively (Table 4). There was a significant difference with the
multivariate F-test also between R. fuscus and R. hartmanii (F=7-:28, p=<0-001), whereas six
characters differed significantly with the single-variate F-tests. R. fuscus was also well separated
from R. pallidus with the multivariate F-test (F=6-93, p<0-001), and yielded significant differences
in six characters with the single-variate F-tests. The last pair of species to compare in this group, R.
hartmanii and R. pallidus, differed considerably both with the multivariate F-test (F=25-77,
p<0-001) and with the single-variate F-tests, yielding seven significantly different characters.

336 T. KRAFT AND H. NYBOM

TABLE 4. BIOMETRICAL CHARACTERS OF RUBUS WHICH SHOWED A _ SIGNIFICANT
VARIATION (P<0-05) WHEN SPECIES WERE COMPARED TO EACH OTHER

Species
Variable fus-pse har-pse_ pal-pse_har-fus___ pal-fus_ _har-pal
Length of the terminal petiolule = = - + 0
Length of the middle petiolule 0 0 0 <P 0 0
Length of the lower petiolule 0 0 0 + - 0
Length of the petiole = 0 - +f 0 +
Length/width ratio of the terminal leaflet 0 - ats 0 ae -
Placement of the lower leaflets on the 0 + 0 + 0 +
petiolules of the middle leaflets
No. of prickles <1-5 mm ar aF + 0 st -
No. of prickles 1-5-3 mm + + =F 0 sp 0
No. of prickles >3 mm - 0 0 EP — +
Distance from inflorescence apex to base of 0 + + 0 0 ~
the uppermost leaf in the inflorescence
No. of prickies on the pedicel 0 0 0 + 0
No. of hairs on pistils 0 - 0 0 0

+ denotes a significantly higher value for the first species, — denotes a significantly lower value for the first
species, and 0 no significant difference. Species: fus, R. fuscus; pse, R. pseudopallidus; har, R. hartmanii; pal, R.
pallidus.

Boe Bee CP Soe SE item oma Pood Ko eo Nie OF iP

Figure 5. DNA samples from leaves of Rubus insularis (A, B, G-J, O, P) and R. septentrionalis (C-F, K-N)
digested with HaelIII and hybridized with the M13 probe. I-P are identical to A-H except for a different
exposure time, A, G-I, O and P represent a genotype, ‘R. insularis IV, which differs somewhat from the
standard genotype, ‘R. insularis 1’. Size markers.(kb) were obtained by digestion of A DNA with HindIII.

DNA FINGERPRINTING, BIOMETRY AND RUBUS B37

R. SEPTENTRIONALIS
Should R. septentrionalis and R. insularis be treated as separate species? Hybridization with the
M13 probe demonstrated the existence of two similar but not identical genotypes of R. insularis in
our material, ‘R. insularis Y and ‘R. insularis IY, differing in two bands (Fig. 5). A comparison of
these two genotypes resulted in a D-value of 0-89. The (AC)/(TG) probe, on the other hand, yielded
no intraspecific variation. ‘R. insularis Y comprises all plants from Scania in southernmost Sweden
and from the Swedish east coast, whereas ‘R. insularis IV comprises material from two close
localities on the Swedish west coast. Intraspecific DNA variation was not encountered in R.
septentrionalis. This species was, however, quite different from R. insularis, with a D-value of 0-56
with the M13 probe for comparisons to both ‘R. insularis V and ‘R. insularis IV , and 0-73 with the
(AC)/)/(TG) probe. All genotypes investigated had nine fragments that could be scored with the M13
probe. With the (AC)/(TG) probe 17 fragments could be scored in R. insularis and 16 in R.
septentrionalis.

The multivariate F-test showed a significant difference between R. insularis and R. septentrionalis
(F=9-30, p<0-001). According to the single-variate F-tests, three characters showed significant
(p<0-05) variation between taxa: R. insularis had fewer prickles in two of the length classes, namely
<4 mm and 4-6 mm and the lower leaflets were attached further up on the petiolules of the middle
leaflets.

9.4 —

6.6 —

4.4 —

0S Vie

FigurE 6. DNA samples from leaves of Rubus infestus from Sweden (A-J) and Germany (K-L) digested with
HaelKI and hybridized with the M13 probe. Size markers (kb) were obtained by digestion of A DNA with
Hindi.

338 TF. KRAFT AND H. NYBOM

R. INFESTUS

Should R. taeniarum and R. infestus be treated as separate species? Both DNA fingerprinting
probes showed the Swedish collections of R. taeniarum to be homogeneous as were also the German
collections of R. infestus (Fig. 6). Moreover, comparison of (AC)/(TG)-hybridized samples from
the two countries yielded identical DNA fingerprints. Somewhat different fragment profiles were,
however, obtained after M13 hybridization resulting in a D-value of 0-90, which indicates that
genotypes differ somewhat between the two countries. Ten fragments could be scored in both sets of
material with the M13 probe and 16 with the (AC)/(TG) probe.

The multivariate F-test showed a barely significant difference between the Swedish and the
German material (F=2-71, p=0-038). Only one character showed significant (p<0-05) variation
with the single variate F-test: the petiolules of the middle leaflets were longer on the German
material.

R. POLYANTHEMUS

Should the Swedish and the German populations of R. polyanthemus be treated as conspecific?
German and Swedish populations of R. polyanthemus had identical DNA-fingerprints with both
probes (Fig. 7). Nine fragments could be scored with the M13 probe and 18 with the (AC)/(TG)
probex 1%:

By contrast, the biometrical data showed a significant difference between the two sets of material
with the multivariate F-test (F=5-29, p=0-005). According to the single-variate F-tests, three
characters showed significant (p<0-05) variation between Swedish and German plants: the
petiolules of the lower and middle leaflets were significantly longer on the German plants and the
lower leaflets were attached further up on the petiolules of the middle leaflets.

FicurE 7. DNA samples from leaves of Rubus polyanthemus from Germany (A, D) and Sweden (B, C) digested
with Haelll and hybridized with the M13 ah Size markers (kb) were obtained by digestion of A DNA with
Hindi.

DNA FINGERPRINTING, BIOMETRY AND RUBUS 339
A BC eS ERIS nena Ae ay Dee ees Ak
9.4 —-

6.6—

44-4

Figure 8. DNA samples from leaves of Rubus scheutzii (A—C, G—I) and R. muenteri (D-F, J—L) digested with
Haelill and hybridized with the M13 probe. G-L are identical to A-F except for a different exposure time. Size
markers (kb) were obtained by digestion of A DNA with HindIII.

R. SCHEUTZII
Should R. scheutzii and R. muenteri be treated as separate species? R. muenteri and R. scheutzii had
identical DNA fingerprints with both probes (Fig. 8). Nine fragments could be scored with the M13
probe and 15 with the (AC)/(TG) probe.

The biometrical data did not show any significant difference between R. muenteri and R.
scheutzii, neither with multivariate (F=1-019, p=0-481) nor with single-variate F-tests.

DISCUSSION

R. PSEUDOPALLIDUS
In 1877 Lund described a new blackberry species from the Swedish east coast, R. mitigatus.
However, he suggested even then that it could perhaps more appropriately be treated as a
subspecies of R. hartmanii, which occurs about 50 km from the new taxon. According to Lund, R.
mitigatus deviates from R. hartmanii by its thinner leaves that are always green underneath,
differently shaped terminal leaflets and inflorescences, and fewer and smaller prickles on the stem.
Areschoug (1886) considered the new taxon to be identical to R. pallidus, Hylander (1941) regarded
it as R. fuscus, and in Gustafsson’s opinion (1935) it is instead most similar to R. menkei Weihe &
Nees. R. pallidus, R. fuscus and R. menkei all occur in Germany and/or Denmark. Since the name
R. mitigatus had already been used for another species, Gustafsson renamed it as R. menkei var.
pseudopallidus. At first Weber (1972) synonymized this taxon with R. fuscus, but later changed his
mind and now regards it as an endemic species, R. pseudopallidus (Weber, pers. comm., 1992).
Surprisingly, R. hartmanii has DNA fingerprints identical to R. fuscus in spite of a significant
biometrical variation between these two species. This discrepancy is probably due to environmen-
tally induced morphological variation, and perhaps also to the occurrence of genetic variation which ©
affects parts of the genome not screened by the DNA fingerprint probes. The inability of DNA
fingerprinting to detect somatic mutations has been reported previously in cultivated apple (Nybom
1990). By contrast, genetic recombination between cross-pollinating genotypes results in easily
detected fingerprint variation (Nybom 1991). Thus R. hartmanii and R. fuscus probably deviate
from each other in somatic mutations only.

340 T. KRAFT AND H. NYBOM

Our study shows that R. pseudopallidus can be distinguished from R. fuscus/R. hartmanii as well
as from R. pallidus by both DNA fingerprint and biometrical data. However, DNA fingerprint
studies in some other Rubus species indicate that intraspecific variation is more common in German
material than in Swedish (Kraft et al., manuscript submitted). Therefore, it appears necessary to
obtain much more data on the variability of R. fuscus in Germany before finally defining this
species. In the meantime, we believe that R. hartmanii and perhaps also R. pseudopallidus are best
treated as synonyms of R. fuscus. R. pallidus, on the contrary, appears to be more dissimilar
according both to morphology and to DNA fingerprint data, and should for the time being retain its
status as a separate species.

R. SEPTENTRIONALIS

In 1883, R. confinis Lindeberg was described from Bro on the Swedish west coast but in 1886 it was
demoted to a subspecies of R. insularis (Areschoug 1886). Watson (1946, 1958) claimed that there
are two different species in the vicinity of Bro; R. septentrionalis, regarded as identical to R.
confinis, with a west European distribution and R. broensis Watson which also occurs sporadically in
England. Weber (1972) at first agreed with Areschoug on R. insularis subsp. confinis and mentioned
both R. septentrionalis and R. broensis as synonyms of this subspecies. Characteristic for subsp.
confinis in comparison to subsp. insularis is, according to Weber, the smaller stature, increased
number of prickles on the stem, more rounded terminal leaflet, narrower inflorescence, white petals

and green pistils. R. broensis is thought to be only a shade form of subsp. confinis. Later Weber

(1984) gave R. septentrionalis species status, and included the plants from Bro in this species, which
also occurs in southern Norway, northern Denmark and Great Britain.

Our study clearly suggests that R. insularis and R. septentrionalis should be treated as two
separate species since they are well distinguished both biometrically and with DNA fingerprinting.
Interestingly, R. insularis is the only taxon to show intraspecific variation within Sweden with DNA
fingerprinting. These differences are, however, quite limited and appeared only after hybridization
with the M13 probe. The genotype ‘R. insularis IV was collected from two localities on the Swedish
west coast, 13 km apart. These localities are isolated by 160-180 km from the remainder of the
Swedish distribution of R. insularis.

R. INFESTUS

Lindeberg (1858) described R. taeniarum as a new species from central Bohuslan on the Swedish
west coast, where it is fairly common in a small area (Oredsson 1974). Both Focke (1877) and
Areschoug (1886) instead treated these populations as identical to R. infestus, although Lindeberg
(1887) never agreed. Thirty years later, Neuman (1915) came to the conclusion that R. taeniarum is
well separated from R. infestus. Gustafsson (1938) also argued that R. taeniarum should be regarded
as a species. Weber (1972), on the other hand, treated R. faeniarum as a synonym of R. infestus.
Apart from a larger stem diameter in German plants, these were almost identical to the Swedish
material in our study, with both biometrical and DNA fingerprint data. It seems likely that the few
recorded differences are due only to environmental influences and/or somatic mutations, and
therefore we conclude that the Swedish populations should be treated as R. infestus.

R. POLYANTHEMUS

R. polyanthemus was described by Lindeberg (1883) from southern Sweden. This species has also
been called R. pulcherrimus Neuman and R. neumanii Focke but these synonyms were due to
differing opinions about nomenclature rather than about the taxonomy. The DNA fingerprints were
identical for the Swedish and German plants investigated, whereas biometrical data showed some
significant differences. We believe that this, still rather restricted, morphological variation can be
explained by differences in growing conditions and/or somatic mutations. In either case, the two
populations must be very closely related and should be treated as a single species.

R. SCHEUTZII

Lindeberg described R. scheutzii as a new species in his exsiccata Herbarium Ruborum Scandinaviae
in 1885. This species is fairly common in a small area on the Swedish east coast (Oredsson 1974).
Areschoug (1886) regarded it as identical to the Central European R. muenteri, but Gustafsson
(1924) claimed that they are well separated with R. muenteri being overall more delicate.

DNA FINGERPRINTING, BIOMETRY AND RUBUS 341

Since we obtained no variation between the two species with biometrical or DNA fingerprint
data, there appears to be no reason to separate R. muenteri and R. scheutzii. This is in agreement
with the most recent taxonomic treatments, where the Swedish populations are included under R.
muenteri (Weber 1985; Pedersen & Schou 1989).

CONCLUSIONS

Blackberry taxonomy is usually based almost exclusively on the subjective evaluation of various
morphological characters. Published descriptions have mentioned how species differ or are identical
for certain characters without any explanation of the basis for these statements. No serious attempts
have been made to investigate the extent to which these characters might be prone to environmen-
tally induced variation. Recently, more objective interpretations of morphological data have
proven useful for the separation of species in critical groups like the apomictic Crataegus (Dickinson
& Phipps 1985). Multivariate statistical methods applied to flowers, fruits and leaves of Crataegus
sect. Crus-galli L. yielded results in close accord with previous classifications. However, comparison
of widespread plants, as in the present study, may be greatly confounded by environmental
influences on the phenotype. Nevertheless, we chose to work exclusively with dried herbarium
specimens in order to determine how valid are these types of data. For future taxonomic work, we
strongly recommend that measurements are taken instead on plants grown in a randomized design
in an experimental garden to circumvent the problem of phenotypic plasticity. Unfortunately this
approach is very time-consuming and expensive.

The recent development of molecular methodology has greatly improved the tools at our
disposal. Estimates of genetic variability can nowadays be obtained that are completely indepen-
dent of where the plants were grown. So far, the most sensitive of these methods in distinguishing
different plant genotypes is DNA fingerprinting by hybridization of DNA samples to minisatellite
DNA probes (reviews in Nybom 1991, 1993). DNA fragment profiles are individual-specific in
cross-pollinated plants, as demonstrated for example in the dioecious North American box elder,
Acer negundo L. (Nybom & Rogstad 1990). Clones may also be identified and delimited as in North
American quaking aspen, Populus tremuloides Michx. (Rogstad et al. 1991). In population studies
of Rubus, varying levels of genetic variation have been encountered, which are closely associated
with the reproductive system in these species (Nybom & Schaal 1990; Antonius & Nybom 1994).

DNA fingerprinting has also yielded results that appear promising for taxonomic applications in
apomictic plant groups. Thus Van Heusden et al. (1991) collected plants of several apomictic species
of Taraxacum from the Netherlands, France and Czechoslovakia. In one species they found the
same DNA fingerprint in material from France and Czechoslovakia and only a slight deviation in
material from the Netherlands. Another DNA fingerprint was found to be identical in material from
three, supposedly different species in the Netherlands. These taxa could be separated on minor
morphological characters, possibly due to mutations within a clonal line or by recombination
between very closely related genotypes.

In our study, intraspecific DNA variation was extremely low. Complete DNA fingerprint
homogeneity in material collected from large parts of the distributional area in Sweden has been
encountered also in other blackberry species like R. nessensis W. Hall, R. grabowskii Weihe ex
Gunther et al. and R. pedemontanus Pinkwart (Kraft et al. 1995; Kraft et al., manuscript submitted).
Apparently, the Swedish populations of each of these species are derived from a single
recombinational event. The resulting genotypes have subsequently spread over large areas,
presumedly by a combination of vegetative reproduction and dispersal of apomictic seed. In some
cases, like R. hartmanii/R. fuscus, morphological variation occurs in spite of identical DNA
fingerprints. Most likely, this variation is caused by environmental influences and/or somatic
mutations. A series of studies on some closely occurring stands of R. nessensis similarly yielded
identical DNA fingerprints in spite of substantial differences in plant demography and flowering
phenology (Kraft et al., manuscript submitted). We believe that variability in morphological
characters caused by somatic mutations is, in most cases, not prominent and consistent enough to
merit species rank for the resulting variants. Plant populations that have identical DNA fingerprints
should, therefore, in general be regarded as conspecific. On the other hand, different DNA

342 T. KRAFT AND H. NYBOM

fingerprints may be encountered within what is commonly accepted as a species. As long as this
species is morphologically well defined, we see no practical reason to split it into several taxa.

Biometrical analysis and DNA fingerprinting yielded consistent results in three of our five case
studies. In the remaining two cases, taxa with identical DNA fingerprints differed in morphological
characters. The additional heterogeneity recorded with biometrical data is probably to a large
extent due to environmental influences. Distinct genetic variation is instead more easily detected by
DNA fingerprinting, as demonstrated by the two deviating plants of R. insularis from the Swedish
west coast. However, DNA fingerprinting is relatively expensive and time-consuming and cannot,
therefore, be used for large series of material. Ideally, a combination of DNA fingerprinting and
biometry should be applied in studies of taxonomic problems in apomictic genera. Such studies
would also increase our understanding of the genetic structure and evolution of these intriguing
plant groups.

ACKNOWLEDGMENTS

Practical help in the field and the herbarium as well as valuable comments on the manuscript were
given by A. Oredsson and H. Weber. Financial support was granted to T. Kraft by Enander’s Fund
and to H. Nybom from the Swedish Natural Science Research Council.

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(Accepted November 1994)

Watsonia 20: 345-349 (1995) 345

Rubus edeesii H. E. Weber & A. L. Bull (Rosaceae), a new
bramble species from Britain and Germany

H. E. WEBER
University of Osnabriick, Dept. Vechta, Driverstrasse 22, D-49364 Vechta, Germany
and
A. L. BULL

Hillcrest, East Tuddenham, Dereham, Norfolk, NR20 3JJ

ABSTRACT

Rubus edeesii H. E. Weber & A. L. Bull is described and illustrated as a new species of subgen. Rubus sect.
Rubus ser. Sylvatici, recorded in Lincolnshire and Westphalia (Germany) and named after the British batologist
EDS: Edees:

Kryworps: Apomictic species, Rubus L., ser. Sylvatici, distribution, bird dispersal.

INTRODUCTION

During the exploration of the vegetation of hedges (Wittig 1975) and the Rubus flora in Westphalia
(Weber 1985) a distinct bramble was found around Minster. Because its distribution area seemed to
be very local, the plant was not treated taxonomically and, like other local or individual
morphotypes, arranged merely under a provisional name and omitted from a book on the
Westphalian bramble flora (Weber 1985). In December 1993, A.L.B. sent to H.E.W. for
examination some sheets with provisionally named brambles gathered in Norfolk and Lincolnshire,
and one of these specimens was easily recognised as the plant collected also in Westphalia. Its
distribution area seems to be very disjunct, like that of other bird-sown bramble species such as
Rubus drejeri Jensen ex Lange and R. tuberculatus Bab.

DESCRIPTION

Rubus edeesii H. E. Weber & A. L. Bull sp. nov. (Figs 1 & 2)

Turio alte arcuatus, viridulus vel parce vinosus, (obtuse) angulatus faciebus planis, pilis plerumque
fasciculatis (vulgo (5—)10—30 ad 1 cm) pilosus. Aculei 4-8 per 5 cm, flavescentes, aequales, e basis 6—
8 mm dilatata paulo reclinati, rarius leviter curvati, 7-8(—8-5) mm longi.

Folia 5-nata (singuli raro 6-7 nata), digitata, supra pilis 15-25 ad cm?, subtus viridia, pilis ad
tactum non vel paulo perceptibilibus pilis stellulatis nullis instructa. Foliolum terminale mediocriter
usque sat longe petiolulatum (longitudo petioluli plerumque 33-41 per centum longitudinis
laminulae), e basi leviter cordata (rarius rotundata) ellipticum vel parce obovatum, apice 15-23 mm
longe acuminatum, dentibus cuspidatis dentibusque principalibus longioribus (non recurvatis) 3—5
mm alte grosse et irregulariter serratum, interdum paulo lobatum. Foliola infima 2-3 mm
petiolulata. Petiolus pilosus, aculeis curvatis 9-13 munitus. Stipulae subfiliformes usque ad 1 mm
latae, margine glandulis (sub)sessilibus instructae.

Inflorescentia ampla, late cylindrica, 3-15 cm infra apicem aphylla, cetera foliis 1—-3-nata,
instructa, ramis usque 70—80° divaricatis plerumque supra mediam earum ramosis. Rachis pilis
minutis fasciculatis pilisque longioribus dense obtecta, glandulis sessilibus multis instructa, aculeis

346 H. E. WEBER AND A. L. BULL

Heinrich F. Weber

van Westisten

Nolotypus!

FiGureE 1. Holotype of Rubus edeesii H. E. Weber & A. L. Bull (MANCH).

sat gracilibus, parce reclinatis, plerumque leviter curvatis, 6-7 mm longis 3-5 per 5 cm armata.
Pedicelli pro maxima parte 10-15 mm long}, pilis usque 0-6—0-8 mm longis hirsuti, glandulis stipitati
0-2 usque 0-3 mm longis instructi, aculeis 3-7, saepe inaequalibus, rectis vel parce reclinatis,

RUBUS EDEESII, A NEW BRAMBLE SPECIES 347

FiGuRE 2. Rubus edeesii at the locus typicus (Germany, Westphalia).

usque (1—2(—3) mm longis muniti. Bracteolae vulgo glandulis breviter stipitatis instructae. Sepala
canoviridia, plerumque inermia, post anthesin reflexa. Petala alba, obovata, unguiculata, apice
parce emarginata vel crenata, 10-15 mm longa. Stamina alba stylos virescentes superantia.
Antherae glabrae. Ovaria glabra. Receptaculum parce pilosum. Fructus bene evoluti. Floret (VI-)
Vil (-VHI).

Rubus e sect. Rubus ser. Sylvatici (P. J. Muller) Focke, honore clarissimi batologii Eric S. Edees
(1907-1993), egregie meritis exploratione generis Ruborum in Britannia, nominatus. Crescit in
Britannia (Lincolnshire) et Germania (Westfalia).

TYPUS: Germany, Westphalia: Margin of wood on the road S.W. of the airfield S. of Telgte, “Im
Berdell”’, 12 August 1994, H. E. Weber no. 94.812.4 (MANCH, holotypus; herb. A.L.B., herb.
H.E.W., isotypi).

Stem high arching, greenish or somewhat reddish, sharply to bluntly angled, with flat sides, each
with (5—)10-30 mostly tufted hairs per 1 cm. Prickles 4-8 per 5 cm, slightly yellowish, equal,
slenderly declining or more rarely slightly curved, 7—8 (—-8-5) mm long with a 6-8 mm broad base.

Leaves 5-nate, digitate (exceptionally and rarely some 6—7-nate), dull green above with 15-25
hairs per cm’, green beneath, moderately hairy with simple hairs usually not or hardly to be felt by
touch. Terminal leaflet elliptical or elliptical-obovate, with a slightly cordate (rarely rounded) base
and a distinct acuminate 15-23 mm long apex and with a petiolule 33-41% as long as the lamina.
Serration coarse, up to 3-5 mm deep, irregular with moderately cuspidate teeth and longer (not
recurved) main teeth. Basal leaflets 2-3 mm, stalked. Petiole hairy, with 9-13 curved prickles.
Stipules nearly filiform, up to 1 mm broad, with subsessile to very shortly stalked glands at the
margin.

Inflorescence large, broadly cylindrical with a broad apex, the upper 3—15 cm leafless, otherwise
with 1—3-nate leaves, with spreading (70—80°) branches divaricately branched in their upper half.
Rachis densely covered with small tufted and also longer hairs and with many sessile glands, prickles
3-5 per 5 cm, slender, slightly declining or a little curved, 6-7 mm long. Pedicels chiefly 10-15 mm
long, with dense small tufted hairs and hirsute with simple hairs up to 0-6-0-8 mm long, stalked

348 H. E. WEBER AND A. L. BULL

glands 0-2, up to 0-3 mm long, prickles 3-7, often unequal, patent or a little declining, 1-2(-3) mm
long. Bracteoles usually with short-stalked glands. Sepals greyish-green, usually without prickles,
reflexed. Petals white, obovate, mostly notched at the apex, 10-15 mm long. Stamens white,
exceeding greenish styles. Anthers and ovaries glabrous, receptacle with few hairs. Fruits well
developed. Flowering time (June—) July (-August).

The plant belongs to subgen. Rubus sect. Rubus ser. Sylvatici (P. J. Miller) Focke. Because of the
yellowish prickles and with some respect also to the leaves it reminds one a little of Rubus
platyacanthus P. J. Muller, but differs much from that species mainly by the panicle which is similar
to that of Rubus gratus Focke. The plant is named after the well known British batologist E. S.
Edees (1907-1993) who published together with A. Newton the monograph Brambles of the British
Isles in 1988.

In 1907 the Rev. Augustin Ley collected a bramble at Apley, Lincolnshire, which he named Rubus
silvaticus Weihe & Nees (Ley 1908), a specimen of which is now in the herbarium at Lincoln
Museum. This was seen around 1936 by W. C. R. Watson who redetermined it as Rubus
macrophylloides Genevier. Edees (1966) saw the specimen in 1965 and stated that “‘it is unlikely that
this determination will stand”. He subsequently referred to the Lincolnshire plant as ‘‘false
macrophylloides” (A. Newton, pers. comm.). Edees stated that he had seen it in the field at
Stainton Wood, Short Wood and Great West Wood (the latter two sites being at Apley) where it
was abundant.

ECOLOGY AND DISTRIBUTION

Hedgebanks, wood margins, shrubberies, clearings, road margins on medium, siliceous (not
calcareous) soils. In Germany it occurs in Westphalia in the plain district around Minster, chiefly
between Telgte and Altenberge, growing mainly together with Rubus gratus Focke, R. plicatus
Weihe & Nees, R. vigorosus P. J. Muller & Wirtgen, R. pyramidalis Kaltenb., R. silvaticus Weihe &
Nees, R. rudis Weihe and R. calvus H. E. Weber. In Britain in the ancient Tilia cordata woods now
known as Bardney Forest, which includes Apley, to the E. of Lincoln, Rubus edeesii is associated
with R. mucronatoides Ley ex Rogers, R. calvatus Lee ex Bloxam, R. gratus Focke, R. vestitus
Weihe and R. lindleianus Lees. To the S.W. andS.E. of Lincoln, it grows on acid sands and gravels
in the area between Woodhall Spa and Newark, in contrast to the boulder clay of Bardney, and here
the associated species include R. gratus, R. nemoralis P. J. Muller, R. lindleianus, R. vestitus Weihe,
R. hylocharis W. C. R. Watson, R. polyanthemus Lindeb., R. pyramidalis and R. platyacanthus P. J.
Muller & Lef.

The great distance between the British and German distribution areas is due to seed transport by
birds (Whitethroats, Sy/via communis, and others) which are able to carry these by non-stop flights
directly from one place to another over distances up to 500 km (Berthold in Weber 1987), thus often
forming discontinuous distribution patterns, or spots (each resulting from a single bird-sown
bramble) along the flight paths of migrating birds. By subsequent continuous spreading such spots
usually later spread to greater distribution areas (Weber 1987). Like other brambles (Rubus
dasyphyllus (Rog.) Marsh., R. tuberculatus Bab.), R. edeesii seems probably to have originated in
Britain and was spread to continental Europe.

REPRESENTATIVE EXSICCATAE

England:

v.c. 53, S. Lincolnshire: Northon Disney, 3 August 1993, SK/88.59, A.L. Bull (herb. A.L.B., herb.
H.E.W.). Stapleford Wood, 3 August 1993, SK/88.56, A.L. Bull (herb. A.L.B.).

v.c. 54, N. Lincolnshire: Ostler’s Plantation, Woodhall Spa, 29 July 1993, TF/21.62, A.L. Bull.
Stainton Wood, 29 July 1993, TF/07.78, A.L. Bull (herb. A.L.B.).

v.c. 56, Nottinghamshire: Crowood Farm Belt, Thorney, 3 August 1994, SK/87.73, A.L. Bull.
Wigsley Wood, 3 August 1994, SK/84.70, A.L. Bull (herb. A.L.B.).

Germany (Westphalia):

(Localities quoted as given on the sheets and arranged according to the grid system used for

mapping in Germany.)

4012.14: Kasewinkel stidlich Handorf bei Munster, 12 August 1994, H.E. Weber no. 94.81/2.7.
Nordwestlich Haus Moilenbeck bei Telgte, July 1975, R. Wittig no. R113 (herb. H.E.W.).

RUBUS EDEESI, A NEW BRAMBLE SPECIES 349

4012.41: “Im Berdel’” zwischen Munster und Telgte, 15 July 1975, E. Foerster no. 750715.4.
Flugplatz Telgte bei Miinster, 15 July 1975, E. Foerster no. 750715.06 (herb. Foerster, herb.
H.E.W.). Wallhecke nérdlich Wolbeck, c. 500 m nordlich B6ckmann, July 1975, R. Wittig no.
R98 (herb. H.E.W.).

4012.42: 1 km 6stlich Flugplatz bei Telgte, July 1975, R. Wittig no. R120 (herb. H.E.W.).

4108.13: NSG Schwarzes Venn nordwestlich GroB Reken, July 1975, R. Wittig s. n. (herb. H.E.W.).

4111.1: ViehstraBe in der Ventruper Heide bei Munster, July 1976, R. Wittig s. n. (herb. H.E.W.).

REFERENCES

EpeEs, E. S. (1966). Brambles of Lincolnshire. Proceedings of the Botanical Society of the British Isles 6: 209-
214.
Ley, A. (1908). Some Lincolnshire Rubi. Journal of Botany 46: 53.

WEBER, H. E. (1985). Rubi Westfalici. Minster.
WEBER, H. E. (1987). Typen ornithochorer Arealentwicklung, dargestellt an Beispielen der Gattung Rubus L.

(Rosaceae) in Europa. Botanischer Jahrbiicher fiir Systematik 108: 525-535.
Wittic, R. (1975). Die Gebtisch- und Saumgesellschaften der Wallhecken in der Westfalischen Bucht.

Abhandlungen aus dem Landesmuseum ftir Naturkunde zu Miinster in Westfalen 38(3): 1-78.

(Accepted February 1995)

Watsonia 20: 351-365 (1995) 351

Eleven new British species of Hieracium L. section Alpina
(Fries) F. N. Williams

Pa) SELLE .C WEST.

The Herbarium, Department of Plant Sciences, Downing Street, The University, Cambridge,
CB2 3EA

and
D. J. TENNANT

Marhead Grange, Arkendale, Knaresborough, North Yorkshire, HG5 ORG

ABSTRACT

Eleven new species of section Alpina (Fries) F. N. Williams of Hieracium L. (Asteraceae), Hieracium calvum P.
D. Sell & D. J. Tennant, sp. nov., H. completum P. D. Sell & C. West. sp. nov., H. kennethii P. D. Sell & D. J..
Tennant, sp. nov., H. leptodon P. D. Sell & D. J. Tennant, sp. nov., H. milesii P. D. Sell & C. West, sp. nov., H.
mundum P. D. Sell & C. West, sp. nov., H. optimum P. D. Sell & C. West, sp. nov., H. pensum P. D. Sell & C.
West, sp. nov., H. perscitum P. D. Sell & C. West, sp. nov., H. probum P. D. Sell & C. West, sp. nov. and H.
subglobosum P. D. Sell & C. West, sp. nov. and three new forms, H. hanburyi forma pusillum P. D. Sell & D. J.
Tennant, forma nova, H. insigne forma celsum P. D. Sell & D. J. Tennant, forma nova and H. marginatum
forma chaetocephalum P. D. Sell & C. West, forma nova, are described and their distributions given. In
addition, a new status is given to two taxa, H. eximium forma tenellum (Backh.) P. D. Sell & C. West, stat. nov.
and H. hanburyi forma atraticeps (Pugsley) P. D. Sell & D. J. Tennant, stat. nov.

Keryworps: hawkweeds, Scotland.

INTRODUCTION

Section Alpina (Fries) F. N. Williams is probably the most difficult group of species of the genus
Hieracium L. occurring in the British Isles. During the 1970s, P.D.S. and C.W. attempted a revision
of the group, based particularly on the fine series of specimens in CGE collected by A. G. Kenneth
and by B. A. Miles. They discovered a number of new species and prepared descriptions of them.
The retirement from active work of C.W. in 1980, followed by his death in 1986, and long periods of
ill-health for P.D.S. since 1981 have delayed publication of the new species. This has been beneficial
in that D.J.T., who has long made a special study of the group, has been able to revise and add to the
descriptions. Alone or with A. G. Kenneth, he has seen all the species in the field, and he has grown
them all side by side in his garden. In the course of his work three more new species have been
discovered.

This paper gives only the Latin diagnoses and descriptions, in order to validate the species. A
second paper will follow in which comparable English descriptions of all the species in section
Alpina found in the British Isles will be given. The distinction of the species in this section is so
critical that the diagnosis should be taken only as a guide; the detailed description should always be ©
checked for certain identification. The chromosome counts given in this paper are derived from
Stace et al. (1995).

Most of the taxa described in this paper are rare plants, some populations being restricted to only
a few specimens. In the interests of their conservation, therefore, it is important that they are not
collected without expert knowledge.

352, P. D. SELL, C. WEST AND D. J. TENNANT
DESCRIPTIONS OF NEW SPECIES AND FORMS

Hieracium calvum P. D. Sell & D. J. Tennant, sp. nov.
Hovotypus: Rock-ledges in east-facing coire of Carn Crom on the west side of Glen Derry, South
Aberdeen, v.c. 92, GR NO/02.95, 6 August 1978, D. J. Tennant no. D1/78 (CGE).

Ab Hieracio larigensi (Pugsley) P. D. Sell & C. West et H. globosifloro Pugsley et H. graniticola W.
R. Linton folio caulino infimo plerumque grandiore, involucro angustiore, involucri squamis paulo
latioribus, caule pilis stellatis paucioribus in parte superiore ornato differt; ab H. backhousei F. J.
Hanb. foliis basalibus integrioribus, folii caulini infimi margine pilis simplicibus densioribus
longioribusque vestito, involucro angustiore villosiore, involucri squamis acutioribus distinguitur;
ab H. milesii P.D. Sell & C. West et H. hanburyi Pugsley involucro pilis glanduliferis brevioribus
sparsioribusque vestito differt; a speciebus ceteris britannicis Sectionis Alpinorum stylis flavis
ornatis folitis suis basalibus in pagina superiore plerumque glabris distinguitur.

Planta phyllopoda. Caulis 13—20(—27) cm altus, interdum 2 vel aliquot, gracilis vel subrobustus,
saepe flexuosus, ad basin interdum rubriusculoviolaceus; in parte inferiore pilis simplicibus
eglanduliferis paucis vel numerosis mediocribus longisque sinuosis albis nigricantibasibus (basin
versus densioribus longissimisque), pilis glanduliferis paucis vel aliquot perbrevissimis flaviusculis,
pilis stellatis paucis vel nullis vestitus; in parte superiore pilis simplicibus eglanduliferis numerosis
mediocribus longisque (vel in parte suprema longissimis) pallidis vel grisetusculis ad bases crassas
nigris, pilis glanduliferis dispersis brevissimis obscuris, pilis stellatis aliquot vel numerosis sed non
densis vestitus. Folia pallidiuscule vel mediocriter vel raro intensiuscule viridia, interiora in pagina
superiore plerumque nitida, in pagina marginibusque interdum rubriusculoviolaceotincta, in pagina
inferiore pallidiora, in pagina tota superiore plerumque glabra, sed ad margines petiolumque pilis
simplicibus eglanduliferis numerosis vel densis longis vel longissimis sinuosis albis nigricantibasibus,
in pagina inferiore pilis simplicibus eglanduliferis dispersis vel numerosis pallidis vel nigricantibasi-
bus, in pagina inferiore et ad margines pilis glanduliferis paucis perbrevissimis flaviusculis, ad
margines interdum pilis stellatis paucis vestita; folia basalia pauca vel numerosa, exteriora 10-45
mm longa, 6-26 mm lata, plerumque late elliptica vel ovata, ad apicem late rotundo-obtusa vel
subacuta vel interdum apiculata, integra vel denticulata, ad basin cuneata vel breviter contracta,
interiora 30-90 mm longa, 5—22 mm lata, rigida et coriacea, saepe canaliculata, elliptica vel anguste
elliptica vel oblanceolata, ad apicem saepe tortum plerumque acuta saepe acuminata aliquando
subacuta, plerumque integra interdum undulata vel serratodentata, dentibus paucis haud profundis
vel anguste mammiformibus, ad basin longe attenuata vel aliquando breviter angustata; petioli
magis minusve (saepe late) alati, basin versus saepe rubriusculoviolaceotincti; folia caulina 1—2(-3),
infimum 10-110 mm longum, 1-5-12 mm latum, semipatens, lineari-oblanceolatum vel anguste
ellipticum, ad apicem acutum vel acuminatum vel raro obtusum, integrum, ad basin attenuatum,
sessile vel petiolatum, in pagina superiore glabrum, sed in marginibus et pagina inferiore pilis
simplicibus eglanduliferis numerosis longiusculis et pilis stellatis aliquot vestitum, cetera folia
caulina plerumque linearia acuta integra vel interdum bracteiformia, unum raro gemma rudimentali
in axilla praeditum. Capitulum plerumque solitarium, interdum 2, vel 2 vel plura a caulibus e rosula
basali exorientibus portata, 40-50 mm diametro; involucrum facie obscure viridiusculogrisea vel
viridiuscula, anguste campanulatum, ad basin breviter angustatum vel rotundatum; pedunculi (si
adsunt) cauli superiori similiter vestiti. Involucri squamae ante anthesin incumbentes, atriusculovir-
ides, interiores marginibus pallidius viridiusculis; omnes pilis simplicibus eglanduliferis longis
sinuosissimis albis vel aliquantum griseiusculis nigricantibasibus inaequaliter distributis (non-
nusquam sparsis sed ad basin semper numerosis crispissimis saepe perlongis), pilis glanduliferis
numerosis brevissimis vel brevibus pallidis vel obscuris, a pilis simplicibus eglanduliferis partim
modo tectis, vestitae, sine pilis stellatis, interiores ad apicem pilis simplicibus nonnullis brevissimis
ornatae; squamae interiores 10-15 mm longae, 1—2 mm latae, appressae vel laxe appressae, lineari-
lancolatae, ad apicem acutissimae vel abrupte acutae vel acuminatae, interdum violaceotinctae;
squamae exteriores breviores, ad apicem subacutae vel acutae. Ligulae mediocriter flavae;
exteriores ad 4(-4-6 cult.) mm latae, ad apicem magis minusve discretae et saepe profunde
dentatae, haud profunde concavae, in pagina inferiore pilis simplicibus eglanduliferis paucissimis
vel aliquot breviusculis pallidis vel obscuriusculis vestitae, ad apicum pilis taiibus brevissimis vel
brevibus paucis vel nullis ultra dentium margines protrudentibus. Sty/i flavi vel flaviusculi, in sicco

NEW BRITISH SPECIES OF HIERACIUM 353

saepe obscuriusculi. Receptaculi alveoli margine breviter dentati. Cypselae 3-8-4-1 mm longae,
purpureiusculonigrae. 2n = 36.

Extremely local and very scarce in crevices and on ledges of shelving coarse-grained granite rocks
and boulders, between 800 and 980 m: on Carn Crom and Derry Cairngorm on the west side of Glen
Derry in South Aberdeen (v.c. 92), and on the east side of Cairngorm in Easterness (v.c. 96).
Endemic.

Hieracium completum P. D. Sell & C. West. sp. nov.
Hototypus: East-facing cliffs, c. 808 m, Coire Etchachan, South Aberdeen, v.c. 92, GR NO/
017.997, 3 August 1966, R. W. Jones & B. A. Miles no. 66/147 (CGE).

A plerisque Sectionis Alpinorum speciebus britannicis maculis suis fusciusculoviolaceis in pagina
superiore foliorum basalium dispersis distinguitur. Ab Hieracio probo P. D. Sell & C. West foliis
basalibus clarioribus nitidioribus plerisque ovatis praecipue differt.

Planta phyllopoda. Caulis 13-30 cm altus, diametro mediocri, interdum robustus et ad basin
lignosus, plerumque infra involucrum incrassatus, striatus, plerumque ad basin rubriusculovio-
laceus et interdum in porcis in parte inferiore guttatus, pilis simplicibus eglanduliferis paucis vel
numerosis pallidis nigricantibasibus (basin versus et ad petiolos plerumque multis longis, in parte
superiore brevioribus sed longius nigricantibasibus), pilis glanduliferis paucis vel numerosis
brevissimis pallidis obscurisque, pilis stellatis nonnullis (utrisque in parte superiore numerosioribus)
vestitus. Folia pallidiuscule vel mediocriter clare viridia, plerumque subnitida raro hebetata,
plerumque per totam paginam superiorem obscure fusciusculoviolaceomaculata vel marmorata, in
pagina inferiore saepe pallidiora, costa rubriusculoviolaceotincta; folia basalia plerumque numer-
osa, aliquando coriacea, pleraque in pagina superiore et ad margines pilis simplicibus eglanduliferis
paucis vel numerosis magis minusve uniformiter dispersis brevibus mediocribusque longiusculisque
graciliusculis subrigidis sinuosis vel ad apicem curvatis albis plerumque pallidibasibus, in pagina
inferiore pilis simplicibus eglanduliferis numerosis brevibus vel mediocribus gracilibus mollioribus
sinuosis, in utraque pagina et in marginibus pilis glanduliferis paucis brevissimis flaviusculis vestita,
exteriora 5—45 mm longa, 3—35 mm lata, subrotunda vel ovata vel elliptica vel raro obovata, ad
apicem rotundo-obtusa vel obtusa saepe mucronulata, denticulata, ad basin plerumque late cuneata
interdum truncata, interiora 10—70(—100) mm longa, 4-30 mm lata, plerumque ovata interdum late
elliptica vel late lanceolata, ad apicem late acuta vel subacuta raro obtusa plerumque mucronata vel
apiculata, interdum convoluta, magis minusve ordinate serratodentata, dentibus anguste mam-
miformibus acutis, interdum denticulata, ad basin cuneata vel attenuata; petioli breves vel longi,
plerumque rubriusculoviolaceotincti praecipue ad basin, pilis simplicibus eglanduliferis paucis vel
numerosis mediocribus vel longis interdum appressis vestiti; folia caulina 1—3(—4), infimum 1-60 mm
longum, 20-30 mm latum, basalia interiora saepe simulans, lanceolatum vel ellipticum vel anguste
ovatum, ad apicem acutomucronatum vel acuminatum, denticulatum vel irregulariter serratodenta-
tum, dentibus paucis angustis acutis, ad basin saepe cuneatum et perspicue petiolatum, cetera
linearia acuta vel magis minusve bracteiformia erecta, summum filamentosum, omnia folia caulina
pilis simplicibus eglanduliferis numerosis, pilis glanduliferis nonnullis minutis, saepe in pagina
inferiore pilis stellatis nonnullis vestita. Capitulum plerumque solitarium, interdum 2-3, 35-50 mm
diametro; involucrum facie atriuscula vel obscure viridiusculogrisea, late campanulatum, ad basin
valde constrictum; pedunculi (si adsunt) cauli superiori similiter vestiti. Involucri squamae ante
anthesin incumbentes, viridiusculoatrae, interiores marginibus indistinctis viridiusculis; omnes pilis
simplicibus eglanduliferis densiusculis mediocribus vel longis (ad 3 mm) gracilibus pallidis vel
griseiusculis ad bases longas incrassatas nigris, pilis glanduliferis numerosis brevissimis vel brevibus
flaviusculis obscurisque, a pilis simplicibus eglanduliferis partim modo tectis, vestitae, sine pilis
stellatis sed interiores ad apicem pilorum ramosorum simpliciumque brevium caespite ornatae;
Squamae interiores 11—15(—18) mm longae, 1-2—1-8(—2-0) mm latae, inaequales, valde appressae,
magis minusve late lineari-lanceolatae, ad apicem abrupte acutae vel subacutae; squamae exteriores ©
breviores erectae, ad apicem magis minusve obtusae. Ligulae clare intensiuscule vel raro
mediocriter flavae; exteriores latiusculae (ad 4-5 mm), ad apicem magis minusve discretae et
profunde dentatae, concavae apicemque versus valde ascendentes, in pagina inferiore pilis
simplicibus eglanduliferis paucis brevissimis vel brevibus pallidis vestitae vel nullis, ad apicem pilis
talibus aliquot perbrevissimis ultra dentium margines protrudentibus. Sty/i uniformiter obscurissimi

354 P. D. SELL, C. WEST AND D. J. TENNANT

(griseiusculi). Receptaculi alveoli margine breviter dentati. Cypselae 3-6—4-5 mm longae, purpur-
eiusculonigrae. 2n = 36.

Rock-ledges, dry gullies and stream-banks on both schist and granite between 600 and 1200 m: on
the Cairnwell in East Perth (v.c. 89); in several localities in and around the heads of Glen Clova and
Glen Isla in Angus (v.c. 90); in Glen Callater, on Lochnagar and at Dubh-loch in South Aberdeen
(v.c. 92); in the Cairngorm Range on Creag an Dail Bheag [Little Craigandail] and in Coire
Etchachan (both in v.c. 92) and on many of the mountains bordering both sides of the Lairig Ghru
Pass (v.cc. 92 and 96); in Glen Einich and Coire Garbhlach in the Cairngorm mountains and on the
adjacent Monadhliath Hills, Easterness (v.c. 96). Endemic.

This species was mistakenly known to F. J. Hanbury, E. F. and W. R. Linton, E. S. Marshall and
other earlier botanists as H. curvatum Elfstrand, a species now considered not to occur in the British
Isles. Most of the specimens later referred to H. pseudocurvatum (Zahn) Pugsley (1948) belong to
H. completum. H. pseudocurvatum is in fact restricted to two localities on the Cairngorms.

Hieracium eximium Backh., Monogr. Brit. Hier. 20 (1856).
Forma eximium.
Styli viridiusculogrisei vel obscurissime grisei. 2n = 36.
Forma tenellum (Backh.) P. D. Sell & C. West, stat. nov.
BasionyM: H. eximium var. tenellum Backh., Monogr. Brit. Hier. 21 (1856).
Styli flavi vel aliquantum sordide flavi. 2n = 36.
Other than style colour there is little to distinguish these two forms, but forma eximium shows a
preference for schistose rock and forma tenellum for granite.

Hieracium hanburyi Pugsley in J. Bot. (London) 79: 178 (1941).

Forma pusillum P. D. Sell & D. J. Tennant, forma nova

Ho otyrus: Rock-ledge on schistose rock, c. 760 m, at head of Canness Glen, Glen Isla, Forfar
[Angus], v.c. 90, GR NO/20.77, 12 July 1982, D. J. Tennant, no. C1/82 (CGE).

Planta parva; folia subintegra vel haud profunde dentata; involucri squamae abrupte acutae vel
subacutae vel obtusae, pilis glanduliferis et simplicibus eglanduliferis numerosis ornatae; styli
obscuri. 2n = 36.

Forma hanburyi

Synonym: H. hanburyi var. humile Pugsley in J. Linn. Soc. London (Bot.) 54: 52 (1948).

Planta saepe major; folia magis dentata; involucri squamae anguste acutae, pilis glanduliferis et
simplicibus eglanduliferis numerosis ornatae; sty/i flavi vel aliquantum discolores. 2n = 36.

Forma atraticeps (Pugsley) P. D. Sell & D. J. Tennant, stat. nov.
BasionyM: H. hanburyi var. atraticeps Pugsley in J. Linn. Soc. London (Bot.) 54: 51 (1948).

Planta saepe magna; folia magis minusve dentata; involucri squamae anguste acutae, pilis
glanduliferis densis, pilis simplicibus eglanduliferis nullis vel sparsis ornatae; styli magis minusve
flavi. 2n = 36.

Hieracium hanbury1is a very variable species whose limits are difficult to define. We recognise the
three most distinct variants as forms. P.D.S. and C.W. once recognised forma atraticeps as a distinct
species because of the characteristic clothing of the heads, but it grows with forma hanburyi and
intermediates occur. Forma pusillum is known only from the Glen Isla and the Clova Mountains,
Angus (v.c. 90). H. perscitum and H. hanburyi can easily be confused on herbarium sheets, but they
look very different in the field.

Hieracium insigne Backh. in Phytologist (Newman), Series 2, 4: 806 (1853).

Lectotypus: Near Loch Ceanndin, Aberdeenshire, v.c. 92, August 1852, J. Backhouse jun. (CGE).
Forma insigne

Capitulum solitarium, 45—65 mm diametro; involucri squamae 13-20 mm longae, 0-7—1-5 mm latae,
laxiores, apicibus angustissimis. 2n = 36.

Forma celsum P. D. Sell & D. J. Tennant, forma nova

Ho.otypus: Schistose cliffs, c. 2400 ft [720 m], south side of Coire Garbhlach, Glen Feshie,
Easterness, v.c. 96, GR NN/880.941, 12 August 1966, R. W. Jones & B. A. Miles no. 66/216 (CGE).

NEW BRITISH SPECIES OF HIERACIUM 355

Capitula 1-3, 25-SO mm diametro; involucri squamae 10-17 mm longae, 1-0-1-5 mm latae,
appressae, apicibus interdum minus angustis. 2n = 27, 36.

Forma insigne is known only from cliffs above Loch Kander (Loch Ceanndin) in Glen Callater,
South Aberdeen (v.c. 92). Forma celsum occurs in Coire an Lochaine Uaine, Cairntoul, in South
Aberdeen (v.c. 92), in Coire Garbhlach, in Glen Feshie and on Creag an Leth-choin in the Lairig
Ghru Pass, both in Easterness (v.c. 96), and in several places on Beinn na Socaich, Glen Spean,
Westerness (v.c. 97). Endemic.

Hieracium kennethii P. D. Sell & D. J. Tennant, sp. nov.
Ho .ortypus: An Teallach, West Ross, v.c. 105, GR NH/055.865, c. 650 m, 1984, A. G. Kenneth no.
5184 (CGE).

Ab Hieracio perscito P. D. Sell & C. West foliis coraceis nitidis, folio caulino infimo magis petiolato
pilisque stellatis paucis vestito, involucri squamis pilis glanduliferis manifestis multo paucioribus
vestitis; ab H. subgloboso P. D. Sell & C. West foliis basalibus exterioribus ad basin magis truncatis
plerumque clarius viridibus, folio caulino infimo majore saepe dentato, involucri squamis minus
acutis, ligulis intensius flavis; ab H. mundo P. D. Sell & C. West foliis basalibus latioribus
clarioribus nitidioribus, exterioribus saepe magis truncatis et dentibus latis obtusis instructis, folio
caulino infimo sine dentibus spinulosis, involucri squamis saepe acutioribus; ab H. penso P. D. Sell
& C. West foliis coriaceis nitidis claris plerumque in pagina superiore uniformiter pilosis, involucri
ad basin angustiore, squamis magis appressis, ligulis intensius flavis distinguitur. Ab omnibus alliis
Sectionis Alpinorum speciebus britannicis aut stylis suis obscuris aut separatione geographica
distinguitur.

Planta phyllopoda. Caulis 10-24 cm altus, gracilis vel mediocris, interdum flexuosus, aliquando
ad basin rubriusculoviolaceotinctus, in parte inferiore pilis simplicibus eglanduliferis paucissimis vel
paucis vel numerosis mediocribus vel longis (ad 3 mm) saepe gracillimis sinuosis albis nigricantibasi-
bus, pilis glanduliferis dispersis brevissimis flaviusculis, pilis stellatis paucissimis vel dispersis
vestitus, in parte superiore pilis simplicibus eglanduliferis paucis vel numerosis mediocribus et
longis (ad 2 mm) pallidis vel griseiusculis nigricantibasibus, pilis glanduliferis numerosis vel
numerosissimis brevissimis vel brevibus vel mediocribus (ad 1-0 mm) obscuris, pilis stellatis
numerosis vel densis vestitus. Folia mediocriter vel intensiuscule clare viridia, supra subtusque
nitida, in pagina inferiore paulo modo pallidiora, in pagina superiore marginibusque pilis
simplicibus eglanduliferis numerosis vel densis magis minusve uniformiter dispersis mediocribus et
longis (ad 2 mm) gracilibus subrigidis albis pallidibasibus (folia exteriora saepe pilis paucis circa
costam vestita), in pagina inferiore pilis simplicibus eglanduliferis paucis vel numerosis vel
numerosissimis, in paginis marginibusque pilis glanduliferis paucis brevissimis flaviusculis, interdum
in utraque pagina circa costam pilis stellatis paucis vestita; folia basalia aliquot vel numerosa,
exteriora 5—35 mm longa, 4-30 mm lata, subrotunda vel late ovata vel obovata, saepe asymmetrica,
ad apicem late rotundo-obtusa interdum retusa vel apiculata, denticulata vel magis minusve
sinuatodentata, dentibus aliquot irregularissimis et interdum dentibus paucis retrorsis late mam-
miformibus (aliquando angustioribus), ad basin late cuneata vel truncata, interiora 6-45 mm longa,
4-25 mm lata, subcoriacea, ovata vel elliptica, ad apicem late acuta apiculataque vel obtusa, sparse
denticulata vel irregulariter serratodentata, dentibus mediocribus vel angustis patentibus mam-
miformibus, ad basin late cuneata vel breviter contracta raro attenuata; petioli breves vel
longiusculi, pilis simplicibus eglanduliferis longis albis dense vestiti; folia caulina 1-5, infimum 13-
60 mm longum, 2-16 mm latum, semipatens, oblanceolatum, aliquando ellipticum, interdum fere
ad basin caulis positum et basalia interiora simulans, ad apicem acutum vel obtusum interdum
acuminatum vel cuspidatum, integrum vel dentibus aliquot irregularibus acutis serratodentatum, ad
basin in petiolum plerumque angustiusculum attenuatum vel anguste contractum, in utraque pagina
et ad margines pilis simplicibus eglanduliferis paucis vel magis minusve irregulariter dispersis
pallidis (in petiolo densioribus, in pagina inferiore nigricantibasibus), in utraque pagina aliquando
pilis stellatis aliquot (praesertim in costa) vestitum, cetera folia caulina parva magis minusve acuta
integra vel bracteiformia. Capitulum plerumque solitarium, interdum 2, 30-48 mm diametro;
involucrum facie griseiusculoviridi vel viridiusculoatra, campanulatum, ad basin angustatum;
pedunculi (si adsunt) cauli superiori similiter vestiti. Involucri squamae ante anthesin incumbentes,
atriusculovirides vel virides; omnes pilis simplicibus eglanduliferis numerosis mediocribus longisque

356 P. D. SELL, C. WEST AND D. J. TENNANT

(ad 3 mm) gracilibus sinuosis pallidis vel griseiusculis nigricantibasibus, pilis glanduliferis numerosis
brevibus brevissimisque atriusculis (a pilis simplicibus eglanduliferis pro parte majore tectis)
vestitae, sine pilis stellatis sed interiores ad apicem pilorum brevium caespite manifesto ornatae;
squamae interiores 11-15 mm longae, 1-2-1-8 mm latae, inaequales, strictim appressae, lineari-
lanceolatae, ad apicem plerumque abrupte acutae interdum subacutae vel obtusae; squamae
exteriores breviores erectae semiappressae. Ligulae uniformiter intensiuscule flavae vel aliquando
mediocriter flavae; exteriores 3-4 mm latae, ad apicem aliquantum discretae et magis minusve
profunde dentatae, fere rectae vel aliquantum concavae, in pagina inferiore pilis simplicibus
eglanduliferis paucissimis vel aliquot brevissimis pallidis vestitae, ad apicem pilis nullis talibus vel
aliquot perbrevissimis vel aliquando paucis brevissimis ultra dentium margines protrudentibus. Styli
magis minusve uniformiter obscure viridiusculogrisei. Receptaculi alveoli margine dentati. Cypselae
3.9-4.3 mm longae, purpureiusculonigrae. 2n = 36.

Rocky knolls and steep grassy slopes from c. 600 to 700 m on Torridonian Sandstone on Mac is
Mathair and elsewhere, rather locally, on the north-eastern and north-western sides of the An
Teallach Range. Dundonnell, in West Ross (v.c. 105); also on Cnoc na Creige, near Glen Coul,
from c. 450 to 550 m, West Sutherland (v.c. 108). Endemic.

Named after Archibald Graham Kenneth (1915-1989), in recognition of his fine work over many
years on the hawkweeds of western Scotland.

Hieracium leptodon P. D. Sell & D. J. Tennant, sp. nov.
Hotortypus: North-east corrie, Ben More, Mid Perth, v.c. 88, August 1978, D. J. Tennant (CGE).

Hieracio calendulifloro Backh., H. eximio Backh. et H. notabili P. D. Sell & C. West simile, sed
folio caulino infimo conspicuo, in plantis bene effectis dentibus multis longissimis gracillimis acutis
prorsum curvatis ornato distinguitur.

Planta phyllopoda. Caulis 12-30 cm altus, medius vel gracilis, interdum flexuosus, ad basin
plerumque rubriusculoviolaceotinctus, in parte inferiore pilis simplicibus eglanduliferis dispersis vel
numerosis mediocribus vel longis vel longissimis (ad 5 mm) gracilibus tenuibus sinuosis pallidis
nigricantibasibus, pilis glanduliferis paucis perbrevissimis vel brevissimis pallidis vel obscurioribus,
pilis stellatis paucis, in parte superiore pilis simplicibus eglanduliferis numerosis longe nigricantiba-
sibus, pilis glanduliferis aliquot vel densis brevissimis vel brevibus nigriusculis, pilis stellatis aliquot
vel numerosis sed non densis vestitus. Folia pallidiuscule vel mediocriter viridia, pruinam levem
caeruleogriseam ferentia, in pagina superiore hebetata vel paulo modo nitida, saepe rubriusculovio-
laceotincta praecipue in marginibus, in pagina inferiore pallidiora; folia basalia numerosa, exteriora
8-50 mm longa, 5—25 mm lata, late elliptica vel ovata, ad apicem rotundo-obtusa raro retusa
interdum apiculata, denticulata vel subintegra, interdum dentibus aliquot mammiformibus, ad
basin rotundata vel cuneata vel breviter angustata interdum attenuata, interiora 25—90 mm longa, 9—
22 mm lata, anguste vel late elliptica raro oblanceolata, ad apicem interdum tortum obtusa vel acuta
saepe apiculata, denticulata vel dentata vel serratodentata, interdum undulata, dentibus regulari-
bus vel irregularibus saepe ascendentibus anguste vel raro late mammiformibus ad apices longe
rotundoapiculatis, in triente superiore saepe integra, ad basin breviter angustata vel attenuata,
omnia pilis simplicibus eglanduliferis numerosis brevibus mediocribusque longisque (ad 3 mm) albis
pallidibasibus vel nigricantibasibus (in paginis ambabus plerumque uniformiter distributis et in
marginibus longis densis), pilis glanduliferis dispersis brevissimis flaviusculis vestita; petioli
plerumque breves, alati, pilis simplicibus eglanduliferis densiusculis longissimis sinuosis vestiti,
subtus saepe rubriusculoviolaceotincti; folia caulina 2-4(—5), infimum 10-90 mm longum, 2—20 mm
latum, semipatens, anguste ellipticum vel elliptico-oblongum, ad apicem acutum vel subacutum,
saepe serratodentatum vel serratum, dentibus irregularibus longissimis (ad 10 mm) angustis saepe
prorsum curvatis apiculatis saepe spinulosis ornatum, ad basin angustatum sessile vel semiamplexi-
caule vel subpetiolatum, folium a basi secundum saepe dentibus paucis acutis angustis ornatum vel
integrum, cetera linearia acuta integra vel bracteiformia vel filamentosa erecta. Capitula 1-3(-4),
35-50 mm diametro: involucrum facie viridiusculogrisea, cylindricum vel campanulatum, ad basin
rotundatum vel truncatum; pedunculi longi graciles, cauli superiori similiter vestiti. Involucri
squamae ante anthesin incumbentes, atriusculovirides; interiores 12-16 mm longae, 0-8-1-0 mm
latae, appressae, lineari-lanceolatae, ad apicem plerumque gradatim acutae vel acutissimae
interdum abrupte acutae, intimae interdum filamentosae; exteriores multo breviores semilaxae

NEW BRITISH SPECIES OF HIERACIUM Spill

erectae, ad apicem subacutae; omnes pilis simplicibus eglanduliferis numerosis vel densis medio-
cribus vel longis (ad 3 mm) pallidis vel griseiusculis nigricantibasibus, pilis glanduliferis numerosis
brevissimis obscuris, a pilis simplicibus eglanduliferis plerumque omnino tectis, sine pilis stellatis,
interiores ad apicem pilis paucis brevissimis vestitae. Ligulae pallide clare vel raro mediocriter
flavae; exteriores numerosae, aliquantum angustae (ad 4 mm), ad apicem plerumque prcfunde et
irregulariter dentatum magis minusve discretae, concavae supraque ascendentes, in pagina
inferiore pilis simplicibus eglanduliferis numerosis vel densis brevissimis pallidis vestitae, ad apicem
pilis talibus numerosis brevissimis nonnullisque brevibus ultra dentium margines protrudentibus.
Styli intense viridiusculogrisei. Receptaculi alveoli margine breviter dentati. Cypselae 3-5—4-0 mm
longae, purpureiusculonigrae. 2n = 36.

Cliff-ledges and crevices in large boulders on muscovite-albite pelitic schist and other hard mica-
schist rocks at 710-890 m in the north-east corrie of Ben More, Mid Perth (v.c. 88). Endemic.

Hieracium marginatum P. D. Sell & C. West in Watsonia 6: 304 (1967).

Forma marginatum

Involucri squamae pilis simplicibus eglanduliferis numerosis vel densis, ad 2-5 mm longis, ornatae.
2n = 36.

Forma chaetocephalum P. D. Sell & C. West, forma nova

Ho.ortypus: Summit corrie of Ben Dearg, East Ross, v.c. 106, 1976, A. G. Kenneth no. 9076 (CGE).

A forma marginato involucri squamis pilis simplicibus eglanduliferis densis, ad 4 mm longis, ornatis
distinguitur. 2n = 36.

These two forms have the same area of distribution — Westerness (v.c. 97), Ross (v.cc. 105 and
106) and Sutherland (v.cc. 107 and 108) — but the single character in which they differ, the woolliness
of the head, is so striking that it is worth distinguishing them by separate names.

Hieracium milesii P. D. Sell & C. West, sp. nov.
Ho .orypus: Cliff-ledges, c. 880 m, due south of Loch Kander, Glen Callater, South Aberdeen, v.c.
92, GR NO/190.807, 6 August 1966, R. W. Jones & B. A. Miles no. 66/162 (CGE).

Ab Hieracio tenuifronti P. D. Sell & C. West foliis caulinis saepe magis effectis, involucri squamis
laxioribus pilisque glanduliferis manifestis ornatis, stigmatibus flaviusculis distinguitur.

Planta phyllopoda. Caulis singularis vel duplicatus, 8—20(—28) cm altus, gracilis vel robustus,
saepe flexuosus, ad basin interdum rubriusculoviolaceus, ubique pilis simplicibus eglanduliferis
paucis vel numerosis brevibus mediocribusque longisque gracilibus sinuosis albis vel griseiusculis ad
bases incrassatas nigricantibus (in parte inferiore modice vestitus, in parte superiore pilis numerosis
brevibus vel mediocribus saepe griseiusculis), pilis glanduliferis aliquot vel numerosis brevissimis
pallidis obscurisque (in parte superiore numerosis vel densis longioribus, ad 1-0 mm), pilis stellatis
multis (in parte superiore densiusculis) vestitus. Folia pallidiuscule vel intensiuscule viridia
aliquando glauciviridia, in pagina superiore hebetata vel nitida, in pagina inferiore pallidiora, saepe
rubriusculoviolaceotincta: folia basalia pauca vel numerosa, exteriora 640 mm longa, 5-26 mm
lata, ovata vel obovata vel late elliptica, interdum asymmetrica, saepe undulata vel convoluta, ad
apicem late rotundo-obtusa, subintegra, denticulata vel serratodentata, dentibus aliquot haud
profundis vel anguste mammiformibus plerumque ad marginum trientes duas inferiores limitatis, ad
basin cuneata vel abrupte contracta, in pagina superiore pilis simplicibus eglanduliferis aliquot
brevibus mediocribusque longisque uniformiter distributis vel aliquando ad margines limitatis, in
pagina inferiore pilis talibus aliquantum brevioribus mollioribusque dispersis vel nullis, in utraque
pagina et in marginibus pilis glanduliferis aliquot perbrevissimis flaviusculis, in marginibus interdum
pilis ramosis paucis brevissimis vestita, interiora 15—75 mm longa, 3-30 mm lata, rigidiuscula et
subcoriacea, anguste vel late elliptica, ad apicem late acuta saepe apiculata aliquando obtusa,
subintegra vel irregulariter sinuatodentata vel dentibus magnis anguste vel late mammiformibus
interdum cuspidatis, ad marginum trientes duas inferiores limitatis, ornata, ad basin cuneata vel
attenuata vel raro subtruncata, in utraque pagina pilis simplicibus eglanduliferis plerumque
numerosis brevibus mediocribusque longisque subrigidis ad bases plerumque bulbosas pallidis vel
discoloribus vel aliquando nigriusculis uniformiter distributis vel prope et ad margines restrictis,
pilis glanduliferis et interdum ramosis foliis exterioribus similiter vestita; petioli breviusculi vel

358 P. D. SELL, C. WEST AND D. J. TENNANT

longi, magis minusve graciles, ad basin saepe valde rubriusculoviolacei, pilis simplicibus eglanduli-
feris paucis vel densiusculis mediocribus vel longis subrigidis pallidis vestiti; folia caulina (1-)2-4
(—5), infimum 10—60(—90) mm longum, 2-18 mm latum, rigide semierectum-vel patens, interdum
undulatum, lineari-oblanceolatum vel anguste ellipticum, ad apicem acutum vel raro rotundo-
obtusum, plerumque integrum, ad basin attenuatum, plerumque valde petiolatum, interdum
aliquantum alatum et semiamplexicaule, in pagina superiore pilis simplicibus eglanduliferis saepe
nigricantibasibus paucis vel nullis, in marginibus et aliquando in pagina inferiore et in paginae
superioris costa pilis talibus aliquot brevibus vel mediocribus, pilis glanduliferis brevissimis pallidis,
pilis stellatis paucis vel numerosis vestitum, folium a basi secundum minus (ad 45 mm longum, 4mm
latum) sed simile, interdum semipatens, cetera minora linearia bracteiformis semierecta vel ad
caulem appressa. Capitulum solitarium, raro 2(—3), aliquando cum altero a secundo caule e basi
exoriente portato, 25-40 mm diametro, interdum nutans; involucrum facie saturate viridiusculogri-
sea vel viridiusculonigra, campanulatum vel late campanulatum, ad basin rotundatum; pedunculi (si
adsunt) cauli superiori similiter vestiti. Jnvolucri squamae ante anthesin incumbentes, atriusculovir-
ides, marginibus pallidioribus; omnes pilis simplicibus eglanduliferis densiusculis mediocribus et
longis (ad 3 mm) gracilibus sinuosis griseiusculis vel intense griseis nigricantibasibus, pilis
glanduliferis numerosis brevissimis brevibusque mediocribusque obscuris, a pilis simplicibus
eglanduliferis plerumque leviter modo tectis, vestitae, sine pilis stellatis sed interiores ad apicem
pilorum brevium pallidorum caespite definitissimo ornatae; squamae interiores 10-17 mm longae,
1-1—-1-8 mm latae, late lineari-lanceolatae, ad apicem abrupte acutae saepe acuminatae interdum
subacutae aliquando convolutae, aliquantum appressae vel sublaxae; squamae exteriores paulo
breviores, interdum laxae, erectae, ad apicem subacutae. Ligulae pallide vel mediocriter flavae;
interiores densae; exteriores mediae vel latiusculae (ad 4-5 mm latae), ad apicem magis minusve
discretae et profunde dentatae dentibus magis minusve gracilibus, rectae vel haud profunde
concavae, in pagina inferiore pilis simplicibus eglanduliferis paucis vel multis brevissimis vel
brevibus pallidis (saepe aliquot aliquantum griseiusculis) vestitae vel nullis, ad apicem pilis talibus
aliquot vel multis perbrevissimis (aliquando cum paucis brevissimis vel brevibus) ultra dentium
margines protrudentibus. Styli flavi vel sordide viridiusculoflavi; stigmata flaviuscula. Receptaculi
alveoli margine breviter dentati. Cypselae 3-2-4-0 mm longae, purpureiusculonigrae. 2n = 36.

Cliff-ledges and rock-crevices on granite, quartz porphyry and quartz-biotite mica-schist from 740
to 1010 m: local and scarce in East Perth (v.c. 89), on Carn nan Sac and Glas Tulaichean near
Glenshee; in Angus (v.c. 90), above Loch Wharral, near Braedownie, at the head of Glen Fee and
in Coire Sharrock, all in Glen Clova, at the heads of Caenlochan and Canness Glens in Glen Isla,
and in Glen Prosen; in South Aberdeen (v.c. 92), not uncommon in the north-east coire of
Lochnagar and in several localities above Loch Kander in Glen Callater, but local and very scarce in
the Cairngorm Mountains in the Lairig Ghru Pass and Glen Geusachan at the southern and south-
eastern foot of Cairntoul and on Creag an Dail Bheag [Little Craigandail]. Endemic.

Named after Beverley Allan Miles (1937-1970) in recognition of his fine set of alpine hawkweeds
collected in the Scottish mountains.

Hieracium mundum P. D. Sell & C. West, sp. nov.
Ho.otypus: On flat rocks at c. 580 m, south-east of Loch a’ Choire Ghranda, Beinn Dearg, East
Ross, v.c. 106, GR NH/273.893, 27 July 1967, R. W. Jones & B. A. Miles no. 67/86 (CGE).

Ab Hieracio leptodonti P. D. Sell & D. J. Tennant (quod facie simulat) facie ubique generaliter
minus dense pilosa, foliis minus pruinosis, involucri squamis minus acutis pilisque glanduliferis
aliquot manifestis vestitis, ligulis intensius flavis; ab H. perscito P. D. Sell & C. West foliis basalibus
angustioribus coriaceioribus ellipticis, exterioribus ad basin minus late cuneatis, folio caulino infimo
dentibus longioribus spinulatis ornato, involucri squamis pilis glanduliferis paucis manifestis
vestitis, pilis brevissimis aliquot vel multis ad ligularum apices protrudentibus; ab H. subgloboso
P. D. Sell & C. West foliis basalibus angustioribus, folio caulino infimo majore latiore spinulatoser-
rato, involucri squamis obtusioribus, ligulis intensius flavis pilosioribusque; ab H. eximio Backh.
planta tota valde rubriusculoviolaceotincta, foliis coriaceioribus minus dense pilosis, folio caulino
infimo spinulatoserrato, involucri squamis obtusioribus pilisque glanduliferis manifestis vestitis,
ligulis intensius flavis distinguitur.

Planta phyllopoda. Caulis 7-24(—30) cm altus, graciliusculus vel subrobustus, saepe flexuosus, ad

NEW BRITISH SPECIES OF HIERACIUM 359

basin valde rubriusculoviolaceotinctus, ubique pilis simplicibus eglanduliferis paucis vel numerosis
brevibus mediocribusque longisque gracilibus sinuosis albis vel aliquantum griseiusculis ad bases
incrassatas nigriusculis (in parte inferiore saepe densis longissimis sinuosissimis interdum retrorsis,
in parte superiore minus densis mediocribus vel longis patentibus), in parte inferiore pilis
glanduliferis paucissimis brevissimis obscuris, in parte superiore numerosis vel densis, ubique pilis
stellatis numerosis, in parte superiore saepe densis. Folia aliquantum triste mediocriter vel
intensiuscule viridia, in pagina superiore hebetata vel paulo modo nitida, in pagina inferiore
pallidiora plerumque valde rubriusculoviolaceotincta vel maculata, praecipue in costa marginibus-
que dentibusque; folia basalia pauca vel numerosa, exteriora 6-55 mm longa, 4-23 mm lata,
pleraque subrotunda vel elliptica, ad apicem rotundo-obtusa, plerumque denticulata, interdum
dentata, ad basin cuneata, interiora 20-80 mm longa, 6-20 mm lata, rigidiuscula et subcoriacea,
saepe subcanaliculata, aliquantum undulata, lanceolata vel oblanceolata vel anguste elliptica, ad
apicem magis minusve obtusa (vel intima acuta), serratodentata vel incisa, dentibus magis minusve
regularibus vel irregularibus angustis saepe apiculatis, ad basin attenuata, omnia folia basalia in
pagina superiore pilis simplicibus eglanduliferis numerosis mediocribus vel longis gracilibus
subrigidis ad bases bulbosas pallidis vel discoloribus (in foliis exterioribus plerumque sparsis, in
interioribus magis minusve densis) uniformiter dispersis vel versus et ad margines restrictis, in
pagina inferiore pilis talibus paucis vel numerosis magis minusve uniformiter distributis brevibus vel
mediocribus saepe nigricantibasibus, in utraque pagina et in marginibus pilis glanduliferis aliquot
perbrevissimis flaviusculis et saepe pilis stellatis paucis vestita; petioli plerumque breviusculi, raro
longi, rigidi, magis minusve alati, rubriusculoviolaceotincti; folia caulina (2—)3—6(—7), infimum 10—
75 mm longum, 1—14 mm latum, basalibus interioribus saepe simile (in plantis robustis saepe usque
ad 4 folia grandia, fere ad basin caulis sita), patens, saepe undulatum, lineare vel lineari-
lanceolatum, ad apicem acutum vel subacutum, serratum vel ordinate spinulatoserratum, dentibus
longis angustis rubriusculoviolaceotinctis, rarissime integrum, ad basin attenuatum, in pagina
inferiore marginibusque pilis simplicibus eglanduliferis densis mediocribus vel longis gracilibus
sinuosis nigricantibasibus, in pagina superiore plerumque nullis, in utraque pagina pilis stellatis
aliquot (praecipue in costa) vestitum; petiolus alatus magis minusve amplexicaulis, pilis simplicibus
eglanduliferis (basin versus densissimis) vestitus; folia caulina superiora parva acuta vel magis
minusve bracteiformia. Capitulum plerumque solitarium, interdum ad 3, 25-40(-47) mm diametro;
involucrum facie obscurissime viridiusculogrisea vel nigriusculogrisea, campanulatum, ad basin
rotundatum; pedunculi (si adsunt) cauli superiori similiter vestiti. Involucri squamae ante anthesin
incumbentes, atriusculovirides vel rubrotinctae vel brunneotinctae; omnes pilis simplicibus eglan-
duliferis numerosis brevissimis vel brevibus vel longis (ad 2:5 mm) gracilibus sinuosis albis vel
aliquantum griseiusculis nigricantibasibus, pilis glanduliferis numerosis brevissimis vel brevibus (ad
0-8 mm) obscuris, a pilis simplicibus eglanduliferis partim modo tectis, vestitae, sine pilis stellatis
sed interiores ad apicem pilorum brevium caespite manifesto ornatae; squamae interiores 11-15 mm
longae, 1-0-1-5 mm latae, magis minusve appressae, lineari-lanceolatae, ad apicem pleraeque
subacutae vel nonnullae obtusoapiculatae, intimae abrupte acutae; squamae exteriores multo
breviores, erectae, ad apicem obtusae. Ligulae uniformiter mediocriter vel intensiuscule flavae;
exteriores latiusculae (ad 4-3 mm), ad apicem discretae et aliquantum vel profunde dentatae, haud
profunde concavae, in pagina inferiore pilis simplicibus eglanduliferis paucissimis vel numerosis
brevissimis vel brevibus pallidis vel aliquantum discoloribus vestitae, ad apicem pilis talibus aliquot
vel multis brevissimis ultra dentium margines protrudentibus. Styli magis minusve uniformiter
obscure viridiusculogrisei. Receptaculi alveoli margine breviter dentati. Cypselae 3—4 longae,
purpureiusculonigrae. 2n = 36.

Rocky knolls, detritus and crevices in shelving rocks, less frequently cliff-ledges, on fine-grained
quartz-biotite and other rock, between 425 and 760 m: on Fannich Forest hills, Braemore, and
Beinn Dearg Range in West and East Ross (v.cc. 105 and 106); on Carn Dearg in East Sutherland
(v.c. 107); on Foinaven, on Ben Hope and near Gien Coul in West Sutherland (v.c. 108). Endemic.

Hieracium optimum P. D. Sell & C. West. sp. nov.
Ho .ortypus: Beinn a’ Chreachain, Argyll, v.c. 98, 9 July 1976, A. G. Kenneth no. 1276 (CGE).

Ab Hieracio grovesii Pugsley foliis basalibus minus dentatis et in pagina superiore pilis stellatis
numerosissimis (sed in pagina inferiore aliquantum paucioribus) vestitis, folio caulino infimo saepe

360 P. D. SELL, C. WEST AND D. J. TENNANT

majore, involucro obscuriore, stylis obscuris differt. Ab omnibus aliis Sectionis Alpinorum
speciebus britannicis pilis stellatis densissimis in utraque foliorum basalium pagina distinguitur.

Planta phyllopoda. Caulis 15—30 cm altus, striatus, ad basin saepe rubriusculoviolaceus, in parte
inferiore pilis simplicibus eglanduliferis sparsis brevibus vel longis albiusculis nigricantibasibus, pilis
glanduliferis sparsis brevissimis flaviusculis, pilis stellatis paucis, in parte superiore pilis simplicibus
eglanduliferis numerosis brevibus vel longis aliquantum griseiusculis nigricantibasibus, pilis
glanduliferis numerosis brevissimis et aliquot brevibus flaviusculis et obscuris, pilis stellatis
numerosis vestitus. Folia tristiuscule mediocriter vel intensiuscule viridia, saepe partim triste
rubriusculoviolaceotincta vel suffusa; folia basalia exteriora 5-60 mm longa, 5-2-5 mm lata,
subrotunda vel late ovata, ad apicem obtusomucronata, integra vel denticulata vel dentibus paucis
haud profundis instructa, ad basin magis minusve cuneata, interiora 25—70 mm longa, 5—25 mm lata,
plerumque late vel anguste elliptica, raro oblanceolata vel obovata, ad apicem obtusomucronata vel
subacuta, integra vel denticulata vel dentibus paucis haud profundis acutis instructa, ad basin
attenuata, in utraque pagina et in marginibus pilis simplicibus eglanduliferis brevibus vel
mediocribus albis pallidibasibus vel nigricantibasibus (in paginis sparsis, in marginibus densioribus),
pilis glanduliferis paucis vel multis brevissimis flaviusculis, pilis stellatis numerosissimis vel densis
vestita; petioli interdum longi, pilis simplicibus eglanduliferis numerosis mediocribus vel longis albis
vestiti; folia caulina 1—2(-3), infimum 10-70 mm longum, 4-10(—20) mm latum, oblanceolatum vel
lanceolatum, ad apicem obtusum vel acutum, integrum, ad basin attenuatum, pilis simplicibus
eglanduliferis pilisque glanduliferis brevissimis nonnullis, pilis stellatis numerosis (praecipue in
pagina inferiore) vestitum, petiolo brevi instructum, cetera magis minusve bracteiformia et similiter
vestita, unum saepe gemma rudimentali in axilla praeditum. Capitulum plerumque solitarium,
interdum 2, circa 30-50 mm diametro; involucrum facie atriusculoviridi, campanulatum, ad basin
late rotundatum, supra basin constrictum; pedunculi (si adsunt) cauli superiori similiter vestiti.
Involucri squamae ante anthesin incumbentes, atriusculovirides; omnes pilis simplicibus eglanduli-
feris densis longis (ad 5 mm) sinuosis albis vel griseiusculis nigricantibasibus, pilis glanduliferis
numerosis brevibus nigriusculistipitatis flavicapitatis, a pilis simplicibus eglanduliferis magis
minusve omnino tectis, vestitae, sine pilis stellatis sed interiores ad apicem pilis ramosis brevibus
nonnullis ornatae; squamae interiores 15-17 mm longae, 1-5—2-0 mm latae, appressae, latiuscule
lineari-lanceolatae, ad apicem acutissimum magis minusve gradatim protractae; squamae exteriores
breviores. Ligulae clare mediocriter vel intensiuscule flavae; exteriores ad apicem magis minusve
discretae et profunde irregulariterque dentatae, concavae, in pagina inferiore pilis simplicibus
eglanduliferis paucis vel multis brevissimis vel brevibus pallidis vestitae, ad apicem pilis aliquot
brevibus ultra dentium margines protrudentibus. Styli obscuri. Receptaculi alveoli margine breviter
dentati. Cypselae 3-5—4-0 mm longae, purpureiusculonigrae.

Rock-ledges on Dalradian mica-schist from 700 to 820 m, on Meall Buidhe and Beinn a’
Chreachain in Argyll (v.c. 98). Endemic.

Hieracium pensum P. D. Seli & C. West, sp. nov.
Ho otypus: East-facing cliffs of Coire Ghranda, Beinn Dearg, East Ross, v.c. 106, GR NH/
267.806, 27 July 1967, R. W. Jones & B. A. Miles no. 67/84 (CGE).

Ab Hieracio mundo P. D. Sell & C. West foliis minus rigidis minus coriaceis minus pilosis, capitulo
saepe majore pilisque glanduliferis manifestioribus ornato; ab H. perscito P. D. Sell & C. West
capitulo multo majore pilisque glanduliferis manifestis multo paucioribus ornato; ab H. subgloboso
P. D. Sell & C. West statura robustiore, involucro majore; ab H. kennethii P. D. Sell & D. J.
Tennant foliorum pagina superiore glabra differt. A ceteris Sectionis Alpinorum speciebus
britannicis aut distributione geographica aut stylis obscuris distinguitur.

Planta phyllopoda. Caulis 10-30 cm altus, mediocris vel robustus, saepe flexuosus, ad basin
interdum rubriusculopurpureotinctus; in parte inferiore pilis simplicibus eglanduliferis numerosis
vel aliquantum densis mediocribus vel longis gracilibus sinuosis albis nigricantibasibus, pilis
glanduliferis paucis brevissimis plerumque pallidis, pilis stellatis aliquantum sparsis vestitus; in
parte superiore pilis simplicibus eglanduliferis aliquot vel multis longis gracilibus griseiusculis
nigricantibasibus, pilis glanduliferis numerosis brevissimis vel brevibus obscuris, pilis stellatis densis
vestitus. Folia pallidiuscule vel mediocriter viridia, pruinam dilute caesiam ferentia, hebetata vel

»

NEW BRITISH SPECIES OF HIERACIUM 361

paulo modo nitida, venis conspicuis ornata, in marginibus et ad apicem interdum leviter
rubriusculoviolaceotincta; folia basalia satis numerosa, exteriora 10-50 mm longa, 8-27 mm lata,
subrotunda vel obovata vel elliptica, ad apicem rotundo-obtusa, acute denticulata vel dentibus
paucis haud profundis vel anguste mammiformibus instructa, ad basin cuneata, interiora 20-80 mm
long, 8-30 mm lata, oblanceolata vel obovata vel elliptica raro ovata, non coriacea, plana vel paulo
canaliculata, raro leviter undulata, ad apicem interdum tortum obtusa perspicueque mucronata vel
subacuta cuspidataque, serrata, dentibus irregularibus patentibus vel retrorsis anguste mammifor-
mibus interdum cuspidatis (aliquando in petiolum descendentibus), ad basin cuneata vel attenuata,
omnia in marginibus pilis simplicibus eglanduliferis numerosis vel densis mediocribus vel longis vel
longissimis gracilibus subrigidis sinuosis pallidibasibus vestita, pleraque in pagina superiore sine
pilis simplicibus eglanduliferis praeter intima (quae interdum uniformiter vestiuntur), in pagina
inferiore pilis simplicibus eglanduliferis brevibus vel longis sinuosissimis interdum nigricantibasibus
modice et uniformiter vel in costa densius vestita, in pagina inferiore marginibusque pilis
glanduliferis aliquot minutis flaviusculis vestita; petioli mediocres vel sublongi graciles alati,
interdum dentibus paucis instructi, plerumque pilis simplicibus eglanduliferis densis longis sinuosis
patentibus vel retrorsis vestiti; folia caulina (2—)3—5(—7), infimum 10—55(—110) mm longum, 2-20
mm latum, interdum prope caulis basin situm, magis minusve patens, interdum undulatum,
oblanceolatum vel anguste ellipticum, ad apicem anguste acutum, saepe dentibus parvis angustis
nonnullis instructum vel denticulatum, ad basin attenuatum saepe petiolatum, in marginibus pilis
simplicibus eglanduliferis densis longis nigricantibasibus (ad basin densissimis longissimis sinuosis),
in pagina superiore pilis talibus paucis vel nullis, in pagina inferiore pilis talibus dispersis, in costa
marginibusque pilis stellatis paucis vel nullis vestitum, cetera folia caulina anguste elliptica vel
linearia integra, vel summum bracteiforme, unum interdum gemma rudimentali in axilla praeditum.
Capitulum plerumque solitarium, aliquando 2 vel plura vel cum altero a secundo caule e basi
exoriente portato, 35-58 mm diametro; involucrum facie obscure viridiusculogrisea vel viridusculo-
nigra, campanulatum, ad basin late rotundatum; pedunculi (si adsunt) cauli superiori similiter
vestiti. Involucri squamae ante anthesin incumbentes, viridiusculoatrae; omnes pilis simplicibus
eglanduliferis densis longis (ad 3-5 mm) albis vel aliquantum griseiusculis nigricantibasibus, pilis
glanduliferis numerosis brevissimis vel brevibus et paucis mediocribus (ad 1-0 mm) obscuris, a pilis
simplicibus eglanduliferis partim modo tectis, vestitae, sine pilis stellatis, interiores ad apicem
pilorum brevium caespite ornatae; squamae interiores 12-20 mm longae, 1-1-1-8 mm latae,
aliquantum laxae paulo curvatae lineari-lanceolatae, ad apicem plerumque acutissimae; squamae
exteriores laxae erectae aliquantum breviores, ad apicem abrupte acutae. Ligulae pallidiuscule vel
mediocriter flavae; exteriores latae (ad 4-7 mm), ad apicem semidiscretae et profunde et saepe
irregulariter dentatae, fere rectae, in pagina inferiore pilis simplicibus eglanduliferis paucis
brevissimis pallidis vestitae vel nullis, ad apicem pilis talibus paucis vel aliquot brevissimis (et
interdum paucis brevibus) ultra dentium margines protrudentibus. Styli uniformiter obscure
viridiusculogrisei. Receptaculi alveoli margine breviter dentati. Cypselae 3-5—4-0 mm longae,
purpureiusculonigrae. 2n = 36.

Cliff-ledges and large rocks, infrequently rock detritus, on quartz-biotite mica-schist and
weathered microgranite between 520 and 915 m: on Beinn Dearg, Seana Bhraigh and Fannich
Forest and Braemore Forest hills in West and East Ross (v.cc. 105 and 106). Endemic.

Hieracium perscitum P. D. Sell & C. West, sp. nov.
Ho .otypus: Rocky knoll, c. 530 m, An Teallach, West Ross, v.c. 105, GR NH/082.864, 29 July
1978, A. G. Kenneth & P. D. Sell no. 78/337 (CGE).

Hieracio hanburyi Pugsley simillimum, a quo foliis basalibus ovatis vel lanceolatis plerumque
dentibus minoribus regularioribus ornatis, folio caulino infimo patente, involucris minoribus
angustioribusque et pilis eglanduliferis multo longioribus quam pilis glanduliferis vestitis, stylis
valde obscurioribus distinguitur.

Planta phyllopoda. Caulis 10-27 cm, plerumque gracilis, saepe flexuosus, infra pallide violaceo-
tinctus, basin versus pilis simplicibus eglanduliferis plus minusve numerosis vel densis mediocribus
et longis vel longissimis gracilibus sinuosis albis plerumque nigricantibasibus, in parte superiore pilis
talibus dispersis vel sparsis brevibus vel longis gracillimis sinuosissimis, in parte inferiore pilis

362 P. D. SELL, C. WEST AND D. J. TENNANT

glanduliferis dispersis brevissimis pallidis, in parte superiore pilis talibus numerosioribus brevissimis
vel brevibus obscuris, in parte inferiore pilis stellatis dispersis, in parte superiore pilis talibus
numerosioribus vestitus. Folia pallidiuscule vel clariuscule mediocriter viridia vel intensius triste
glauciusculoviridia, dilute nitida vel hebetata, in utraque pagina et in marginibus plerumque
sordidoviolaceotincta vel notata (vel in pagina inferiore pallidiora); folia basalia pauca vel aliquot,
exteriora 540 mm longa, 8-25 mm lata, subrotunda vel late ovata vel late elliptica, ad apicem late
rotundo-obtusa, integra vel denticulata vel haud profunde sinuatodentata, ad basin cuneata vel
rotundata, interiora 15-50 mm longa, 6-20 mm lata, interdum leviter undulata, subrigida sed non
(vel leviter modo) coriacea, ovata vel ovatolanceolata vel elliptica, ad apicem rotundo-obtusa vel
subacuta vel acuta saepe apiculata, remote denticulata vel irregulariter serratodentata, dentibus
angustis vel latis saepe ascendentibus rubriusculoviolaceis mammiformibus, ad basin cuneata vel
aliquando attenuata, folia basalia omnia pilis simplicibus eglanduliferis dispersis vel satis densis
brevibus et mediocribus et aliquando longis gracilibus mollibus vel subrigidis sinuosis albis ad bases
bulbosas pallidis vel obscuris (in foliorum interiorum pagina superiore saepe uniformiter distributis,
in marginibus densis sinuosissimisque, in pagina inferiore aliquantum paucioribus sed in costa saepe
densis; in foliis exterioribus sparsius distributis), in utraque pagina et in marginibus pilis
glanduliferis paucis perbrevissimis flaviusculis vestita; petioli plerumque breviusculi, graciles, ad
basin saturate rubriusculoviolaceotincti, pilis simplicibus eglanduliferis numerosis vel densis longis
vestiti; folia caulina (0—)1-3(-5), infimum 10-50(—70) mm longum, 2-15(—20) mm latum, rigide
patens, saepe undulatum, ad trientem vel dimidium inter caulis basin et apicem positum, plerumque
lanceolatum, aliquando anguste ovatum, saepe folia basalia intima simulans, ad apicem gradatim
acutum vel acuminatum vel magis minusve cuspidatum, denticulatum vel irregulariter serratum,
dentibus paucis acutis vel anguste mammiformibus, ad basin saepe sessilem aliquando petiolatam
semiamplexicaulem cuneatum vel attenuatum, in pagina superiore pilis simplicibus eglanduliferis
sparsis interdum nigricantibasibus, in pagina inferiore marginibusque pilis talibus densioribus, in
pagina inferiore pilis stellatis nonnullis vestitum, folium a basi secundum semipatens lineare vel
lineari-lanceolatum integrum sessile, vel bracteiforme, in pagina superiore glabrum, in pagina
inferiore pilis stellatis densis vestitum, cetera folia caulina (si adsunt) perminuta filamentosa acuta
appressa vel semipatentia, unum interdum gemma rudimentali in axilla praeditum. Capitula 1-2,
23-42 mm diametro; involucrum facie viridiusculonigra vel nigriuscula, anguste campanulatum, ad
basin rotundatum vel subtruncatum; pedunculi (si adsunt) cauli superiori similiter vestiti. Involucri
squamae ante anthesin incumbentes, atriusculovirides, interiores marginibus aliquantum pallidiori-
bus: omnes pilis simplicibus eglanduliferis numerosis mediocribus vel longis (ad 2 mm) gracilibus
aliquantum griseiusculis nigricantibasibus, pilis glanduliferis numerosis brevissimis brevibusque
obscuris (quorum pauci a pilis simplicibus eglanduliferis teguntur), pilis stellatis paucis vel nullis
vestitae sed interiores ad apicem pilorum brevium caespite ornatae; squamae interiores 10-13 mm
longae, 1-2-1-7 mm latae, arcte appressae, latiuscule lineari-lanceolatae, ad apicem pleraeque
obtusae nonnullae subacutae; squamae exteriores breviores, magis minusve appressae. Ligulae
clare mediocriter flavae; exteriores ad apicem magis minusve discretae et profunde dentatae vel
laciniatodentatae, rectae vel aliquantum concavae, in pagina inferiore pilis simplicibus eglandulifer-
is paucis vel nullis pallidis vestitae, ad apicem pilis talibus brevissimis paucis vel nullis ultra dentium
margines protrudentibus. Sty/i obscure viridiusculogrisei. Receptaculi alveoli margine interdum
fimbriatodentati. Cypselae 3:9-4-7 mm longae, purpureiusculonigrae. 2n = 36.

Rocky knolls, from 530 to 760 m, on Torridonian Sandstone and Moine Schist: An Teallach
Range, Ben More Coigach, Braemore Forest and Fannich Forest hills in West and East Ross (v.cc.
105 and 106). Endemic.

Hieracium probum P. D. Sell & C. West, sp. nov.
Ho torypus: Cliffs at north-west side of Coire Ardair, near Loch Laggan, Westerness, v.c. 97, GR
NN/430.884, 25 July 1966, R. W. Jones & B. A. Miles no. 66/68 (CGE).

A speciebus omnibus britannicis Hieracii Sectionis Alpinorum stylis obscuris ornatis (praeter
quasdam geographice disjunctas) caulibus foliisque suis sparsius pilosis, involucri squamis minus
acutis minusque sericeis, ligulis saturate flavis distinguitur.

Planta phyllopoda. Caulis 13-35 cm altus, singularis vel interdum plures, gracilis vel robustus,

.

NEW BRITISH SPECIES OF HIERACIUM 363

saepe flexuosus, valde striatus, in parte inferiore pilis simplicibus eglanduliferis pilisque glandulifer-
is pilisque stellatis sparsissimis, in parte superiore pilis simplicibus eglanduliferis paucis vel multis
brevibus vel longis gracilibus sinuosis albiusculis ad bases longas aliquantum tumidas nigris, pilis
glanduliferis plus minusve numerosis brevissimis vel brevibus (ad 0-5 mm) nigriusculis, pilis stellatis
sparsis vel densis vestitus. Folia pallidiuscule vel saturate tristiuscule viridia, subnitida vel hebetata,
interdum subcoriacea, raro rubriusculoviolaceomaculata; folia basalia numerosa (8-14), exteriora
10-30 mm longa, 7—25 mm lata, interdum undulata, obovata vel subrotunda vel elliptica, ad apicem
rotundo-obtusa interdum apiculata raro retusa, integra vel denticulata, ad basin attenuata vel
cuneata vel subtruncata, interiora 35-70 mm longa, 13-25 mm lata, undulata, obovata vel
oblanceolata vel elliptica, ad apicem saepe obtusa interdum subacuta plerumque apiculata
leviterque torta, denticulata vel incisodentata vel serratodentata, dentibus irregularibus late vel
anguste mammiformibus vel aquilinomammiformibus, ad basin cuneata vel attenuata, omnia in
pagina superiore (vel modo ad vel prope margines) pilis simplicibus eglanduliferis paucis vel multis
mediocribus vel longis (ad 2 mm) subrigidis sinuosis albis ad bases bulbosas pallidiusculis, in pagina
inferiore pilis talibus paucis vel multis brevioribus magis minusve uniformiter distributis, in utraque
pagina et in marginibus pilis glanduliferis paucis brevissimis gracilibus flaviusculis vestita; petioli
aliquantum graciles, longi (ad 7 cm), late alati, interdum dentibus paucis instructi, ad basin
rubriusculoviolacei, in marginibus pilis simplicibus eglanduliferis paucis vel multis mediocribus vel
longiusculis interdum appressis vestiti vel fere glabri; folia caulina 1-3(—5), infimum 10-60 mm
longum, 2-12 mm latum, erectum appressumque vel patens, interdum undulatum, anguste
ellipticum vel lineari-oblanceolatum, ad apicem abupte acutum vel obtusum saepe tortum,
subintegrum vel dentatum dentibus paucis, ad basin breviter cuneatum vel attenuatum interdum
magis minusve petiolatum saepe alatum, in pagina superiore (saepe modo margines versus) pilis
simplicibus eglanduliferis multis mediocribus pallidis, in pagina inferiore interdum pilis stellatis
paucis vestitum, cetera folia caulina linearia abrupte acuta vel bracteiformia vel filamentosa erecta.
Capitulum plerumque solitarium, interdum 2(-5), circa 35-45 mm diametro; involucrum facie
obscure viridiusculogrisea vel viridiusculonigra, campanulatum vel late campanulatum; pedunculi
(si adsunt) cauli superiori similiter vestiti. Involucri squamae ante anthesin incumbentes, atriusculo-
virides, interiores marginibus indistinctis pallidioribus; omnes pilis simplicibus eglanduliferis
mediocribus vel longis (ad 3 mm) gracilibus pallidis vel aliquantum griseiusculis ad bases longas
incrassatas nigris, pilis glanduliferis aliquot vel numerosis manifestis brevissimis vel brevibus (ad 0-4
mm) obscuris vestitae, sine pilis stellatis sed interiores ad apicem pilorum ramulosorum simplicium-
que caespite definito ornatae; squamae interiores 15-17 mm longae, 1-0—1-7 mm latae, inaequales
appressae linearilanceolatae, ad apicem interdum incurvatum plerumque abrupte acutae vel
subacutae interdum obtusae vel acuminatae vel mucronatae, intimae saepe angustae acutae;
Squamae exteriores breviores appressae vel semilaxae. Ligulae saturate tristiuscule flavae; inter-
iores densae; exteriores ad 4-2 mm latae, ad apicem discretae vel valde discretae, aliquantum
concavae vel ad apicem profunde dentatum perspicue concavae, raro non evolutae (ad apicem
convexe involutae), in paginae inferioris parte superiore pilis simplicibus eglanduliferis paucis
pallidis vestitae vel nullis, pilis talibus paucis perbrevissimis vel brevibus ultra dentium margines
protrudentibus. Sty/i obscure viridiusculogrisei; stigmata aliquantum pallidiora. Receptaculi alveoli
margine breviter dentati. Cypselae 3-2-4-2 mm longae, purpureiusculonigrae. 2n = 36.

Rocks and ledges between 550 and 915 m: on hard mica-schist on Creagan Liatha, Stob Binnein
and An Caisteal in West Perth (v.c. 87), on Cruach Ardrain in (?) Mid Perth (v.c. 88), and in Moy
Corrie and Coire Ardair, both on Creag Meagaidh, and in the adjacent Coire nan Gall, Westerness
(v.c. 97); on granitic rock on Aonach Mor and Aonach Beg, Westerness (v.c. 97); on Bidean nam
Bian, Glen Coe, in Main Argyll (v.c. 98). Endemic.

Hieracium subglobosum P. D. Sell & C. West, sp. nov.
Ho otypus: Cliff-ledges, c. 640 m, south of lochan in north-east corrie of Ben Hope, West
Sutherland, v.c. 108, GR NC/484.503, 22 July 1966, R. W. Jones & B. A. Miles no. 66/40 (CGE).

Ab Hieracio globosifloro Pugsley (quod est facie simillimum) statura saepe minore, foliis clarius
viridibus, ligulis minus densis, stylis obscuris differt.
Planta phyllopoda. Caulis 6-20(—30) cm altus, gracilis vel mediocris, saepe flexuosus, ad basin

364 P. D. SELL, C. WEST AND D. J. TENNANT

rubriusculoviolaceus, ubique pilis simplicibus eglanduliferis paucis vel multis brevibus mediocribus-
que longisque gracilibus sinuosis albis vel griseiusculis ad bases incrassatas nigriusculis, in parte
inferiore pilis talibus dispersis pallidis interdum retrorsis, in parte superiore pilis talibus aliquot vel
multis mediocribus vel longis (ad 4 mm) patentibus saepe griseiusculis ad bases longas nigris, in
parte inferiore pilis glanduliferis dispersis brevissimis flaviusculis, in parte superiore pilis talibus
numerosis vel densis brevibus mediocribusque (ad 1-0 mm) nigriusculis, in parte inferiore pilis
stellatis dispersis, in parte superiore pilis talibus numerosis vel densis vestitus. Folia mediocriter
saepe clare viridia, in pagina superiore nitida, in pagina inferiore pallidiora, saepe triste
rubriusculoviolaceotincta (praecipue in pagina dentium marginumque superiore et in costa subtus);
folia basalia aliquantum pauca vel numerosa, saepe parva, in foliorum satis regularium rosula,
exteriora 3-50 mm longa, 3-25 mm lata, subrotunda vel ovata vel obovata, ad apicem late rotundo-
obtusa, integra vel denticulata, ad basin cuneata, interiora 15-90 mm longa, 2-30 mm lata,
aliquantum tenuia vel coriacea, aliquando canaliculata, anguste vel late elliptica vel obovato-
lanceolata, ad apicem acuta vel obtusa saepe apiculata, subintegra vel denticulata interdum
undulata vel irregulariter serratodentata vel incisa, dentibus aliquando magnis irregularissimis
patentibus vel ascendentibus anguste mammiformibus vel aquilinomammiformibus, ad basin
cuneata vel breviter attenuata, folia interiora in pagina superiore plerumque fere glabra sed versus
et ad margines pilis simplicibus eglanduliferis aliquot vel numerosis mediocribus vel longis subrigidis
albis ad bases bulbosas saepe discoloribus vel nigris (in foliis intimis pilis longioribus densioribus-
que), folia omnia in pagina inferiore pilis talibus dispersis mollioribus (in costa densis), in utraque
pagina pilis glanduliferis paucis brevissimis flaviusculis (in marginibus nonnullis) pilisque ramosis
paucissimis brevissimis vestita; petioli breves vel longi, graciles, saepe basin versus rubriusculovio-
laceotincti, in marginibus interdum dentibus paucis pilisque simplicibus eglanduliferis dispersis vel
densis (praecipue basin versus) mediocribus vel longis subrigidis interdum sinuosis ornati; folia
caulina (0—)1—3(—5), infimum. 6-60 mm longum, 1—7 mm latum, semipatens vel erectum, lineari-
ellipticum, ad apicem attenuatoacutum vel rarissime obtusum, plerumque integrum raro remote
denticulatum vel dentibus paucis acutis ornatum, ad basin attenuatum, in pagina superiore glabrum,
in marginibus et pagina inferiore pilis simplicibus eglanduliferis mediocribus subrigidis saepe
nigricantibasibus (basin versus longis saepe densis) pilisque stellatis dispersis vestitum, cetera folia
caulina parva acuta vel filamentosa vel bracteiformia appressa. Capitulum solitarium, saepe
parvum, raro 2, 25-40 mm diametro; involucrum facie nigriuscula vel raro obscure viridiusculogri-
sea, campanulatum vel anguste campanulatum, in pedunculum decrescens, sed ad basin rotunda-
tum, supra basin constrictum; pedunculi (si adsunt) cauli superiori similiter vestiti. /nvolucri
squamae ante anthesin incumbentes, atriusculovirides, interiorum nonnullae marginibus inconspi-
cuis viridiusculis; omnes pilis simplicibus eglanduliferis aliquot vel densis mediocribus vel longiuscu-
lis (ad 4 mm) patentibus sinuosis albis vel frequentius griseiusculis ad bases longissimas incrassatas
nigris, pilis glanduliferis numerosis brevissimis pallidis et brevibus nigriusculis, a pilis simplicibus
eglanduliferis partim modo tectis, sine pilis stellatis sed ad apicem pilis ramosis nonnullis brevibus
vestitae; squamae interiores 9-16 mm longae, 0-9-1-7 mm latae, inaequales, saepe curvatae, raro
numerosissimae, satis arcte appressae, lineari-lanceolatae, intimae angustissimae, ad apicem
acutissimum vel acuminatum attenuatae, raro subacutae vel obtusae; squamae exteriores brevius-
culae, erectae, ad apicem acutae. Ligulae clare pallide flavae, raro mediocriter flavae; exteriores ad
apicem saepe valde discretae et profunde interdumque irregulariter dentatae, rectae vel haud
profunde concavae, sine pilis vel interdum pilis simplicibus eglanduliferis paucissimis brevibus
pallidis in pagina inferiore vestitae et aliquando pilis talibus paucis brevissimis magis minusve
griseiusculis ultra dentium margines protrudentibus. Sty/i magis minusve uniformiter obscurissime
grisei vel mediocriter viridiusculogrisei. Receptaculi alveoli margine breviter dentati. Cypselae 3-5—
4-2 mm longae, purpureiusculonigrae. 2n = 27.

Mainly on gentle rocky slopes or rocky knolls, occasionally on rock-ledges, on Moine and other
schists, quartzite, microgranite and Torridonian Sandstone, usually between 550 and 850 m, but
descending to 350 m just south of Cape Wrath. Widespread in the north-western Highlands, but
often local in its individual localities: near Loch Cluanie and in Glen Affric and Glen Cannich near
the western border of Easterness (v.c. 96), and on several hills in the northern part of Westerness
(v.c. 97); on numerous hills in West Ross (v.c. 105) and several in East Ross (v.c. 106) including
Seana Bhraigh and Beinn Dearg and in Glen Carron (Achnasheen); on hills south and east of Loch
More in East Sutherland (v.c. 107) and’‘many.hills throughout West Sutherland (v.c. 108), Endemic.

NEW BRITISH SPECIES OF HIERACIUM 365
ACKNOWLEDGMENTS

We are greatly indebted to Philip Oswald not only for correcting the Latin grammar but also for
making the descriptions more lucid. We thank Clive Stace for allowing us to include the
chromosome numbers counted at the School of Biological Sciences of the University of Leicester.

REFERENCES

Pucs.ey, H. W. (1948). A prodromus of the British Hieracia. Journal of the Linnean Society of London
(Botany) 54: 1-356.

Stace, C. A., GORNALL, R. J., SQUIRRELL, J. & SHI, Y. (1995). Chromosome numbers in Hieracium L. section
Alpina (Fries) F. N. Williams. Watsonia 20: 367-377.

(Accepted March 1995)

Watsonia 20: 367-377 (1995) 367

Chromosome numbers in Hieracium L. section Alpina (Fries)
F. N. Williams

C. A. STACE, R. J. GORNALL, J. SQUIRRELL and Y. SHI
Department of Botany, University of Leicester, Leicester, LEI 7RH

ABSTRACT

Cytological study of 34 of the 35 named British taxa of Hieracium section Alpina, plus two un-named taxa,
showed that 24 are tetraploids (2n = 36), six are triploids (2n = 27), one is a pentaploid (2n = 45), one (H.
macrocarpum) is a tetraploid or hypertetraploid (2n = 36 or 37), and four occur as both triploids and tetraploids.
Of the latter four, H. tenuifrons is represented by a western tetraploid, a central triploid and an eastern
tetraploid race; H. insigne f. celsum is represented by a triploid western and a tetraploid eastern race (but only
one plant of the former was counted); and material of the other two taxa was insufficient to draw any firm
conclusions. The pentaploid taxon (un-named, from An Teallach, W. Ross, v.c. 105) appears to be the first
count for any taxon of Hieracium sensu stricto above the level of tetraploid.

KEYworDs: apomixis, hawkweeds, polyploidy, Britain.

INTRODUCTION

TAXONOMY

The genus Hieracium L. (Asteraceae) is well known as one in which apomixis is widespread, giving
rise to a very large number of variants that have been described as species and to a lesser extent at
other ranks. In the older literature the genus was divided into subgenus Hieracium and subgenus
Pilosella (Hill) Gray, but most modern works now treat the latter as the separate genus Pilosella
Hill. This separation is based on a range of morphological, biochemical, cytological and genetical
characteristics. Nevertheless, in most of the literature the generic name Hieracium remains
ambiguous; it might or might not include Pilosella, or indeed concern Pilosella alone. An idea of the
number of taxa in these two genera is provided by the fact that under Hieracium and Pilosella there
are 11,827 entries in Index Kewensis (CD-ROM version, 1993).

British hieraciologists have traditionally divided Hieracium sensu stricto into up to 16 sections.
The latest work (Kent 1992) recognized twelve, which follows the views of Sell & West (1968).
Section Alpina (Fries) F. N. Williams is relatively easily recognized by its short scapose stems
bearing usually only a single terminal capitulum. The species in Britain mostly occur in rock crevices
and on rock ledges and other rather bare rocky places at over 600 m altitude. Sell & West (1976), in
their account of Hieracium (incl. Pilosella) for Flora Europaea, did not divide the genus into
sections, but instead recognized 260 main taxa at the level of species or species groups. Section
Alpina in the sense of British authors corresponds with Sell & West’s ‘H. alpinum group’ and ‘H.
nigrescens group’ (nos. 158 and 159).

Kent (1992) listed 19 microspecies within section Alpina, to which must be added the eleven new
species and five extra formae described by Sell et al. (1995). There still remain about half a dozen
other variants in Scotland that merit names (D. J. Tennant, pers. comm., 1994). The 35 named taxa |
are distributed as follows: 28 are endemic to Scotland; H. subgracilentipes is endemic to the English
Lake District; five species (H. alpinum, H. calenduliflorum, H. eximium, H. globosiflorum and H.
tenuifrons) occur in Scotland and European mountains, the first-named extending to Greenland;
and H. holosericeum occurs in Scotland, the English Lake District, Snowdonia and the European
mountains.

368 C. A. STACE, R. J. GORNALL, J. SQUIRRELL AND Y2SHI

TABLE 1. SUMMARY OF CHROMOSOME COUNTS OF SPECIES OF
HIERACIUM AND PILOSELLA

Ploidy Level No. of Taxa
Diploid (2n = 2x = 18) 83
Triploid (2n = 3x = 27) 110
Tetraploid (2n = 4x = 36) 66
Pentaploid (2n = 5x = 45) 21

Hexaploid (2n = 6x = 54) 6
Heptaploid (2n = 7x = 63) 4
Octoploid (2n = 8x = 72) 1
1
2

Nonaploid (2n = 9x = 81)
Decaploid (2n = 10x = 90)

See text for sources. Aneuploid counts have been rounded off to the nearest
euploid level.

CHROMOSOME NUMBERS

We have carried out a literature survey of the chromosome numbers of the genera Hieracium and
Pilosella reported in the standard abstracting references, covering the years up to 1989, to which we
have added relevant data in The cytological catalogue of the British and Irish flora (Bailey & Gornall
ined.). The results are summarized in Table 1. The total number of taxa cannot be obtained by
summing the totals, because some occur at more than one ploidy level. Species of both Hieracium
and Pilosella contribute to the diploid, triploid and tetraploid totals, but we have been unable to
separate the totals for these two genera. However, in the pentaploid and higher levels all reports but
one refer to the genus Pilosella. The single exception is actually somewhat spurious, since it refers to
the presence of 54 chromosomes in certain pollen mother cells of a plant of H. umbellatum with 2n =
27 (Rosenberg 1927b). It is thus the case to date that all known chromosome numbers of Hieracium
sensu stricto are diploid, triploid or tetraploid (with some aneuploids). As far as we are aware, the
diploids represent sexual and the triploids and tetraploids apomictic plants (Rosenberg 1927a).

There have been relatively few chromosome studies of British or Irish Hieracium sensu stricto.
Mills & Stace (1974) reported counts for 21 and Morton (1974) for twelve taxa, only one being
common to the two lists. We know of British counts for only two other species (Bailey & Gornall
ined.). These 34 taxa cover eleven of the twelve sections of the genus. Eleven of the species are
tetraploid and 21 triploid; the other two taxa (H. umbellatum L. subsp. umbellatum and subsp.
bichlorophyllum (Druce & Zahn) Sell & C. West) exist as both diploid and triploid cytotypes.
Hence in the British Isles only the two subspecies of H. umbellatum are known to be sexual.

In section Alpina there have hitherto been only two British counts. Mills & Stace (1974) found H.
hanburyi to be tetraploid (2n = 36) and Morton (1974) reported the same for H. calendulifiorum.
Elsewhere there are counts for H. alpinum (2n = 27 for material from Greenland, Iceland,
Fennoscandia, Komi ASSR, Siberian Arctic, Poland, Slovakia and Switzerland) and for the non-
British H. nigrescens Willd. (2n = 27 from Komi ASSR).

As part of a molecular genetic investigation of section Alpina, we have made a detailed
cytological investigation of our material. Here we report the chromosome numbers of 152 plants (34
of the 35 named taxa, plus two un-named ones and four ‘intermediates’) and discuss their
significance.

MATERIALS AND METHODS

Mr D. J. Tennant has made a detailed study of Hieracium section Alpina over the past 25 years.
Uniquely, he has seen all the taxa in their native situation, and has grown them all in his Yorkshire
garden. Most of his collection (c. 150 pots), including all but two of the 35 taxa, was transferred to
Leicester in 1993 and 1994 for safe-keeping, propagation and study. This has been supplemented by
some of our own collections, plants having been grown from wild-collected seed, and by F, plants

CHROMOSOME NUMBERS IN HIERACIUM L. SECTION ALPINA (FRIES) F. N. WILLIAMS 369

grown from seed produced by Mr Tennant’s plants. The collection now contains all the 35 named
taxa except H. optimum Sell & C. West, as well as six other problematical un-named taxa (Table 2).
All determinations have been made by Mr D. J. Tennant. At present the vouchers are the living
plants, which are too valuable to press. However, F, plants are being grown as vouchers.

Plants are kept in an unheated greenhouse, being grown in 12-5 cm pots containing equal parts of
potting compost and terrace gravel (derived from granite) collected in the Cairngorms, Scotland.
Seed is germinated on 1% agar after two weeks in tap-water at 4°C. Plants are transferred to pots in
compost as soon as they are large enough to move from the agar, and potted on into the compost-
gravel mixture when they are fully established.

Cytological studies were made on the pot-grown plants. Actively growing roots were placed into a
pretreatment solution of 0.002M 8-hydroxyquinoline and kept at 4°C for 24 hours. Subsequently the
pre-treatment solution was replaced by fixative (3 parts absolute ethyl alcohol: 1 part glacial acetic
acid). Roots were hydrolysed for 10 minutes at room temperature in 5M HCl, which was then
removed and replaced by 70% IMS (industrial methylated spirit).

Dissection of the root was carried out in a drop of 40% acetic acid on a microscope slide. The
removed root tip was then transferred into a drop of 2% certified aceto-orcein. Epidermal tissue was
removed and discarded thus revealing the meristematic regions, which were then macerated and
finally squashed.

The resultant slides were viewed under a high power microscope for suitable, countable
metaphases. Chromosome counts were made from at least three root tips for each accession.

RESULTS

The chromosome counts made for 152 plants are listed in Table 2. These cover 34 named taxa, plus:
1. Plants apparently intermediate between H. eximium and H. calenduliflorum, now thought by Mr
Tennant to be a variant of H. leptodon (EXI/CLD-1);

2. Plants apparently intermediate between H. eximium and H. memorabile (EXI/MEM-51, 52 &
53).

. Plants apparently intermediate between H. hanburyi and H. tenuifrons (HAN/TNF-1 & 2);

. Plants apparently intermediate between H. tenuifrons and H. memorabile (TNF/MEM-51);

An un-named, probably new, taxon from Ben Dearg, E. Ross, v.c. 106 (XYZ-2); and

An un-named, probably new, taxon from An Teallach, W. Ross, v.c. 105 (XYZ-1).

The first five of these are tetraploids (2n = 36), the last a pentaploid (2n = 45).

Of the 34 named taxa, six are triploids (2n = 27), 24 are tetraploids (2n = 36), and the other four

gave both triploid and tetraploid counts. Two plants of H. macrocarpum (MAC-24 & 25) were

found to be hypertetraploids (2n = 4x + 1 = 37); otherwise all the plants were euploids.

Nn &

DISCUSSION

The marked prevalence of tetraploid over triploid counts for British species of section Alpina
(28:10) is in strong contrast to the previously published figures for Hieracium (including some
Pilosella species) overall world-wide (66:110, see Table 1) or for Hieracium sensu stricto in Britain
alone (13:23). This is particularly surprising since all the previous counts (c. 12) for non-British
plants of section Alpina (involving two species, H. alpinum and H. nigrescens) were triploid. Our
counts of British H. alpinum are also triploid (Fig. 1a).

The existence of a pentaploid (X YZ-1 from An Teallach, W. Ross) is unique (Fig. 1d); we have
been able to discover no previous counts for plants of Hieracium sensu stricto above the level of
tetraploid.

There are many reports of aneuploid counts in the literature. In Table 1 these figures were ~
subsumed into the nearest euploid level. Without detailed population and karyotypic studies it is
difficult to be sure of the significance of aneuploids. They could refer to odd individual plants, odd
cells of euploid individuals, or simply to errors. On the other hand they might represent whole
‘populations’ of plants, or whole taxa. Examples of all of these situations are well known in
apomicts. For example, most taxa of the Limonium binervosum (G. E. Sm.) Salmon aggregate are

370 C. A. STACE, R. J. GORNALL, J. SQUIRRELL AND Y. SHI
TABLE 2. CHROMOSOME NUMBERS IN HIERACIUM SECTION ALPINA

H. alpinum L.

ALP-1. V.c. 92, S. Aberdeen: Glen Derry, NO/0.9. 2n = 27.

ALP-22. V.c. 92, S. Aberdeen: Glen Derry, NO/0.9. 2n = 27.
ALP-23. V.c. 88, Mid Perth: Ben More, Crianlarich, NN/4.2. 2n = 27.

H. backhousei F. Hanb.
BAC-1. V.c. 90, Angus: Glen South Esk, above Glen Clova, NO/2.7. 2n = 36.
BAC-4. V.c. 92, S. Aberdeen: Allt an Dubh Loch, above Glen Muick, Ballater, NO/2.8. 2n = 36.

H. calendulifiorum J. Backh.

CLD-1. V.c. 97, Westerness: Ben na’ Socaich, Glen Spean, NN/2.7. 2n = 36.

CLD-2. V.c. 92, S. Aberdeen: Lochnagar, above Glen Muick, Ballater, NO/2.8. 2n = 36.

CLD-8. V.c. 92, S. Aberdeen: Allt an Dubh Loch, above Glen Muick, Ballater, NO/2.8. 2n = 36.
CLD-13. V.c. 92, S. Aberdeen: Allt an Dubh Loch, above Glen Muick, Ballater, NO/2.8. 2n = 36.
CLD-30. V.c. 92, S. Aberdeen: Allt an Dubh Loch, above Glen Muick, Ballater, NO/2.8. 2n = 36.
CLD-48. V.c. 92, S. Aberdeen: Allt an Dubh Loch, above Glen Muick, Ballater, NO/2.8. 2n = 36.
CLD-65. V.c. 92, S. Aberdeen: Allt an Dubh Loch, above Glen Muick, Ballater, NO/2.8. 2n = 36.
CLD-86. V.c. 92, S. Aberdeen: Allt an Dubh Loch, above Glen Muick, Ballater, NO/2.8. 2n = 36.
CLD-150. V.c. 97, Westerness: Ben na’ Socaich, Glen Spean, NN/2.7. 2n = 36.

CLD-151. V.c. 90, Angus: Craig of Gowal, head of Glen South Esk, above Glen Clova, NO/2.8. 2n = 36.
CLD-152. V.c. 97, Westerness: Ben na’ Socaich, Glen Spean, NN/2.7. 2n = 36.

CLD-153. V.c. 90, Angus: Bachnagairn, head of Glen South Esk, above Glen Clova, NO/2.7. 2n = 36.
CLD-154. V.c. 90, Angus: Loch Brandy, Glen Clova, NO/3.7. 2n = 36.

CLD-155. V.c. 97, Westerness: Ben na’ Socaich, Glen Spean, NN/2.7. 2n = 36.

CLD-158. V.c. 97, Westerness: Aonach Mor, Ben Nevis range, above Glen Spean, NN/1.7. 2n = 36.

H. calvum Sell & D. Tenn.
CLV-1. V.c. 92, S. Aberdeen: Glen Derry, NO/0.9. 2n = 36.
CLV-2. V.c. 92, S. Aberdeen: Glen Derry, NO/0.9. 2n = 36.

CLV-3. V.c. 92, S. Aberdeen: Falls, Coire an Lochain Uaine, Derry Cairngorm, Glen Derry, NO/0.9. 2n = 36.

CLV-4. V.c. 92, S. Aberdeen: Glen Derry, NO/0.9. 2n = 36.

CLV-5. V.c. 92, S. Aberdeen: Glen Derry, NO/0.9. 2n = 36.

CLV-6. V.c. 92, S. Aberdeen: Glen Derry, NO/0.9. 2n = 36.

CLV-7. V.c. 96, Easterness: East side of Cairn Gorm, NJ/0.0. 2n = 36.
CLV-8. V.c. 92, S. Aberdeen: Glen Derry, NO/0.9. 2n = 36.

CLV-9. V.c. 96, Easterness: East side of Cairn Gorm, NJ/0.0. 2n = 36.
CLV-10. V.c. 96, Easterness: East side of Cairn Gorm, NJ/0.0. 2n = 36.
CLV-11. V.c. 96, Easterness: East side of Cairn Gorm, NJ/0.0. 2n = 36.
CLV-12. V.c. 96, Easterness: East side of Cairn Gorm, NJ/0.0. 2n = 36.

H. completum Sell & C. West

COM-1. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 36.
COM-25. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 36.
COM-3S. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 36.
COM-36. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 36.
COM-37. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 36.

H. eximium J. Backh.

EXI-24. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 36.

EXI-54. V.c. 92, S. Aberdeen: Creag an Dail Bheag, Cairngorms, NO/1.9. 2n = 36.
EXI-55. V.c. 90, Angus: Glen Doll, Glen Clova, NO/2.7. 2n = 36. An F, plant.
EXI-56. V.c. 90, Angus: Glen Doll, Glen Clova, NO/2.7. 2n = 36. An F, plant.

H. eximium f. tenellum (J. Backh.) Sell & C. West
TNE-1. V.c. 92, S. Aberdeen: Creag an Dail Bheag, Cairngorms, NO/1.9. 2n = 36. Three F, plants.
TNE-2. V.c. 92, S. Aberdeen: Creag an Dail Bheag, Cairngorms, NO/1.9. 2n = 36.

H. eximium/H. calenduliflorum intermediate
EXI/CLD-1. V.c. 98, Main Argyll: Allt nan Giubhas, Meall a’ Bhuiridh, Glencoe, NN/2.5. 2n = 36.

CHROMOSOME NUMBERS IN HIERACIUM L. SECTION ALPINA (FRIES) F. N. WILLIAMS 371

TABLE 2. continued

H. eximium/H. memorabile intermediate

EXI/MEM-S51. V.c. 92,S. Aberdeen: Coire an Lochain Uaine, Derry Cairngorm, Glen Derry, NO/0.9. 2n = 36.
EXI/MEM-S2. V.c. 92, S. Aberdeen: Coire an Lochain Uaine, Derry Cairngorm, Glen Derry, NO/0.9. 2n = 36.
EXI/MEM-SS3. V.c. 92, S. Aberdeen: Coire an Lochain Uaine, Derry Cairngorm, Glen Derry, NO/0.9. 2n = 36.

H. globosiflorum Pugsley
GLO-1. V.c. 92, S. Aberdeen: Coire an Lochain Uaine, Cairn Toul, Cairngorms, NN/9.9. 2n = 27.
GLO-19. V.c. 92, S. Aberdeen: Creag an Dail Bheag, Cairngorms, NO/1.9. 2n = 27.

H. graniticola W. R. Linton

GRA-1. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 27. Also an F, plant.
GRA-3. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 27.

GRA-4. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 27.

GRA-5. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 27.

H. grovesii Pugsley
GRO-1. V.c. 92, S. Aberdeen: Beinn a’ Bhuird, Cairngorms, NO/0.9. 2n = 36. Also an F, plant.

H. hanburyi Pugsley

HAN-1. V.c. 88, Mid Perth: Ben More, Crianlarich, NN/4.2. 2n = 36.
HAN-23. V.c. 90, Angus: Head of Glen Fee, Glen Clova, NO/2.7. 2n = 36.
HAN-26. V.c. 90, Angus: Head of Glen Fee, Glen Clova, NO/2.7. 2n = 36.

H. hanburyi f. atraticeps (Pugsley) Sell & D. Tenn.

ATR-1. V.c. 90, Angus: Head of Glen Fee, Glen Clova, NO/2.7. 2n = 36. Also three F, plants.
ATR-2. V.c. 90, Angus: Above lowest falls, Glen Fee, Glen Clova, NO/2.7. 2n = 36.

ATR-3. V.c. 90, Angus: Head of Glen Fee, Glen Clova, NO/2.7. 2n = 36.

ATR-4. V.c. 90, Angus: Glen Fee falls, Glen Clova, NO/2.7. 2n = 36.

ATR-5. V.c. 90, Angus: Head of Glen Fee, Glen Clova, NO/2.7. 2n = 36.

ATR-6. V.c. 90, Angus: Head of Glen Fee, Glen Clova, NO/2.7. 2n = 36.

ATR-7. V.c. 90, Angus: Head of Glen Fee, Glen Clova, NO/2.7. 2n = 36.

ATR-8. V.c. 90, Angus: Upper falls, Glen Fee, Glen Clova, NO/2.7. 2n = 36.

H. hanburyi f. pusillum Sell & D. Tenn.

PUS-2. V.c. 90, Angus: Canness Glen, Glen Isla, NO/2.7. 2n = 36.
PUS-24. V.c. 90, Angus: Canness Glen, Glen Isla, NO/2.7. 2n = 36.
PUS-25. V.c. 90, Angus: Boustie Ley, Glen Clova, NO/3.7. 2n = 36.

H. hanburyi/H. tenuifrons intermediate
HAN/TNF-1. V.c. 96, Easterness: Glen Einich, Cairngorms, NN/9.9. 2n = 36.
HAN/TNF-2. V.c. 96, Easterness: Glen Einich, Cairngorms, NN/9.9. 2n = 36.

H. holosericeum J. Backh.

HOL-13. V.c. 92, S. Aberdeen: Allt an Dubh Loch, above Glen Muick, Ballater, NO/2.8. 2n = 27.
HOL-33. V.c. 97, Westerness: Aonach Mor, Ben Nevis range, above Glen Spean, NN/1.7. 2n = 27.
HOL-36. V.c. 96, Easterness: Bynack More, Strath Nethy, Cairngorms, NJ/0.0. 2n = 36.

HOL-47. V.c. 49, Caernarvonshire: Summit of Craig yr Ysfa, south-east of Carnedd Llywelyn, SH/3.6. 2n = 27.

H. insigne J. Backh.

INS-1. V.c. 92, S. Aberdeen: Coire Kander, head of Glen Callater, NO/1.8. 2n = 36.
INS-3. V.c. 92, S. Aberdeen: Coire Kander, head of Glen Callater, NO/1.8. 2n = 36.
INS-4. V.c. 92, S. Aberdeen: Coire Kander, head of Glen Callater, NO/1.8. 2n = 36.

H. insigne f. celsum Sell & D. Tenn.

CEL-1. V.c. 96, Easterness: Creag an Lethchoin, Lairig Ghru, NH/9.0. 2n = 36.

CEL-2. V.c. 97, Westerness: Ben na’ Socaich, Glen Spean, NN/2.7. 2n = 27.

CEL-3. V.c. 92, S. Aberdeen: Coire an Lochain Uaine, Cairn Toul, Cairngorms, NN/9.9. 2n = 36.
CEL-4. V.c. 96, Easterness: Creag an Lethchoin, Lairig Ghru, NH/9.0. 2n = 36.

CEL-5. V.c. 96, Easterness: Coire Garbhlach, Glen Feshie, Cairngorms, NN/8.9. 2n = 36.

a2 C. A. STACE, R. J. GORNALL, J. SQUIRRELL AND Y. SHI
TABLE 2. continued

H. kennethii Sell & D. Tenn.

KEN-1. V.c. 105, W. Ross: An Teailach, Dundonnell, NH/0.8. 2n = 36.
KEN-3. V.c. 105, W. Ross: An Teallach, Dundonnell, NH/0.8. 2n = 36.
KEN-4. V.c. 105, W. Ross: An Reallachs Dundonnell, NH/0.8. 2n = 36.

H. larigense (Pugsley) Sell & C. West
LAR-1. V.c. 96, Easterness: Creag an Lethchoin, Lairig Ghru, NH/9.0. 2n = 36. An F, plant.
LAR-2. V.c. 96, Easterness: Creag an Lethchoin, Lairig Ghru, NH/9.0. 2n = 27.

H. leptodon Sell & D. Tenn.

TND-1. V.c. 88, Mid Perth: Ben More, Crianlarich, NN/4.2. 2n = 36. An F, plant.
TND-2. V.c. 88, Mid Perth: Ben More, Crianlarich, NN/4.2. 2n = 36. Also an F, plant.
TND-3. V.c. 88, Mid Perth: Ben More, Crianlarich, NN/4.2. 2n = 36.

TND-S. V.c. 88, Mid Perth: Ben More, Crianlarich, NN/4.2. 2n = 36.

H. macrocarpum Pugsley

MAC-1. V.c. 92, S. Aberdeen: Creag an Dail Bheag, Cairngorms, NO/1.9. 2n = 36. Six F, plants.
MAC-19. V.c. 106, E. Ross: Carn Liath, Strath Carron, NH/1.5. 2n = 36.

MAC-21. V.c. 92, S. Aberdeen: Coire an t-Saighdeir, Cairn Toul, Cairngorms, NN/9.9. 2n = 36.
MAC-22. V.c. 96, Easterness: Coire Chuirn, south of Dalwhinnie, NN/6.7. 2n = 36.

MAC-23. V.c. 96, Easterness: Coire Chuirn, south of Dalwhinnie, NN/6.7. 2n = 36.

MAC-?24. V.c. 96, Easterness: Coire Chuirn, south of Dalwhinnie, NN/6.7. 2n = 37.

MAC-25. V.c. 96, Easterness: Coire Chuirn, south of Dalwhinnie, NN/6.7. 2n = 37.

MAC-27. V.c. 96, Easterness: Outflow shingle, N. end of Loch Einich, NN/9.9. 2n = 36.

H. marginatum Pugsley

MAR-2. V.c. 106, E. Ross: Coire Ghranda, Ben Dearg, NH/2.8. 2n = 36. Also an F, plant.
MAR-4. V.c. 105, W. Ross: Meall Doire Faid, Braemore, NH/2.7. 2n = 36.

MAR-S. V.c. 106, E. Ross: Coire Ghranda, Ben Dearg, NH/2.8. 2n = 36.

H. marginatum f. chaetocephalum Sell & C. West
CHA-1. V.c. 97, Westerness: Coire Ardair, Craig Meagaidh, Loch Laggan, NN/4.8. 2n = 36. Also three F,
plants.

H. memorabile Sell & C. West

MEM-6. V.c. 97, Westerness: Aonach Mor, Ben Nevis range, above Glen Spean, NN/1.7. 2n = 36. An F, plant.
MEM-10. V.c. 92, S. Aberdeen: Head of Glen Callater, NJ/1.7. 2n-= 36.

MEM-11. V.c. 96, Easterness: Coire Garbhlach, Glen Feshie, Cairngorms, NN/8.9. 2n = 36.

MEM-13. V.c. 92, S. Aberdeen: Head of Glen Callater, NO/1.8. 2n = 36.

MEM-14. V.c. 96, Easterness: Coire Garbhlach, Glen Feshie, Cairngorms, NN/8.9. 2n = 36.

MEM-15. V.c. 96, Easterness: Creag an Lethchoin, Lairig Ghru, NH/9.0. 2n = 36.

H. milesii Sell & C. West

MIL-1. V.c. 90, Angus: Canness Glen, Glen Isla, NO/2.7. 2n = 36.

MIL-2. V.c. 90, Angus: Caenlochan, NJ/1.7. 2n = 36.

MIL-3. V.c. 92, S. Aberdeen: Loch Kander, head of Glen Callater, NO/1.8. 2n = 36.
MIL-S. V.c. 90, Angus: Caenlochan, NJ/1.7. 2n = 36.

MIL-6. V.c. 90, Angus: Caenlochan, NJ/1.7. 2n = 36.

H. mundum Sell & C. West
MUN-1. V.c. 106, E. Ross: Coire Ghranda, Ben Dearg, NH/2.8. 2n = 36.

H. notabile Sell & C. West
NOT-1. V.c. 97, Westerness: Coire Ardair, Craig Meagaidh, Loch Laggan, NN/4.8. 2n = 36. Three F; plants.
NOT-3. V.c. 97, Westerness: Coire Ardair, Craig Meagaidh, Loch Laggan, NN/4.8. 2n = 36.

H. pensum Sell & C. West
PEN-1. V.c. 106, E. Ross: Coire Ghranda, Ben Dearg, NH/2.8. 2n = 36. Also two F, plants.

CHROMOSOME NUMBERS IN HIERACIUM L. SECTION ALPINA (FRIES) F. N. WILLIAMS 373
TABLE 2. continued

H. perscitum Sell & C. West

PER-1. V.c. 105, W. Ross: An Teallach, Dundonnell, NH/0.8. 2n = 36. Also an F, plant.
PER-2. V.c. 105, W. Ross: An Teallach, Dundonnell, NH/0.8. 2n = 36.

PER-S. V.c. 105, W. Ross: An Teallach, Dundonnell, NH/0.8. 2n = 36.

PER-7. V.c. 105, W. Ross: An Teallach, Dundonnel!, NH/0.8. 2n = 36.

H. probum Sell & C. West

PRO-2. V.c. 97, Westerness: Aonach Mor, Ben Nevis range, above Glen Spean, NN/1.7. 2n = 36.
PRO-24. V.c. 87, W. Perth: Stob Binnein, Crianlarich, NN/4.2. 2n = 36.

PRO-25. V.c. 87, W. Perth: Stob Binnein, Crianlarich, NN/4.2. 2n = 36.

H. pseudocurvatum (Zahn) Pugsley

PCU-1. V.c. 92, S. Aberdeen: Creag an Dail Bheag, Cairngorms, NO/1.9. 2n = 27.
PCU-2. V.c. 92, S. Aberdeen: Creag an Dail Bheag, Cairngorms, NO/1.9. 2n = 27.
PCU-3. V.c. 92, S. Aberdeen: Creag an Dail Bheag, Cairngorms, NO/1.9. 2n = 27.

H. pseudopetiolatum (Zahn) Roffey

PPE-1. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 27.

PPE-17. V.c. 97, Westerness: Coire Ardair, Craig Meagaidh, Loch Laggan, NN/4.8. 2n = 27.
PPE-18. V.c. 97, Westerness: Coire Ardair, Craig Meagaidh, Loch Laggan, NN/4.8. 2n = 27.

H. subglobosum Sell & C. West
SGL-1. V.c. 106, E. Ross: Coire Ghranda, Ben Dearg, NH/2.8. 2n = 27. Also two F, plants.
SGL-2. V.c. 106, E. Ross: Coire Ghranda, Ben Dearg, NH/2.8. 2n = 27.

H. subgracilentipes (Zahn) Roffey

SGR-1. V.c. 69, Westmorland: Dollywaggon, Helvellyn, NY/3.1. 2n = 36.

SGR-8. V.c. 69, Westmorland: Rampsgill Head, above Haweswater, NY/4.1. 2n = 36.
SGR-11. V.c. 69, Westmorland: Dollywaggon, Helvellyn, NY/3.1. 2n = 36.

H. tenuifrons Sell & C. West

TNF-1. V.c. 98, Main Argyll: Allt nan Giubhas, Meall a’ Bhuiridh, Glencoe, NN/2.5. 2n = 36.
TNF-2. V.c. 88, Mid Perth: Creag Roro, Ben Lawers, Glen Lyon, NN/6.4. 2n = 27. Also an F, plant.
TNF-4. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 36.

TNF-44. V.c. 97, Westerness: Coire Ardair, Craig Meagaidh, Loch Laggan, NN/4.8. 2n = 27.
TNF-45. V.c. 97, Westerness: Coire Ardair, Craig Meagaidh, Loch Laggan, NN/4.8. 2n = 27.
TNF-46. V.c. 88, Mid Perth: Creag Roro, Ben Lawers, Glen Lyon, NN/6.4. 2n = 27.

TNF-47. V.c. 97, Westerness: Coire Ardair, Craig Meagaidh, Loch Laggan, NN/4.8. 2n = 27.
TNF-48. V.c. 98, Main Argyll: Allt nan Giubhas, Meall a’ Bhuiridh, Glencoe, NN/2.5. 2n = 36.
TNF-49. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 36.

TNF-50. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 36.

TNF-52. V.c. 88, Mid Perth: Ben Heasgarnich, Glen Lyon, NN/4.3. 2n = 36. Two F, plants.
TNF-53. V.c. 92, S. Aberdeen: Coire Etchachan, NO/0.9. 2n = 36.

TNF-54. V.c. 97, Westerness: Coire Ardair, Craig Meagaidh, Loch Laggan, NN/4.8. 2n = 27.

I

H. tenuifrons/H. memorabile intermediate
TNF/MEM-51. V.c. 96, Easterness: Coire Garbhlach, Glen Feshie, Cairngorms, NN/8.9. 2n = 36.

Un-named new taxa
XYZ-1. V.c. 105, W. Ross: An Teallach, Dundonnell, NN/O.8. 2n = 45.
XYZ-2. V.c. 106, E. Ross: Summit coire, Ben Dearg, NH/2.8. 2n = 36.

hypotetraploids (2n = 4x — 1 = 35) (Ingrouille & Stace 1986). One previous count of H. alpinum
from the Siberian Arctic (Sokolovskaya & Strelkova 1960) is a hypotriploid (2n = 3x — 1 = 26), but
must be viewed in the light of the above comments. In the present study two plants of H.
macrocarpum (MAC-24 & 25) from Coire Chuirn, Easterness, were found to be hypertetraploids
(2n = 4x + 1 = 37) (Fig. Ic). Two other plants (MAC-22 & 23) from the same locality were
tetraploids (2n = 36).

374 C. A. STACE, R. J. GORNALL, J. SQUIRRELL AND Y. SHI

FicureE |. Root-tip squashes of (a) Hieracium alpinum (ALP-22) from Glen Derry, S. Aberdeen, 2n = 3x = 27;
(b) H. calenduliflorum (CLD-152) from Ben na’ Socaich, Westerness, 2n = 4x = 36; (c) H. macrocarpum
(MAC-24) from Coire Chuirn, Easterness, 2n = 4x + 1 = 37; and (d) Un-named new taxon (XYZ-1) from An
Teallach, W. Ross, 2n = 5x = 45. All x 1250.

The caveats concerning the interpretation of aneuploid counts mentioned above must be repeated
when considering taxa in which two ploidy levels have been found. However, in such a highly critical
group as Hieracium sect. Alpina, there are two taxonomic problems as well. Firstly, it is possible
that in fact the two cytotypes should be recognized as different taxa, not one. Secondly, pots might
have become mislabelled due to clerical or horticultural errors or due to misidentification. The last is
areal possibility even when identification has been made by the acknowledged experts in the group,
since several of the microspecies are very difficult to distinguish if one or two characters (e.g. style
colour) are missing. In section Alpina this problem is exacerbated because frequently several
(sometimes more than ten) species grow in one locality, and in any one locality only a small
proportion of the plants flower in any given year.

Apart from the case of H. macrocarpum mentioned above, four taxa were found with two

chromosome numbers:
a. H. tenuifrons. D. J. Tennant has for long recognized three non-overlapping regional races within
this species: a western race found in Westerness (v.c. 97) , Main Argyll (v.c. 98) and just into Mid
Perth (v.c. 88); a central race found in W., Mid and E. Perth (v.cc. 87-89) and just into Westerness;
and an eastern race found in the Cairngorms (S. Aberdeen and Easterness: v.cc. 92, 96). The
chromosome numbers of these three races are 2n = 36 (three plants), 27 (six plants) and 36 (four
plants) respectively. In terms of gross morphology, the western race (which includes the type) is far
more distinct from the other two than the latter two are from each other (D. J. Tennant, pers.
comm., 1994). Although the cytological data do not fully support the morphological data, the two
together do provide evidence that the three races of H. tenuifrons are distinct entities. Whether they
are worth recognizing taxonomically, and if so at what ranks, are moot points.

Chromosome numbers throw no light on the identity of the H. hanburyi/H. tenuifrons

CHROMOSOME NUMBERS IN HIERACIUM L. SECTION ALPINA (FRIES) F. N. WILLIAMS 375

‘intermediate’ (HAN/TNF-1 & 2), since it, H. hanburyi and the eastern race of H. tenuifrons are all
tetraploids. The same is true of the H. tenuifrons/H. memorabile ‘intermediate’ (TNF/MEM-S1).
b. H. insigne f. celsum. The single triploid (CEL-2) comes from the western area (Glen Spean,
Westerness, v.c. 97), while the four tetraploids are from the eastern area (S. Aberdeen, v.c. 92 and
Easterness, v.c. 96). These five plants come from four localities which represent much of the total
range of this rare taxon. The different chromosome number of the western plant clearly requires
further investigation.

c. H. larigense. Both plants are from the area of the type locality (Lairig Ghru, Easterness, v.c. 96)
and the existence of two ploidy levels in H. /arigense from this area requires confirmation. A
misidentification is one possible explanation, which will be investigated the next time the plants
flower at Leicester and by molecular studies.

d. H. holosericeum. Although this is the most widespread British species, and is often common in
Scottish localities, it is one of the most difficult to keep healthy in cultivation. The single tetraploid
and two of the triploid plants come from three rather widely separated Scottish localities. The single
Welsh count was also triploid. Clarification of the cytological pattern in this species must await more
counts from the same and other localities in Scotland as well as from the Lake District, but clearly
the Easterness plant might be misidentified. This will be checked by molecular studies.

Comments on some of the other taxa are also desirable.

H. calenduliflorum is a somewhat variable species. D. J. Tennant recognises two races — a western
one from Westerness (v.c. 97) and Main Argyll (v.c. 98), which shows some resemblances to H.
notabile; and an eastern one from Angus (v.c. 90) and S. Aberdeen (v.c. 92). In addition to this,
CLD-151 is an unusually highly pubescent plant, and CLD-158 an unusually small, narrow-leaved
plant. All these plants are tetraploids (Fig. 1b), as was the previously counted plant from
Lochnagar, S. Aberdeen (Morton 1974). Preliminary results from molecular analyses of the two
races have, however, revealed some differences.

The H. eximium/H. calenduliflorum ‘intermediate’ plant (EXI/CLD-1) is, like the two species,
tetraploid, as is H. leptodon, to which D. J. Tennant now refers it. The same is true of the H.
eximium/H. memorabile ‘intermediate’ (EXI/MEM-S1, 52 & 53).

The three plants of H. eximium (EXI-51, 52 & 53) from Derry Cairngorm, S. Aberdeen, might
represent a distinct taxon, having more coriaceous leaves and being more obviously glandular (D. J.
Tennant, pers. comm., 1994), but all have the same chromosome number.

Our plants of H. hanburyi are tetraploids, as was the previously counted plant from River Eidart,
Easterness (Mills & Stace 1974). The H. hanburyi/H. tenuifrons ‘intermediate’ has been commented
upon already.

H. macrocarpum is virtually confined to the Cairngorms except for the distant locality in E. Ross,
where the plants (MAC-19) differ in some respects from those in the Cairngorms and might
represent a different taxon. The chromosome numbers, however, do not differ.

The two plants of H. pseudopetiolatum from Westerness differ slightly from that from Coire
Etchachan, but not in chromosome number.

Since there are in the main only two ploidy levels involved, it is not surprising that they throw
rather little ight on taxonomic problems. However, all the five formae newly recognized by Sell e¢
al. (1995) have the same chromosome number as the respective type formae, and none of the other
variants mentioned above differs in chromosome number from the respective typical species. This
indicates that there is in fact rather little (if any) variation in chromosome number within each
microspecies as delimited by traditional taxonomists. A corollary of this conclusion is, therefore,
that those cases where both triploid and tetraploid counts are found in one microspecies constitute
evidence that the two cytological races do, in fact, represent different taxa. This is clearly true of H.
tenuifrons and perhaps of H. celsum. Moreover, it suggests that the pentaploid un-named plant from
An Teallach (XY Z-1) is indeed a new species.

The use of chromosome numbers as taxonomic evidence in this way is justifiable in an obligately _
apomictic group, but much less so in a sexual or facultatively apomictic one. Without the existence
of sexual reproduction (i.e. meiosis followed by fertilization) the evolution of a triploid from a
tetraploid, or vice versa, although conceivable, is a most unlikely event. It would involve the chance
inclusion of 27 chromosomes in a cell destined to become an embryo from a plant with 36
chromosomes, or vice versa. If such an event is at all likely (i.e. if the requisite mechanisms exist), it
is surely unlikely that the only new chromosome numbers to arise would be euploids; one would

376 C. A. STACE, R. J. GORNALL, J. SQUIRRELL AND Y. SHI

expect aneuploids to be formed at least as frequently (perhaps much more frequently) than
euploids. There is no evidence that this occurs; only two of 152 plants so far counted are aneuploids,
and none of the F, progeny of these 152 plants differed from their parent in chromosome number. A
far more likely cause of the existence of both triploids and tetraploids (and higher euploid levels) is
their independent origins from sexual events. The only exceptions to this hypothesis would be
chromosome number doubling (e.g. triploids giving rise to hexaploids) or the occasional origin of
aneuploids, both of which are possible by unexceptional mitotic events.

The published lists used to compile Table 1 show several examples of species represented by more
than one ploidy level. Many of these, however, are members of the genus Pilosella, in which sexual
diploids, tetraploids and hexaploids exist and where mechanisms for the production of plants of
different ploidy level (up to decaploid) are well known (Gadella 1991). Of the species of Hieracium
sensu stricto in which two or more polyploid levels have been recorded, some (e.g. H. umbellatum L.)
exist as sexual diploids as well and others may well be cases of misidentification (e.g. H.
pulmonarioides Villars) or varied use of the same name (e.g. H. murorum L. and H. villosum Jacq.).

Sell & West (1976), in Flora Europaea, segregated British Hieracium into two species groups (H.
alpinum group and H. nigrescens group), giving chromosome numbers of 2n = 27 and 2n = 36
respectively. The former was based upon the [Continental] counts for H. alpinum and the latter
upon the British counts for H. hanburyi and H. calenduliflorum (Moore 1982). The only British
species included in the H. alpinum group were H. alpinum and H. holosericeum (both with triploid
counts in Britain), while the other 15 British species mentioned were all placed in the H. nigrescens
group. Besides the special case of H. tenuifrons four of these are triploids, and recent counts of H.
nigrescens from Komi ASSR are also triploid (Lavrenko et al. 1988). Furthermore, one of our three
counts of H. holosericeum is tetraploid. Hence there is no chromosome number difference between
the two species groups, whose descriptions in Flora Europaea differ only by few relatively trivial
features (viz. leaf width and three characters all prefixed by “‘often’”’). Moreover, although Pugsley’s
(1948) definition of the two species groups was very similar, he placed five of the species that Sell &
West included in the H. nigrescens group in the H. alpinum group (H. grovesii, H. eximium, H.
calenduliflorum, H. macrocarpum and H. graniticola), the first four of these five being tetraploids.

We are carrying out further chromosome studies of Hieracium section Alpina, including
karyotype analyses, in parallel with molecular (DNA and enzyme) methods.

ACKNOWLEDGMENTS

We are greatly indebted to Mr David Tennant for providing us with most of the material, for guiding
and tutoring two of us (Y.S. and C.A.S.) ona trip to the Scottish Highlands in 1993, and for sharing
much of his unparalleled knowledge with us. We are also grateful to Rebecca Smith, Jacqueline
Clayton, Peter Hollingsworth and Lisa Page for making some of the chromosome counts, to Dr
John Bailey for help with cytological techniques, and to Mr Jim Bevan for putting us in touch with
Mr Tennant in the first place.

REFERENCES

BaiLey, J. P. & GorNALL, R. J. (ined.) Cytological catalogue of the British and Irish flora.

GADELLA, T. W. J. (1991). Variation, hybridization and reproductive biology of Hieracium pilosella L.
Proceedings, Koninklijke Nederlandse Akademie van Wetenschappen 94: 455-488.

INGROUILLE, M. J. & Stace, C. A. (1986). The Limonium binervosum aggregate (Plumbaginaceae) in the British
Isles. Botanical journal of the Linnean Society 92: 177-217.

KENT, D. H. (1992). List of vascular plants of the British Isles. B.S.B.1., London.

Lavrenko, A. N., Serpirov, N. P. & ULLE, Z. G. (1988). Chromosome numbers in some species of
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605-607.

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(Botany) 54: 1-356.

ROSENBERG, O. (1927a). Die semiheterotypische Teilung und ihre Bedeutung fiir die Entstehung verdoppelter
Chromosomenzahlen. Hereditas 8: 305-338.

ROSENBERG, O. (1927b). Homoeotypic division in uninucleate pollen mother cells. Hereditas 9: 285-288.

SELL, P. D. & West, C. (1968). Hieracium, in PERRING, F. H. & SELL, P. D., eds. Critical supplement to the Atlas
of the British flora, pp. 75-131. E. P. Publishing Ltd, London.

SELL, P. D. & WEsrt, C. (1976). Hieracium, in TutTin, T. G. et al., eds. Flora Europaea 4: 358-410. Cambridge
University Press, Cambridge.

SELL, P. D., West, C. & TENNANT, D. J. (1995). Eleven new British species of Hieracium L. section Alpina
(Fries) F. N. Williams. Watsonia 20: 351-365.

SOKOLOvVSKAYA, A. P. & StTRELKOVA, O. S. (1960). Geographical distribution of polyploid plant species in
Eurasian arctic. Botaniceskij Zurnal 45: 370-381.

(Accepted March 1995)

NOTE ADDED IN PROOF
Recent work on sect. Alpina by J. Chrtek jun. in eastern Europe (Folia geobotanica et phytotaxonomica
Bohemoslavaca, Praha 29: 91-100 (1994), and unpublished) has shown that:

1. Hieracium alpinum sensu stricto from Czechoslovakia, Poland and the Ukraine exists mostly as triploid (2n =
27) but also as diploid (2n = 18) cytotypes. This is the first diploid count for sect. Alpina.

2. All the other (non-British) taxa in sect. Alpina from these areas are triploids or tetraploids.

3. Continental material named H. calenduliflorum and H. eximium does not belong to these species but to non-
British taxa. It is possible that H. alpinim itself is in fact the only British species that is not endemic.

We are grateful to Dr Chrtek for these data.

Watsonia 20: 379-384 (1995) 379

Sorbus domestica L., new to Wales and the British Isles

M. HAMPTON
9 Adenfield Way, Rhoose, South Glamorgan, CF62 3EA
and
Q.O.N. KAY

School of Biological Sciences, University of Wales, Swansea, SA2 &PP

ABSTRACT

The discovery of two native or long-established populations of the mainly southern European tree Sorbus
domestica L. (Rosaceae) in sea-cliff sites on the coast of southern Wales is reported. The True Service-tree had
not been previously known as a native species or persistent introduction in Britain, with the exception of a single
tree that formerly grew in the Wyre Forest in central England. Identification characters, including previously
undescribed epidermal features, isozyme variation, and its ecology and associates at the two sites are described.
Its occurrence and possible history in Britain are briefly discussed.

KeEyworbs: Sorbus aucuparia, isozyme analysis, ecology, distribution, disjunction.

INTRODUCTION

In May 1983 Marc Hampton observed a number of wind-stunted trees of a Sorbus L. species
(Rosaceae) with pinnate leaves, resembling Sorbus aucuparia L. but apparently distinct from that
species, growing on inaccessible ledges on a south-facing limestone sea-cliff in southern Glamorgan,
Wales (v.c. 41). The site is in an area which had been well studied botanically and it seemed unlikely
that any long-established but undiscovered species of tree could occur there. During subsequent
brief visits to the site he observed several distinctive features of the trees, all consistent with the
possibility that they might be S. domestica L., notably their strikingly silvery young leaves in early
spring and, in some years, their display of flowers 16-18 mm in diameter, larger and more
conspicuous than in S. aucuparia and with pink-flushed petals. In addition, the branches of the
inflorescence, especially the final pedicels, were comparatively lax and straight rather than curved as
in S. aucuparia, forming a more or less pyramidal inflorescence which differed in shape from the
flat-topped corymbs of S. aucuparia, and the inflorescences enlarged simultaneously with the leaves,
not after them as in S. aucuparia. The limestone cliff habitat, with strongly basic soil, would be a
most unusual one for S. aucuparia, but was similar to the cliff refuges of other Sorbus species. It was
not possible for M.H. to make a closer study of the population until early 1993, when his searches of
similar cliffs in the area revealed another, rather larger population of the same taxon on higher cliffs
about 4 km to the east of the first site, again growing on inaccessible ledges, and again apparently
overlooked in an area with an even longer history of botanical study. The precise localities of the
two sites are here deliberately withheld in order to prevent damage to the sites, but they have been
confidentially notified to the relevant vice-county recorders and to the regional and national officers
of the Countryside Council for Wales with responsibility for species conservation.

IDENTIFICATION, MORPHOLOGY AND VARIATION

Confirmation of the identity of the taxon forming the Glamorgan cliff populations presented
considerable problems. Sorbus domestica is similar to S. aucuparia in many vegetative characters.

380 M. HAMPTON AND Q. O.N. KAY

Although several morphological characters of the Glamorgan cliff taxon strongly suggested from
the outset that it might be S. domestica, this species had not previously been found either in natural
habitats or as a persistent introduction in Britain, with the exception of a single ancient tree that
formerly grew in the Wyre Forest in Worcestershire (Lees 1867; Amphlett & Rea 1909). The largest
trees in the Glamorgan cliff populations are evidently of considerable age (ring-counts suggest
possible ages of several hundred years for the larger trunks), but are wind-trimmed, partly prostrate
and only c. 3-5 m high. Immature fruits, pyriform and pubescent rather than globose and
glabrescent as in S. aucuparia, are formed in some years, and a single green, pyriform, mature but
seedless fruit was seen on one of the western trees in the autumn of 1991, but full fruiting has not yet
been observed and probably takes place only in exceptionally favourable years, if at all. A similar
shrub-like habit, with fruiting rare or absent, has been reported for populations of S. domestica
growing on dry, warm slopes at the northern limit of the range of the species in mainland Europe,
which lies in southern Germany (Gerstberger 1980).

In contrast, the Wyre Forest tree was tall and apparently fruited freely before it became decrepit
(it was locally known as the ‘Witty Pear’). It was first described in 1678, became decrepit during the
mid-nineteenth century and was finally destroyed by fire in 1862 (Lees 1867; Amphlett & Rea 1909).
Although it grew in a remote part of the Wyre Forest it was ’regarded as a probable introduction
(e.g. Clapham, Tutin & Warburg 1962). It has left descendants including one of the two tall (17 m
and 24 m, Bean 1980) and free-fruiting trees of S. domestica cultivated since the mid-nineteenth
century in the Oxford Botanic Garden (T. Walker in litt.), and could, like similar S. domestica trees
cultivated in modern arboreta in Britain, have been descended from cultivars grown in mediaeval
orchards. Apart from the very obvious distinction provided by the fruits (c. 2-2-5 cm in diameter,
greenish-brown and pear- or less often apple-shaped with numerous stone-cells in S$. domestica, 0-6-
0-9 cm in diameter, globose, bright red and soft-fleshed when mature in S. aucuparia), the other
characters given in Floras for distinguishing the two species are inconsistent and potentially
unreliable (Gerstberger 1980), especially for trees growing in exposed habitats. Consequently, non-
fruiting trees cannot be certainly identified from the keys published in major Floras.

As fruits were not available, a range of other characters, including previously uninvestigated
micromorphological and isozyme characters, were investigated in the Glamorgan cliff taxon and
compared with cultivated material of S. domestica and with wild and cultivated material of S.
aucuparia during the summer of 1993 (Table 1). This comparison confirmed that the Glamorgan cliff
taxon is undoubtedly S. domestica. Isozyme comparisons were made by Rosemary John and Sarah
Martin. Five of the six trees from which leaf samples were taken, all from the western population,
were isozymically identical to one another at the nine loci that were investigated (GOT-1 and 2,
G6PDH, IDH, MDH, PGI-1 and 2, PGM, SOD); the sixth was less well characterised, but may
have differed at one locus (GOT-1). All were clearly distinct from S. aucuparia at several loci.

TABLE 1. CHARACTERS USED TO DISTINGUISH SORBUS DOMESTICA FROM S. AUCUPARIA

Sorbus domestica

(Glamorgan cliff taxon)

Styles 5-(6)'

Stipules 6-14 mm, forked at or below half-way, soon
falling, with little green tissue*

Epidermal cells around stomata on lower leaf epi-
dermis with sinuous median ridges but lacking
papillae; stoma with ‘collar’ ridge*

Buds glabrous and viscid*, green or pale brown!’
Bark on trunk and older branches deeply and nar-
rowly fissured”, flaking!

Sorbus aucuparia

Styles 3-4"

Stipules 4-8 mm, simple or fan-shaped, persistent
and green on long shoots?

Epidermal cells around stomata on lower leaf epi-
dermis with central papillae flanked by radiating
ridges, so that 5—6 papillae surround each stoma;
no conspicuous ‘collar’ ridge* around stoma.

Buds tomentose*, brown

Bark on trunk and older branches remaining
smooth’ or with fissures at wide intervals

All characters were determined in fresh material and compared with fresh material of known identity. The
Glamorgan cliff taxon agreed with S. domestica in all characters that were investigated. Additional characters
are described in the first paragraph of this paper. References: 'Warburg & Karpati 1968; *Gerstberger 1980;
*Kay unpublished; *Burnat 1899; °Stace 1991.

SORBUS DOMESTICA L. IN BRITAIN 381

Isozyme comparisons with fresh leaf material of cultivated S. domestica trees from Oxford Botanic
Garden (single trees of the apple-fruited and pear-fruited varieties) and Mount Pleasant Trees of
Berkeley, Gloucestershire (two cultivars, one from a long-established local garden) showed
polymorphism within the species at all loci except MDH; the Glamorgan cliff taxon differed from
the Oxford trees in PGI and SOD, and from the Gloucestershire garden cultivars in PGI, SOD and
GOT-1; PGI alleles differed amongst the cultivars. In practice the leaf epidermis characters
(apparently previously unobserved) and (when still present) the stipule characters (Table 1),
combined with the bud characters in fresh material, are likely to be most useful for the separation of
vegetative material of S. domestica from S. aucuparia.

ECOLOGY AND HISTORY

At the first (western) Glamorgan site, S$. domestica grows as scattered small trees, branching from
the base and up to about 3-5 m in height, on a vertical sea-cliff of horizontally bedded Lias limestone
c. 25 m high, with a S.S.W. aspect. The trees are rooted into softer beds of marly calcareous shale
which form recessed ledges between harder, projecting limestone strata; 13-14 separate trees can be
counted, most on the higher inaccessible ledges. The three westernmost S. domestica trees grow in
dense cliff-scrub with Prunus spinosa, Hedera helix, Clematis vitalba, Rubus ulmifolius and
Crataegus monogyna* forming a thicket 2-2-5 m deep, but most of the other trees grow in more
open sites, often with the lower part of the trunk prostrate and appressed to the cliff surface. Trunk
diameters at the base of the tree ranged from 10 cm to 23 cm in four accessible trees, with similar
dimensions estimated for the trees on the higher inaccessible ledges. Conspicuously fissured, flaking
bark (a characteristic of S. domestica, illustrated photographically by Step (1905)) covers the lower
parts of the older trunks. Several trees show the remains of dead older trunks or branches embedded
in the base of the present trunk, suggesting that the living trunks may have arisen as basal shoots
from older trunks that are now dead. A section of one of these older dead trunks 6 cm in diameter
examined by M.H. showed 85-90 annual rings, indicating an age of about 300-400 years for the
largest living trunks. Bean (1980) states that there is little doubt that trees of Sorbus domestica can
live for 500-600 years. If the trees in the Glamorgan cliff populations have periodically regrown
from basal shoots or from suckers after the death and decay of an older main trunk, considerably
greater ages seem possible. A pH of 7-55 was found for the stony calcareous marl into which one of
the westernmost trees was rooted. Associated species are listed in Table 2. No introduced species
grow in association with the S. domestica population here, although a number of weedy species grow
on or near the cliff-top and on the nearby maritime drift-line. The population extends along about
100 m of cliff-face, bounded to the east by cliffs that are probably now too exposed to salt spray for
S. domestica to survive and to the west by sloping degraded cliffs covered by denser, largely closed
woodland and scrub.

At the second site, 4 km east of the first, S. domestica again grows as scattered small trees and
shrubs, on the upper, open, vertical or near-vertical part of a south-facing horizontally bedded Lias
limestone sea-cliff about 50 m high, with softer strata, into which S. domestica is probably rooted,
alternating with projecting ledges of harder limestone. The lower part of the cliff here is steeply
sloping and fairly densely wooded, with ash, elm and some yew. The largest S. domestica trees here
are about 5 m high, but most are smaller, spreading along horizontal ledges, and with many
apparent saplings or small sucker shoots growing in unshaded and largely unvegetated open vertical
areas. A total number of about 70 apparently separate S$. domestica trees, shrubs and saplings or
sucker shoots was estimated, extending along about 230 m of cliff; as at the western site, the
population is bounded to the west by denser, closed woodland on less steeply sloping cliffs, and to
the east by more exposed and unstable vertical cliffs. The cliffs on which S. domestica grows here are
entirely inaccessible or, at best, extremely hazardous; voucher specimens could be obtained only
from a few trees at the edge of the population. At this second site, the introduced species Quercus
ilex was present nearby but was not seen growing with S. domestica (Table 2).

The south-facing Lias limestone cliffs and valley-slopes of southern Glamorgan provide
favourable microhabitats for thermophilous calcicole species, a number of which are at or near the

* All nomenclature follows Stace (1991), except where stated otherwise.

382 M. HAMPTON AND Q. O.N. KAY

TABLE 2. ABUNDANCE OF SPECIES ASSOCIATED WITH SORBUS DOMESTICA AT CLIFF SITES
IN SOUTHERN GLAMORGAN, WALES

Site

Species Western site Eastern site

Sorbus domestica 3) la
Adiantum capillus-veneris
Brachypodium sylvaticum
Brassica nigra

Centaurea scabiosa

Clematis vitalba

Cornus sanguinea

Crataegus monogyna

Dactylis glomerata

Daucus carota

Festuca rubra

Fraxinus excelsior

Hedera helix

Helianthemum nummularium
Inula conyzae

Tris foetidissima

Ligustrum vulgare
Lithospermum purpurocaeruleum
Ononis repens

Origanum vulgare

Parietaria judaica

Pilosella officinarum ~
Pimpinella saxifraga -
Plantago lanceolata ~
Prunus spinosa HESS
Quercus ilex ~
Q. robur

Rosa canina agg. 1-4
Rubia peregrina , 4
Rubus ulmifolius 1-4-5
Sambucus nigra -
Sanguisorba minor

Senecio erucifolius -
S. jacobaea -
Taxus baccata -
Teucrium scorodonia ~
Ulmus sp. ~
Viburnum lantana 1

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The lists are composite estimates for the areas in which S. domestica grows. Because of the inaccessibility of the
greater part of both populations, some less conspicuous species are likely to have been missed. At the eastern
site, frequencies of the species growing on vertical cliffs were estimated from a range of c.80—-200 m using
binoculars. The figures for the western site show Domin cover-abundance scores. Letter abbreviations: la,
locally abundant; If, locally frequent; 0, occasional; r, rare; x, present nearby but not seen with or under S.
domestica.

limits of their European ranges, for example Adiantum capillus-veneris, Campanula glomerata,
Cirsium eriophorum, C. acaule and Lithospermum purpurocaeruleum. Elsewhere in the Bristol
Channel area, the hard Carboniferous Limestone cliffs of the Gower Peninsula in western
Glamorgan and of the Avon Gorge, Brean Down and the southern Mendip Hills in Somerset have
acted as refuges for a greater number of calcicole species, including the vast majority of the British
apomictic Sorbus microspecies (Proctor & Groenhof 1992; Stace 1991). However, the friable, marly
calcareous shales which are interbedded with the Lias limestone strata provide special cliff
microhabitats, which do not occur in these Carboniferous Limestone districts, and the Lias

SORBUS DOMESTICA L. IN BRITAIN 383

limestone itself is relatively soft, maintaining a higher pH in the soils of cliff-tops and valley-slopes
than on Carboniferous Limestone. The warm, south-facing ledges on which S. domestica grows are
an exceptionally favourable environment for a thermophilous calcicole species at the northern limit
of its range; the Mediterranean species Adiantum capillus-veneris grows as an undoubted native on
the same ledges, in places where water seepages emerge from the permeable calcareous marl.

It seems quite possible that S. domestica may be native in the Glamorgan sites. Ring-counts
indicate that the oldest trees at both sites are a minimum of several hundred years old, and
introduced species are absent from the cliff refuges on which S. domestica grows, with the exception
of Quercus ilex at the eastern site, where it was not seen with S. domestica (Table 2). There are no
records of past cultivation or planting of S. domestica in southern Wales. Similar disjunctions
between Continental and British populations occur in the European ranges of other calcicole
species with outlying localities in the Bristol Channel area, for example Arabis scabra, Draba
aizoides and Koeleria vallesiana. The full extent of the native distribution of S. domestica in Europe
is hard to determine, because it has been widely cultivated for its fruits in the past, especially in
Germany and France, and often occurs as scattered, isolated trees which seem likely to be
descended from cultivated trees (e.g. Burnat 1899; Hegi 1922). It typically grows on calcareous
soils. Wild populations are apparently widespread in southern Europe from Spain to the Balkans
(Hegi 1922; Warburg & Karpati 1968), and also in Turkey (Davis 1972), extending northwards to
southern Germany, where S. domestica is an indicator species of thermophilous mixed-oak
woodland (Quercetalia pubescenti-petraeae) on dry slopes (Gerstberger 1980). In France, S.
domestica is often of doubtful spontaneity, but occurs in possibly native populations in woods and
hedgerows and calcareous soils in the Midi and in central and western France as far north as
Morbihan (Coste 1901). The discovery of S. domestica in Glamorgan suggests that it could have
been similarly overlooked or confused with S. aucuparia at other possible refuge sites in southern
Britain and perhaps elsewhere in north-western Europe. The close superficial vegetative similarity
between S. domestica and S. aucuparia has apparently not been realised by most botanists working
in the British Isles, and non-fruiting populations of pinnate-leaved Sorbus on south-facing
calcareous cliffs or open limestone slopes, if found, should be critically examined.

ACKNOWLEDGMENTS

We are very grateful to the University of Liverpool Botanic Garden at Ness, the University of
Oxford Botanic Garden, and Mount Pleasant Trees of Berkeley, Gloucestershire for generously
providing fresh material of cultivated $. domestica, to Gwynn Ellis and George Hutchinson of the
National Museum of Wales and to Philip Nethercott for their taxonomic advice and assistance, to
Colin Pascoe for historical research, to Rosemary John and Sarah Martin for expertly carrying out
isozyme analyses, and to Peter Jones for his help in the field.

REFERENCES

AmMPHLETT, J. & REA, C. (1909). The botany of Worcestershire. Cornish Brothers Ltd, Birmingham.

BEAN, W. J. (1980). Trees and shrubs hardy in the British Isles, ed. 8. John Murray, London, (4 vols. &
supplement. Sorbus, vol. 4, 1980, pp. 399-464).

Burnat, E. (1899). Flora des Alpes Maritimes, 3 (1). H. Georg, Genéve, Bale & Lyon.

CLAPHAM, A. R., Tutin, T. G. & WarBurRG, E. F. (1962). Flora of the British Isles. Cambridge University Press,
Cambridge.

Coste, H. J. (1901). Flore descriptive et illustrée de la France, de la Corse et des contrées limitrophes, 2 (1). Paul
Klincksieck, Paris.

Davis, P. H., ed. (1972). Flora of Turkey and the East Aegean Islands, 4 (Rosaceae to Dipsacaceae). Edinburgh |
University Press, Edinburgh.

GERSTBERGER, P. (1980). Neue Merkmale zur Unterscheidung von Sorbus domestica L. und Sorbus aucuparia L.
GOttinger floristische Rundbriefe 14: 1-5.

HE, G. (1922). Illustrierte Flora von Mittel-Europa, 4 (2). J. F. Lehmanns Verlag, Miinchen.

Legs, E. (1867). The botany of Worcestershire. Worcestershire Naturalists’ Club, Worcester.

384 M. HAMPTON AND Q. O.N. KAY

Proctor, M. C. F. & GRoENHOF, A. C. (1992). Peroxidase isoenzyme and morphological variation in Sorbus L.
in South Wales and adjacent areas, with particular reference to S. porrigentiformis E. F. Warb. Watsonia
19: 21-37.

Stace, C. A. (1991). New flora of the British Isles. Cambridge University Press, Cambridge.

STEP, E. (1905). Wayside and woodland trees. Frederick Warne & Co., London.

WARBURG, E. F. & KARPATI, Z. E. (1968). Sorbus L., in Tutin, T. G. et al., eds., Flora Europaea 2: 67-71.
Cambridge University Press, Cambridge.

(Accepted February 1995)

Watsonia 20: 385-390 (1995) 385

Elytrigia repens (L.) Desv. ex Nevski subsp. arenosa (Spenner)
A. Love (Poaceae) in north-western Europe

P. J. O. TRIST

Glovers, 28 High Street, Balsham, Cambridge CBI 6DJ

ABSTRACT

Elytrigia repens subsp. arenosa differs from subsp. repens in being a much smaller plant with shorter ribbed
cauline leaves and shorter panicles and spikes. It is recorded from the east, south and south west coasts of
England and the Channel Islands on maritime sand. Its nomenclature is summarised, specimens seen are listed
and the north western European distribution is given and mapped.

KEyworDs: nomenclature, European specimens, distribution, maritime sands, sandy heaths.

INTRODUCTION

Elytrigia repens (L.) Desv. ex Nevski subsp. arenosa (Spenner) A. Léve has had a chequered
history. It is an undistinguished grass which has had little attention. It was first recognised by Koch
& Ziz (1814) in Germany, later by Jansen & Wachter (1933) in the Netherlands and by Holmberg
(1926) and Hylander (1953) in Denmark, Finland and Sweden. Korneck (1966) and Hecker (1987)
both record it in studies on the Mainz sands in Germany. In Britain it was not recorded in Bentham
& Hooker (1858) or subsequently. As far as I know, Melderis (1980) was the first to report this taxon
from Britain. It is likely that Stace (1991) carried its first description in a British Flora. Of this taxon
he says “‘its distribution and taxonomic status are very uncertain’. I here attempt to clarify this
uncertainty.

The late C. E. Hubbard had early knowledge of subsp. arenosa and had been alerted by a
specimen he collected in 1936 from reclaimed land near The Wash, W. Norfolk, v.c. 28. The
specimen is in K, labelled ““Agropyron repens var.?”. | have examined it and consider it to be
Elytrigia repens subsp. arenosa. Further evidence of his early interest in this taxon comes from a
sheet recently found at NWH. The label records ““Agropyron maritimum (Triticum maritimum)
Koch & Ziz, sea bank, Burnham Overy Staithe, Norfolk, 28 July 1967: collected by C. E. Hubbard
and E. L. Swann’’. I have seen this specimen and agree the determination with the nomenclature
updated to Elytrigia repens subsp. arenosa. Swann also annotates the sheet “‘like A. repens but
glaucous with glabrous leaves, convolute when dry, with stout smooth ribs, spikelets small’’. This
agrees with my own description.

Hubbard (in litt. 1972) informed me “I have a living plant of the plant originally named Triticum
repens var. maritimum by Koch & Ziz in Germany’’. He had also received six sheets of specimens of
var. maritimum from K. Watermann of Ingelheim which were collected from the type locality on the
Mainz sands in Germany; these were later incorporated at K. Of these sheets, Hubbard also
commented “these I hope to pass to Melderis for study”. Hubbard’s health intervened and Melderis
never received the sheets. When the latter included Elymus repens subsp. arenosus in Flora
Europaea (Melderis 1980) I was interested to see a specimen. In 1986 I asked Melderis if he had a list
of British localities for this taxon. He did not reply to my question but said that the inclusion of this
taxon in Melderis & McClintock (1983) “‘was simply on the authority of C. E. Hubbard’’. He said he
would shortly go to Kew and study the Watermann specimens of E. repens var. maritimum and later
let me have a report. However, this intention, written into his last letter which was addressed to me
and in his pocket at the time of his death, was never realised. Melderis had also referred (in litt.
1986) to specimens at BM, but in my subsequent researches I found no annotations on any sheets
referring to. Elymus repens subsp. arenosus. From these events it seems clear that Melderis was

386 P. J. O. TRIST

unable to complete his proposed study and had accepted Hubbard’s opinion in the listing of subsp.
arenosus. It is not known which of the two proposed the rank of subspecies.

The specimens at K from the Mainz area of Germany subsequently proved of guidance to me in
the primary field identification. In 1986 on coastal dunes on L’Ancresse Common, Guernsey I found
my first living specimens of subsp. arenosa. Subsequent searches at CGE and LTR yielded several
British specimens which matched those I had taken in Guernsey. I then made further successful
searches on the east coast of England and was satisfied that this taxon could be included in the
British list and its description could be improved.

NOMENCLATURE

The original description of this taxon was made under Triticum repens var. maritimum Koch & Ziz
in 1814. It will later be shown that this name is unacceptable. Smith (1800) in recording Triticum
repens var. y did not give a description of the plant but as his authority he cited Ray and Withering
and referred to material in herb. Lightfoot. Roth (1802) validly published a description of Smith’s
plant under Triticum repens var. maritimum Sm. ex Roth. It is evident that the name var. maritimum
Koch & Ziz published in 1814 was not based on var. maritimum Sm. ex Roth (1802), but was a later
homonym and a nom. illeg.

It is likely that the Smith plant was based on British material and the Koch & Ziz plant was
probably based on a different type as the authors were accounting for plants on the Continent and
published in Catalogus plantarum Palatinatus.

The synonym Triticum repens var. maritimum Koch & Ziz non Sm. ex Roth is retained in the
summary of the nomenclature and confirms that the plant Triticum repens var. maritimum Sm. ex
Roth (1802) was unrelated to Triticum repens var. maritimum Koch & Ziz published in 1814.

The first valid and legitimate name for our plant is Triticum repens var. arenosum Spenner
published in 1825. This latter was elevated to Elymus repens subsp. arenosus by Melderis in 1978
although he used a later publication of the epithet by Petif (1830) as the basis for his new
combination (Melderis 1978).

Stace (1991) has corrected the nomenclature and returned the species to Elytrigia.

The following summarises the nomenclature. i
Elytrigia repens (L.) Desv. ex Nevski subsp. arenosa (Spenner) A. Léve in Taxon 29: 351 (1980).
Triticum repens L. var. maritimum Koch & Ziz, Catalogus plantarum quas in ditione florae

Palatinatus legerunt 5: 17 (1814) nom. illeg., non Smith ex Roth (1802).

Triticum repens var. arenosum Spenner, Florae Friburgensis 1: 162 (1825).
Triticum repens var. arenosum Petif, Enumeratio plantarum in ditione florae Palatinatu sponte

crescentium 5: 16 (1830).

Triticum maritimum Jansen & Wachter in Nederlandsch kruidkundig archief 43: 178 (1933), non L.

(1762).

Agropyron maritimum Jansen & Wachter, Flora Neerlandica 1 (2): 116 (1951), non (L.) P. Beauv.

(1812).

Elytrigia repens var. maritima N. Hylander in Botaniska notiser: 357 (1953).
Elymus repens subsp. arenosus (Petif) Melderis in Botanical journal of the Linnean Society 76: 379

(1978).

DESCRIPTION OF ELYTRIGIA REPENS SUBSP. ARENOSA
Perennial, usually forming small patches but sometimes as a single-culmed plant, with rhizomes;
whole plant glaucous-green. Culms (16—)28-65 cm high, 0-4-1-0(—1-6) mm wide below the spike,
erect, slender, sometimes geniculate at or near the base; nodes 2-3, light to dark pink. Sheaths
glabrous, rounded on the back with well developed auricles; ligules less than 0-5 mm. Lower leaves
few, (4-)7-14(-20) cm x 2-4 mm, rigid, finely pointed at apex. All leaves with prominently raised,
broad, whitish or green veins c. 0-15—0-2 mm wide on the adaxial surface, glabrous or with a few
scattered hairs, with margins sometimes minutely scabrid, all leaves involute or quickly becoming so
on drying. Upper cauline leaves 1-5—6(—9-5) cm Xx 1—2(-3-4) mm, long-acuminate at apex. Spikes

ELYTRIGIA REPENS SUBSP. ARENOSA IN N.W. EUROPE 387

(2-5—)4-9 cm long, erect, short and narrow, rhachis strap-shaped, with scabrid margins. Spikelets
few, (7-)9-14 mm long, sessile, 2-6 flowered. Glumes (4-4-)6-9(-10) mm long, lanceolate to
lanceolate-oblong, strongly keeled, scabrid, slightly unequal, blunt or mucronate and sometimes
with awns (0-2—)1-0-2:3 mm at apex; veins 3(—7). Lower lemmas (5-5—)7—10(-12) mm long,
lanceolate-oblong, blunt or mucronate at apex, sometimes with an awn (0-3—)1-8—2-8(—4-6) mm. In
both glumes and lemma the awn is an extension of the central nerve and is sometimes scabrid
distally. Palea shorter than the lemma, with ciliate margins. Anthers 4-0-5-3 mm.

The incidence of awns on subsp. arenosa is variable. On British material I found 22 spikes awned
and 25 awnless. All 26 spikes of Swedish specimens were awned but only two out of 23 in German
material. Those from Finland and Spain were awned but in French specimens the incidence was only
two out of ten. The incidence of hairs on the adaxial leaf surfaces on subsp. arenosa is of little help in
effecting determination. Four specimens of 50 British plants had leaf hairs, one in 26 of German and
15 of 18 in Swedish material. The occasional presence of hairs on subsp. arenosa leaves should not
lead one to confuse the leaves with those of subsp. repens, as in the latter the leaf surface is flat and
the leaf is flaccid and longer than in subsp. arenosa. Another possible confusion in determination
may arise from the rare occurrence of cilia on the sheath margin of Elytrigia repens subsp. arenosa
which is a character of Elytrigia atherica (Link) Kerguélen ex Carreras Mart. and its hybrids. Where
a specimen with sheath cilia also has narrow and stiff leaves with involute margins and raised veins,
it should be determined as Elytrigia repens subsp. arenosa.

An examination of a wide range of European specimens has enabled me to expand and modify the
previous descriptions given by Melderis (1978, 1980) and Stace (1991). As examples I consider that
all leaf margins of subsp. arenosa are involute or become so soon after collection; that geniculation
at lowest nodes is infrequent; that leaves are found with either flat or round topped ribs and that
none have been found more than 4 mm wide; and that lemmas are sometimes awned.

A comparison of the two subspecies is given in Table 1.

TABLE 1. A COMPARISON OF ELYTRIGIA REPENS SUBSP. REPENS AND SUBSP. ARENOSA

Elytrigia repens
subsp. repens

Elytrigia repens

subsp. arenosa

Culm length
Culm
geniculation
Culm nodes
Leaves

Upper leaves
Basal leaves

Spike length

Spike habit

Spikelet length

Number of
florets

Glume length

Glume: no. of
veins

Glume awns

Lemma length
Lemma awns

Anthers

30-120 cm
Sometimes at or near the base

Light to dark brown

Generally flat and flaccid, veins narrow;
usually with dispersed hairs on adaxial
surface

8-16(—25) cm X 2-8-5 mm

8-30 cm X 3-10 mm

5—20(-30) cm

Erect to nodding with up to c. 25
spikelets

10-20 mm

3-8

7-12 mm
3-7

Sometimes mucronate and rarely awned
in British specimens

8-13 mm

Usually absent in British material but in
var. aristatum are thin and weak up to
15 mm long

3-5-6 mm

(16-)28-65 cm
Infrequent

Light to dark pink

Thick, rigid; margins involute, veins
broad, rounded or flat, 0-15—0-2 mm
wide; usually glabrous

1-5-6(-9-5) cm Xx 1-2(-—3-4) mm

(4-)7-14(—20) cm X 2-4 mm

(2:5—)4-9 cm

Short and strict often with <10 spikelets

(7-)9-14 mm
2-6

(4-4-)6-9(-10) mm
3(-7)

(0-2—)1-0-2:3 mm

(5:5—)7-10(-12) mm

Thick, straight and rigid, (0-3—)1-8-2-8
(—4-6) mm long. Where the lemma is
mucronate, the tip is corneous and
often scabrous

4-5-3 mm

388 P. J. O. TRIST
HABITAT

Elytrigia repens subsp. arenosa occurs on maritime sand and dunes and can be seen within 30 m of
the tide but is more often at least 70 m from the sea. It is also found at the rear of partially colonized
sand beaches where infrequent tidal overflows encourage the growth of a salt tolerant flora. More
observations and sampling for salinity are necessary before an opinion can be given on the salt
tolerance of subsp. arenosa. The pH of the sands examined ranges from 6 to 7.

To the east of Caen, Normandy, it is seen on high dunes of mobile fine sand which are partially
colonized with Ammophila arenaria (L.) Link, Ononis repens L. and Tamarix gallica L. At Sizewell
E. Suffolk (v.c. 25) it grows on similar dunes with Elytrigia atherica, Rumex crispus L. and Carex
arenaria L. At both of these sites subsp. arenosa only colonizes the perimeter of the associated
vegetation. At L’Ancresse Common, Guernsey it grows as single-culmed plants widely spaced in
semi-open consolidated dunes with Agrostis stolonifera L., Cynosurus cristatus L., Gaudinia fragilis
(L.) P. Beauv. and Vulpia bromoides (L.) Gray.

In the vicinity of the Old Dunwich River at Walberswick, E. Suffolk (v.c. 25) it is found on
consolidated sand subject to occasional tidal overflow, the soil of which is a mixture of fine sand and
undecomposed organic matter derived from plant remains of Festuca rubra L., Glaux maritima L.,
Atriplex portulacoides L. and Spergularia marina (L.) Griseb. indicating a low salinity. On the beach
at Thorpness, E. Suffolk (v.c. 25) it is found in small shingle and coarse sand with Silene uniflora
Roth, Senecio vulgaris L. and Ononis repens L. On the same coast at Dunwich it is sparingly seen in
small isolated colonies in sandy areas in large shingle on the back of the sea defences with Lathyrus
japonicus Willd. It is recorded from the sand banks of the tidal estuary of the River Parret, S.
Somerset (v.c. 5).

In Germany subsp. arenosa occurs on sandy heaths and in sandy arable land in the Mainz area c.
400km inland (Korneck 1966; Hecker 1987). The protologue of this taxon by Koch & Ziz (1814)
reads ‘Triticum repens maritimum in sabulosis prope Moguntiam (Mainz) cum aliis plantis salinis
copiose occurit”’. Hecker (1987) lists 217 taxa on the Mainz sands, and 159 of these are also found on
the sands in the West Suffolk Breckland. Subsp. arenosa has not yet been reported from the latter,
but it could well occur there. Recalling the above “‘plantis salinis copiose occurit”’, the Breckland
sands support some maritime taxa including Corynephorus canescens (L.) P. Beauv., Carex arenaria
L., Phleum arenarium L., Trifolium scabrum L. and T. suffocatum L.; the area is today c. 40 km
from the tidal bay of The Wash. The inland stations of Elytrigia repens subsp. arenosa in the Rhine
locality of Mainz suggests that it requires open sandy habitats rather than specifically maritime ones.
It may well be a taxon like Hippophaé rhamnoides L. which was once more widespread and now
occurs predominantly on the coast of Britain as its former inland sand habitats have been eroded
(Godwin 1975). Both subsp. arenosa and H. rhamnoides are included in Hecker’s (1987) list.

DISTRIBUTION OF ELYTRIGIA REPENS SUBSP. ARENOSA IN NORTH-WESTERN EUROPE
The distribution of Elytrigia repens subsp. arenosa in north-western Europe is shown in Fig. 1.

ENGLAND

S. Somerset, v.c. 5. Banks of River Parret between Stert Point and Combwich, 1907, E. S. Marshall
(CGE, OXF).

W. Sussex, v.c. 13. Sea shore at Pagham, 1875, H. E. Fox (OXF).

S. Essex, v.c. 18. Foot of sea wall, Sandbeach Farm, Bradwell Juxta Mare, TM/030.053, 1986, P. J.
O. Trist (herb. P.J.O.T.).

E. Suffolk, v.c. 25. The Dunes, Walberswick, 1950, E. K. Horwood (LTR); near the Old Dunwich
River, The Flats, Walberswick, TM/501.749, 1989, P. J. O. Trist (herb. P.J.O.T.); sand erosion
from shingle sea defence, Corporation Marshes, Walberswick ,"TM/495.740, 1991, P. J. O. Trist
(herb. P.J.O.T.); foot of shingle bank sea defences, Dunwich, TM/479.707, 1989, P. J. O. Trist
(herb. P.J.O.T.); fixed dunes, Sizewell beach, TM/475.629, 1989, P. J. O. Trist (herb. P.J.O.T.);
sand and shingle beach, N. of Haven House, Thorpness, TM/470. 589, 1989, P. J. O. Trist (herb.
P.J.O.T.).

|

ELYTRIGIA REPENS SUBSP. ARENOSA IN N.W. EUROPE 389

Ficure 1. European distribution of Elytrigia repens subsp. arenosa.

E. Norfolk, v.c. 27. Sandy waste, Gt Yarmouth, 1953, B. A. Poulton (E).

W. Norfolk, v.c. 28. Hunstanton, 1935, T. G. Tutin (LTR); near the river by the paper mill, West
Newton, 1936, C. E. Hubbard (K); sea defence bank, Burnham Overy Staithe, 1967, C. E.
Hubbard & E. L. Swann (herb. E.L.S. in NWH).

Channel Islands v.c. S. Consolidated dunes by the Doyle Rock, L’Ancresse Common, Guernsey,
WV/348.834, 1986, P. J. O. Trist (herb. P.J.O.T.); coastal sand dunes, Port Soif, Guernsey, WV/
304.818, 1991, Patience Ryan (det. P.J.O.T.).

SPAIN
San Sebastian, 1895, M@. Gandoger (E).

FRANCE

Dyke bank on sea shore, west of St Valery-sur-Somme, 1959, D. P. Young (BM); sand dunes at rear
of beach, E. of Le Home, West of Cabourg, Calvados, Normandy, 1989, P. J. O. Trist (herb.
P.J.O.T.); high sand dunes at rear of beach, N.E. of Merville-Franceville, Calvados, Normandy,
1989, P. J. O. Trist (herb. P.J.O.T.).

THE NETHERLANDS

Oostvoorne, 1914, A. W. Kloos & J. W. Henrard (L); Oostvoorne, 1917, J. T. Henrard (L); Hook
of Holland, 1910, P. Jansen & W. H. Wachter (L); Noordwolde, Hemelumer Oldeferd, 1929, A.
N. Koopmans (L).

390 P. J. O. TRIST

GERMANY

Sandy area near Cleves, E.S.E. of Nijmegen, 1866, Florae rhenanae Fasc. 6 (BM); Offenbach am
Main, undated, C. B. Lehmann (K); heathland on ‘Mainzer sand’, Mainz, 1971, Kurt Watermann
(K); sandy arable field, Weilersberg near Mainz, 1969, 1971, Kurt Watermann (K); Priwall, near
Traveminde, N.E. of Lubeck, 1845, G. R. Haecker (E).

FINLAND
Upper part of the sea shore meadow, west of Kuljunmaa in the Tirkkale group, off Lokalahti, 1973,
Sakari Hinneri (E).

SWEDEN

Nyk6ping, Sodermanland, 1921, Carl Blom (K); sandy shore south of Kristineberg, Skaft6,
Bohuslan, 1947, K. H. Mattesson (NMW); Norrvreda, Singd, Upplandia, 1928, G. A. Ringselle
(NMW); Tranvik, Sing6, Upplandia, 1928, G. A. Ringselle (NMW); Lund, Skane, 1901, O. R.
Holmberg (BM); Rundskar, Nyk6ping archipelago, 1882, Hugo Samzelius (E).

I have been unable to identify any specimens from collections sent from Trondheim (TRH) and
Tromso (TROM) in Norway and it is likely that these areas are too far north for subsp. arenosa to
occur. Dr Sivertsen of Trondheim suggests that this taxon might be found on the sands around Lista
and in the Jaeren district in the south of Norway. I was surprised to find no specimens of subsp.
arenosa in a large collection from Copenhagen (C).

ACKNOWLEDGMENTS

I thank the curators of BM, C, CGE, E, K, L, LTR, NUW, NWH, OXF, RNG, TRH and TROM for
search and loan of specimens. I am also indebted to Mrs G. Beckett, A. O. Chater, Dr T. A. Cope,
the late R. W. David, R. J. Driscoll, R. G. Ellis, Dr P. Grubb, D. R. McKean, C. D. Preston, Mrs
P. Ryan, P. D. Sell, Professor C. A. Stace, Dr S. M. Walters and Dr P. Yeo for various assistance
and guidance. I am particularly grateful to Dr C. E. Jarvis and D. H. Kent for search and guidance
with nomenclature. I also acknowledge the help received from Dr U. Hecker, Dr D. Korneck, Dr
R. van der Meijden, Dr M. Morkved, J. Reiersen, Dr S. Sivertsen and Dr J. F. Veldkamp.

REFERENCES

Gopwin, H. (1975). Hippophaé rhamnoides, in History of the British flora, p. 208. Cambridge University Press,
Cambridge.

Hecker, U. (1987). Agropyron maritimum, in Die Farn-und Blitenpflanzen des Mainzer Sandes. Mainzer
naturwissenschaftliche archiev 25: 85-133.

HotmBereG, O. R. (1926). Scandinavian Flora 2: 274. Stockholm.

Hy anper, N. (1955). List of plants of north west Europe. 1. Vascular plants, 26. Lund.

Kocu, W. D. J. & Ziz, J. B. (1814) Triticum repens var. maritimum, in Catalogus plantarum quas in ditione
florae Palatinatus legerunt 5: 17. Mainz.

Korneck, D. (1966). Agropyron repens var. maritimum, in Wiederfund von Kochia arenaria Roth im
Naturschutzgebeit ‘Mainzer Sand’. Hessische Floristische Briefe 178: 49-56.

MELpDE_RIS, 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.

MELpe_RIS, A. (1980). Elymus L.,in Tutin, T. G. etal., eds. Flora Europaea 5: 196. Cambridge University Press,
Cambridge.

ME tperis, A. & McCuintock, D. (1983). The genera Elymus L. and Leymus Hochst. in Britain. Watsonia 14:
391-395.

Petir, C. (1830). Triticum repens var. arenosum. Enumeratio plantarum in ditione florae palatinatu sponte
crescentium 5: 16. Zweibrucken.

Rotn, A. W. (1802). Triticum repens var. maritimum Smith ex Roth. in Neue Beitrage zur Botanik, 136-137.
Frankfurt.

SmitH, J. E. (1800). Triticum repens var. y. Flora Britannica 1: 158. London.

STAcE, C. A. (1991). Elytrigia repens (L.) Desv. ex Nevski subsp. arenosa (Spenner) A. Love in New Flora of the
British Isles, p. 1066. Cambridge University Press, Cambridge.

(Accepted November 1994)

|

Watsonia 20: 391-396 (1995) 391

Puccinellia distans (Jacq.) Parl. subsp. borealis (O. Holmb.)
W. E. Hughes (Poaceae) in mainland Scotland and the
Outer Isles

P. J. O. TRIST
Glovers, High St, Balsham, Cambridge, CBI 6DJ
and
J. K. BUTLER

Seaside Cottage, Thurso East, Caithness, KW14 8NH

ABSTRACT

Puccinellia distans subsp. borealis (Northern Saltmarsh Grass) is distinguished from subsp. distans by its shorter
culms and narrower leaves and in its native habitat the culms of the former are frequently procumbent. It has
mostly contracted, dense and narrow panicles as opposed to the open character of the panicles of subsp. distans.
Refliexed branches are frequent in subsp. distans but infrequent in subsp. borealis. The lemmas of subsp. distans
are marginally smaller. The predominant natural habitat is identified as cracked seashore pavement of Old Red
Sandstone rock and the importance of the man-made habitat on harbour and slipway stone structures is
emphasised. The latter habitat probably controls the distribution of the plant. It is only rarely found in the
saltmarsh habitat.

KEyworbDs: taxonomy, habitat, phenology, distribution.

INTRODUCTION

It is considered that the first British collection of Puccinellia distans subsp. borealis was made by
Ralph Tate under the name of Sclerochloa distans from Balta Voe, Unst, Shetland in 1865 (BM). E.
S. Marshall collected it as Glyceria distans Wahl. on “‘cliffs c. half a mile south of Wick, Caithness”
and also at Melvich, West Sutherland, both sheets of 1886 in BM and CGE. In 1888 W. H. Beeby
collected it as Glyceria distans Wahl. ‘“‘on the stoney shores of Hildasay Is., Shetland’”’ (BM) and in
E. S. Marshall’s collection in CGE a sheet is labelled “ccomm. W. H. Beeby leg. 1888, E. S.
Marshall’. Further collections were made by F. J. Hanbury at Uyeasound, Unst, Shetland, 1894,
who also gave a sheet to Marshall (CGE). One of us (P.J.O.T.) has considered these specimens and
is satisfied that they are all the same taxon, namely Puccinellia distans subsp. borealis. In Hubbard &
Milne-Redhead (1968) the title “A Puccinellia new to Britain” is misleading. Credit can be given to
Beeby for publishing a short account under Glyceria distans var. prostrata in 1895 but he was not the
first to collect the taxon. He had noted the prostrate growth in his first collection and later obtained
plants from F. J. Hanbury from which he took seed. From his plants in cultivation Beeby (1895)
recorded “‘the prostrate habit is not so pronounced as in the wild specimens’. Nevertheless he
mistakenly published Glyceria distans var. prostrata Beeby.

Subsp. borealis was not included in Hubbard (1954) and first appeared under Puccinellia capillaris —
(Liljebl.) Jansen in Hubbard (1968). He was first alerted to this taxon in 1955 when he received
specimens from the Isle of May off the coast of Fife (v.c. 85). In 1962 Elaine Bullard sent a specimen
from Sule Skerry, Orkney to Hubbard who identified it as Puccinellia capillaris. Subsequently
Bullard (1968) found the same taxon on the coast of all Orkney Islands. Hubbard & Milne-Redhead
(1968) reported it from the coasts of Caithness, West Sutherland, Banff and Moray.

392 P. J. ©) TRIST ANDY BUTLER
DESCRIPTION OF PUCCINELLIA DISTANS SUBSP. BOREALIS, NORTHERN SALTMARSH GRASS

A loose to compact perennial. Culms spreading, procumbent or erect, 2-40 cm long, 2-4 noded,
smooth. In natural habitats the culms seldom exceed 28 cm long. Leaves grey-green, glabrous,
sheaths smooth, slightly inflated, striate; ligules 0-4-2-2 mm long, membranous; blades blunt-
tipped, 3-12 cm long, seldom flat and mostly plicate, 1-2 mm wide but width in small plants is <1-0
mm. Panicles mostly contracted, erect and dense throughout, 2-12 cm long; when panicles exert,
the branches at the lower nodes are patent; pedicels 0-8-4-8 mm long, scabrid.

Spikelets oblong, 2-5-9 mm X 1-5-2 mm, 2-8 flowered. Glumes slightly unequal, ovate to elliptic;
lower 0-5-2 mm long, 1|-nerved; upper 1-3—2-8 mm long, 3-nerved. Lower lemmas oblong to elliptic,
blunt, (1-8—)2-2-2-8 mm long, minutely hairy at the base. Paleas about as long as lemma, hairy on
the keels. Anthers 0-4-1 mm long.

The following summarises the nomenclature:

Puccinellia distans (Jacq.) Parl. subsp. borealis (O. Holmb.) W. E. Hughes in Botanical journal of

the Linnean Society 76: 363 (1978).

Glyceria distans var. prostrata Beeby in Journal of botany (London) 33: 315-316 (1895).
Poa retroflexa Curtis subsp. borealis O. Holmb. in Botaniska notiser 26: 182 (1926).
Puccinellia distans var. prostrata (Beeby) Jansen & Wachter in Nederlandsch kruidkundig archief

40: 248 (1930).

Puccinellia capillaris (Liljebl.) Jansen in Flora Neerlandica 1: 2, 69 (1951).
Puccinellia borealis (O. Holmb.) A. Love & D. Léve in Botaniska notiser 128: 498 (1976).

HABITAT

THE NATURAL HABITAT

The most convincing natural habitat of Puccinellia distans subsp. borealis is at a low level on a rocky
seashore platform. On the British mainland it occurs around the coast of Caithness (v.c. 109),
having a specific preference for the flagstone pavement of the Old Red Sandstone. This fine-grained
sedimentary rock forms gently sloping flat surfaces fractured at intervals by long fissures typically a
few mm wide.

This grass occupies these fissures (as shown in Fig. 1) at the lowest level on the shore capable of
accommodating higher plants, i.e. 4m above Ordnance Datum. Here there is a sparse community in
which the constant species are Armeria maritima (Miller) Willd., Plantago maritima L., Puccinellia
distans subsp. borealis and Sagina maritima G. Don. Occasional associates are Aster tripolium L.
and Plantago coronopus L. This community occupies about 1 m of vertical height and gives way at
higher levels to a similar but richer community in which the dominant grass is Festuca rubra subsp.
juncea (Hackel) K. Richter which is responsible for some displacement of the P. distans. In this
higher zone Agrostis stolonifera L. and Tripleurospermum maritimum (L.) Koch occur in addition
to Carex distans L. and Euphrasia foulaensis F. Towns. ex Wettst. at the top of the zone some 7 m
above Ordnance Datum level.

On the Scottish mainland good examples of this community in platform cracks were found at
Proudfoot ND/383.510 (see Fig. 2) and at Thurso East ND/143.701 (both v.c. 109, Caithness); such
communities occur all round the coast from Reay in north-west Caithness to Helmsdale on the east
coast of Sutherland wherever the exacting conditions of sloping pavement at the correct intertidal
height occur. It is also the dominant natural habitat on seashores in Orkney.

Subsp. borealis is also to be found in saltmarshes but its occurrence in this habitat is uncommon.
Saltmarsh itself is rare due to the combination of a rocky wave-exposed shore and the prevalence of
coarse sand on the sea bottom. On this account subsp. borealis is seldom found in association with
Puccinellia maritima (Hudson) Parl.; where it does occur on Orkney the former has a hazardous
chance of withstanding the competition of a close sward created by the stolons of P. maritima. This
association in saltmarsh is known in Orkney and Shetland. It is reported by Bullard (pers. comm..,
1987) ‘“‘in saltmarsh with open areas of bare mud”’ at Westayre Loch, Sanday, 1966; “‘in muddy
saltmarsh at the edge of a small loch’, Sandsend, Shapinsay, 1966 with Juncus bufonius L. and
Spergularia marina (L.) Griseb.; and on saltmarsh at Sebay Mill, Tankerness, 1966. In 1982 Bullard
sent specimens (now in herb. P.J.O.T.) of subsp. borealis from “open areas of saltmarsh on

PUCCINELLIA DISTANS SUBSP. BOREALIS IN SCOTLAND 393

SSG

NAW
S

N
WS WW NN SN
RN

eS
Ss

Figure |. Puccinellia distans subsp. borealis in association with Armeria maritima in the cracks of Old Red
Sandstone natural paving at Proudfoot, Wick.

s

Y SRO a
&
%

Sg

M2

SS

| FIGURE a Individual

a

.
SN

SS

plants of Puccinellia distans subsp. borealis in crevices. The rock is partly coated with tar.

394 P. J.O. TRIST AND J. K. BUTLER

the east side of Cata Sound, Orkney” which were associated with Agrostis capillaris L., Armeria
maritima, Juncus gerardii Loisel., Plantago maritima and Puccinellia maritima.

This uncommon association between the two species of Puccinellia is also recorded by Scott &
Palmer (1987) ‘“‘about the tiny brackish pool behind the jetty at Grutness’’, Shetland and by Gilbert
& Holligan (1979) on Rona, North Ebudes, v.c. 104.

Subsp. borealis is less common on sand and fine stones, but does occur on such substrates, for
example on the sloping banks of the tidal River Halladale at Bighouse, West Sutherland NC/
890.649. E. S. Marshall also records finding occasional plants ‘“on muddy decomposed boulder
clay” south of Wick in 1886 (CGE) and also “‘in mud among stones at the rivermouth at Dunbeath”’
ND/165.295, both in Caithness. It also occurs on the stony shore on the western side of Loch Eriboll,
v.c. 108, W. Sutherland. There are two examples, at Janetstown and Lybster on the east coast of
Caithness of the plant being found among larger (c. 150 mm) waveworn boulders which did not have
a matrix of silt or other fine material; this situation is, however, uncommon.

MAN-MADE HABITAT

The natural habitat occurs around 1 m above mid tide level, which is also the level at which the top of
a harbour quay is constructed. This proves to be a very good habitat for Puccinellia distans subsp.
borealis. There are many records from small quays, slipways and stonework on larger harbour walls.
The quay top is often a stone pavement and the pattern of local distribution suggests that the seed is
carried by waves flooding over the quay. Plants are to be found mainly in the crevices of the
pavement and also on tracks and rough ground within reach of the swell. Bullard (pers. comm.,
1987) found the plant in two locations amongst Leymus arenarius (L.) Hochst. in coastal dunes some
distance from the swell region. The plant may occasionally act as a short-lived ruderal when carried
by vehicle tyres from a harbour area.

In the quay top community which ts on the horizontal, the plant association is richer than in the
natural sloping pavement habitat. Armeria maritima, Plantago maritima and Sagina maritima are
the constant associates but Aster tripolium, Cochlearia anglica L., Matricaria discoidea DC.,
Plantago coronopus, Potentilla anserina L., Spergularia marina and Tripleurospermum maritimum
typically occur. In this habitat the Puccinellia distans subsp. borealis is able to survive alongside
Festuca rubra subsp. juncea, whereas in the natural habitat of the sloping rock platform they tend to
occupy different levels. A good example of a rich community is the harbour quay at Keiss in
Caithness ND/351.609 where thousands of plants of subsp. borealis were seen in July 1993.

A comparison of the vertical distribution of the plants in the natural and man-made habitats
shows that the horizontal surface of a quay allows the species a better foothold at higher elevation
above mid-tide.

The success of the man-made habitat is clear. It can be said that almost every quay and slipway on
the far north and east coast of the mainland and on Orkney and Shetland are candidate sites for at
least a few plants of subsp. borealis. This habitat is the main source of new seed and probably the
basis of the present distribution. Numerically the distribution seems to be centred on north-east
Caithness, Orkney and Shetland where the population is sufficiently dense to provide seaborne seed
for wider distribution.

PHENOLOGY

Davis (1983) records that in species of Puccinellia ‘growth and flowering are continuous throughout
the summer and simultaneous among varied microhabitats within populations”. The various
maritime habitats in which subsp. borealis is found shows its ability to adapt to environmental
variability.

The sea shore by the harbour at Lybster, Caithness, v.c. 109 has a man-made earth-stone boat
slipway. A single plant of subsp. borealis collected on 16 July 1982 was 20 cm high overall with 17
fresh green panicles. All culms were erect and panicles were in varying stages of development from
first break of sheath to open panicles. Within a few metres five plants of 4-8 cm high had 1-3 panicles
per plant, all of which had one or more dead panicles and others in stages of dying or development.
In another collection taken the same day at Janetstown on the east coast of Caithness, where the
large-shingle beach was similar, subsp. borealis was found between stones on a bank above and on

PUCCINELLIA DISTANS SUBSP. BOREALIS IN SCOTLAND 395

the harbour wall. Four plants of 6-10 cm in height were taken and three had single dead panicles and
leaves; on the other plant there were two dead panicles and three panicles in stages of dying. These
records illustrate the continuous growth of individual plants of subsp. borealis which may be seen
from mid-June to mid-August in all the types of maritime habitats.

Davis (1983) also discusses the periods of moisture stress which arise from varying frequencies of
tidal wash. In addition there are variations in salt stress on plants where salt is retained in crevices as
opposed to direct drainage through shingle and sand beaches at river mouths. There may also be a
difference in salt retention in natural platform crevices compared to quay top crevices because the
latter lie on a rubble base.

Procumbent growth is noticeable on beach habitats subject to tidal flow and this appears as a
reaction to the tidal thrust over the rocks. Plants out of reach of seawater flush, but within the spray
and swell of high tides, frequently have erect growth.

Davis (1983) also records that species of Puccinellia “‘are patchily distributed due to the
discontinuous and restricted occurrence of suitable habitats”. This is demonstrated on the north
coast of the Scottish mainland. Between Loch Eriboll in West Sutherland and Reay in Caithness the
rocks are from the Moine Succession or igneous intrusions related to it. Here, Puccinellia spp. are
very sparse. Between Reay and Scrabster the shore is mostly vertical cliff. But between Scrabster
and John o’Groats there is an almost continuous occurrence of flat rocky platforms of Old Red
Sandstone where subsp. borealis can be found whenever these platforms have the preferred
intertidal height. On the east coast of Caithness from John o’Groats to Dunbeath there are many
stretches of vertical cliff but any occurrence of suitable rocky platforms is likely to have some
specimens of the plant. South of Dunbeath there are few suitable natural habitats since cliff, boulder
beach or fine sand prevail.

DISTRIBUTION

Puccinellia distans subsp. borealis is reported from the north of the Outer Hebrides, v.c. 110 on the
Butt of Lewis, Uig and at Stornoway, but there are no records for the North Ebudes, v.c. 104 or the
west coast of Scotland. It occurs frequently on the coasts of the Orkney Isles (v.c. 111), on North
Rona (v.c. 110) to their west, on Fair Isle, Foula and Shetland (v.c. 112). On the north coast of
Scotland it is found on the shores of Loch Eriboll and then eastward at intervals to Bighouse (v.c.
108) almost at the western boundary of Caithness. It is seen at Brimsness and Scrabster. Eastward
from Scrabster it is locally plentiful along to John o’Groats. South from John o’Groats it occurs at
intervals, being plentiful around Wick and occurring at Lybster, Dunbeath, Helmsdale and as far as
Golspie. Then again it can be found at Hopeman and Peterhead on the coast of Banff (v.c. 93, N.
Aberdeen) and at intervals down to Fife (v.c. 85), where it occurs frequently between Crail and
Pittenweem and on the Isle of May to the east. It is recorded from Rosyth, Fife and at Leith Docks,
(v.c. 83, Midlothian), also on the Bass Rock in the Firth of Forth and at Tyninghame, East Lothian
(v.c. 82).

The plant is known from the Faeroes, Iceland, Greenland, Norway and on the coast of the
Netherlands.

The hybrid between Puccinellia distans (Jacq.) Parl. subsp. borealis (O. Holmb.) W. E. Hughes
and P. maritima (Huds.) Parl., named Puccinellia X mixta O. Holmb., has been found on the
Fianuis peninsula on North Rona, Outer Hebrides, v.c. 110 by Gilbert & Holligan (1979). It is also
recorded from Denmark, Holland, Iceland, Norway and Sweden (Jones & Stace 1975).

ACKNOWLEDGMENTS

We are grateful to the curators of BM and CGE for access to specimens and information. We thank
D. H. Kent for assistance with nomenclature and Elaine Bullard for field notes and discussions and
also Elizabeth Norman for help with field work over several years. Charles Turner kindly
commented on the manuscript.

396 P. J.O. TRIST AND J. K. BUTLER
REFERENCES

Beesy, W. H. (1895). Glyceria distans var. prostrata Beeby. Journal of botany (London) 33: 315-316.

BULLARD, E. R. (1968). Notes on the occurrence of Puccinellia capillaris (Liljebl.) Jansen. Bulletin of Orkney
Field Club 4: 5.

Davis, J. J. (1983). Phenotypic plasticity and the selection of taxonomic characters in Puccinellia (Poaceae).
Systematic botany 8: 341-353.

GILBERT, O. L. & HOLLIGAN, P. M. (1979). Puccinellia capillaris (Liljebl.) Jansen x P. maritima (Huds.) Parl.
on North Rona, Outer Hebrides. Watsonia 12: 338-339.

HuBBARD, C. E. (1954). Grasses, 1st ed. Penguin, Harmondsworth.

HussarpD, C. E. (1968). Grasses, 2nd ed., p. 199. Penguin, Harmondsworth.

Hupparb, C. E. & MILNE-REDHEAD, E. (1968). A Puccinellia new to Britain. Proceedings of the Botanical
Society of the British Isles 7: 507.

Jones, B. M. G. & Stace, C. A. (1975). Puccinellia capillaris (Liljebl.) Jans. x P. maritima (Huds.) Parl. = P. x
mixta Holmberg in Stace, C. A. ed. Hybridisation and the flora of the British Isles, p. 559. Academic Press,
London.

Scott, W. & PALMER, R. C. (1987). Puccinellia distans (Jacq.) Parl. subsp. borealis (O. Holmb.) W. E. Hughes
in Scott, W. & PALMER, R. C. The flowering plants and ferns of the Shetland Islands, p. 357. The Shetland
Times, Lerwick.

(Accepted September 1994)

Watsonia 20: 397-404 (1995) 397

A graphical analysis of the characters of Calamagrostis stricta
(Timm) Koeler, C. canescens (Wigg.) Roth and their hybrid
populations in S. E. Yorks., v.c. 61, northern England

F. E. CRACKLES

143 Holmgarth Drive, Bellfield Avenue, Hull, HU& 9DX

ABSTRACT

Population samples of Calamagrostis stricta (Timm) Koeler, C. canescens (Wigg.) Roth and the hybrids C.
canescens X C. stricta (Poaceae), 2n = 28 and 2n = 56, as well as a strongly introgressant C. canescens population
along Leven Canal, S.E. Yorks., v.c. 61 are described using methods of polygraphic analysis: pictorial scatter
diagram, hybrid index analysis and polygonal graphs. A polygonal graph is also given for a recombinant
represented by a single plant. The overall intermediate nature of the hybrids is demonstrated as well as the loose
non-random association of characters. The nature of the hybrids, the effect of polyploidy and the relative merits
and limitations of the different methods of polygraphic analysis used are discussed. Evidence for a C. canescens
population being strongly introgressed is assembled.

KEyworpbs: Poaceae, introgression, multivariate analysis.

INTRODUCTION

In an earlier paper (Crackles 1994) I described a diverse group of Calamagrostis populations along
the Leven Canal in S.E. Yorkshire (v.c. 61) and showed that these could be assigned to six ‘taxa’: C.
canescens (Wigg.) Roth, C. stricta (Timm) Koeler, C. canescens X C. Stricta 2n = 56 (H,
population), C. canescens X C. stricta 2n = 28 (H2 population), a recombinant individual plant (H3)
and an introgressed C. canescens (1, population). These could all be identified in the field.

To explore in more detail the relationship between these various populations and the characters
by which they differed, I undertook a comparison of population samples by graphical methods of
analysis — using pictorial scatter diagrams, hybrid index analysis (Anderson 1949) and polygonal
graphs. The results provide some further insights into the relationships of the Leven Calamagrostis
populations, as well as illustrating the continuing value of these relatively simple graphical methods
and indicating their relative weaknesses and strengths.

METHOD OF STUDY

25 shoots of C. stricta, C. canescens and the putative hybrid populations H, and H; and 20 shoots of
the I, population (Crackles 1994) were collected at random. Material was collected at the end of July
or in early August 1970, when the panicles were in their after-flowering condition so that
comparison of the maximum number of floret characters could be made. The range of measure-
ments, the mean and the standard deviation for various characters are given in the previous paper
(Crackles 1994).

As axes for each scatter diagram (Figs 1 & 2), the ratio of bract length to bract width was plotted
against the ratio of panicle length to length of basal branch of panicle. Ratios were used to minimise
the possible effect of varying environmental conditions on lengths of structures. Other information
was added to the scatter diagram to produce metroglyphs.

In the case of the ratio of glume length to glume width, and awn length, the intermediate range
was Calculated by dividing the range between the standard deviations for the two species by four and
taking the median two quarters as the intermediate range.

398 F. E. CRACKLES

Length of panicle
Length of basal branch of panicle

6.0
e@
e C. stricta
o C. canescens
w Hybrid 1
® O Hybrid 2
5.0
om e 6
e
ats e e
e
. oO
=
<= °
4.0 e@ @

“i

+ $
Gree ee
ee, oe grt open

2.0 3.0 4.0 5.0 6.0 7.0 8.0

Ficure 1. Scatter diagram for population samples of Calamagrostis stricta, C. canescens and C. canescens X C.
stricta, Hybrid 1 and Hybrid 2 (H, and Hj populations).

Callus hairs: shorter than lemma @; + equal to lemma @; longer than lemma 9.

Awn insertion from: bottom 37-:5% of lemma @; 37-5%-85-0% way up lemma @; near tip of lemma ©.
Awn length: >1-:7 mm @; 0-9-1-7 mm @; <0-9 mm @.

Glume length/glume width: <4-5 @; 4-5-6-0-@; >6-0 -®.

A more arbitrary method had to be used to determine the intermediate range for the position of
awn insertion on the lemma. It was known from observation that the awn is inserted just below the
middle of the lemma in the hybrid, C. canescens X C. stricta. In C. canescens the awn is almost
apical, while in C. stricta it is inserted 25%-35% along the length of the lemma from its base. The
intermediate range was defined by dividing the range 25%-S0% into two and taking the lower half
as C. stricta and the upper half as intermediate. |

The hairs surrounding the floret (callus hairs) are generally at least 0-7 mm longer than the lemma |
in C. canescens and substantially shorter than the lemma in C. stricta. Callus hair length between 0.4
mm longer and 0.4 mm shorter than the lemma was taken to be an intermediate character.

The method of hybrid index analysis is described by Anderson (1949) and consists of selecting a
number of characters with respect to which the species differ, and recording each character of each
plant as identical with that of one of the species or as intermediate between them. Each character is
then assigned an arbitrary value. When each of the characters of the plant listed has received a
score, the total index score of the plant is obtained by summing the scores of all its characters.

In selecting characters the following principles were borne in mind:

a

1. in obtaining an index score for an individual plant or a population it is essential that as many |
characters as possible are taken into account; |

2. an index score for each character used must be obtainable for each population analysed;

3. characters which can be observed with the naked eye or with a hand lens and structures which can

ANALYSIS OF THE CHARACTERS OF CALAMAGROSTIS 399

Length of panicle
Length of basal branch of panicle

5.0 A
a a C. canescens
a 14 population
ray
4.0 ,
ad
A
a
A ad S
A A
a fas a
A
3.0 A A A A
A «& A A
a A AA
A 4
A A 2 Bract length
rs A A A Bract width

2.0 3.0 4.0 5.0 ee eS

Figure 2. Scatter diagram for population samples of Calamagrostis canescens and a suspected C. canescens
introgressant (I, population).

be measured without the use of a microscope are most useful and were used whenever possible;
and

4. floret characters were included in order to achieve coverage of important features although
measurements had to be made using a binocular microscope.

TABLE 1. METHOD OF SCORING THE HYBRID INDEX FOR CALAMAGROSTIS TAXA

C. stricta Intermediate C. canescens

Character Score 0 Score 1 Score 2
Panicle length (cm) <14-0 14-0-15-5 S155
Panicle width (in fruit) (cm) <1-6 1-6-2:5 >2°5
Length of basal branch of panicle (cm) <4-0 4-0-5-1 >5-1
Culm rough near panicle + =
Culm width at 2nd node (mm) <1:6 1-6-2-0 >2:0
Bottom leaf sheath hairy te =
Glume length (mm) <4-0 4-0-4-5 >4:5
Glume width (mm) >0-:90 0-55—0-90 0-60—0-70
Glume length/glume width <4-5 4-5-6-0 >6:0
Callus hairs <floret + = floret >floret
Awn length (mm) 5/187/ 0-9-1-7 <0:9
Awn insertion** <37-5% 37:5%-85-0% >85%
Ligule length (mm) <3-0 3-0-3-6 >3-6

*scored as 1.
** awn insertion measured from base of floret as % of floret height.

400 F. E: CRACKLES

12 H, population
10

8

6

4

2

0

Hybrid index score

12 Hz population
10

8

6

4

v2

0

Hybrid index score

Number of individuals

Number of individuals

12 1, population

Number of individuals

2 4 6 8) 10)" (12 (14 Ge aS 20)— 22524
Hybrid index score

Ficure 3. Hybrid index scores for Leven Calamagrostis canescens X C. stricta, H, and H>, and the C. canescens I,
population samples.

A list of characters used and the method of scoring are given in Table 1. The intermediate range
for each character was determined in the same way as for the scatter diagrams. The frequency
distribution of hybrid index scores for the H;, H2 and I, populations are shown in Fig. 3.

A polygonal graph was constructed for each of the Leven Calamagrostis populations studied and
for a single plant, the only one of its kind found, H; (Fig. 4) The method of scoring is given in Table
2. In using this method, characters in which a hybrid shows a range of measurements which is not
intermediate between those of the parents can be included and I have selected lemma length as one
of the eight characters. The lemma of individuals of the H; population is longer than that of both
parents (Crackles 1994). ;

ANALYSIS OF THE CHARACTERS OF CALAMAGROSTIS 401

Pre.

A.L.
C. stricta C. canescens
Hz population H, population
H3 plant 1, population

Figure 4. Variation in eight characters of the Leven Calamagrostis taxa. Each radius has six divisions (see Table
2). In the case of each Leven Calamagrostis population sample, except the H;, the polygonal graph was drawn
using the mean of the measurements for each character; standard deviations are shown along each axis. The
polygonal graph for the H3; was drawn using measurements for one plant only. Abbreviations are given in
Table 2.

RESULTS

The overall intermediate morphological nature of the H, and H; populations between that of the
two species is demonstrated by the intermediate shape of the polygonal graphs for the two hybrids
(Fig. 4) and by the range of hybrid scores for both hybrid populations (Fig. 3).

The tendency of intermediate characters of the H,; and H; populations to be inherited together,
but not invariably so, is shown for four characters by the use of intermediate length arms attached to
dots on the scatter diagram (Fig. 1). By examining these four characters, a pair at a time (Table 3), it
is seen that such intermediate characters have a strong tendency to remain together, but that the

402 F.-E} CRACKLES

TABLE 2. METHOD OF SCORING FOR POLYGONAL GRAPHS OF CALAMAGROSTIS TAXA

Radius division from the centre of a circle*

Character 0 1 2 3 4 5
Panicle length (P.L.) (cm) 5-9-8-9 9-0-12-0 12-1-15-1 15-2-18-2 18-3-21-3 21-4-24-4
Length of basal branch

of panicle (P.Br.) (cm) 1-2-2-4 2:5-3-7 3-8-5-0 5-1-6:3 6:4-7-6 7:7-8-9
Glume length (G.L.) (mm) 2-6-3-1 3-2-3-7 3-8-4-3 4-4-4.9 5-0-5-5 5-6-6:1
Lemma length (L.L.) (mm) 2:5—2-6 2-7-2°8 2-9-3-0 3-1-3-2 3-3-3-4 3-5-3-6
Awn length (A.L.) (mm) 0-3-0-6 0-7-1-0 1-1-1-4 1-5-1-8 1-9-2-2 2-3-2-6
Awn insertion** (A.1.) 27%-38% 39%-5S0% 51%-62% 63%-74% 715%-86% 87%-98%
Ligule length (Li.L.) (mm) 1-3-1-8 1-9-2-4 2:5-3-0 3-1-3-6 3-7-4-2 4-3-4-8
Callus hairs (C.H.) (mm) 1-9-2-1 2-2-2:5 2-6-2:9 3-0-3-3 3-4-3-7 3-8-4-1

*see Fig. 4.
** measured from the base of the floret as a % of floret height.

TABLE 3. THE FREQUENCY OF ASSOCIATION OF CHARACTERS WITHIN TWO POPULATIONS
(H, AND H,) OF CALAMAGROSTIS, EXAMINED TWO AT A TIME

Percentage association*

Associated character pair H, H, H, and H,
Callus hairs and awn length 80 84 82
Awn length and awn insertion 64 84 74
Callus hairs and awn insertion 1: Fe 72
Glume ratio and awn insertion 76 60 68
Glume ratio and awn length 80 64 IP
Callus hairs and glume ratio 96 60 78

* percentages calculated using the number of short ‘arms’, representing intermediate states, on the scatter
diagram shown in Fig. 1.

strength of the affinity between these varies not only from one pair of characters to another, but also
usually from one hybrid population to the other.

The H, and H, populations were found to be morphologically distinct (Crackles 1994).
Differences between these two populations are further demonstrated by:

a. the fact that most individuals of the H population appear in a different part of the scatter diagram
from those of the H, population (Fig. 1); and
b. differences between the polygonal graphs for the two populations.

The differences between the polygonal graphs for the hybrid populations H, and Hj is mainly one
of size, there being a general similarity of shape except for the difference caused by the much greater
lemma length in the H, population.

Morphological evidence has already been given to support the view that the C. canescens |,
population is introgressed (Crackles 1994). Further evidence of this view is provided by graphical
analysis:

a. over half of the individuals of the population occur outside the area occupied by typical C.
canescens individuals on the scatter diagram (Fig. 2);

b. a hybrid index range of 13-19 was obtained for the I, population where C. stricta = 0 and C.
canescens = 23; and

c. a polygonal graph for the I, population (Fig. 4) while of the same shape as that for typical C.
canescens shows a shift towards the hybrid range for some characters.

ANALYSIS OF THE CHARACTERS OF CALAMAGROSTIS 403

The polygonal graph for the Leven H3, represented by a single plant, confirms that this is a
recombinant. The position of awn insertion was as in C. canescens and the awn length as in some C.
canescens I, individuals while the panicle and ligule length were as in C. stricta. The basal branch of
the panicle and the glume were intermediate in length, being within the range for these characters in
the C. canescens X C. stricta H, population. The callus hairs tended to be shorter than in C. stricta.

DISCUSSION

The loose non-random association of characters in both the H, and H populations demonstrated by
the scatter diagrams (see Table 3) is considered by Anderson (1949) to be critical evidence of
hybridization. This evidence together with the intermediate nature of many characters of these taxa
(Crackles 1994) and their overall intermediacy demonstrated by the polygonal graphs and their
hybrid index range, (H; 10-15, mode = 15; Hz 8-14, mode = 13; C. stricta = 0, C. canescens = 23),
as well as the fact that they occur in the same locality as both parental species, leave no doubt that
both the H, and H; populations are the hybrid C. canescens X C. stricta.

Comparison of the polygonal graphs for the H, and H, populations suggest that the main
difference between the two taxa is one of size of several characters.

The chromosome number of the Leven C. stricta, C. canescens and the H, C. canescens X C.
stricta populations is 2n = 28 (Crackles 1994), a number Nygren (1946) found to be constant for the
two species. The chromosome number for the Leven H, C. canescens X C. stricta is, however,
2n = 56 so that this taxon is an octoploid (Crackles 1994).

The question of the likely effects of polyploidy on characters of the H, population thus arises and
whether such effects may be the full explanation of differences between the H, and H; populations.
Stebbins (1971) drew attention to the varying effects of polyploidy in different genotypes and also to
the fact that gigas effects are seen most often and most strongly in organs with a determinate type of
growth. It would seem that polyploidy is the explanation of increased lemma length which is greater
than in either parent. It is also the probable explanation of the longer and wider glumes while an
intermediate shape is retained and of greater callus hair length which approaches that of C.
canescens while remaining more or less the same length as the lemma as in other hybrid individuals.
A tendency to greater plant height, greater panicle length and greater awn length would also seem
to be due to polyploidy. Features frequently resulting from polyploidy and not occurring in the H,
population are later flowering and reduction of branching (Stebbins 1971): The H, C. canescens xX
C. stricta individuals have a branching culm as does C. canescens and the flowering time is
intermediate between that of the two species while the H, population flowers later, only slightly
earlier than C. canescens or at the same time (Crackles 1994). The H, population is vegetatively
vigorous and this is thought to be a result of polyploidy as the H, population is not notable in this
respect.

There is some overlap of C. canescens X C. stricta H, and C. canescens individuals on the scatter
diagram (Fig. 1) and this might be taken to indicate some backcrossing to C. canescens, but the
hybrid index scores point to the strictly intermediate nature of this taxon.

The position of most individuals of the H, population on the scatter diagram mainly away from
individuals of other Calamagrostis taxa sampled is of considerable interest, although the significance
of this is obscure. This positioning of H, individuals on the scatter diagram results partly from the
fact that their bract shape resembles that of C. stricta while H, individuals may resemble C.
canescens in this respect. The ratio of panicle length to length of basal branch of panicle in the H,
individuals is similar to that for most C. canescens individuals while H, individuals tend to be
intermediate in this respect. The low figure for this ratio in H, individuals is mainly due to a
tendency for the basal branch of the panicle to be markedly longer than in H, individuals and a
possible explanation of this is that the C. canescens parent had an exceptionally long lowest branch
to the panicle. The scatter diagram proved to be a powerful tool in separating individuals of the two
species and most of those of the two hybrids.

It is a limitation of hybrid index analysis that individuals with the same hybrid score may have
different genetic constitutions and that this is true both for members of the same population and for
those of different populations. However the hybrid index range in this study is important in pointing

404 F. E. CRACKLES

to the overall intermediacy of the two hybrids. The slightly higher hybrid score for H, compared
with H, individuals is likely to be due to polyploidy.

The polygonal graph describes a population as a characteristic shape which the eye can take in ata
glance. By comparing different polygonal graphs differences between taxa with regard to a fixed and
limited number of characters can be seen at the same time. Polygonal graphs in this study were
particularly important in demonstrating the effect of polyploidy in the H, population.

ACKNOWLEDGMENT

I am indebted to Dr D. W. Shimwell for advice and encouragement during the early stages of the
research.

REFERENCES

ANDERSON, E. (1949). Introgressive hybridisation. Wiley, London and New York.

CRACKLES, F. E. (1994). Calamagrostis stricta (Timm) Koeler, C. canescens (Wigg.) Roth and their hybrids in
S.E. Yorks., v.c. 61, northern England. Watsonia 20: 51-60.

NyYGREN, A. (1946). The genesis of some Scandinavian species of Calamucroue Hereditas 32: 131-262.

STEBBINS, G. L. (1971). Chromosomal evolution in higher plants. Arnold, London.

(Accepted January 1995)

Watsonia 20: 405-418 (1995) 405

Notes

PETRORHAGIA PROLIFERA (L.) P. W. BALL & HEYWOOD (CARYOPHYLLACEAE),
AN OVERLOOKED NATIVE SPECIES IN EASTERN ENGLAND

Recent years have seen several additions to the native flora of Britain, either as a result of new
geographical records or the elucidation of taxonomic problems. Petrorhagia prolifera (L.) P. W.
Ball & Heywood, Proliferous Pink, reported here as a native species, represents a mixture of both.
Floras published before 1962 recorded P. prolifera, under the names Dianthus prolifer L., Tunica
prolifera (L.) Scop. or Kohlrauschia prolifera (L.) Kunth, as a native plant, at least on the south
coast.

However, plants on coastal shingle beaches from Hampshire to Kent have been shown
subsequently to belong to the closely related P. nanteuilii (Burnat) P. W. Ball & Heywood, Childing
Pink (Ball & Heywood 1962). This species is now restricted in Britain to W. Sussex (v.c. 13).
P. nanteuilii can be distinguished from P. prolifera by several small but constant morphological
features, notably the tuberculate rather than reticulate seed testa, and by a chromosome number of
2n = 60 as opposed to 2n = 30 (Ball & Heywood 1962, 1964).

Evidence derived from morphological, cytological and geographical data and from hybridization
experiments suggests strongly that P. nanteuilii is an allotetraploid derived from P. prolifera and
another diploid species, P. velutina (L.) P. W. Ball & Heywood (Akeroyd 1975; Thomas 1983).
P. velutina, which has smaller, echinate seeds and a chromosome number of 2n = 30, is widespread
in the Mediterranean region and southern Europe, but does not occur in Britain. Two other closely
related species are endemic to the Balkan Peninsula. The most recent revision of the genus (Ball &
Heywood 1964) and the second edition of Flora Europaea Volume 1 (Ball & Akeroyd 1993) include
these five annual species within Petrorhagia, as section Kohlrauschia. However, some continental
botanists retain Koh/rauschia as a distinct genus.

It has recently become clear that two species of Petrorhagia section Kohlrauschia are present in
Britain. Each of us had concluded independently, together with Dony & Dony (1986), that a
Petrorhagia species is native inland in eastern England. Beckett (1992) reported on the status of an

- extant population of P. nanteuilii in West Norfolk (v.c. 28), suggesting that it was native. The plant

had been reported, as Tunica prolifera (L.) Scop., by Trimmer (1866) from between Stanhoe and
Bircham and from Fincham in the same part of the county, and was familiar to Norfolk botanists up
until 1950, although not seen again until 1985. Examination by J.R.A. of collections of Petrorhagia
in the herbaria of the Universities of Cambridge (CGE) and Reading (RNG), the Natural History
Museum, London (BM) and the Castle Museum, Norwich (NWH), including a comparison of seed
testas with a sample collected by G.B. in Norfolk in 1992, has confirmed that the Norfolk plant is
indeed P. prolifera. A preliminary note of these observations has been published elsewhere
(Akeroyd 1993).

P. prolifera, recorded mostly in eastern England, had long been regarded as a casual or locally as
an established alien. The species, now apparently reduced in Britain to two populations, has its
British headquarters in a few adjacent parishes in the Breckland of West Norfolk (v.c. 28).
P. prolifera is a species with a central to south-eastern European distribution, extending northwards

' to Denmark and the Swedish Baltic islands of Oland and Gotland. Its presence in eastern England is
’ therefore not unexpected, especially in the Breckland, famous for its native flora of species of

central European affinity (Trist 1979). A second population in Bedfordshire (v.c. 30) is less

| convincingly native, but may derive from one or more extinct populations, or from seed introduced
_ from Norfolk.

P. nanteuilii has a western Mediterranean and Lusitanian distribution, reaching its northernmost
limit in the Channel Islands and on the southern coast of England. A further station recently

406 NOTES

reported from a railway embankment in Giamorgan (v.c. 41) probably represents an introduction

via Cardiff Docks, where the species has occurred as a casual (Dawson 1988).

The habitats of both species in Britain, as on the continent, are dry, open or stony places and dry
grasslands. At Pagham Harbour, Sussex (fide J.R.A.), P. nanteuilii grows in sparse, open plant
communities on stabilized shingle. In Norfolk P. prolifera grows in dry, rather sparse grassland on a
sandy soil; in Bedfordshire on open ground on sand and railway ballast.

We have seen the following putative native specimens of P. prolifera from Britain:

W. Norfolk (v.c. 28): Cockford Heath, 30 September 1835, K. Trimmer, CGE; Northwold, gravel
pit, W. J. Cross, 9 August 1889, BM; Stoke Ferry, W. J. Cross, July 1890, August 1891, BM; nr
Northwold, J. E. Little, 19 September 1927, BM, CGE, det. P. W. Ball; Cranwich, ‘ground
reverting to breck’, E. L. Swann 2127, 28 July 1950, NWH. There is a record from Mundford,
reported by Mrs Gomershall in her 1951 Wildflower Society Diary (fide G.B.); also, probably in
v.c. 27 (E. Norfolk), from ‘nr Norwich, Dr. Smith’ [Sir J. E. Smith (1759-1828)], BM.

Beds. (v.c. 30): Potton, abundant for over 0-5 mile (1 km) by disused railway, J. E. Lousley, 4
September 1974, RNG; Potton, south-facing bank and track of dismantled railway, J. G. & C. M.
Dony, LTN (Dony & Dony 1986); Potton (TL/210.489), old railway, on cinders and heaps of
ballast, G. Crompton, 23 July 1980, CGE. This population has been extensively damaged by sand
extraction (C. R. Boon, pers. comm., 1994).

The BM specimens from Norfolk were cited, as casuals, by Petch & Swann (1968). Two of them

had been determined as P. nanteuilii by P. W. Ball (in litt. to E. L. Swann, fide G.B.), but | |

examination of the seed testa by J.R.A. confirmed that they do indeed belong to P. prolifera.
P. prolifera was rediscovered in the county in 1985 by J. E. Gaffney at Cranwich, where 70 plants
were counted in 1992. The Bedfordshire population was observed until 1991 by C. M. and J. G.
Dony, who also regarded it as a probable British native (Dony & Dony 1986). The plants are
inconspicuous, of slender habit and with only one or two flowers within each inflorescence out at a
time, so may survive elsewhere undetected.

P. prolifera has also undoubtedly been introduced into Britain from time to time, behaving as a
casual, for example in railway sidings at Richborough, E. Kent (J. E. Lousley, 16 August 1936,
RNG). Herbarium specimens from Galashiels, Selkirk (M. McC. Webster 14263, 1970, CGE) and
Blackmoor, Hants. (e.g. J. E. Lousley, 5 October 1968, RNG) are all P. nanteuilii. These were
probably alien plants — part of the wool shoddy flora for which Blackmoor was famous during the
1960s to early 1970s (Ryves 1974, 1988).

P. nanteuilii is included on Schedule 8 of the Wildlife and Countryside Act 1981, which gives it full
protection in Britain. P. prolifera at present has no legal protection.

ACKNOWLEDGMENTS

We are grateful to Chris Boon, Mrs Chris Dony, the late Dr John Dony and Mrs Jean Gafiney for
providing data on P. prolifera. We also thank the curators of BM, CGE, RNG and NWH for allowing
us access to herbarium material of Petrorhagia. Chris Boon and Chris Preston kindly commented on
a draft of the manuscript.

REFERENCES

AKERoyD, J. R. (1975). The possible allopolyploid origin of Petrorhagia nanteuilit (Burnat) P. W. Ball &
Heywood. Pp. 26. Unpublished B.Sc. Thesis, University of St Andrews.

AKEROYD, J. R. (1993). Wildlife Reports. Flowering Plants. British wildlife 4: 261.

BALL, P. W. & AkeERoypD, J. R. (1993). Petrorhagia (Ser. ex DC.) Link., in Tutin, T. G. et al., eds. Flora
Europaea 1 (2nd ed.): 224-227.

BALL, P. W. & Heywoop, V. H. (1962). The taxonomic separation of the cytological races of Kohlrauschia
prolifera (L.) Kunth sensu lato. Watsonia 5: 113-116.

BALL, P. W. & Heywoop, V. H. (1964). A revision of the genus Petrorhagia. Bulletin of the British Museum
(Natural History). Botany 3: 121-172.

Beckett, G. (1992). Childing Pink, a Norfolk plant. B.S.B./. News 62: 20-21.

|
gE
)
.
5

NOTES 407

Dawson, H. (1986). Petrorhagia nanteuilii (Burnat) P. W. Ball & Heywood in Mid-Glamorgan. Watsonia 16:
174-175.

Dony, J. G. & Dony, C. M. (1986). Further notes on the flora of Bedfordshire. Watsonia 16: 163-172.

Petcu, C. & Swann, E. L. (1968). The flora of Norfolk. Jarrolds, Norwich.

Ryves, T. B. (1974). An interim list of the wool-alien grasses from Blackmoor, North Hants., 1969-1972.
Watsonia 10: 35-48.

Ryves, T. B. (1988). Supplementary list of wool-alien grasses recorded from Blackmoor, North Hants., 1959-
1976. Watsonia 17: 73-79.

Tuomas, S. M. (1983). A taxonomic clarification of Petrorhagia section Kohlrauschia (Caryophyllaceae).
Botanical journal of the Linnean Society 87: 55-75.

TrimM_R, K. (1866). Flora of Norfolk. London.

Trist, P. J. O., ed. (1979). An ecological Flora of Breckland. E.P. Publishing, Wakefield.

J. R. AKEROYD

Lawn Cottage, Fonthill Gifford, Tisbury, Wilts., SP3 6SG
G. BECKETT

Bramley Cottage, Stanhoe, King’s Lynn, Norfolk, PE31 8QF

RUBUS PERCRISPUS D. E. ALLEN & R. D. RANDALL (ROSACEAE) IN DORSET
(V.C. 9)

After the paper describing this new species (Allen 1994) had gone to press, it was discovered that
records of various Rubus taxa from one locality in the far east of v.c. 9 (Dorset) — “‘near Foxholes
Wood” — all relate to R. percrispus too, thus adding a ninth vice-county to its known British Isles
range.

When first collected there, by E. F. Linton in 1890 (BM), it was labelled ““R. radula near type’’, a
determination subsequently confirmed by Rogers in 1903. In 1891 and 1892 R. P. Murray revisited
the locality, probably following directions given to him by Linton (and improving on the latter’s data
by identifying the habitat as a roadside). After deciding that the plant was R. anglosaxonicus Gelert
(a species now known as R. micans Godron), he distributed material under that name in the second
of those years through the Botanical Exchange Club (B.E.C.), only to have Rogers pronounce this
intermediate between R. anglosaxonicus and R. raduloides (Rogers) Sudre but nearer the latter.
Two examples of the B.E.C. gathering later passed into herb. Barton & Riddelsdell as their nos.
7376 and 10385 (now in BM), one of which was redetermined by Barton as R. anglosaxonicus X
R. echinatus Lindley and later still by Watson as R. aspericaulis Lef. & P. J. Mueller (a species not
now accepted as British). Another example of the same B.E.C. gathering in LIV has been referred
to R. raduloides pure and simple.

In 1936, this time from a spot yet more precisely identified as a hedge to the north of the wood, N.
Douglas Simpson collected (no. 36.1030, now in BM) in company with Watson a specimen which the
latter considered a white-flowered form of R. radula Wethe ex Boenn. Watson had apparently
collected this on his own there some years earlier, for he had recorded (Watson 1932) sowing seeds
of it in order to test whether the flower colour in this species is independent of soil influences.
Subsequently, however, he must have had second thoughts, for he was to omit R. radula from the
list of all v.c. 9 Rubus species that he compiled for Good (1949).

With the aid of Simpson’s more precise localization I succeeded in July 1994 in refinding what
proved to be just a single clump under the east hedge of the A350 road just to the north of the wood
(SY/950.984). R. D. Randall concurs with my determination of this, as well as all the other
specimens referred to above, as R. percrispus.

R. radula has never seemed very likely to occur in Dorset and the sole evidence of its occurrence is
thus now shown to be ill-founded. R. raduloides, in turn, now has its supposed Dorset localities cut
back to a single wood near Sturminster Newton, in the far north of the county (where many
batologists have collected it from 1889 onwards), which is more in line with the rest of its range in
Wessex. Given the known preference of that species for basic soils, its presence in the
neighbourhood of a wood renowned for Rubus species characteristic of acid soils appeared
additionally anomalous. Foxholes Wood, near Wimborne, in the south-east of v.c. 9, constitutes a
natural extension, rather, of the chain of localities for R. percrispus already known along the coastal

408 NOTES

hinterland of the western half of South Hampshire, v.c. 11. The discrimination of this new species
has thus had the happy effect in this particular instance of enabling a whole cluster of long-standing
puzzles and anomalies to be resolved.

REFERENCES

ALLEN, D. E. (1994). Four new Anglo-Sarnian species of Rubus L. (Rosaceae). Watsonia 20: 143-151.
Goon, R. (1949). A geographical handbook of the Dorset flora. Dorchester.
Watson, W. C. R. (1932). Report of the Botanical Society and Exchange Club of the British Isles 9: 826.

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

TYPIFICATION OF RUBUS PULLIFOLIUS W. C. R. WATSON (ROSACEAE)

It has been apparent for some time that the holotype of this species (in BM) is not as it should be.
The inflorescence and one of the two stem-pieces accompanying it on the same sheet patently belong
to some other species of Rubus. That that is R. oxyanchus Sudre is better revealed by a specimen
(no. 36.1013) collected by N. Douglas Simpson on the same occasion, doubtless under Watson’s
guidance, which is now in BM also. That species occurs in some quantity in the type locality,
Southampton Common, S. Hants.. v.c. 11, especially in its east section, from the central part of
which (‘‘near the Tram Dep6ét’’) the greater precision of Simpson’s label shows that at least the
latter’s specimen came. Watson evidently did not know very well R. oxyanchus, a deceptively
variable species mainly confined in Britain to the Bournemouth area, as suggested by his later
erroneous determination of shade-grown Dorset material of that (in BM and SLBI) as a non-British
species, R. majusculus Sudre. But it was in any case rash of him to have collected on Southampton
Common, a locality apparently unknown to him till then and one exceptionally rich in Rubus
species, on a date as late in the season as 8 September. Southampton has one of the hottest summer
climates in Britain and at least in most years brambles there have virtually all wholly shed their
petals by mid-August. In the circumstances it is consequently not surprising that he mixed up two
species. The fact that he noted the petals on the inflorescence that he clipped as “‘pinkish” ought,
however, to have given him pause, if only in subsequent years, for the petals of R. pullifolius are
liable to be that colour only on first opening, before turning to pure white.

The second stem-piece on the sheet could well be that of R. pullifolius, however. Although the
main range of this similarly Bournemouth area species does not extend eastwards further than
Lymington, there have been one or two outlying finds of it in and around Southampton and one
bush was seen on the Common there in 1974 — though repeated subsequent searches, especially on
the site of the former tram depot, have failed to turn up more. Rather than dislodge a well-
established name, the best course would seem to be to give the stem-piece the benefit of the doubt,
and I accordingly here designate it as the lectotype.

It is desirable in a case such as this that the name be reinforced by the designation of an epitype.
Fortunately there is a specimen in BM (Alum Chine, Bournemouth, S. Hants., v.c. 11, 27 July 1907,
W. Moyle Rogers s.n., as R. leucandrus) which bears a label in Watson’s handwriting showing that
he determined it in 1948 as R. pullifolius, a name which he does on the whole appear to have applied
consistently. As that specimen can conveniently be filed alongside tlhe sheet bearing the lectotype, I
accordingly here select it for this purpose.

ACKNOWLEDGMENTS

I wish to thank Dr C. E. Jarvis for nomenclatural advice and A. Newton for a helpful critique of a
preliminary review of the problem.

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

NOTES 409

ADDITIONAL SETS OF CONTINENTAL RUBUS EXSICCATAE IN BRITISH HERBARIA

The recent monograph of Rubus in the British Isles by Edees & Newton (1988) most usefully
includes an appendix listing the principal sets of relevant exsiccatae and the British institutions in
which these are to be found.

Since that list was published several additional sets have come to light, and the location of these
seems worth placing on record:

BAENITZ, Herbarium Europaeum. MANCH; OXF.

BRAUN, Herbarium ruborum Germanicorum. Also SLBI.

FRIDERICHSEN & GELERT, Rubi exsiccati Daniae et Slesvigiae. Also BM.

SUDRE, Batotheca Europaea. BM possesses two sets.

WIRTGEN, Herbarium ruborum Rhenanarum. Portions also in BM and SLBI.

The following Rubus sets not listed by Edees & Newton are also represented in British herbaria at
least in part:

BILLOT, Flora Galliae et Germaniae exsiccata. OXF (via herb. F. Stratton); SLBI (via herb.

F. Townsend).

LETENDRE, Rubus de la Seine-Inférieure. MANCH.

SCHULTZ, Herbarium normale. OXF; SLBI.

WIRTGEN, Herbarium plantarum selectarum florae Rhenanae. Ed. 2. OXF (a few only).

REFERENCE

Epes, E. S. & Newron, A. (1988). Brambles of the British Isles. Ray Society, London.

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

LECTOTYPIFICATION OF ROSA ROTHSCHILDITI DRUCE (ROSACEAE)

George Claridge Druce (1850-1932) was an enthusiastic, if uncritical, rhodologist who named four
taxa in the genus, but only one (Rosa rothschildii) at specific rank. Of the six collections labelled
R. rothschildii in the Oxford herbarium (OXF), only one is suitable to be considered as a lectotype
(Druce 4821, September 1910). Firstly, it matches the protologue, both morphologically and
geographically. Druce’s mention of resinous scent implies that the subfoliar glands are not of the
R. rubiginosa type, and the protologue suggests that he was particularly basing the species on the
plants seen while botanizing with the Hon. N. Charles Rothschild at Ashton (Druce 1924;
Rothschild 1983: 174). Secondly, the specimen has Druce’s own protologue attached. It is therefore
likely that he made the most use of this specimen in drawing up his description. The only other
specimen that might be considered a candidate (Druce s.n., June 1911) has no protologue appended
and is too immature to have been of much assistance in drawing up the description.

Subsequent to the lectotypification, the six specimens were identified critically by A. L. Primavesi
and G. G. Graham. The lectotype is R. canina X R. sherardii. It is clear that Druce, and later
authors, intended the epithet rothschildii to apply also to certain nothomorphs of R. canina xX
rubiginosa (R. X nitidula of Besser (1815) has priority for hybrids of this parentage (Kent 1992)),
and the other specimens labelled R. rothschildii fall broadly into this category. However, the
lectotypification of Rosa rothschildii (and the fact that Druce based part of his description on
material that is unambiguously R. canina X sherardii) allows us to resurrect the name from obscurity
to be used for this hybrid, a practice already adopted in Graham & Primavesi (1993). R. x |
rothscnildii combines the habit of R. canina with the stipitate glands and resinous scent of
R. sherardii. The word “acicles”’, in both the Latin and English parts of the protologue, is clearly
intended by Druce to refer to stipitate glands. These are abundant on the lectotype specimen, but
there are no acicles in the conventional sense of small slender prickles. It is extraordinary that
Wolley-Dod should have linked R. rothschildii to R. obtusifolia Desv. (= R. tomentella Lem.;

410 NOTES

R. borreri Woods) as none of the specimens in OXF (except possibly that of A. Ley which is
inadequate for determination) has any relationship to this taxon.

Rosa x rothschildii Druce, Rep. B.E.C. 3: 157-158 (1913), emend. Graham & Primavesi, Roses of
Great Br. and Ireland: 98 (1993). Hybrid formula: R. canina L. X R. sherardii Davies.
Synonyms: R. tomentella Lem. var. rothschildii (Druce) W.-Dod, Roses of Britain: 71 (1924);
R. obtusifolia Desv. var. rothschildii (Druce) W.-Dod, Revis. Brit. roses (Suppl. J. Bot.): 73 (1931).
Misapplied names: R. verticillacantha sensu Druce p.p., Journal of botany 42: 6 (1880), J. Northants
nat. Hist. Soc. 1: 273 (1881); R. caryophyllacea sensu Druce p.p., B.E.C. Rep. [1911]: 87 (1912),
W.-Dod, List Brit. roses: 37 (1911). [The synonyms and misapplied names probably refer to
extended elements of the taxon.|

Protologue (extract): “‘740 (2). Rosa Roruscuitpu, Druce ... 2-3 m. Rami aculeis falcatis
horrentes. Caules floriferi aciculati, aciculis infra inflorescentiam numerosis . . . glandulis sub-
foliaribus sat numerosis . . . Odor foliorum ei gregis Mollissimae similis, et odor florum ei gregis

Caninae similis. Habitat: Northamptonshire — Dane’s Camp, 1878, 1896; Farthinghoe; Ashton,
near Oundle, 1910, G. C. Druce; Geddington Chase, Waddenhoe, Ley teste Wolley-Dod; Hunts. —
Catsworth, Ellington, Ley teste Wolley-Dod; Surrey - Coombe (No. 786); Malden (No. 838), C. E.
Britton, 1912. The Surrey plants have fruits slightly more spherical, and leaflets somewhat shorter
and broader, but the acicular branches and glandular foliage bring them under Rothschildii . . . In
the seventies I found a rose on Hunsbury Hill — the Dane’s camp — near Northampton . . . However
in August 1910 when staying at Ashton, I saw a rose in the very luxuriant hedgerows bordering the
road leading to the Hon. N. Charles Rothschild’s house, which at once reminded me of the
Hunsbury Hill plant. . . In June 1911 I went to Ashton again in order to obtain flowering specimens
when I found the rose in several places in the vicinity . . . The plant forms tall handsome bushes with
conspicuous flowers of a brighter pink than normal canina, while the acicular branches, naked fruit,
the very glandular, nearly glabrous leaves, the acicular petioles and peduncles are distinguishing
characters which separate it from its allies. I have associated the plant with the name of my friend on
whose estate it grows: and who has done so much to forward the study of Natural Science.”
Specimens in OXF: Druce no. 4821, Ashton Wold, Northants, August 1910. [R. canina L. (foem.) Xx
R. sherardii Davies (masc.)], LECTOTYPUS, hic desig.

Excluded specimens at OXF: Druce, Danes Camp, Northants, September 1889 [R. rubiginosa
hybrid indet.]; Druce, Ashton, Northants, June 1911 [too young for determination but not the same
as 4821]; A. Ley [B.E.C. 596], hedges, Wadenhoe, Northants, July 1910 [as R. borreri Woods var.
[inadequate for determination]; C. E. Britton, open ground, Malden, Surrey, 5 September 1911
[R. canina X rubiginosa]; C. E. Britton [B.E.C. 838], open ground, Malden, Surrey, 19 August
1912[R. canina X rubiginosa]; J. P. M. Brenan 6950, rough pasture v.c. 23, near Woodeaton by the
road to Marston, Oxon, 27 August 1943 (fruits), 11 June 1944 (flowers) [R. canina with some
introgression with R. rubiginosa or micrantha].

REFERENCES

Druce, G. C. (1924). Obituary. The Hon. N. Charles Rothschild. Journal of the Northamptonshire Natural
History Society 22: 135-141.

Druce, G. C. (1930). The Flora of Northamptonshire. T. Buncle & Co., Arbroath.

Kent, D. H. (1992). List of vascular plants of the British Isles. Botanical Society of the British Isles, London.

GrauanM, G. G. & Primavesi, A. L. (1993). Roses of Great Britain and Ireland, Botanical Society of the British
Isles, Handbook no. 7, London.

ROTHSCHILD, M. (1983). Dear Lord Rothschild. Philadelphia & London.

Q. C. B. Cronk, S. K. MARNER

Department of Plant Sciences, University of Oxford, South Parks Road, Oxford, OXI 3RB
A. L. PRIMAVESI

Ratcliffe College, Fosse Way, Ratcliffe-on-the-Wreake, Leicester, LE7 4SG

NOTES 41]
FURTHER DRUCE ROSA TAXA (ROSACEAE)

Eponymy: Rosa X drucei W.-Dod, Journal of Botany 62: 205 (1924), as R. canina X rubiginosa,
later suppressed by Wolley-Dod (1931) under R. canina var. latebrosa (Déséglise) N. E. Br. R. x
nitidula Besser is an earlier name for the hybrid combination.

Druce’s infraspecific taxa have been typified as follows and the types and associated specimens
critically examined by G. G. Graham and A. L. Primavesi. All the taxa are probably best
suppressed, although R. arvensis var. suberecta is likely to be the valid name for “‘cristate”’
R. arvensis (probably deriving from introgression with R. canina), in the (perhaps unlikely) event of
any botanist wishing to use a name for this phenotype.

Rosa arvensis Huds. var. suberecta Druce, B.E.C. Rep. 5: 559 (1920) [R. a. var. cristata Druce, Fi.
Berks: 206 (1897), as ‘*?var. cristata”, nomen confusum; R. a. var. subcristata Druce ined.}.
Protologue: “923. Rosa arvensis Huds., var. suberecta mihi. This differs from the type in the erect,
persistent sepals. Greenham, Berks, 1893, G. C. Druce. see Fl. Berks 206, 1897”.

Notes: Not recognised by Wolley-Dod and probably not worthy of recognition above the level of
form. Druce (1897) contends that although Crépin considers it only an accidental condition, he
himself noticed it for four consecutive years in ‘‘these localities’ (apparently ‘“‘By the Emborne
[Enborne] Stream near Greenham Common and near Sandleford”’). This variant (“‘with ascending
and semipersistent sepals’) has also been recorded from three localities in Hertfordshire (Purchas
& Ley 1889). The sepal character is likely to be under genetic control and provides a parallel to the
subcristate forms of R. canina.

Specimen in OXF: Druce, var. subcristata, Greenham, Berks, September 1893 [HOLOTYPE].

Rosa eglanteria L. var. corstorphinae Druce, B.E.C. Rep. 4: 195 (1916) [Rosa rubiginosa L. f.
corstorphinae (Druce) W.-Dod, Roses Br.: 92 (1924)].

Protologue (extract): “937. Rosa Eglanteria L., var. Corstorphinae mihi. Bush tall, stem prickles
distant, long based, uncinate, of the flowering shoots crowded, nearly straight, 2-3 mm. long, of the
peduncles crowded, straight slender. Leaves broadly ovate, densely glandular above and below,
biserrate. Flowers in dense umbellate clusters, 8-10, dark rose-red, fragrant, very showy. . . Near
Duninald, Forfar, in plenty. Shown me by Mrs Corstorphine . . . This handsome and very distinct-
looking plant is quite new to me. I saw nothing in its vicinity which could suggest a hybrid origin, but
the bushes were remarkably constant. Major Wolley-Dod, too, says he has seen nothing like it. Ifa
hybrid, it is almost certainly R. Eglanteria X gallica, the armature recalling that of the latter species.
Grer-Driuce!’

Notes: Wolley-Dod (1924) wrote that he could “‘see nothing in this but a very luxuriant form of the
type’, and that since its discovery in Forfar he had seen similar specimens from W. Kent and E. &
W. Ross. We agree with Wolley-Dod that Druce’s specimens are straightforward R. rubiginosa.
The variety is not worth recognizing even at forma level. The Glassford specimens, distributed as f.
corstorphinae, are hybrids.

Specimens in OXF: Druce (B.E.C. 937), Dunninald, Angus, August 1915 [HOLOTYPE]; Druce
(B.E.C. 937), near Dunning, Perth, August 1916; Druce (BEC 937), near Montrose, Forfar,
August 1916; Druce, inter Montrose and Arbroath, Forfar.

Excluded specimens in OXF: J. G. Glassford, Aberfeldy, mid-Perth, 20 August 1928 (three
specimens ex herb. Wolley-Dod as R. rubiginosa t. corstorphinae W.-Dod); J. G. Glassford,
Aberfeldy Mid-Perth, 20 August 1928 (ex B.E.C.) (both specimens are R. rubiginosa xX
pimpinellifolia).

Rosa mollissima Willd. f. alba Druce, Journal of Botany 40: 184 (1902), nomen nudum.
Protologue: ‘Rosa mollissima Willd., (R. tomentosa Sm.) f. alba. Near Llanerchymedd, Anglesey.”
Notes: although this appears to be a Druce name, he did not append his name to it or provide a
description. No specimen has been found in OXF.

REFERENCES

Druce, G. C. (1897). The Flora of Berkshire. Oxford.
Purcuas, W. H. & Ley, A. (1889). A Flora of Herefordshire. Hereford.

412 NOTES

Wo LEyY-Dop, A. H. (1924). The roses of Britain. London.
Wo tey-Dop, A. H. (1931). Revision of British roses. Journal of botany (suppl.): 43-44.

Q. C. B. Cronk

Department of Plant Sciences, University of Oxford, South Parks Road, Oxford, OX1 3RB
A. L. PRIMAVESI

Ratcliffe College, Fosse Way, Ratcliffe-on-the-Wreake, Leicester, LE7 4SG

TWO SUBSPECIES OF FESTUCA RUBRA L. NEW TO ENGLAND

While preparing the Poaceae account for the Flora of Cumbria, 42 collections of Festuca rubra s.1.
were submitted to Dr A. K. Al-Bermani and Prof. C. A. Stace for identification. The material was
not representative as half the specimens were from coastal sites and a quarter from the Lake District
and Pennine hills.

Three of the specimens proved to be F. arenaria Osbeck, a species largely restricted to the east
coast of Britain and not previously recorded in the west further north than south Lancashire. It was
collected from St Bees Head, Cumberland (v.c. 70, GR NX/9.1, C. W. Muirhead, 1949, PLYP) and
the Duddon estuary, Westmorland (v.c. 69, Sandscale Haws, SD/1.7, 1992; Askam-in-Furness,
SD/2.7, 1991, both P. Burton, LANC).

The remaining specimens included all seven subspecies of F. rubra L. currently recognised as
occurring in the British Isles. The commonest was subsp. juncea, which appears to be frequent
around the entire coast. |

Of particular interest are the records of the montane subspecies arctica and scotica, both new to
England. The former was previously known south of the Scottish Highlands only from Snowdonia.
The Cumbrian records are from rock ledges in the Lake District: near Fleetwith Pike (v.c. 70, NY/
2.1), Hart Crag, Fairfield and Dollywaggon Pike (v.c. 69, NY/3.1), Red Screes (v.c. 69, NY/3.0) and
High Street (v.c. 69, NY/4.1), four sites in the Pennines: two around Cross Fell (v.c. 70, NY/6.3)
and others on limestone scars in upper Teesdale almost on the Durham border (v.c. 70, NY/7.3) and
at High Cup Nick (v.c. 69, NY/7.2), and one from a lane in the upper Eden valley (v.c. 69, SD/7.9).
The earliest record for Cumberland is that from near Fleetwith Pike (C. W. Muirhead, 1952, PLYP)
and for Westmorland that from Hart Crag (G. Halliday, 1981, LANC). This subspecies will
probably prove to be quite widely distributed in the Lake District and the Pennines.

The only records of subsp. scotica are from Cumberland, from limestone at 610 m on the north
side of Crowdundle Beck, Cross Fell (NY/6.3, C. W. Muirhead, 1949, PLYP), in the Pennines, and
in the Lake District from the north-east slopes of Pillar at 730 m (NY/1.1, G. Halliday, 1993,
LANC). It was formerly unknown south of Argyllshire.

ACKNOWLEDGMENTS
I am grateful to Dr A. K. Al-Bermani and Prof. C. A. Stace for identification of the material.

G. HALLIDAY
Department of Biological Sciences, The University, Lancaster, LAI 4YQ

RUMEX FRUTESCENS THOUARS xX R. OBTUSIFOLIUS L. (POLYGONACEAE), A
PREVIOUSLY UNDESCRIBED HYBRID DOCK, AND NEW RECORDS OF
R. X WRIGHTII LOUSLEY IN WEST CORNWALL (V.C. 1)

Argentine Dock, Rumex frutescens Thouars, has been established at Phillack Towans, W. Cornwall
(v.c. 1, SW/56.39), since at least 1921 (Thurston & Vigurs 1922; Margetts & David 1981; Margetts &
Spurgin 1991). In late August 1994 it was locally plentiful there, with many hundreds of plants, some
of which formed large patches. These were growing on calcareous dune-sand at the edges of a sand
quarry and on banks and in grassland,nearby.

NOTES 413

Other species of dock found in the same area were Rumex crispus L. subsp. crispus (plentiful),
R. obtusifolius L. subsp. obtusifolius (plentiful), R. conglomeratus Murray (locally plentiful),
R. sanguineus L. (four plants) and the hybrid R. crispus x R. obtusifolius (R. X pratensis Mert. &
Koch, four plants). A few plants of R. pulcher L. were found 300 m away.

In addition to these, a number of plants that were sterile and intermediate between R. frutescens
and R. obtusifolius (nine plants) and R. conglomeratus (two plants) were presumed to be hybrids.
This note describes R. frutescens X R. obtusifolius, a hybrid which has not been reported before,
and gives details of R. frutescens X R. conglomeratus (R. X wrightii Lousley), which has been
reported only once before (Lousley 1953; Lousley & Kent 1981).

Rumex X cornubiensis D. T. Holyoak, hybr. nov.
(Rumex frutescens Thouars X R. obtusifolius L. subsp. obtusifolius)

Hybrida inter Rumex frutescens Thouars et R. obtusifolius L. subsp. obtusifolius genita, characteri-
bus variabilis et inter parentes media (Fig. 1), ab ambobus fructibus abortivis differt.

A hybrid between R. frutescens and R. obtusifolius L. subsp. obtusifolius, found within a few
metres of colonies of the parent species. Although rather variable, it is intermediate between them
in most characters (Fig. 1) and almost, if not completely, infertile.

A robust creeping perennial, spreading by underground rhizomes (mostly shorter than the far-
creeping rhizomes of R. frutescens), so that it forms more spreading clumps than those of R.
obtusifolius. Shoots arise from the rhizomes at intervals and attain a maximum height of 105 cm
(nearly as tall as R. obtusifolius at this site, and distinctly taller than the maximum of 70 cm reached
by R. frutescens). Lower leaves with lamina up to 16 X 6.7 cm, thicker than that of R. obtusifolius,
but not as thick and leathery in texture as that of R. frutescens. Lamina often broader than in
R. frutescens, with its greatest width around the middle and the base mostly truncate to weakly
cordate; resembling R. frutescens in having the leaf-margin more or less crenulate, but the back of
the midrib and main veins weakly scabrid with small papillae as in R. obtusifolius. Stem leaves much
smaller, narrower and with more acute apices.

Panicle with branches arising at c. 40° from the main stem. Branches more numerous than is usual
in R. frutescens, but fewer than in well-grown plants of R. obtusifolius. Whole inflorescence often
with conspicuous deep red coloration. Whorls of inflorescence often less congested than in
R. frutescens, but typically closer to each other than in R. obtusifolius. Pedicels mostly 2-5 mm (0-3-
2 X length of inner perianth-segments when in fruit), most of these being distinctly longer than in
R. frutescens but shorter and thicker than in R. obtusifolius. Inner perianth-segments up to 6 mm in
length when fruits form, but mostly shorter and withering where fruits fail to develop. Well-formed
inner perianth-segments varying in shape from narrowly ovate-triangular with rather acute apex (as
in R. frutescens) to broader, triangular, with obtuse apex (as in R. obtusifolius); many with 2 or 3
short teeth, less than one-quarter of width of segment, on each margin at widest, basal part. When
well-formed, all three inner perianth-segments with a prominent tubercle along the mid-vein,
although the tubercle is typically larger and longer on one perianth-segment (tubercles with
punctulate surface in fresh material). All of the few nutlets found were shrunken when dried and
apparently infertile, 2-3-5 mm long, ovoid, and trigonous, brown, glossy, with acute angles.

Ho.otypus: W. Cornwall, v.c. 1, Phillack Towans (SW/568.392), edge of sand quarry, 21 August
1994, D. T. Holyoak (RNG).

The largest patch of R. X cornubiensis covers an area of some 12 X 9 m on top of a low bank.
Elsewhere, a single plant has spread to form a roughly circular patch 2 m diameter. Hence it is likely
that this hybrid has been established and spreading vegetatively at Phillack Towans for some years.

414 NOTES

\

FiGurE 1. Representative lower leaves (underside) and fruits (side view and t.s) of Rumex from Phillack Towans,
W. Cornwall. A. R. frutescens, B. R. X cornubiensis, C. R. obtusifolius subsp. obtusifolius. Scale lines are
marked at intervals of 1 cm (leaves) or 1 mm (fruits).

Rumex X wrightii Lousley
(R. conglomeratus Murray X R. obtusifolius L. subsp. obtusifolius)

Two plants were found in grassland, close to populations of both parents (specimens lodged at
RNG). One had five groups of stems close together and linked by underground rhizomes, implying
that this hybrid is a long-lived perennial that can spread vegetatively. Both plants were short, not
exceeding 30 cm in height, with a rather untidy appearance due to numerous short, leafy branches
and the infertile inflorescences. Most branches were at angles of c. 30° to the main stem, but a large
branch on one specimen was at c. 80° to the main stem. Some of the larger leaves were obovate and
thicker than those of R. conglomeratus, with crenate margins and truncate or subcordate bases. The
whorls of the inflorescence are mostly remote and the lower whorls are subtended by bracts. Both

plants appeared to be completely infertile, with only a minority of the inner perianth-segments ©

enlarging as fruits began to develop. ‘These enlarged inner perianth-segments are narrowly ovate

< Se -

NOTES 415

and up to 5 mm in length, all three segments with a long tubercle. On one specimen the pedicels are
short (mainly 1-2 mm) on the other longer (up to 4-5 mm).

This hybrid has been reported only once before, from Braunton Burrows, N. Devon (v.c. 4), in
1952 (Lousley 1953; Lousley & Kent 1981). Descriptions of the Devon plants indicate that they are
similar to those at Phillack Towans, although somewhat taller, up to 40 cm.

R. crispus is abundant close to the colonies of R. frutescens at Phillack Towans, but no hybrids
between these species have been found, despite an extensive search. However, R. crispus there
probably flowers earlier than R. frutescens, since by 21-31 August 1994 many of the R. crispus plants
had ripe nutlets, whereas those of R. frutescens were either flowering or had mainly unripe nutlets
and those of R. conglomeratus and R. obtusifolius mostly had ripening nutlets.

ACKNOWLEDGMENTS

Thanks are due to Paul Gainey, Geraldine Griffiths and Rose Murphy for helpful discussions in the
field and to Dr John Akeroyd for commenting on specimens and a draft of this note.

REFERENCES

LousLey, J. E. (1953). Rumex cuneifolius and a new hybrid. Watsonia 2: 394-397.

Lous.ey, J. E. & Kent, D. H. (1981). Docks and knotweeds of the British Isles. B.S.B.1., London.

Marcetrs, L. J. & Davin, R. W. (1981). A review of the Cornish flora 1980. Institute of Cornish Studies, Pool,
Cornwall.

Marcetrs, L. J. & SpurGIN, K. L. (1991). The Cornish Flora supplement 1981-1990. Trendrine Press, Zennor,
Cornwall.

Tuurston, E. & Vicurs, C. C. (1922). A supplement to F. Hamilton Davey’s Flora of Cornwall. Truro.

D. T. HoLyoak
8 Edward Street, Tuckingmill, Camborne, Cornwall, TR14 8PA

POTAMOGETON X COGNATUS ASCH. & GRAEBN. AT LOCH BORRALIE, WEST
SUTHERLAND (V.C. 108), SCOTLAND

Potamogeton X cognatus Asch. & Graebn. (P. perfoliatus L. x P. praelongus Wulfen) (Potamoge-
tonaceae) was originally described from Germany. It was discovered in Britain by J. M. Taylor, who
found it in drains at Belton and Crowle, N. Lincolnshire (v.c. 54), in 1943. Both parents grew in the
vicinity. An illustrated account of the Lincolnshire plant (which was the first flowering example of
this hybrid to be discovered anywhere) was provided by Taylor & Sledge (1944). I am not aware of
any records of the hybrid from Lincolnshire after Taylor’s collections, which were made in 1943 and
1944, and I was unable to find either P. praelongus or P. X cognatus when I visited the area with Mrs
I. Weston and others in 1989.

The only other locality for the hybrid in the British Isles is Loch Borralie, a loch on the Durness
limestone of W. Sutherland, where it was collected by Sir George Taylor in 1948, Mrs B. Welch in
1951 and D. Dupree in 1970 (specimens in BM; records also in J. E. Dandy’s card index at BM). The
locality was published by Dandy (1975). In a detailed survey of Loch Borralie, Spence et al. (1984)
refound all the aquatic plants previously recorded from the loch except this hybrid. Similarly, the
hybrid was not recorded by the Nature Conservancy Council’s Scottish Loch Survey team when they
visited Loch Borralie on 29 June 1988. On 21 August 1993 I visited Loch Borralie with I. M. & Mrs
P. A. Evans and D. A. & Mrs A. Pearman in an attempt to refind P. X cognatus. To my surprise, we.
found it at two places in the loch.

In view of the lack of published information on the hybrid at Loch Borralie, and the fact that some
crucial characters are difficult to interpret on herbarium specimens, notes on its morphology and
habitat are provided below. Voucher specimens of P. X cognatus and its putative parents (Preston
93/51—56) will be deposited in CGE and E.

416 NOTES

MORPHOLOGY
The following description is based on fresh material of P. X cognatus collected at Loch Borralie.

Stems to 1-2 m, 2-2—3-5 mm in diameter, terete. Submerged leaves 45-80 x 14-23 mm, 2-6-4-0
times as long as wide, translucent, green, often with a brownish tinge on the upper leaves and
becoming brown with age, ovate-oblong, sessile, clasping the stems at the base for more than half
the diameter of the stem but with a broad gap between the two edges of the leaf on the far side of the
stem, tapering to a slightly hooded apex, denticulate and plane or undulate at the margin, the teeth
10(—25) wm long and 250-500 um apart towards the leaf apex, just visible with a X20 lens, more
distant further from the apex and very distant towards the base, consisting of a single cell with an
obtuse apex; midrib bordered on each side by a narrow band of lacunae, the lateral veins 6-9 on
each side, 1-3 of which are more strongly developed than the others, the secondary veins transverse
or ascending between the midrib and the inner lateral veins, more or less transverse elsewhere, all
the veins with a dark tinge so that the leaf has a net-like appearance. Floating leaves absent. Stipules
12-5-18 mm, flexible, translucent with a milky or a slight pinkish tinge, rounded at the apex,
persisting for several nodes behind the apex, two of the veins slightly more prominent than the
others but not forming distinct ridges. Inflorescences 8-11 x 4-5-6-5 mm; peduncles 62-254 mm,
2:2-3-5 mm in diameter, of uniform diameter throughout their length, terete. Flowers 16-24,
usually with 4 carpels (single flowers seen with 1, 3 and 5), the dark brown stigmas protruding from
tightly closed green tepals.

When fresh material of P. x cognatus and its putative parents was compared side by side, the hybrid
was clearly intermediate in vegetative characters (Table 1). It differed from both parents in its short
inflorescences with closed tepals. A comparison of the Loch Borralie P. x cognatus with the published
description of the Lincolnshire plant (Taylor & Sledge 1944) suggests that they are essentially similar.
The main difference lies in the length of the peduncles, 45—75 mm in Lincolnshire compared to 62-254
mm at Loch Borralie. The long peduncles of the Borralie plant probably reflect the fact that the water
was high following a wet season, and are unlikely to indicate a genetic difference between the plants.
The fact that both the Lincolnshire and the Borralie plants had denticulate leaf margins is interesting:
the original material of P. X cognatus had toothed margins (Ascherson & Graebner 1897) but
Hagstr6m (1916) described plants with entire leaves from a lake in Denmark.

HABITAT

A detailed description of Loch Borralie is given by Spence eral. (1984). It is 1-2 km long and 0-2-0-5
km wide, with an area of 36 hectares. It lies in a shallow basin in the Cambrian Durness limestone,

TABLE 1. CHARACTERS OF POTAMOGETON PERFOLIATUS, P. x COGNATUS AND P.
PRAELONGUS FROM LOCH BORRALIE

P. perfoliatus P. X cognatus P. praelongus
Length of main stem 25-48 45-80 125-155
leaves (mm)
Leaf length: breadth ]-3—2-5 2:6-4-0 6-7-7-4
ratio
Leaf base Clasping stem with edges = Clasping stem with broad Slightly clasping stem
on far side almost gap between edges on
meeting or overlapping far side
Leaf margin Denticulate Denticulate Entire
Leaf apex Scarcely hooded Slightly hooded Markedly hooded
Stipules Fugacious Persisting on upper Persistent
nodes
Inflorescence length 12-16 8-11 37-40
(mm)
Tepals Open Closed Open

_ All observations based on small samples collected on 21 August 1993. The quantitative characters show the
difference between the taxa at Loch Borralie, but should not be used to identify plants from other sites.

NOTES 417

and has a small catchment of 154 hectares. The water of the loch is calcareous (pH 8.5) and
remarkably clear, with low levels of nitrogen and available phosphorus and very low plankton
densities. The shallow water at the edge of the lake has an open plant community in which the main
species are Chara aspera and Littorella uniflora, with Potamogeton filiformis and Myriophyllum
alterniflorum. In deeper water the vegetation is dense, and is dominated by Hippuris vulgaris,
Myriophyllum spicatum, Potamogeton naians, P. pectinatus, P. perfoliatus and P. praelongus.
Below 4-5 m there is a deep-water charophyte sward dominated by Chara globularis.

Around much of the edge of Loch Borralie the water shelves gradually, and the Potamogeton-
dominated community is inaccessible to the observer on the shore, especially when the water level is
high. We detected the hybrid at two points where the water shelves much more steeply, and where
the dense macrophyte-dominated community was visible from the shore or could be sampled by
grapnelling. At the N.W. side of the loch, grid reference NC/382.673, P. x cognatus, P. perfoliatus
and P. praelongus were dredged up together. At the S.E. side of the loch the hybrid was visible just
offshore at a point where the limestone outcrops at the edge of the loch, grid reference NC/383.668.
It grew in vegetation dominated by the submerged shoots of Hippuris vulgaris, in water c.1.5 m
deep. The other species growing here were P. perfoliatus and P. praelongus. The presence of the
hybrid in two localities 0-6 km apart suggests that it may be widespread in the vegetation in which its
parents occur. The Lincolnshire population of P. X cognatus reproduced vegetatively by buds at the
end of short stolons which arose at the nodes of the non-flowering shoots (Taylor & Sledge 1944).

Potamogeton perfoliatus and P. praelongus may be closely related (Haynes 1985). Their hybrid,
P. X cognatus, has been recorded from only a few localities in northern Europe. The rarity of the
hybrid has been commented on by Hagstrém (1916), who suggested that the earlier flowering time
of P. praelongus restricted the opportunities for hybridisation. He contrasted the rarity of P. x
cognatus with the frequency of P. X nitens, the hybrid between P. perfoliatus and P. gramineus.
Although P. gramineus is morphologically dissimilar to P. perfoliatus, the two species ‘scarcely can
grow together without producing crosses’. The presence of P. < cognatus in Loch Borralie adds to
the interest of this remarkable site, which is classified as an area of international importance in the
Nature Conservation Review (Ratcliffe 1977).

ACKNOWLEDGMENTS

Iam very grateful to D. A. and Mrs A. Pearman for joining me on the trip to Scotland, andtoI. M.
and Mrs P. A. Evans for meeting us at Loch Borralie. Scottish Natural Heritage kindly provided
copies of the records from the 1988 visit of the Scottish Loch Survey. A. O. Chater and P. T.
Harding kindly commented on a draft of this note.

REFERENCES

ASCHERSON, P. & GRAEBNER, P. (1897). Synopsis der Mitteleuropdischen Flora, 1. Wilhelm Engelmann, Leipzig.

Danpy, J. E. (1975). Potamogeton L., in Stace, C. A., ed., Hybridization and the flora of the British Isles,
pp. 444-459. Academic Press, London.

Hacstrom, J. O. (1916). Critical researches on the Potamogetons. Kongliga Svenska Vetenskapsakademiens
Handlingar n.s. 55(5): 1-281.

_ Haynes, R. R. (1985). A revision of the clasping-leaved Potamogeton (Potamogetonaceae). Sida 11: 173-188.

— RatcuiFFe, D. A. (1977). A nature conservation review. 2 vols. Cambridge University Press, Cambridge.

.. Spence, D. H. N., Barciay, A. M. & ALLEN, E. D. (1984). Limnology and macrophyte vegetation of a deep,

ft clear limestone lake, Loch Borralie. Transactions of the Botanical Society of Edinburgh 44: 187-204.

| Taytor, J. M. & SLEDGE, W. A. (1944). x. Potamogeton cognatus Asch. and Graeb. in. Britain. The Naturalist

t 1944: 121-123.

C. D. PRESTON
[.T.E., Monks Wood, Abbots Ripton, Huntingdon, Cambs., PE17 2LS

418 NOTES

AN EARLY SCOTTISH RECORD OF RUBUS ARCTICUS L. (ROSACEAE)

The small Northern bramble Rubus arcticus L. is considered to have become extinct in Britain in the
mid-nineteenth century (Harley 1956). In the previous hundred years there were several reported
occurrences, and several specimens were collected from widely separated localities (Harley 1956;
Edees & Newton 1988). Some of the occurrences possibly resulted from seeds brought in by migrant
birds or from cultivation in gardens (Harley 1956), and unfortunately no wild colony was regularly
recorded, the localities of specimens being imprecise so that later botanists could not refind the
populations.

Rubus arcticus was listed in the “Catalogue of British Plants in Dr Hope’s Hortus Siccus, 1768”
(Balfour 1907). However the entry is marked with a sign denoting “‘plant not yet found in Scotland,
and that the specimen I had from England’’, and the source of the entry is given as “from Mr
Gordon’.

John Hope was Regius Keeper of the Royal Botanic Garden in Edinburgh from 1760 to 1786
(Balfour 1907) and corresponded with Dr David Skene, an Aberdeen medical practitioner and
botanist (Welch 1989, 1993); much of this correspondence concerned new species being found in
Scotland and exchanges of specimens. In a letter to Dr Skene dated 31 August 1765 (Skene MS 38*)
Dr Hope wrote *. . . Mr Freer has added 4 score Plants to his collection, the last plants were the
Rubus arcticus and Osmunda crispa. A list of them shall be sent you . . . Mr Freer I imagine may be
ready again next spring to publish his list. . .”

Clearly this statement is in contradiction to the 1768 catalogue entry, and I suspect that errors
occurred in its compilation or transcription. For some species two or more localities are given by
Hope, so mention of Freer was not precluded by the Gordon source. Moreover two of the entries
originating from information supplied by David Skene are dubious viz. the source of Arenaria
laricifolia (sic) (= Minuartia verna (L.) Hiern) is given as “near Tongue, Aberdeenshire Dr Skene’”’,
and the source of Chelidonium majus is ‘“‘at Revelston in Aberdeenshire Dr D. Skene’’. Places
named Tongue and Revelston do not occur in Aberdeenshire, and we know that the Minuartia
grows only on serpentine rocks in a very restricted district around Cabrach (Welch 1993); David
Skene in an undated list (MS 482 p. 11*) accurately gave its locality as ““Betwixt Clova & Craig’.

According to Kent & Allen (1984) 40 of Adam Freer’s specimens passed to Dr Hope, but Hope’s
herbarium is believed to have been destroyed around 1840, so there is little chance of finding the
locality from which Freer obtained Rubus arcticus.

REFERENCES

BaAFrour, L. B. (1907). Eighteenth century records of British plants. Notes from the Royal Botanic Garden,
Edinburgh 18: 123-190.

EpeEs, E. S. & Newron, A. (1988). Brambles of the British Isles. Ray Society, London.

Har ey, R. M. (1956). Rubus arcticus L. in Britain. Watsonia 3: 237-238.

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

WE LcH, D. (1989). Dr Skene’s botanical writings c. 1760-1770. B.S. B.I. Scottish Newsletter 11: 12.

WE LcH, D. (1993). Flora of North Aberdeenshire. Banchory.

D. WELCH
East Fernbank, Woodside Road, Banchory, Kincardineshire, AB31 3XL

*These numbers are from the catalogue of David Skene’s papers held in the Aberdeen University Library
(Special Collections).

ee . =

EO LER TNE SS 3 =

ee See

Watsonia 20: 419-437 (1995) 419

Plant Records

Records for publication must be submitted to the appropriate Vice-county Recorder (see B.S.B.I. Year Book
for 1995), and not the Editors. The records must normally be of species, hybrids or subspecies of native or
naturalized plants belonging to one or more of the following categories: 1st or 2nd v.c. record; Ist post-1930 v.c.
record; only extant v.c. locality, or 2nd such locality; a record of an extension of range by more than 100 km.
Such records will also be accepted for the major islands in v.cc. 102-104 and 110. Only Ist records can normally
be accepted for Rubus, Hieracium and hybrids. Records for subdivisions of vice-counties will not be treated
separately; they must therefore be records for the vice-county as a whole. Records of Taraxacum are now being
dealt with separately, by Dr A. J. Richards, and will be published at a later date.

Records are arranged in the order given in the List of vascular plants of the British Isles by D. H. Kent (1992),
from which the species’ numbers, taxonomy and nomenclature are taken. 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: 1-4, 9-14, 17-19, 25-27, 29, 35, 38-
44, 46-50, 52, 53, 57-59, 64, 67-70, 73, 75, 77-81, 91, 93, 94, 98-100, 102, 108, 110.

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.

1/2.1. LYCOPODIELLA INUNDATA 70, Cumberland: Wasdale, NY/1.0. J. Hooson, 1994. 2nd
extant locality.

1/3.1. LycOPODIUM CLAVATUM 58, Cheshire: Peaty bank, Coombes Clough, Crowden, SK/
053.994. G. M. Kay, 1994, herb. G.M.K. Only extant locality.

1/3.2. LYCOPODIUM ANNOTINUM *108, W. Sutherland: W. of Gobernuisgach Lodge, NC/4.4.
D.C. Lang, R. McBeath & G. P. Rothero, 1988. Ungrazed tall herb community on ledge just above
water level, Loch Bealach a’Bhuirich, NC/262.280. G. P. Rothero, 1994. Ist and 2nd records. ~

4/1.1. EQUISETUM HYEMALE *2, E. Cornwall: Marshy corner of field, Portquin, SW/973.804.
C. H. Watts, 1994.

11/1.3. POLYPODIUM CAMBRICUM 98, Main Argyll: Base of cliff in woodland between Port
Kilcheran and Lochan Treshtil, Lismore, NM/822.382. A. McG. Stirling & B. H. Thompson, 1994.
2nd record.

14/1.1. THELYPTERIS PALUSTRIS *57, Derbys.: Near Baslow, SK/2.7. I. D. Rotherham & P. A.
Ardron, 1991. 69, Westmorland: Storth, Arnside, SD/4.7. F. W. Grayson, 1994, LANC. 2nd
extant locality.

17/1.1. POLYSTICHUM SETIFERUM *91, Kincardines.: Den of Fenella, NO/77.66. C. H.
Gimingham, 1947, ABD, det. C. N. Page. Deciduous woodland in den, Morphie, NO/708.645. D.
Welch, 1994, ABD. Ist and 2nd records.

17/1.1 X 2. POLYSTICHUM SETIFERUM X P. ACULEATUM (P. X BICKNELLII) *46, Cards.: With
both parents on wall of ruined leadmine, Esgair Hir, SN/727.910. S. P. Chambers, 1993, herb.
S.P.C., det. A. M. Paul.

17/1.2. POLYSTICHUM ACULEATUM 2, E. Cornwall: Walls of old mine buildings, Phoenix United ©
Mine near Minions, SX/267.720. M. C. G. & A. A. Atkinson, 1994, conf. A. C. Jermy. Ist
authenticated record since 1868.

+20/5.2. LARIX KAEMPFERI *77, Lanarks.: Caldercruix, NS/825.666. P. Macpherson, 1993,
herb. P.M., det. J. R. Akeroyd.

420 PLANT RECORDS

+28/4.1. ERANTHIS HYEMALIS “46, Cards.: Well established in lawns, Welsh Agricultural
College, Llanbadarn Fawr, SN/602.811. S. P. Chambers, 1994.

28/9.1. ANEMONE NEMOROSA 102, S. Ebudes: Oronsay, NR/3.8. A. Walker, 1992. Under
Gunnera, Colonsay House Woods, Colonsay, NR/394.966. R. L. Gulliver, 1994. 1st Oronsay and
Colonsay records.

28/13.11. RANUNCULUS SCELERATUS 102, S. Ebudes: Flush above shore, Port Mor, Colonsay,
NR/361.949. M. C. Gulliver, 1993. Ist post-1930 Colonsay record.

28/13.20. RANUNCULUS TRIPARTITUS 41, Glam.: Trampled mud in shallow pool, Pitton Cross
Moor, SS/434.880. Q. O. N. Kay & R. Wright, 1994, det. Q.O.N.K. & R. John. Only extant
locality.

28/13.25b. RANUNCULUS PENICILLATUS subsp. PSEUDOFLUITANS 46, Cards.: Shallow stream in
pasture, Afon Dulas at Ffynnon Wen, Llangybi, SN/605.528. A. O. Chater, 1993, NMW, det. S. D.
Webster as var. pseudofluitans. 2nd record.

28/17.4. THALICTRUM MINUS *78, Peebless.: Old dyke by road S. of Newhall, NT/317.320. A.
McG. Stirling & B. H. Thompson, 1994.

+29/1.9. BERBERIS DARWINII *50, Denbs.: Roadside, Capel Garmon, SH/80.56. W. McCarthy,
1994. “77, Lanarks.: Craigton, NS/53.64. P. Macpherson & E. K. Lindsay, 1994, herb. P.M..,
det. D. R. McKean.

+30/1.1. PAPAVER ORIENTALE *$8, Cheshire: Spreading on dunes, Hoylake, SJ/208.875. G. M.
Kay & P. D. Stanley, 1994. :

+30/1.2. PAPAVER ATLANTICUM *58, Cheshire: Wall to canal, Higher Poynton, SJ/944.831. G.
M. Kay, 1994.

31/5.1. FUMARIA CAPREOLATA *81, Berwicks.: Ayton, NT/92.60. F. M. Norman, 1878, herb.
Berwickshire High School, Duns, det. M. E. Braithwaite.

31/5.6. FUMARIA PURPUREA *100, Clyde Is.: Rubbish tip, Brodick, Arran, NS/02.35. M.
McCallum Webster, 1969, E, det. M. Lidén.

+36/3.1. SOLEIROLIA SOLEIROLII 26, W. Suffolk: Old wall, Brandon Country Park, TL/785.852.
E. M. Hyde, 1981. 2nd record. *77, Lanarks.: Walls and paths, Botanic Gardens, Glasgow, NS/
56.67. P. Macpherson & A. McG. Stirling, 1994, herb. P.M.

+40/2.2. ALNUS INCANA *26, W. Suffolk: Verge of A45, Exning, TL/62.65. Verge of A45 E. of
bridge over River Kennett, TL/704.672. Both M. D. Crewe, 1993. 1st and 2nd records. STlhs
Lanarks.: Damp ground by ponds, Hamilton, NS/72.55. P. Macpherson, 1993, herb. P.M. Bare
waste ground, Glasgow, NS/56.65. P. Macpherson & E. K. Lindsay, 1993, herb. P.M., det J. R.
Akeroyd. 1st and 2nd records.

+40/2.3. ALNUS CORDATA *58, Cheshire: Edge of wood, High Lane, Disley, SJ/950.849. G. M.
Kay, 1994.

43/1.13. CHENOPODIUM FICIFOLIUM 46, Cards.: Disturbed refuse tip, Pendinas, Aberystwyth,
SN/585.799. J. P. Woodman & A. O. Chater, 1994, NMW. 2nd record.

+43/3.1. ATRIPLEX HORTENSIS *38, Warks.: Side of lane, Farnborough, SP/435.498. P. J.
Copson, 1984, WAR, det. J. C. Bowra. Heaps of topsoil, Long Itchington, SP/404.668. J. W.
Partridge, 1987, det. J. C. Bowra. 1st and 2nd records.

43/3.4. ATRIPLEX LONGIPES *46, Cards.: Top of salt marsh, S. side of Dyfi estuary E. of mouth
of Afon Leri, SN/620.936. A. O. Chater, 1993, NMW, det. J. R. Akeroyd.

+46/1.4. ARENARIA BALEARICA 40, Salop: Bank, Church Stretton, SO/456.951. J. Warren,
1990. 2nd record.

46/5.5. STELLARIA HOLOSTEA 102, S. Ebudes: Edge of bracken stand, Scallasaig, Colonsay,
NR/394.943. R. L. Gulliver, 1994. 1st post-1930 Colonsay record.

PLANT RECORDS 421

46/7.11. CERASTIUM PUMILUM 49, Caerns.: Limestone grassland, Nant y Gamar, Llandudno,
SH/80.81. G. Battershall, 1992. Only extant locality.

46/8.1. MYOSOTON AQUATICUM +*49, Caerns.: Flower beds, Llandudno, SH/780.827. W.
McCarthy, 1994, NMW.

46/9.1. MOENCHIA ERECTA 46, Cards.: Dry slope below coastal footpath W. of Banc,
Llanrhystud, SN/534.703. J. Turner & J. Woodman, 1994. Only extant locality.

46/10.3. SAGINA SUBULATA 42, Brecs.: Tirabad Cross, Cefngorwydd, SN/8.4. R. G. Woods,
1994. 2nd record.

46/10.8. SAGINA MARITIMA 81, Berwicks.: Concrete by sea, Burnmouth harbour, NT/959.608.
M. E. Braithwaite, 1994, herb. M.E.B. 2nd record, Ist since 1896. Concrete by sea, Glensgreen,
Eyemouth, NT/946.645. M. E. Braithwaite, 1994, herb. M.E.B. 2nd extant locality.

46/11.2. SCLERANTHUS ANNUUS 99, Dunbarton: Shandon, NS/25.86. J. H. Penson, 1971. Ist
record since 1886. Abandoned sand quarry, Callendoun Quarry, Dumfin, NS/33.84. K. Futter,
1994. 2nd post-1930 record.

746/18.1. LYCHNIS CORONARIA *12, N. Hants.: Well naturalized in rough grassland E. of
Blackbushe, SU/812.594. C. R. Hall, 1992. 46, Cards.: Rough ground by St Mary’s College,
Bryn Road, Aberystwyth, SN/586.823. S. P. Chambers, 1994. 2nd record. 69, Westmorland:
Coal yard, Arnside, SD/461.789. M. Baecker, 1993. 2nd record.

+46/20.9a X 10. SILENE LATIFOLIA subsp. ALBA X S. DIOICA (S. X HAMPEANA) *73, Kirkcud-
brights.: With both parents on embankment of disused railway, Summerville, Dumfries, NX/95.76.
O. M. Stewart, 1994.

46/20.12. SILENE CONICA +*40, Salop: New bank by Oswestry Bypass, SJ/295.272. M.
Wainwright, 1987, det. P. M. Benoit.

+47/1.2. PERSICARIA CAMPANULATA *12, N. Hants.: Towpath, Basingstoke Canal, Pondtail,
SU/82.53. C. R. Hall, 1986, det. A. R. G. Mundell.

47/4.2a. POLYGONUM OXYSPERMUM Subsp. RAII *102, S. Ebudes: Rocky shore just above strand
line, Kiloran Bay, Colonsay, NR/396.979. R. L. & M. C. Gulliver, 1993. 1st localized record.

47/4.3. POLYGONUM ARENASTRUM 102, S. Ebudes: Disturbed ground in croftyard, Colonsay,
NR/372.954. R. L. & M. C. Gulliver, 1991. 1st Colonsay record. 3

47/4.5. POLYGONUM BOREALE 73, Kirkcudbrights.: Mound of earth, Southerness, NX/975.542.
O. M. Stewart, 1994. 2nd record.

+47/5.1 X 2. FALLOPIA JAPONICA X F. SACHALINENSIS (F. X BOHEMICA) *38, Warks.: Riverside
scrub, Leamington Spa, SP/308.656. J. W. Partridge, 1993, WAR, det. J. C. Bowra.

47/5.4. FALLOPIA CONVOLVULUS 102, S. Ebudes: Barley field W. of road to Kiloran Bay,
Colonsay, NR/397.976. R. L. Gulliver, 1992. 1st post-1930 Colonsay record.

47/8.8. RUMEX LONGIFOLIUS 81, Berwicks.: Verge of A6089'N. of Gordon, NT/629.451. R. W.
M. Corner, 1994. 2nd extant locality. *100, Clyde Is.: Strandline, Lamlash, Arran, NS/028.311.
R. W. M. Corner, 1989, herb. R.W.M.C., det. J. R. Akeroyd.

+47/8.11. RUMEX CRISTATUS *12, N. Hants.: Growing through tarmac on side of B3011,
Heckfield, SU/733.608. D. L. Brookman, 1993, herb. Lady A. Brewis.

+47/8.12. RUMEX PATIENTIA *12, N. Hants.: Station yard, Aldershot, SU/86.50. A. C. Leslie,
1983, herb. Lady A. Brewis.

50/1.1. ELATINE HEXANDRA *42, Brecs.: Margin of reservoir emptied for repair, Dolymynach
Reservoir, SN/907.617. C. Mosscroft, 1994.

*51/1.1. HYPERICUM CALYCINUM *77, Lanarks.: Wood, Cambuslang, NS/63.60. P. Macpher-
son, 1993, herb. P.M., det. N. K. B. Robson. 1st record of established population.

422 PLANT RECORDS

+51/1.6 X 7. HYPERICUM PERFORATUM X H. MACULATUM (H. X DESETANGSII) *98, Main Argyll:
By gravel track at head of Loch Feochan, NM/876.246. B. H. Thompson, 1994, det. N. K. B.
Robson as probably nothovar. carinthiacum.

51/1.14. HYPERICUM HIRSUTUM *2, E. Cornwall: Open scrub on disused railway embankment
S. of Bagbury Farm near Bude, SS/213.051. T. J. Dingle, 1994.

753/1.6. MALVA PUSILLA 40, Salop: Garden weed, Wentnor, SO/383.926. S. Kingsbury, 1991.
2nd record.

+52/2.3. LAVATERA OLBIA *35, Mons.: Rough vegetation at side of woodland track, Ifton
Great Wood, ST/460.895. T. G. Evans, 1994, NMW.

57/1.9b. VIOLA PALUSTRIS subsp. JURESSI *46, Cards.: Alder carr, Rhos Pil-bach D.W.T.
Reserve, Plwmp, SN/367.529. A. O. Chater & S. P. Chambers, 1994.

+57/1.10. VIOLA CORNUTA 44, Carms.: Roadside hedgebank, Blaendyfnant near Llanfyrnach,
SN/236.294. G. Hutchinson & R. D. Pryce, 1994, NMW. 3rd Welsh record.

61/1.41 xX 2. PopULUS ALBA X P. TREMULA (P. X CANESCENS) *77, Lanarks.: Rubbly waste
ground, Dalmarnock, Glasgow, NS/61.63. P. Macpherson, 1993, herb. P.M., det. R. D. Meikle. 1st
record of established population.

61/1.3a. POPULUS NIGRA subsp. BETULIFOLIA “46, Cards.: Bank of Afon Llanfihangel,

Llanfihangel-y-Creuddyn, SN/658.762. A. O. Chater, 1992, NMW, det. E. Milne-Redhead.
Wooded bank of dry gulley, Penparc, SN/212.485. A. O. Chater, 1994, NMW. 1st and 2nd records
of native trees. |

+61/1.4. PoPULUS TRICHOCARPA *58, Cheshire: Canal bank, Wolverham, SJ/41.75. G. M. Kay,
1994. Suckering freely. *77, Lanarks.: Old quay, Glasgow, NS/57.64. P. Macpherson, 1994,
herb. P.M., det. R. D. Meikie.

61/2.12 X 16. SALIX AURITA X S. REPENS (S. X AMBIGUA) *80, Roxburghs.: Mire, Blind Moss,
Ashkirk, NT/458.185. R. W. M. Corner, 1994, herb. R.W.M.C., det. R. D. Meikle.

62/11.1. BARBAREA VULGARIS *100, Clyde Is.: Streamside, Torrylinn Shore, Kilmory, Arran,
NR/957.207. T.. Smith, 1988, det. T. C. G. Rich.

¥62/11.3. BARBAREA INTERMEDIA “78 Peebless.: Roadside near Walkerburn, NT/351.374.
Mrs Grant, 1981, herb. M. E. Braithwaite, det. M.E.B. Still present in Walkerburn in 1994. Near
foot of Holm’s Water, Rachan, NT/12.34. D. J. McCosh, 1994. Ist and 2nd records.

62/12.3. RORIPPA ISLANDICA *41, Glam.: Near River Loughor, Garnswllt, SN/62.10. R. G.
Ellis, 1981, NMW, det. T. C. G. Rich.

62/12.5. RORIPPA SYLVESTRIS *91, Kincardines.: By River Dee, Banchory, NO/795.957. D.
Welch, 1978.

+62/15.5. ARABIS CAUCASICA *39, Staffs.: Road verge under hedge, Yewtree Lane, Tetten-
hall, SJ/871.003. B. R. Fowler, 1994, herb. B.R.F.

62/15.6. ARABIS HIRSUTA 102, S. Ebudes: Balnahard Dunes, NR/427.999. R. L. & M. C.
Gulliver, 1993. Ist post-1930 Colonsay record.

+62/18.2. ALYSSUM SAXATILE *42,. Brecs.: River bank N.W. of Gilwern, SO/242.154. M.
Porter, 1992.

62/22.1. EROPHILA MAJUSCULA *38, Warks.: Garden weed, Warwick, SP/281.657. K. Davies,
1987, WAR, det. T.‘T. Elkington.

62/22.3. EROPHILA GLABRESCENS “12, N. Hants.: Sandy forest land, Longmoor Inclosure,
Woolmer, SU/783.294. Sandy verge, N. of Woolmer Road E. of Greatham, SU/789.313. Both F.
Rose, 1992, herb. Lady A. Brewis, det. T. T. Elkington. Ist and 2nd records. *38, Warks.:
Quarry, Alderminster, SP/255.505. Car park, Leamington Spa, SP/327.645. Both J. W. Partridge,
1987, WAR, det. T. T. Elkington. Ist and 2nd records. *43, Rads.: Bank of rocky track,

PLANT RECORDS 423

Cascob, $O/234.669. D. R. Humphreys, 1994, conf. R. G. Woods. *52, Anglesey: Sand dunes,
Aberffraw, SH/35.68. J. Lowell, 1994. *77, Lanarks.: Rocky streamside, Daer Water, NS/
97.09. P. Macpherson, 1994, herb. P.M., conf. T. T. Elkington. *98, Main Argyll: Pathside,
Dunstaffnage Castle, NM/88.34. P. Macpherson, 1957, herb. P.M., det. T. T. Elkington.

62/23.5. COCHLEARIA DANICA +*42, Brecs.: Roadside N.W. of Erwood, SO/075.444. R. G.
Woods, 1994.

62/30.3. LEPIDIUM HETEROPHYLLUM 26, W. Suffolk: Meadow near River Lark, Bury St
Edmunds, TL/858.650. N. Gibbons, 1993, conf. G. Beckett. 2nd extant locality.

62/30.5. LEPIDIUM RUDERALE *26, W. Suffolk: Verge of slip road at junction of A45 and A142,
Newmarket, TL/633.659. M. D. Crewe, 1993. 40, Salop: Festival Square, Oswestry, SJ/
282.294. E. Roberts, 1989. Ist post-1930 record.

62/30.6. LEPIDIUM LATIFOLIUM +*40, Salop: Near River Onny at Craven Arms, SO/437.826.
W. Prestwood, 1989. Oswestry, SJ/294.299. E. Townsend, 1993. Ist and 2nd records.

62/31.1. CORONOPUS SQUAMATUS 81, Berwicks.: Path through arable field, Barefoots, Eye-
mouth, NT/941.648. M. E. Braithwaite, 1994, herb. M.E.B. Path through arable field above
Hallydown shore, NT/927.653. M. E. Braithwaite, 1994. Ist and 2nd post-1930 records.

762/34.4. BRASSICA JUNCEA 25, E. Suffolk: Waste land alongside railway, Nacton, TM/
217.406. E. M. Hyde, 1993, conf. T. C. G. Rich. 2nd record.

+62/36.1: ERUCASTRUM GALLICUM *49, Caerns.: Tarmac on promenade, Llandudno, SH/
792.821. W. McCarthy, 1994.

162/40.1. RAPISTRUM RUGOSUM *50, Denbs.: New roadside by car park, Ysbytty Ifan, SH/
842.488. W. McCarthy, 1994, NMW. Field edge, Glan Conwy, SH/802.766. W. McCarthy, 1994. Ist
and 2nd records.

+62/42.1a. RAPHANUS RAPHANISTRUM 102, S. Ebudes: Kale field, Uragaig, Colonsay, NR/
388.981. R. L. Gulliver, 1994. 1st post-1930 Colonsay record.

64/1.1a. EMPETRUM NIGRUM subsp. NIGRUM 52, Anglesey: Maritime heath, Ynys Llanddwyn,
SH/3.6. M. & L. Howe, 1994. 2nd record.

+65/8.1. GAULTHERIA SHALLON *46, Cards.: Well naturalized in woodland, Furnace, SN/
684.952. A. O. Chater, 1993. Extensive thickets in woodland and scrub, Glandyfi Castle, SN/
692.965. A. O. Chater & W. M. Condry, 1994. 1st and 2nd records.

+65/8.3. GAULTHERIA MUCRONATA 58, Cheshire: Heath, Caldy Hill, SJ/224.858. D. P. & J. B.
Earl, 1994. 2nd record.

69/1.1 X 3. PRIMULA VULGARIS X P. VERIS (P. X POLYANTHA) *73, Kirkcudbrights.: Wood near
shore, St Mary’s Isle, NX/671.850. O. M. Stewart, 1994.

+69/1.9. PRIMULA JAPONICA *77, Lanarks.: Damp wood, Busby, NS/58.55. P. Macpherson,
1989, herb. E. J. Clement, det. E.J.C. as ‘Postford White’. Still present in 1994.

69/2.1. HOTTONIA PALUSTRIS +*77, Lanarks.: Abundant in pond, Hamilton, NS/724.566. K.
Watson, 1993, GL.

+69/3.2. CYCLAMEN COUM “14, E. Sussex: Naturalized in churchyard, Balcombe, TQ/307.309.
Peeiaeare, 1994° cont. P. A. Harmes.

69/6.3. ANAGALLIS MINIMA *40, Salop: Wimper Hill Wood, Wyre Forest, SO/732.763. J.
Bingham, 1991.

771/1.1. PHILADELPHUS CORONARIUS *38, Warks.: Railway embankment, Warwick, SP/
294.654. J. C. Bowra, 1983, WAR, det. P. J. Copson. Still present in 1993.

424 PLANT RECORDS

+71/1.1 X mic. X pub. PHILADELPHUS CORONARIUS X P. MICROPHYLLUS X P. PUBESCENS (P. xX
VIRGINALIS) “77, Lanarks.: edge of rough grassland, Burnside, Glasgow, NS/61.59. P. Macpher-
son, 1994, herb. P.M.

72/2.6. RIBES ALPINUM +58, Cheshire: Hedgerow in green lane, Disley, SJ/986.846. G. M.
Kay, 1994. Ist record this century, from site where species was recorded last century. +73,
Kirkcudbrights.: Embankment of A75, Ramscale Bridge, NX/792.665. O. M. Stewart & A. White,
1994. Ist record since 1887.

73/1.1. CRASSULA TILLAEA *19, N. Essex: Floor of gravel pit, Alphamstone, TL/872.353. T.
Tarpey, 1994.

+73/1.3. CRASSULA HELMSII *26, W. Suffolk: Farm pond, Long Melford, TL/879.445. D.
Casey, 1983. *40, Salop: Pool, Haughton, Shawbury, SJ/554.165. W. Prestwood, 1988, det. C.
A. Sinker. Brown Moss, SJ/563.394. I. C. Trueman, 1990. Ist and 2nd records.

+73/5.4. SEDUM SPECTABILE *25, E. Suffolk: Grassy strip between houses and shingle beach,
Shingle Street, TM/367.426. M. D. Crewe, 1993.

+73/5.7. SEDUM SPURIUM *58, Cheshire: Wasteland, Brinnington, Stockport, SJ/916.929. E.
Kearns, 1993, det. G. M. Kay.

+74/4.1. DARMERA PELTATA “98, Main Argyll: Roadside near Invereck, Kilmun, NS/142.830.

A. McG. Stirling, 1983.

+74/5.6. SAXIFRAGA ROTUNDIFOLIA *77, Lanarks.: Well established on river shingle, Kelvin-
side, NS/56.67. A. Walker et al., 1986, GL, still present in 1994. River shingle, Kelvindale, NS/
56.66. K. Watson, 1991. Ist and 2nd records.

74/5.16. SAXIFRAGA HYPNOIDES 50, Denbs.: Wet cliffs, Craig Berwyn, SJ/076.333. J. A.
Green, 1994, NMW. Only extant locality.

+74/7.1. TOLMIEA MENZIESII 46, Cards.: Frequent dense patches in woodland, Cwm Wyre E.
of Llanrhystud, SN/556.698. J. Davies, 1993, det. A. O. Chater. 2nd record.

+74/8.1. TELLIMA GRANDIFLORA 79, Selkirks.: Side of River Yarrow, Bowhill, NT/432.281. R.
W. M. Corner, 1994, herb. R.W.M.C. 2nd record. ;

+75/3.1 X 2. SPIRAEA SALICIFOLIA X S. ALBA (S. X ROSALBA) *50, Denbs.: Alwen Reservoir,
SH/953.534. A. P. Jones, 1994, NMW.

+75/3.1 < 3. SPIRAEA SALICIFOLIA X S. DOUGLASII (S. X PSEUDOSALICIFOLIA) *50, Denbs.:
Cadney Moss, SJ/469.348. K. Watson, 1994.

+75/5.1. HOLODISCUS DISCOLOR *73, Kirkcudbrights.: By River Nith, Maxwelltown, NX/
967.760. O. M. Stewart & A. White, 1993.

+75/8.5 X 6. RUBUS ODORATUS X R. PARVIFLORUS 13, W. Sussex: Traffic island S.E. of
Chichester, SU/878.041. A. C. Leslie, G. H. Foster, P. A. Harmes et al., 1985, det. A. C. Leslie.
Still present in 1994,

75/8.51. RUBUS ERRABUNDUS “64, Mid-W. Yorks.: Riverside, Dunsop Bridge, SD/659.503.
Dr Bane 90S: *79, Selkirks.: Roadside, Yarrow Kirk, NT/35.27. A. Newton, 1993, herb.
A.N.

+75/8.55. RUBUS LACINIATUS *42, Brecs.: Woodland, Llangynidr, SO/164.200. M. Porter &
L. Kellaway, 1994. “44, Carms.: Base of wall, lane N. of Cedric Street, Llanelli, SN/512.001. I.
K. Morgan, 1994, NMW, det. G. Hutchinson.

75/8.81. RUBUS SCIOCHARIS “14, E. Sussex: Heath, ‘Gills Lap’, Ashdown Forest, TQ/46.31.
A. Bull & A. Newton, 1994.

75/8.92. RUBUS BOUDICCAE *53,S. Lincs.: Roadside, Doddington, SK/9.7. L. Cumming & A.
Ley, 1907, OXF, det. D. E. Allen:

PLANT RECORDS 425

+75/8.102. RUBUS ELEGANTISPINOSUS *13, W. Sussex: Roadside, Stoneleigh Forest near
Emsworth, SU/748.087. D. E. Allen, 1993, BM, conf. A. Newton.

75/8.105. RUBUS INCURVATIFORMIS *49, Caerns.: Hedgerow, Ty’n-y-Groes, SH/78.71. G.
Battershall, 1993, det. A. Newton. *50, Denbs.: Hedge, Bryn y Maen, SH/836.765. G.
Battershall, 1993, det. A. Newton.

75/8.109. RUBUS LINDEBERGII *79, Selkirks.: Roadside, Foulshiels, Yarrow, NT/42.29. A.
Newton, 1993, herb. A.N.

75/8.111. RUBUS MILFORDENSIS *11, S. Hants.: Gateway by Dunwood near Romsey, SU/
306.230. D. E. Allen, 1985, BM, conf. A. Newton.

75/8.132. RUBUS SPRENGELII *44, Carms.: Edge of forestry plantation, Forestry Picnic Site
near Cenarth, SN/262.408. B.S.B.I. meeting, 1994, NMW, det. A. Newton.

75/8.134. RUBUS ARMENIACUS *§2, Anglesey: Car park, Beaumaris, SH/60.76. D. P. Earl,
1994.

75/8.140. RUBUS ROSSENSIS *1, W. Cornwall: Scrub, Tregarne Beacon, The Lizard, SW/
78.23. L. J. Margetts, 1983, det. A. Newton.

75/8.143. RUBUS WINTERI *10, Wight: Chalk scrub, Tennyson Down, SZ/332.857. D. E.
Allen, 1993, BM, conf. A. Newton.

75/8.144. RUBUS ADSCITUS *44, Carms.: Field hedge, Springfield Road, Carmarthen, SN/
412.212. B.S.B.I. meeting, 1994, NMW, det. A. Newton.

75/8.174. RUBUS WIRRALENSIS *64, Mid-W. Yorks.: Laneside, Great Mitton, SD/712.415. D.
P. Earl, 1994.

75/8.176. RUBUS AEQUALIDENS *9, Dorset: Heathy hill slopes, Pilsdon Pen, Broadwindsor,
ST/41.01. D. E. Allen, 1994, BM, det. A. Newton.

75/8.180. RUBUS DIVERSUS *12, N. Hants.: Old planted-over common, Fox Plantation,
Tangley, SU/33.52. D. E. Allen, 1993, BM, conf. A. Newton.

75/8.193. RUBUS MELANODERMIS *44, Carms.: Roadside hedge, Cwm Cathan, Garnswllt, SN/
628.099. B.S.B.I. meeting, 1994, det. A. Newton.

75/8.194. RUBUS MICANS *35, Mons.: Trackside in Hale Wood, ST/412.974. T. G. Evans,
1994, herb. T.G.E., det. A. Newton.

75/8.200. RUBUS TRICHODES *10, Wight: Rough ground, Tennyson Down, SZ/332.857. D. E.
Allen, 1993, BM, det A. Newton.

75/8.214. RUBUS FORMIDABILIS *11, S. Hants.: Rides, Parnhold Wood, SU/37.28. D. E.
Allen, 1994, BM, conf. A. Newton.

75/8.218. RUBUS LEYANUS *26, W. Suffolk: Assington Thicks, TL/92.38. A. L. Bull, 1979,
herb. A.L.B., conf. A. Newton.

75/8.222. RUBUS ADENANTHOIDES *79, Selkirks.: Side of B7039 E. of Bowhill, NT/43.27. A.
Newton, 1993, herb. A.N. *99, Dunbarton: Trackside by margin of wood, Rosneath, NS/
273.819. A. McG. Stirling, 1994, GLAM.

75/8.231. RUBUS ECHINATOIDES *79, Selkirks.: Roadside near Yarrow Kirk, NT/35.27. A.
Newton, 1993, herb. A.N.

75/8.234. RUBUS FLEXUOSUS _ *46, Cards.: Dominant in oak/ash wood W. of Nant Adal,
Llanilar, SN/622.743. D. E. Allen & A. O. Chater, 1994, NMW, det. D.E.A.

75/8.264. RUBUS ATREBATUM *9, Dorset: Hedges, Pilsdon Pen, ST/413.011. D. E. Allen,
1994, BM, conf. A. Newton. Extension of range 130 km westwards.

426 PLANT RECORDS

75/8.305. RUBUS BRITANNICUS *12, N. Hants.: Clay copse, Clanville Lodge near Andover,
SU/322.483. D. E. Allen, 1993, BM, conf. A. Newton.

75/8.310. RUBUS HEBRIDENSIS *94, Banffs.: Dufftown, NJ/323.392. D. Welch, 1992, ABD,
det. A. Newton.

75/8.313. RUBUS NEMOROSUS *46, Cards.: Thickets on stream bank, Afon Wyre E. of
Llanrhystud church, SN/538.696. D. E. Allen & A. O. Chater, 1993, BM, conf. A. Newton.

75/8.314. RUBUS PICTORUM *42, Brecs.: Woodland, Cusop Dingle, SO/2.4. A. Ley, 1896,
BIRM, det. M. Porter.

75/8.321. RUBUS CAESIUS *67, S. Northumb.: By disused railway track near Wylam, NZ/
125.649. Shaded bank of River Tyne near Riding Mill, NZ/017.618. Both G. A. Swan, 1994, herb.
G.A.S., det. A. Newton. Ist and 2nd confirmed records. *68, Cheviot: Open wood above
Howick Burn, NU/251.172. G. A. Swan, 1994, herb. G.A.S., det. A. Newton. Ist confirmed record.

75/8.ar1. RUBUS ARICONIENSIS *44, Carms.: Edge of forestry plantation, Forestry Picnic Site
near Cenarth, SN/262.406. B.S.B.I. meeting, 1994, NMW, det. A. Newton.

75/8.vag. RUBUS VAGENSIS *42, Brecs.: Woodland, Park Wood, Talgarth, SO/168.338. M.
Porter, 1991, herb. M.P.

75/13.2. GEUM URBANUM 102, S. Ebudes: By unmetalled road through Colonsay House
Woods, Colonsay, NR/393.966. R. L. & M. C. Gulliver, 1993. 1st post-1930 Colonsay record.

75/15.2. AGRIMONIA PROCERA 50, Denbs.: Laneside bank, Nantglyn, SJ/009.636. W.
McCarthy, 1994. 1st post-1930 record.

+75/17.3b. SANGUISORBA MINOR subsp. MURICATA 26, W. Suffolk: Bank by A45, Bury St
Edmunds, TL/862.644. M. D. Crewe, 1993. Ist post-1930 record. *35, Mons.: Grassy bank of
Nant Hafod-Tudur, Wattsyille, ST/203.916. T. G. & U. T. Evans, 1986, herb. T.G.E. Bank of River
Ebbw, Risca, ST/2.9. J. Harper, 1992. 1st and 2nd records. *77, Lanarks.: Grassy area, New
Stevenston, NS/761.598. P. Macpherson, 1993, herb. P.M. 1st record of established population.

+75/18.1. ACAENA OVALIFOLIA *77, Lanarks.: Long abandoned railway platform, Hillhead,
NS/56.67. P. Macpherson, 1994, herb. P.M., conf. P. F. Yeo. -

+75/18.4. ACAENA INERMIS *44, Carms.: Disturbed river shingle, Carreg Sawdde Common,
Llangadog, SN/706.277. I. K. Morgan & R. D. Pryce, 1987, NMW, det. G. Hutchinson. 1st Welsh
record. :

+75/19.15. ALCHEMILLA MOLLIS *40, Salop: By path, Marrington Dingle, SO/27.98. F. H.
Perring, 1988. *48, Merioneth: Bushy lake shore, Llyn Tegid near Llanycil, SH/9.3. P. M.
Benoit & A. O. Chater, 1993, NMW.

75/21.5 X 17. ROSA PIMPINELLIFOLIA X R. MOLLIS (R. X SABINII) *98, Main Argyll: Foot of cliff
of raised beach S.E. of Castle Coeffin, Lismore, NM/855.436. A. McG. Stirling & B. H. Thompson,
1994, det. G. G. Graham.

+75/21.6. ROSA RUGOSA *80, Roxburghs.: Steep bank above Kale Water near Corbet Tower,
NT/778.238. R. W. M. Corner, 1991, herb. R.W.M.C. Hedge N. of Lochside, Yetholm, NT/
800.283. R. W. M. Corner, 1994, herb. R.W.M.C. Ist and 2nd records.

75/21.12. ROSA CANINA 102, S. Ebudes: Scrubby margin of road, W. side of Kiloran Bay,
Colonsay, NR/397.978. C. Wilson, 1993. 1st post-1930 Colonsay record. 108, W. Sutherland:
Rocky trackside, Ardroe, NC/071.246. P. A. Evans, 1994, det. G. G. Graham as Group
Transitoriae. 1st confirmed record.

75/21.12 X 5. ROSA PIMPINELLIFOLIA X R. CANINA (R. X HIBERNICA) *49, Caerns.: Roadside
hedge, Porth Neigwl, SH/262.292. A. Burns & A. P. Conolly, 1994, det. A. L. Primavesi. 2nd

~ Welsh record.

PLANT RECORDS 427

75/21.12 X 13b. ROSA CANINA X R. CAESIA subsp. GLAUCA (R. X DUMALIS) *1, W. Cornwall:
Scrub on bank near Rosehill, Penzance, SW/462.306. D. T. Holyoak, 1994, conf. A. L. Primavesi.

75/21.13b. ROSA CAESIA subsp. GLAUCA 3, S. Devon: Top of beach, Strete Gate, SX/835.456.
R. E. N. & C. J. Smith, 1994, det. G. G. Graham. Ist confirmed record. 108, W. Sutherland:
Roadside scrub, Achmelvich Road, NC/082.247. P. A. Evans, 1994, det. G. G. Graham. 2nd post-
1930 record.

75/21.15. ROSA TOMENTOSA *68, Cheviot: Hedge near Alnmouth, NU/239.107. G. A. Swan,
1994, herb. G.A.S., det. G. G. Graham.

75/21.16 X 5. ROSA SHERARDII X R. PIMPINELLIFOLIA (R. X INVOLUTA) *46, Cards.: Grassy
coastal heath, R.A.E. site, Aberporth, SN/245.524. A. O. Chater, 1994, NMW, det. G. G. Graham
as hybrid with R. sherardii as female parent.

75/21.18. ROSA RUBIGINOSA *67, S. Northumb.: Under deciduous trees on E. bank of River
South Tyne, NY/683.518. G. A. & M. Swan, 1994, herb. G.A.S., det. G. G. Graham. Ist confirmed
record. 73, Kirkcudbrights.: Bank of river, Holm of Dalry, NX/614.810. O. M. Stewart, 1993,
E. Only extant locality.

75/21.19. ROSA MICRANTHA *68, Cheviot: Hedge by road near Weldon, NZ/139.993. G. A.
Swan, 1994, herb. G.A.S., det. G. G. Graham. Ist confirmed record.

75/21.20. ROSA AGRESTIS 11, S. Hants.: One shrub in scrub on chalk grassland, Gallows Hill,
SU/134.212. P. D. Stanley, 1994, herb. R. P. Bowman, conf. R.P.B. 2nd record.

+75/28.7. SORBUS INTERMEDIA *40, Salop: Lynclys, SJ/273.240. J. Pedlow, 1990, det. P. J. M.
Nethercott.

+75/30.1. AMELANCHIER L AMARCKII “77, Lanarks.: Old industrial site, Dalmarnock, NS/
61.63. P. Macpherson, 1993, herb. P.M., conf. R. D. Meikle.

+75/32.9. COTONEASTER FRIGIDUS 25, E. Suffolk: Long established in grounds of old isolation
hospital, Ufford, TM/288.518. M. D. Crewe, 1993. 2nd record. *44, Carms.: Hedgerow,
Kidwelly, SN/399.063. G. Hutchinson, 1990, NMW, det. J. Fryer.

+75/32.9 X 10. COTONEASTER FRIGIDUS X C. SALICIFOLIUS (C. X WATERERI) *50, Denbs.:
Woods, Colwyn Bay, SH/844.786. G. Battershall, 1993, det. J. Fryer.

+75/32.10. COTONEASTER SALICIFOLIUS *49, Caerns.: Hedgerow, Little Orme, SH/81.81. G.
Battershall, 1993, det. J. Fryer.

175/32.11. COTONEASTER DAMMERI *58, Cheshire: Riverbank, Wilmslow, SJ/838.822. D.
Shaw, 1993. *77, Lanarks.: Breast wall of River Clyde near Govan, NS/56.65. P. Macpherson,
1994, herb. P.M., det. J. Fryer. 2nd record.

175/32.13. COTONEASTER LACTEUS *47, Monts.: Limestone scree, Llanymynech Rocks, SJ/
261.218. M. Wainwright & J. Pedlow, 1993, det. J. Fryer.

¥75/32.22. COTONEASTER HJELMQVISTII *47, Monts.: Limestone scree, Llanymynech Rocks,
SJ/261.218. M. Wainwright & J. Pedlow, 1993, det. J. Fryer. *49, Caerns.: Edge of path, West
Shore, Llandudno, SH/769.822. W. McCarthy, 1994, det. G. M. S. Easy.

+75/32.26. COTONEASTER DIVARICATUS *58, Cheshire: Wood on lime waste, Witton, SJ/
656.752. G. M. Kay & E. & O. Kearns, 1994, det. G. M. S. Easy.

175/32.29. COTONEASTER VILLOSULUS *77, Lanarks.: Wood near Busby, NS/58.55. P. Mac-
pherson, 1993, herb. P.M., det. J. Fryer.

775/32.33. COTONEASTER SIMONSII *108, W. Sutherland: Rocky banks of burn, Tumore, NC/
186.269. P. A. Evans, 1994.

+75/32.34. COTONEASTER BULLATUS *50, Denbs.: Limestone rocks, Llandulas, SH/914.777.
W. McCarthy, 1994, det. J. Fryer. *69, Westmorland: Coppice behind Arnside railway station,
SD/462.788. M. Baecker, 1994, LANC, det. J. Fryer.

428 PLANT RECORDS

+75/32.39. COTONEASTER FRANCHETII *26, W. Suffolk: Roadside at junction of A45 and A120
N. of Newmarket, TL/633.660. M. D. Crewe, 1993.

+75/32.40. COTONEASTER STERNIANUS *35, Mons.: Top of railway embankment off Spytty
Road, Newport, ST/332.868. G. Hutchinson, 1990, NMW. *44, Carms.: Steep hedgebank S. of
Llanybyther, SN/52.42. G. Hutchinson, 1991, NMW, conf. J. Fryer. 1st Welsh records.

+75/35.5. CRATAEGUS PERSIMILIS *35, Mons.: Hedgerow, Y Sguborwen, ST/411.958. T. G.
Evans & P. Macpherson, 1993, NMW, conf. C. A. Stace. 1st Welsh record.

+75/35.10. CRATAEGUS LACINIATA *57, Derbys.: Disused railway sidings, Rowsley, SK/26.65.
Various recorders from 1970s onwards. Railway bank, Butterley, Ripley, SK/407.519. J. P. Bailey,
1994, LTR. Ist and 2nd records.

77/4.2. ASTRAGALUS DANICUS 108, W. Sutherland: Limestone grassland, Ardvreck Castle,
NC/238.237. T. Fowler, 1987; refound by P. A. Evans in 1994. 2nd record.

+77/14.11¢c. VIcIA SATIVA subsp. SATIVA *12, N. Hants.: Roadside below Noar Hill, SU/75.32.
Lady A. Brewis, 1982. Field edge, North Warnborough, SU/728.517. C. R. Hall, 1993, herb. Lady
A. Brewis. 1st and 2nd records. 46, Cards.: Reconstructed roadside slope by Glanyrafon
Industrial Estate, Llanbadarn Fawr, SN/609.805. S. P. Chambers, 1994, conf. A. O. Chater. Only
extant locality. *81, Berwicks.: Reseeded verge of Al, English Border, NT/974.570. M. E.
Braithwaite, 1994, herb. M.E.B. 1st localized record.

+77/15.7. LATHYRUS GRANDIFLORUS *12, N. Hants.: Scrub, Norris Hill, Fleet, SU/832.533. C.
R. Hall, 1993, herb. Lady A. Brewis. :

77/15.12. LATHYRUS NISSOLIA *40, Salop: Rough grassland, Randlay, Telford, SJ/712.072.
Roadside, Stirchley Grange, -Telford, SJ/702.070. Both J. Box & J. Mincher, 1985. Ist and 2nd
records.

77/19.1. TRIFOLIUM ORNITHOPODIOIDES 12, N. Hants.: Very sparse short vegetation on sandy
soil, Kennels Lane near Southwood, SU/840.548. T. Dove, 1992, herb. Lady A. Brewis, conf. A. R.
G. Mundell. Only extant locality.

+77/19.19a TRIFOLIUM INCARNATUM subsp. INCARNATUM *40, Salop: Banks of new A5 bypass,
SJ/618.107. J. Martin, 1990.

77/19.25. TRIFOLIUM SUBTERRANEUM *39, Staffs.: Sparse, rabbit-grazed turf, Kinver Edge,
SO/837.829. E. Blunt, 1993, det. M. Dowlen.

+77/22.1. LABURNUM ANAGYROIDES *49, Caerns.: Waste ground, Penrhyn Bay, SH/82.81. G.
Battershall, 1992.

+77/23.3. CYTISUS STRIATUS *35, Mons.: Side of A4042, Pontypool, SO/299.004 & 300.004. T.
G. Evans, 1990. Steep roadside bank N. of Goytre Farm, ST/477.977. T. G. Evans, 1991, NMW.
Still present at both localities in 1994. 1st and 2nd records.

+77/24.1. SPARTIUM JUNCEUM *50, Denbs.: Railway bank, Llandulas SH/894.784. G. Hutchin-
son, 1994.

77/26.3. ULEX MINOR 98, Main Argyll: Stony verge of forest road, A’Bheinn near Tibertich,
NM/846.033. B. H. Thompson, 1994, herb. B.H.T., conf. A. McG. Stirling. lst post-1930 record.

+79/2.2. MyRIOPHYLLUM AQUATICUM *35, Mons.: Pond, Little Campston, SO/37.24. T. G.
Evans, 1994. *40, Salop: Ditch near Fenemere, Eyton, SJ/443.232. R. Wells, 1990.

81/1.2. LYTHRUM HYSSOPIFOLIA +*35, Mons.: Weed of potato crop near pond, The Nurtons,
Tintern, SO/536.011. A. & E. Wood, 1994, NMW, conf. T. G. Evans.

84/1.3 X +8. EPILOBIUM MONTANUM X E. CILIATUM “47, Monts.: Gravel car park, Caersws,
S$O/031.918. P. M. Benoit, 1993.

+84/1.8. EPILOBIUM CILIATUM 102, S. Ebudes: Oronsay, NR/3.8. A. Walker, 1992. 1st record
from Colonsay or Oronsay.

PLANT RECORDS 429

+84/1.12. EpILOBIUM BRUNNESCENS 12, N. Hants.: Disturbed ground, recently planted with
shrubs, beside Zebon Coppice, Crookham, SU/802.518. C. R. Hall, 1991, herb. Lady A. Brewis.
2nd record. 102, S. Ebudes: Disturbed ground, Ballarulin, Colonsay, NR/384.963. R. L.
Gulliver, 1994. 1st Colonsay record.

+84/4.1 x 3. OENOTHERA GLAZIOVIANA X QO. BIENNIS *46, Cards.: Refuse tip, S.W. side of
Pendinas, Aberystwyth, SN/584.799. S. P. Chambers, 1994, det. J. C. Bowra.

84/6.1 x 2. CIRCAEA LUTETIANA X C. ALPINA (C. X INTERMEDIA) 50, Denbs.: Bank of River
Dee, Llangollen, SJ/208.431. J. A. Green, 1994. 2nd record.

+94/1.2. LINUM_ USITATISSIMUM *50, Denbs.: Dawn, Betws-yn-Rhos, SH/874.732. W.
McCarthy, 1994, NMW.

+103/1.6. GERANIUM PSILOSTEMON *77, Lanarks.: Grassy waste ground, site of Garden
Festival, Glasgow, NS/56.65. P. Macpherson, 1993. Edge of rough grassland, Burnside, Glasgow,
NS/61.59. P. Macpherson, 1994. Both herb. P.M., conf. P. F. Yeo. 1st and 2nd records.

+103/1.8. GERANIUM HIMALAYENSE *77, Lanarks.: Rubbly waste ground, site of Garden
Festival, Glasgow, NS/56.65. P. Macpherson, 1992, herb. P.M., det. E. J. Clement.

103/2.2. ERODIUM MOSCHATUM 46, Cards.: Verge of B4321, Llangranog, SN/312.541. A. O.
Chater & S. P. Chambers, 1994, NMW. 2nd record, Ist since 1941.

+105/1.2. IMPATIENS CAPENSIS *40, Salop: N. of Calcott Moss, SJ/448.142. P. Parker & J. Ing,
1984. Bank of River Tern S. of Market Drayton, SJ/671.334. F. H. Perring, 1988. 1st and 2nd
records. °

+105/1.3. IMPATIENS PARVIFLORA *91, Kincardines.: Bank of River North Esk, Kinaber, NO/
724.621. D. Welch, 1994, ABD.

+106/1.1. HEDERA COLCHICA *77, Lanarks.: Wood, Calder, NS/61.72. P. Macpherson & E. L.
S. Lindsay, 1991, herb. P. M., det. A. McG. Stirling as var. dentata. Grassy bank, Kelvinside, NS/
56.67. A. McG. Stirling, K. Watson & P. Macpherson, 1994. Ist and 2nd records. *98, Main
Argyll: Well established and spreading in disused garden, Ederline, Ford, NM/870.030. B. H.
Thompson, 1993, herb. B.H.T., det. A. Rutherford.

+107/5.1. CHAEROPHYLLUM HIRSUTUM *77, Lanarks.: River bank, Kelvinside, NS/57.67. A.
Walker, 1989, GL, conf. E. J. Clement. Still present in 1994.

107/6.2. ANTHRISCUS CAUCALIS 58, Cheshire: Dunes, Wallasey, SJ/284.931. G. M. Kay, 1994.
Only extant locality.

7107/10.1. SMyRNIUM OLUSATRUM *57, Derbys.: Roadside verge between Ingleby and Swar-
kestone, SK/361.274. C. Higginbottom, 1993, DBY. 1st localized record.

7107/20.1b. AETHUSA CYNAPIUM subsp. AGRESTIS *4, N. Devon: Waste ground, Torrington,
SS/492.196. W. H. Tucker, 1994, herb. W.H.T.

7107/21.1. FOENICULUM VULGARE *73, Kirkcudbrights.: Top of steep embankment by A755
W. of Kirkcudbright, NX/676.518. A. White & O. M. Stewart, 1994.

107/28.4. APrluUM INUNDATUM *78, Peebless.: Oxbow of River Tweed, Putts’ Pool, Inner-
leithen, NT/34.37. M. E. Braithwaite et al., 1994.

+107/32.1. AMMI MAJUS *40, Salop: Garden weed, Chapel Lane, Trefonnen, SJ/259.271. R.
Davies, 1988.

7107/33.1. FALCARIA VULGARIS - “14, E. Sussex: On railway bank and arable margin, Tidemills
near Seaford, TQ/463.003. D. J. Manning, 1993, herb. P. A. Harmes. 1st record of established
population.

+107/41.2. HERACLEUM MANTEGAZZIANUM 50, Denbs.: Old castle, Holt, SJ/412.537. J. Osley,
1993. 2nd record.

430 PLANT RECORDS

110/8.1. SOLANUM NIGRUM 102, S. Ebudes: Above strand line on shore N.W. of Rubha Dubh,
Colonsay, NR/393.916. R. L. & M. C. Gulliver, 1991. 1st Colonsay record.

7111/3.4. CALYSTEGIA SILVATICA *80, Roxburghs.: Garden hedge, Dean Road, Denholm, NT/
568.182. M. E. Braithwaite, 1988, herb. R. W. M. Corner. Roadside bank and hedge above E. end
of Linton Loch, Morebattle, NT/788.253. R. W. M. Corner, 1994, herb. R.W.M.C. ist and 2nd
records. *108, W. Sutherland: Well established garden weed, Lochinver, NC/093.223. P. A.
Evans, 1994.

112/1.3. CUSCUTA EPITHYMUM +59, S. Lancs.: On Cytisus scoparius in shrub bed planted two
years ago, Notre Dame Hospital, Taggart Avenue, Liverpool, SJ/406.875. S. Edmondson, 1994. ist
post-1930 record.

4115/1.1. PHACELIA TANACETIFOLIA *50, Denbs.: Weedy field, Dawn, Betws-yn-Rhos, SH/
874.733. W. McCarthy, 1994, NMW.

+116/3.4. PULMONARIA ‘MAWSON’S BLUE’ *39, Staffs.: Damp woodland, Patshall, SJ/800.007.
S. O’Donnell, 1994, herb. B. R. Fowler.

116/4.3. SYMPHYTUM TUBEROSUM +*46, Cards.: Hedgebank of lane to Y Felin, Ader-Mad, SN/
596.763. A. O. Chater, 1994, NMW.

+116/4.7. SYMPHYTUM CAUCASICUM 26, W. Suffolk: Roadside verge, Stoke-by-Nayland, TL/
998.378. F. W. Simpson, 1993. 2nd record.

¥116/12.2. AMSINCKIA MICRANTHA *40, Salop: Beach Bank Dingle, SJ/429.024. K. Thorn,
1991. *59, S. Lancs.: Weed in sugar beet field, Ince Blundell, SD/324.036. V. Gordon, 1994.

118/1.1. STACHYS OFFICINALIS 77, Lanarks.: Rough grassland near Stevenston, NS/76.59. P.
Macpherson, 1994, herb. P.M. Ist record this century.

4118/1.2. STACHYS BYZANTINA *35, Mons.: Waste ground in disused railway station, Golivon,
SO/27.13. R. Fraser, 1989. Well naturalized on waste ground, Comin, Coed-y-moeth, SO/158.017.
P. A. Smith, 1994. Ist and 2nd records. *77, Lanarks.: Grassy bank, Old Dock, Glasgow, NS/
57.64. P. Macpherson, 1994, herb. P.M.

118/1.5 X 6. STACHYS SYLVATICA X S. PALUSTRIS (S. X AMBIGUA) 102, S. Ebudes: Roadside by
turning to Balnahard from road to Kiloran, NR/399.974. R. L. Gulliver, 1993. 1st post-1930
Colonsay record.

118/1.8. STACHYS ARVENSIS 81, Berwicks.: Arable edge above Fleurs Dean, NT/924.653. M.
E. Braithwaite, 1994, herb. M.E.B. 2nd extant locality.

7118/4.1c. LAMIASTRUM GALEOBDOLON subsp. ARGENTATUM *57, Derbys.: Wooded trackside
by Pitt Wood, Alfreton, SK/412.547. R. Smith, 1993, DBY. 80, Roxburghs.: Woodland,
Maxton Woods, NT/609.303. E. G. Cutter, 1994. 2nd record.

118/5.4. LAMIUM HYBRIDUM *100, Clyde Is.: Waste ground on roadside, An Garradh,
Largybeg, Arran, NS/049.230. A. R. Church, 1988. 108, W. Sutherland: Recently disturbed
ground in car park, Lochinver, NC/094.224. P. A. Evans, 1994. 2nd record.

118/5.6. LAMIUM AMPLEXICAULE 102, S. Ebudes: Weed in cabbage field S.E. of Balnahard
Dunes, Colonsay, NR/401.975. R. L. Gulliver, 1994. 1st post-1930 record.

118/6.2. GALEOPSIS ANGUSTIFOLIA *40, Salop: Foot of scree, Jones’s Rough, SJ/247.247. C.
Johnson, 1987. British Telecom Yard, Llanymynech, SJ/266.210. F. H. Perring & M. Wainwright,
1992. 1st and 2nd records.

118/6.5. GALEOPSIS BIFIDA 102, S. Ebudes: Cabbage field, Colonsay, NR/372.954. R. L.
Gulliver, 1991. Ist Colonsay record.

118/23.2 X +3. MENTHA AQUATICA X M. sPICATA (M. X PIPERITA) 102, S. Ebudes: Sandy banks
beside burn through Kiloran Dunes, er NR/399.975. C. Wilson, 1993. Ist post-1930
Colonsay record.

PLANT RECORDS 43]

+118/23.3. MENTHA SPICATA 102, S. Ebudes: Marshland, Port Mor, Colonsay, NR/362.947. C.
Wilson, 1993. 1st Colonsay record.

118/23.+3 x 4. MENTHA SPICATA X M. SUAVEOLENS (M. X VILLOSA) *47, Monts.: Car park,
Caersws, SO/031.918. P. M. Benoit, 1993.

118/23.43 < lon. MENTHA SPICATA X M. LONGIFOLIA (M. X VILLOSONERVATA) “3,5: Devon:
Waste marshy ground, Torcross, Slapton Ley, SX/823.420. L. J. Margetts & L. M. Spalton, 1994,
herb. L.J.M., det. R. M. Harley. Ist certain record. *81, Berwicks.: Damp bank by burn,
Milldown Burn, Coldingham, NT/913.661. P. A. Green, 1954, E, det. D. R. McKean.

120/1.4. CALLITRICHE PLATYCARPA *75, Ayrs.: Wet depressions, Garnock Floods, Irvine New
Town, NS/30.41. K. Watson, 1992. Marshy depression, Knadgerhill Pond, Irvine New Town, NS/
33.40. K. Watson, 1992. Ist and 2nd records.

121/1.4. PLANTAGO MEDIA 77, Lanarks.: Grassy area, New Stevenston, NS/76.59. P. Macpher-
son, 1993, herb. P.M. Ist post-1930 record.

124/1.7 x 11. VERBASCUM THAPSUS X V. PULVERULENTUM (V. X GODRONII) *26, W. Suffolk: In
large colony of V. pulverulentum by side of A45, Higham junction, TL/748.663. M. D. Crewe, 1993.

124/5.1. LIMOSELLA AQUATICA 73, Kirkcudbrights.: Damp mud, Southwick Merse, NX/
918.564. O. M. Stewart, 1994. 2nd extant locality.

7124/15.1. ERINUS ALPINUS *77, Lanarks.: Well naturalized on walls where first planted c.
1967, Coulter, NT/02.33. D. J. McCosh, 1994.

+124/16.9. VERONICA SCUTELLATA *102, S. Ebudes: Margin of shallow pool, Lochan Chille
Moire, NR/361.888. A. Walker, 1992. 1st post-1930 record from Colonsay or Oronsay.

124/16.24. VERONICA HEDERIFOLIA *102, S. Ebudes: Disturbed ground at base of tree in
woodland, Colonsay House Woods, Colonsay, NR/393.966. R. L. Gulliver, 1993.

7124/17.1. HEBE SALICIFOLIA *73, Kirkcudbrights.: Abundant on wall by River Nith, Max-
welltown, NX/967.760. A. White & O. M. Stewart, 1993.

124/20.7 X 9. EUPHRASIA NEMOROSA X E. CONFUSA *25, E. Suffolk: Chalk pit, Blakenham
Chalk Pit Nature Reserve, TM/109.490. P. G. Lawson, 1993, det. A. J. Silverside. *47, Monts.:
Mortar-rich turf below tumbledown building, Dylife, SN/862.941. S. P. Chambers, 1993.

124/23.1. PARENTUCELLIA VISCOSA +46, Cards.: Reconstructed roadside slope by Glanyrafon
Industrial Estate, Llanbadarn Fawr, SN/609.805. S. P. Chambers, 1994. 2nd record.

¥125/1.2. LATHRAEA CLANDESTINA *40, Salop: Hardwick estate, Clive, SJ/510.222. A. John-
son, 1991.

7125/2.7. OROBANCHE CRENATA *18, S. Essex: One plant at edge of rape field, Cranham
Marsh, TQ/574.852. A. Dunton, 1994, conf. in field by K. J. Adams. Re-occurrence at only site in
Britain where this species has become established.

7129/1.9. CAMPANULA PORTENSCHLAGIANA *49, Caerns.: Wall between road and railway,
Cadnant Park, Conway, SH/777.777. R. Lewis, 1994, NMW.

130/6.5b. GALIUM PALUSTRE subsp. ELONGATUM *46, Cards.: Stream in wooded dingle E.N.E.
of Nantyronnen, Aber-ffrwd, SN/667.778. S. P. Chambers, 1993.

130/6.7. GALIUM MOLLUGO *78, Peebless.: Roadside near Broughton, NT/11.38. D. J.
McCosh, 1994, herb. D.J.McC.

130/7.1. CRUCIATA LAEVIPES 93, N. Aberdeen: Abandoned railway, Maud, NJ/930.481. M.
Faulkner & D. Potter, 1993, herb. D. Welch. 2nd record.

7131/1.3. SAMBUCUS CANADENSIS *77, Lanarks.: Old Canting Basin, NS/56.64. P. Macpher-
son, 1994, herb. P.M., det. R. D. Meikle as var. acutiloba.

432 PLANT RECORDS

*131/2.3. VIBURNUM TINUS 12, N. Hants.: Railway cutting, Overton, SU/51.50. J. & P.
Rollinson, 1993. 2nd record. *30, Denbs.: Woodland, Bryn Euryn, SH/83.80. Roadside, Tal-y-
cafn, SH/79.72. Both R. Lewis, 1993. 1st and 2nd records.

+131/3.orb. X mic. SYMPHORICARPOS ORBICULARIS X S. MICROPHYLLUS (S. X CHENAULTII) *44,
Carms.: Forge Quarry N. of Cwmdwyfran, SN/408.257. G. Hutchinson, 1992, NMW, det. R. M.
Burton. 1st Welsh record.

+131/5.1. LEYCESTERIA FORMOSA 50, Denbs.: Plas Muriau plantation, Capel Garmon, SH/
80.56. W. McCarthy, 1994. 2nd record.

+131/6.1. LONICERA PILEATA *77, Lanarks.: Waste ground, Hutchestown, Glasgow, NS/59.64.
P. Macpherson, 1988, herb. P.M., conf. E. J. Clement. Rubbly waste ground, Dalmarnock, NS/
61.63. P. Macpherson, 1993, herb. P.M., conf. D. R. McKean. Ist and 2nd records.

7131/6.3. LONICERA INVOLUCRATA “77, Lanarks.: Damp wood near Busby, NS/58.55. P.
Macpherson, 1992, herb. P.M., det. E. J. Clement. By River Kelvin, Kelvinside, NS/56.67. K.
Watson, 1993. Ist and 2nd records.

+131/6.4. LONICERA XYLOSTEUM 77, Lanarks.: Rough grassland at edge of wood, Burnside,
Glasgow, NS/61.59. P. Macpherson, 1994, herb. P.M. 2nd record.

+131/6.6. LONICERA JAPONICA “12, N. Hants.: By railway, Alice Holt, SU/8.4. Lady A. |
Brewis, 1971. Thoroughly established in scrub by road, Pondtail, Fleet, SU/823.535. C. R. Hall,
1993. 1st and 2nd records.

133/1.2. VALERIANELLA CARINATA *27, E. Norfolk: Top and base of garden wall, Kenninghall,
TM/043.859. J. Hawksford, 1994, herb. J.H.

134/1.1. DipsAcUS FULLONUM “91, Kincardines.: Waste ground next to gravel pit, Edzell, NO/
631.670. D. Welch, 1994. |

135/6.6 X 9. CIRSIUM HETEROPHYLLUM X C. PALUSTRE (C. X WANKELII) *78, Peebless.: With
both parents on bank of River Lyne near West Linton, NT/14.52. D. J. McCosh, 1994.

135/11.7. CENTAUREA NIGRA 75, Ayrs.: Light soil amongst boulders near the lighthouse, Ailsa
Craig, NX/02.99. B. Zonfrillo, 1994, GL. Ist Ailsa Craig record. )

135/16.3. LEONTODON SAXATILIS +81, Berwicks.: Old lawn, Mellerstain House, NT/648.389.
R. W. M. Corner, 1994. 2nd record.

+135/19.2. TRAGOPOGON PORRIFOLIUS 35, Mons.: Rough wet grassland, Alpha Steelworks,
Newport, ST/337.844. T. G. Evans & M. Jones, 1994, herb. T.G.E. 2nd record.

135/26.1. CREPIS PALUDOSA 102, S. Ebudes: Oronsay, NR/3.8. A. Walker, 1992. Ist record
from Colonsay or Oronsay.

7135/28.137. HIERACIUM SEVERICEPS *98, Main Argyll: Near Kentallen, NN/003.578. E.
Norman, 1993, BM, det. P. D. Sell.

135/28.147. HIERACIUM CALEDONICUM “94, Banffs.: Serpentine outcrop, Craigs of Succoth,
NJ/435.363. D. Welch, 1992, ABD, det. D. J. McCosh.

¥135/32.1. ANAPHALIS MARGARITACEA *12, N. Hants.: Anton Lakes Local Nature Reserve,
SU/357.466. M. F. Wildish, 1992. *40, Salop: Corbets Park, Wyre Forest, SO/740.778. J.
Bingham, 1990. 1st certain record.

+135/40.4a. SOLIDAGO GIGANTEA subsp. SEROTINA *50, Denbs.: Waste ground, Cadole, SJ/
205.626. K. Watson, 1993. 81, Berwicks.: Grass by Milldown Burn, Coldingham, NT/913.661.
M. Little, 1994, herb. M. E. Braithwaite. 2nd record.

7135/41.3. ASTER LAEVIS *69, Westmorland: By reservoir, Barrow-in-Furness, SD/19.70. P.
Burton, 1988, LANC, det. P. F. Yeo.

PLANT RECORDS 433

+135/41.3 x 4. ASTER LAEVIS X A. NOVI-BELGII (A. X VERSICOLOR) *73, Kirkcudbrights.: By
River Nith, Nithside, Dumfries, NX/970.765. O. M. Stewart, 1993.

+135/41.5. ASTER LANCEOLATUS 69, Westmorland: By River Bela, Milnthorpe, SD/496.813.
C. E. Wild, 1989, LANC, det. P. F. Yeo. 2nd record.

+135/43.4. ERIGERON KARVINSKIANUS *42, Brecs.: Pavement and walls, Crickhowell, SO/
216.183. M. Porter, 1993. *44, Carms.: Street weed, Queen Street, Carmarthen, SN/414.200. J.
Davies, 1994.

7135/43.5. ERIGERON ANNUUS *77, Lanarks.: Rough grassland, Cessnock, Glasgow, NS/
56.64. P. Macpherson & E. K. Lindsay, 1993, herb. P.M., det. P. F. Yeo.

135/43.6. ERIGERON ACER +*75, Ayrs.: Sandy waste ground, site of former Barassie Works,
Troon, NS/329.322. E. Nugent, 1994, GLAM, conf. A. McG. Stirling.

7135/44.2. CONYZA SUMATRENSIS *11, S. Hants.: Bare ground amongst shrubs by A3,
Portsmouth, SU/645.017. D. P. J. Smith, 1994. Derelict building site, Keswick Road, Woolston,
SU/436.110. P. D. Stanley, 1994. 1st and 2nd records, both conf. E. J. Clement. *13, W. Sussex:
Garden weed, Pagham, SZ/891.973. A. C. Leslie, 1994. *14, E. Sussex: Garden weed, Upper
Lewes Road, Brighton, TQ/319.057. Waste ground on quayside, Newhaven, TQ/448.011. Both A.
Spiers, 1994, herb. P. A. Harmes. ist and 2nd records.

135/50.3. ARTEMISIA ABSINTHIUM +81, Berwicks.: Rocky grassland, Coldingham Law, NT/
907.657. M. E. Braithwaite, 1994. Only extant locality.

+135/50.4. ARTEMISIA STELLERIANA *75, Ayrs.: Sandy shore between Barassie and Gailes, NS/
32.34. A. McG. Stirling, 1994, E.

+135/51.1. SANTOLINA CHAMAECYPARISSUS *77, Lanarks.: Rough railway bank, Old Dock,
Glasgow, NS/57.64. P. Macpherson, 1994, herb. P.M.

7135/58.lac. X max. LEUCANTHEMUM LACUSTRE X L. MAXIMUM (L. X SUPERBUM) *26, W.
Suffolk: Roundabout on A45 at Westley, TL/82.65. M. D. Crewe, 1993.

135/59.1. MATRICARIA RECUTITA *108, W. Sutherland: Disturbed ground in new car park,
Lochinver, NC/094.224. P. A. Evans, 1994.

135/62.12. SENECIO ERUCIFOLIUS *48, Merioneth: Roadside bank, Pont Islyn, Trawsfynydd,
SH/7.3. P. M. Benoit, 1994, NMW. Ist certain record.

+135/62.13. SENECIO SQUALIDUS *91, Kincardines.: Railway yard, Laurencekirk, NO/
717.718. D. Welch, 1994, ABD. *98, Main Argyll: Waste ground, former livestock mart, Oban,
NM/859.296. A. McG. Stirling & B. H. Thompson, 1994, herb. B.H.T.

135/62.+13 X 18. SENECIO SQUALIDUS X S. viscosuUS (S. X SUBNEBRODENSIS) *47, Monts.:
Derelict railway land, Caersws station, SO/029.919. P. M. Benoit, 1993.

+135/67.1.-SINACALIA TANGUTICA 40, Salop: Croesau Bach, SJ/239.272. F. H. Perring, 1988.
2nd record.

135/69.2. DORONICUM PLANTAGINEUM *40, Salop: Near Great Ness Church, SJ/397.190. J.
Hooson, 1987.

+135/71.3. PETASITES ALBUS 50, Denbs.: Hedge, Bontuchel, SJ/094.584. I. Evans, 1994, conf.
J. A. Green. Only extant locality.

7135/81.4. BIDENS FRONDOSA *29, Cambs.: With B. tripartita amongst flood debris in ditch E.
of River Great Ouse, Brownshill Staunch, Over, TL/370.729. C. D. Preston, 1994.

+138/4.2. ELODEA NUTTALLII *40, Salop: River Severn, Atcham, SJ/540.093. C. D. Preston &
T. C. G. Rich, 1987, CGE, conf. D. A. Simpson. Pool, Lynclys Common, Pant, SJ/271.236. F. H.
Perring, 1993. Ist and 2nd records.

434 PLANT RECORDS

+147/3.1. CALLA PALUSTRIS *59, S. Lancs.: Quarry pool, Hunter’s Hill Quarry, Parbold, SD/
501.122. P. Jepson, 1994. .

148/1.1. SPIRODELA POLYRHIZA *46, Cards.: Amongst Lemna minor in ox-bow pond in
pasture, S. side of Afon Teifi W. of Cenarth, SN/253.413. A. O. Chater, 1994, NMW.

148/2.3. LEMNA TRISULCA *78, Peebless.: Ornamental pond, Baddingsgill, West Linton, NT/
13.54. D. J. McCosh, 1991. Appeared spontaneously after weed clearance.

7135/2.4. LEMNA MINUTA *40, Salop: Montgomery Canal near Queen’s Head, SJ/335.264.
N.C.C. Field Course, 1989. Merrington Green, SJ/466.210. J. Martin, 1989. ist and 2nd
records. *42, Brecs.: Edge of lake, Llangasty Tal-y-llyn, SO/133.262. M. Porter, 1993. Garden
pond, Cathedine, SO/146.231. M. Porter, 1994. 1st and 2nd records.

151/1.3. JUNCUS COMPRESSUS *41, Glam.: Upper limit of saltmarsh, The Groose, Whiteford
Point, SS/444.945. R. Murphy, F. H. Perring & T. G. Evans, 1993, det. C. A. Stace.

151/1.8. JUNCUS AMBIGUUS *81, Berwicks.: Brackish mud, mouth of Eye Water, Eyemouth,
NT/945.640. M. E. Braithwaite, 1994, herb. M.E.B., det. T. A. Cope.

151/1.11. JUNCUS SUBNODULOSUS 68, Cheviot: Marshy edge of pool, Newton Pool, NU/
241.240. M. Jeffries, 1993, herb. G. A. Swan. 2nd record.

151/1.13 X 14. JUNCUS ARTICULATUS X J. ACUTIFLORUS (J. X SURREJANUS) *79, Selkirks: Mire,
W. end of Akermoor Loch, NT/404.208. R. W. M. Corner, 1993, herb. R.W.M.C., det. C. A.
Stace.

151/1.25 X 26. JUNCUS EFFUSUS X J. INFLEXUS (J. X DIFFUSUS) *46, Cards.: Slope between fen
and pasture, W. of Maesymeillion, Penparc, SN/196.483. J. P. Woodman & A. O. Chater, 1994,
NMW.

152/3.4. ELEOCHARIS MULTICAULIS *35, Mons.: Stream N. of The British, Abersychan, SO/
249.046. T. G. Evans & R. Hewitt, 1987, herb. T.G.E. Wet heath S. of Mountain Ash, $O/141.062.
T. G. Evans & R. Fraser, 1988, herb. T.G.E. 1st and 2nd records.

152/16.9a. CAREX DIVULSA subsp. DIVULSA 58, Cheshire: Road bank, Shotwick, SJ/339.719.
G. M. Kay, 1994. Ist post-1930 record.

152/16.15. CAREX REMOTA 102, S. Ebudes: Gully, Choille Mhor Wood, Colonsay, NR/
414.965. R. L. Gulliver, 1993. Ist post-1930 Colonsay record.

152/16.29. CAREX VESICARIA 108, W. Sutherland: Carex rostrata mire, Lon na Braclaich, NC/
193.147. R. E. C. Ferreira, 1985. 2nd record.

152/16.42. CAREX PUNCTATA 3.S. Devon: Low cliff slope, Wadham Rocks, SX/578.468. R. E.
N. & C. J. Smith, 1994, det. L. J. Margetts. 1st post-1930 record.

152/16.46c. CAREX VIRIDULA subsp. VIRIDULA 40, Salop: Around largest pool, Brown Moss,
SJ/563.394. I. C. Trueman, 1985, det. A. O. Chater. 2nd record and only extant locality.

7153/4.1. SASAELLA RAMOSA *46, Cards.: Extensive thickets in woodland, grounds of Glandyfi
Castle, SN/692.966. A. O. Chater & W. M. Condry, 1994, NMW.

153/11.1. MILIUM EFFUSUM *79, Selkirks.: Edge of woodland E. of Upper Loch, Bowhill, NT/
430.277. R. W. M. Corner, 1994, herb. R.W.M.C. tst localized record.

¥153/12.5. FESTUCA HETEROPHYLLA 78, Peebless.: Light shade in garden, Glen House near
Innerleithen, NT/300.329. D. J. McCosh, O. M. Stewart et al., 1994, det. P. J. O. Trist. 1st
confirmed post-1930 record.

+153/12.7d. FESTUCA RUBRA subsp. COMMUTATA *35, Mons.: Waste ground, Alpha Steel-
works, Newport, ST/338.846. T. G. Evans & M. Jones, 1994, herb. T.G.E.

153/12.8c. FESTUCA OVINA subsp. OPHIOLITICOLA *77, Lanarks.: Upland heath S. of Daer
Reservoir, NS/96.04. P. & A. C. Macpherson, 1994, herb. P.M., det. C. A. Stace.

PLANT RECORDS 435

153/12.14. FESTUCA LONGIFOLIA *3, S. Devon: Clefts in rock near coastal footpath, Gara
Rock, SX/750.369. L. J. Margetts & R. Takagi-Arigho, 1993, K, det. T. A. Cope.

153/12.15. FESTUCA BREVIPILA *12, N. Hants.: Well-drained acid slope, Farnborough, SU/
868.540. C. R. Hall, 1993, herb. Lady A. Brewis, det. P. J. O. Trist.

153/12.7 x 14.1. FeEsTUCA RUBRA X VULPIA FASCICULATA (X FESTULPIA HUBBARDII) *69,
Westmorland: Sand dunes, Sandscale Haws, Dalton-in-Furness, SD/18.75. P. Burton, 1994,
LANC, det. C. A. Stace.

153/14.3. WULPIA MYUROS +*98, Main Argyll: Ballast in siding, Oban Railway Station, NM/
857.298. A. McG. Stirling & B. H. Thompson, 1994, herb. B.H.T.

153/14.4b. VULPIA CILIATA subsp. AMBIGUA *18, S. Essex: Short turf on fixed dunes, Shoebury
Old Ranges Nature Reserve, TQ/92.84. K. J. Adams ef al., 1994, herb. K.J.A., det. A. Copping.

153/16.2. PUCCINELLIA DISTANS +26, W. Suffolk: Verge of A45 S. of Exning, TL/61.64. M. D.
Crewe, 1993. 2nd post-1930 record.

153/18.12. POA NEMORALIS 102, S. Ebudes: Woodland, Colonsay House Woods, NR/372.954.
R. L. Gulliver, 1993. Ist post-1930 Colonsay record.

153/20.1. CATABROSA AQUATICA 78, Peebless.: Newly cleaned peaty ditch, Kippit Farm,
Dolphinton, NT/11.47. D. J. McCosh, 1993, herb. D.J.McC. Ist record since 1930.

¥153/28.4a. AVENA STERILIS subsp. LUDOVICIANA *40, Salop: Hodnet area, SJ/60.28. S.
Kingsbury, 1991.

153/30.1. TRISETUM FLAVESCENS 102, S. Ebudes: Dry turf at foot of ancient cross, Oronsay
Priory, NR/349.889. R. L. Gulliver, 1993. Ist record from Oronsay or Colonsay.

153/46. 1. POLYPOGON MONSPELIENSIS +*12, N. Hants.: Near dumped soil on sandy heathland,
Hawley Hill, SU/845.588. C. R. Hall, 1991. Sandy track near tip, Bramshill Common, SU/752.618.
Hampshire Flora Group, 1993. Both herb. Lady A. Brewis, det. E. J. Clement. Ist and 2nd records.

153/47.1. ALOPECURUS PRATENSIS 102, S. Ebudes: Mown grassland around ruins of Oronsay
Priory, NR/3.8. R. L. & M. C. Gulliver, 1991. 1st post-1930 record from Colonsay or Oronsay.

153/47.1 x 2. ALOPECURUS PRATENSIS X A. GENICULATUS (A. X BRACHYSTYLUS) *4,N. Devon:
Road verge, Mole’s Chamber, SS/717.392. I. P. Green, 1994, det. T. A. Cope. *77, Lanarks.:
Rough grassland, Mount Vernon, Glasgow, NS/65.62. P. Macpherson, 1993, herb. P.M., conf. H.
J. M. Bowen.

153/47.2 X 3. ALOPECURUS GENICULATUS X A. BULBOSUS (A. X PLETTKEI) *44, Carms.:
Saltmarsh of Afon Taf near Mwcha Farm W. of Llanstephan, SN/320.112. M. Lloyd, 1994, NMW,
det. T. A. Cope. Ist Welsh record.

153/50.4. BROMUS HORDEACEUS 102, S. Ebudes: Dune grassland, Ardskenish Dunes, Colon-
say, NR/352.922. R. L. Gulliver, 1991. 1st post-1930 Colonsay record.

153/52.3 ANISANTHA STERILIS 73, Kirkcudbrights.: Farmyard, Auchinleck, NX/449.709. O.
M. Stewart, 1994. 2nd record.

+153/53.3. CERATOCHLOA CATHARTICA *44, Carms.: Garden weed, Dylan Thomas Boathouse
garden, Laugharne, SN/307.111. J. Rees, 1992. NMW, det. G. Hutchinson.

153/54.1. BRACHYPODIUM PINNATUM +*79, Selkirks.: Large colony on side of track, Bowhill
estate, NT/427.272. R. W. M. Corner, 1994, herb. R.W.M.C.

7153/59.3. HORDEUM JUBATUM “*75, Ayrs.: Road verge by A71 near Riccarton interchange,
Kilmarnock, NS/44.36. P. J. Cook, 1994.

153/63.1b. MOLINIA CAERULEA subsp. ARUNDINACEA 44, Carms.: Flush at edge of willow scrub,
Lower Lliedi Reservoir, Llanelli, SN/519.033. I. K. Morgan, 1992, NMW, conf. G. Hutchinson.
2nd record.

436 PLANT RECORDS

155/1.1. TYPHA LATIFOLIA 102, S. Ebudes: Margin of lochan, Colonsay, NR/368.960. P. M.
Clarke, 1993, det. D. R. McKean. Ist Colonsay record.

155/1.1 X 2. TYPHA LATIFOLIA X T. ANGUSTIFOLIA (T. X GLAUCA) *35, Mons.: Edge of sludge
pond, Newport, ST/33.84. T. G. Evans & M. Jones, 1993, NMW.

+158/6.2. KNIPHOFIA PRAECOX *49, Caerns.: Roadside, Bodafon, Llandudno, SH/801.816. W.
McCarthy, 1994.

158/7.1. COLCHICUM AUTUMNALE *49, Caerns.: Meadowland, Bryn Pydew, Llandudno, SH/ | |

81.79. G. Battershall, 1992.

+158/19.3. SCILLA SIBERICA *49, Caerns.: Woodland, Gloddaeth Woods, SH/80.80. G.
Battershall, 1993.

158/20.2 X +3. HYACINTHOIDES NON-SCRIPTA X H. HISPANICA *49, Caerns.: Hedgerow, Nant y
Gamar, Llandudno, SH/80.81. G. Battershall, 1991.

+158/22.1. CHIONODOXA FORBESII *77, Lanarks.: Rough grass bank, Botanic Gardens,
Glasgow, NS/56.67. K. Watson, 1993, GL.

+158/23.2. MUSCARI ARMENIACUM *49, Caerns.: Grassy waste ground near Conway, SH/
771.764. R. Lewis, 1994, NMW.

158/24.1. ALLIUM SCHOENOPRASUM +46, Cards.: Well established on hedgebank S.W. of
Penlan-y-Mor, Gilfachreda, SN/416.594. A. O. Chater, 1994, NMW. 2nd record.

158/24.14. ALLIUM SCORODOPRASUM 58, Cheshire: Side of ditch, Norton Marsh, Runcorn, SJ/
569.857. G. M. Kay, 1994. 1st record this century, at a site from which the species was recorded last
century.

158/31.1. LEUCOJUM AESTIVUM +79, Selkirks.: Margin of pond E. of Lower Loch, Bowhill, NT/
431.271. R. W. M. Corner, 1994, Ist post-1930 record, at same site as earlier record.

+158/33.5b. NARCISSUS PSEUDONARCISSUS subsp. OBVALLARIS *46, Cards.: Grassy roadside
slope, Banc y Warren, Penparc, SN/204.476. A. O. Chater & J. R. Akeroyd, 1994. Well established
on hedgebank near The Woodlands, Ciliau Aeron, SN/509.586. A. O. Chater, 1994. Ist and 2nd
records. .

159/2.1. SISYRINCHIUM BERMUDIANA **12, N. Hants.: Bramshott Common, SU/85:32. C.
Darter, 1992, herb. Lady A. Brewis, det. A.B. Ist confirmed record.

159/5.8. IRIS FOETIDISSIMA 73, Kirkcudbrights.: Edge of wood N. of St Mary’s Isle, Kirkcud-
bright, NX/680.562. O. M. Stewart, 1994. Ist post-1930 record.

+159/8.1. CROCUS VERNUS *46, Cards.: Extensively naturalized in rough grass, Penparc chapel
graveyard, SN/212.478. A. O. Chater, 1994. Abundant in rough grass, Tregroes churchyard, SN/
406.447. A. O. Chater, 1994, NMW. 2nd record.

159/9.2b. GLADIOLUS COMMUNIS subsp. BYZANTINUS *17, Surrey: Rough grassland, Wood-
cote, Epsom, TQ/196.593. J. F. Leslie, K. Page & J. E. Smith, 1994.

+159/13.1. CROCOSMIA PANICULATA *46, Cards.: Hedgebank E. of Llancynfelyn church, SN/
648.922. A. O. Chater, 1994, NMW. Grassy bank of Afon Rheidol, Penparcau, SN/592.804. A. O.
Chater, 1994. 1st and 2nd records.

+159/13.3 X aur. CROCOSMIA POTTSII X C. AUREA (C. X CROCOSMIIFLORA) *81, Berwicks.: Sea
braes below Barefoots, Eyemouth, NT/937.647. M. E. Braithwaite, 1994. Flush on sea braes,
Gunsgreen, Eyemouth, NT/950.646. M. E. Braithwaite, 1994, herb. M.E.B. Ist and 2nd records.

162/7.3. SPIRANTHES ROMANZOFFIANA 110, Outer Hebrides: Molinia caerulea lawn, Kilpheder,
South Uist, NF/737.196. F. Horsman, 1994. ist South Uist record.

162/10.1. HAMMARBYA PALUDOSA 2, E. Cornwall: Wet heath, Rentire Common, SX/004.631.
I. Bennallick, 1993, conf. R. J. Murphy, Only extant locality.

PLANT RECORDS 437

162/16.1b. GYMNADENIA CONOPSEA subsp. DENSIFLORA *46, Cards.: Base-rich flush fen S. E. of
Afon Mwldan, Penparc, SN/195.483. A. O. Chater, 1986, det. F. Rose.

162/16.1c. GYMNADENIA CONOPSEA subsp. BOREALIS *46, Cards.: Heathy slope, Banc Cwmer-
fyn, SN/704.824. A. O. Chater, 1988, det. F. Rose.

162/17.1. COELOGLOSSUM VIRIDE 3, S. Devon: Dunnabridge, SX/644.743. R. Keedle, 1994. 1st
post-1930 record.

162/18.3c. DACTYLORHIZA INCARNATA subsp. PULCHELLA 2, E. Cornwall: Wet flush in culm
grassland, Otterham valley near Otterham Station, SX/144.896. T. J. Dingle, 1994, det. L. J.
Margetts. 2nd record.

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Book Reviews

Greek wild flowers and plant lore in ancient Greece, H. Baumann, translated and augmented by W.
T. Stearn & E. R. Stearn. Pp. 252; with 481 illustrations (mostly colour). Herbert Press, London.
1993. Price £16.95 (ISBN 1-871569-57-5S).

This book is translated from the second (1986) edition of Baumann’s 1982 work Die griechische
Pflanzenwelt in Mythos, Kunst und Literatur. It is clearly a labour of love compiled over many years
in the field throughout Greece. Its strength is twofold; the profusion of colour photographs, and the
references to the literary sources. There are five main sections: the Homeric Landscape, Cults and
Myths, Medicinal and Magic Plants, the Gifts of Demeter and the Lotus and Acanthus in art. These
form the core of the book and are the most solid. Less central are sections on lowland and mountain
flora, orchids, introduced plants and the ancient awareness of botany.

There are many delights in the illustrations, images of landscape, people or artefacts, scattered
amongst the flowers, viz. plate 340 which shows two octopi in a market, to parallel the design on a
late Minoan vase. The format of the book is small, less than A5, which makes it a handy travelling
companion but robs the photographs of the chance to dazzle the reader which would have happened
in a larger format. Some pages have nine small photographs and this leaves one peering at the page
to see the detail. That said, the quality and variety of the photographs is outstanding.

The Ancient Greek doctors, scientists, poets and historians had a tremendous awareness of the
botanical diversity of their country. Baumann has trawled the literature for references most
assiduously; examples from art and architecture are also included. References to archaeological
evidence, however, tend to be general rather than to specific sites or publications and this aspect is
less useful to the student. Environmental Archaeology has come late to the Classical world and it
would be unfair to dwell on the lack of scientific evidence from recent excavations which one would
be expecting in a new work rather than a revised edition.

Whilst the book is a pleasure to dip into, it can feel heavy at times. This is inevitable due to its
fundamental nature as a list of plants. The structure also allows for some repetition. The use of the
teasel in textiles is referred to on p. 77, plate 128, as a plant of the gods and again in plate 301 in the
context of textile preparation. There is no cross-reference in the text but it is picked up in the index.
In the same way the olive is discussed in the cults and myths section and again as an economic plant
without cross-reference. These are not faults, however, and probably go unnoticed by the student or

' browser.

This is a very enjoyable survey of the subject and a good blend of economic botany with history. It
will be widely read, probably serving to attract visitors to the country and increase their awareness
not just of the diversity of the flora but of its long and distinguished heritage.

E. C. SoUTHWORTH

Vascular plant families and genera. A listing of the genera of vascular plants of the world according to
their families, as recognised in the Kew Herbarium, with an analysis of relationships of the flowering

. plant families according to eight systems of classification. R. K. Brummitt. Pp. viii + 804. Royal

| Botanic Gardens, Kew. 1992. Price £24.00 (ISBN 0 947643 43 5).

The subtitle of this impressive work gives a clear indication of its aims and contents. Even in the
three years since it made its appearance, it has become a standard reference work for anyone
curating herbarium collections, large or small, or whose work requires up-to-date knowledge of

440 BOOK REVIEWS

generic and family names. Its predecessors are the various editions of A. J. Willis’s A dictionary of
flowering plants and ferns and D. J. Mabberley’s The plant-book: a portable dictionary of the higher
plants (1987). In contrast to them, the compiler of this lexicon sought advice from a large number
(over 160) of specialists throughout the world. The result is a consensus of informed opinion on
modern generic and family nomenclature with Kew having acted as arbiter in cases of disagreement
or uncertainty. Even with the sophisticated facilities of present-day data bases, the complexities
involved in producing such a compendium are obvious and manifold. Over 30,000 generic names
have been published over the last two and a half centuries; and where Bentham & Hooker in their
classic Genera plantarum recognised 200 plant families, current thinking is that more than twice that
number should be accepted. It is probably a reasonable generalisation to say that, with exceptions,
as one progresses from species to genera to families and higher groupings, their circumscription
becomes increasingly beset with problems and subjectivity. Usually botanists agree more about
species and generic limits and nomenclature than higher categories; or at least they are more
concerned with the former. Decisions about plant families to be recognised resulted from the
setting-up of a ‘family planning unit’ comprising botanists from Kew and the Natural History
Museum, London. Their decisions, not surprisingly, are a middle course between the broad
concepts of Bentham & Hooker and the apparently narrower views of some present-day authors.
Yet in the Kew list there are a surprising number of mono- or oligotypic families recognised, mostly
tropical: from the first letter of the alphabet are Aextoxicaceae, Agdestidaceae, Akaniaceae,
Alseuosmiaceae, Aizateaceae, Amborellaceae, Aralidiaceae and Asteropeiaceae, strange-sound- -
ing names for at least European botanists. Nearer home, Illecebraceae, perhaps surprisingly, are
kept separate from Caryophyllaceae. British botanists will be relieved to learn that the status quo of
most currently recognised families is maintained, but the Orobanchaceae, surely correctly when the
world-wide scenario is taken into account, are subsumed into Scrophulariaceae, Pyrolaceae into
Ericaceae, and Ruppiaceae into Potamogetonaceae. They will also be happy that Compositae,
Cruciferae, Gramineae and Labiatae are preferred to Asteraceae, Brassicaceae, Poaceae and
Lamiaceae, more favoured by continental botanists. The traditional boundaries between such
clearly very closely allied families as Araliaceae and Umbelliferae, and Verbenaceae and Labiatae
are also maintained.

More than half of the dictionary comprises a straight-forward listing of generic names, their
authors, and the families they belong to: generic synonyms are indicated by italics. Again the
approach is conservative; not accepted, for example, are the recently published segregate genera of
Himalayan Gentianaceae, and the splitting-up of Thlaspi. Inevitably, not everybody is going to be
happy with all the names given authoritative blessing. For example, in the Chenopodiaceae, the
strange-looking Iranian Esfandiaria is surely not different from Anabasis, Hammada is virtually
identical with Haloxylon, as is Climacoptera with Salsola. But these are small quibbles.

Part II lists the families in alphabetical sequence with their constituent genera also alphabetically;
the numbering of the 454 families recognised are those of the herbarium at Kew. Genera not
represented at Kew are indicated by asterisks and it is a clear measure of the enormous wealth of
their collections to see how relatively few asterisks there are. And some of the genera asterisked are
certainly present at Kew, presumably lurking under another name. For each family is also given an
indication of where it fits in within eight other frequently used systems of classification, from
Bentham & Hooker about the turn of the last century, through to Cronquist (1981) and Takhtajan
(1987). The final Part III gives a synoptic listing of these eight systems which is particularly useful in
comparing and contrasting them; for many, the ‘Young’ system will not strike a chord of
recognition.

Throughout the entire 800 pages of this sturdily bound, clearly printed and easy-to-use dictionary,
there is little to criticise and much to respect. No doubt there will be botanists and herbarium
curators who will find little items to niggle about or dispute, but they will mainly be most happy to
possess this invaluable information-source. A small errata slip draws attention to a few omissions
and a printing sin. Kew’s resources of specimens and expertise are probably unrivalled throughout
the world and they have been used in the best possible modern way, giving, with this publication, a
lead into the next century. Dr Brummitt and his core of Kew-based colleagues have done a splendid
job.

I. C. HEDGE

BOOK REVIEWS 44]

The West Yorkshire plant atlas. Edited by J. C. Lavin & G. T. D. Wilmore. Pp. 287, six colour plates,
700 maps. City of Bradford Metropolitan Council. 1994. Price £25.00 (ISBN 0—907734-39-1).

Jack Lavin was a pioneer of the local, computerised, biological records centre movement. In the
early 1970s his centre was a model used to convince others to go and do likewise — and Jack himself
was a splendid advocate, not only outside his county but within where he was able to convince four
other local authorities to fund what started as a District centre at Keighley, as a unit to cover the
whole of the new county of West Yorkshire when it was created in 1974: it later became their
Ecological Advisory Service. His influence with local authorities has continued and this handsome
volume carries the imprint of the City of Bradford Metropolitan Council and claims to be the first
Flora to be produced by a Department of Local Government.

The primary objective of the Ecological Advisory Service was to organise surveys of all the major
semi-natural habitats in the county and the 1-km square was adopted as the recording unit. This has
strongly influenced the collection of the data for the Flora and its presentation. The idea of
producing a Flora did not come until 1986 when it was realised that the 100,000 plant records
collected during surveys in the preceding 12 years couid be the basis of a West Yorkshire plant atlas.
With the help of about 100 individuals and societies an extra 150,000 1-km square records were
added in the next 33 years with over 90% of the squares finally covered. But it is still a by-product of
the site surveys and it shows.

First, it is not a Flora of the former West Riding or of v.cc. 63 & 64: the area covered comprises
the northern and western half of v.c. 63, v.c. 64 between the Aire and the Wharfe, and some at least
of the Pennines in v.c. 59. There may also be overlap with v.c. 58 S.W. of Holmfirth, but no map
showing the v.c. boundaries is provided and v.c. distribution is not given in the text. The only map of
the area appears in the end-papers and is limited to major rivers and towns, 10-km squares, District
boundaries, and a shaded area which may be land over 300 m, though there is no legend to indicate
this.

Secondly it is a very large area, with over 2000 1-km squares, and an attempt was made to cover
them all. It says a great deal for the enthusiasm of the team that so much was achieved but, with 10%
not visited, it suggests that a scheme aimed at covering 500 tetrads would have given a more
complete picture without detracting from the interpretation of the results.

However ‘interpretation’ is not a strong point of this Atlas. In a book with that word in the title a
range of maps to assist in understanding distribution patterns would surely be expected. Here is an
area with stark contrasts between the Magnesian Limestone of the east and the Millstone Grit in the
west, between the lowlands of the Vale of York and the Pennines rising to 560 m S.W. of Holmfirth,
and with rainfall varying from below 800 mm to over 1250 mm/annum, yet there is no geological map
(just an E.-W. section across the county), the one topographical map showing ‘Pennine Uplands’ in
the end papers, and no climatic maps whatsoever though half of the single page given to ‘Climate’ is
a blank.

There are maps in the introductory section on ‘The vegetation of West Yorkshire’ by John
Rodwell. This is a valuable and extensive account of the habitats classified in the National
Vegetation Classification (N.V.C.) which have been recognised in West Yorkshire. The Ecological
Advisory Service provided many original stands for the N.V.C. and this is reflected in the detail in
these maps — but even these are difficult to interpret with as many as ten different symbols per map.

This vegetation information is usefully carried into the short accounts of each of the 686
commoner species mapped and other native species: many include lists of 4-6 associated species.
Rarer species and aliens are less well served and, though reported as occurring, often neither
localities nor grid squares are given.

The book is well produced: the colour plates are ravishing but it would have been so much more
useful with those maps. Is it too late to suggest the production of a set of transparent overlays? It has
been done before!

F. H. PERRING |

442 BOOK REVIEWS

Catalogue of the Brambles of Britain and Ireland in the herbarium of Liverpool Museum (LIV).
Compiled by M. Palmer, edited by J. Edmondson. Pp. viii + 48. National Museums & Galleries on
Merseyside, Liverpool. 1994. Price £12.00 (ISBN 0-906367—70-0).

A catalogue of the Herbarium of the British Flora collected by Margaret Stovin (1756-1846). M.
Simmons. Pp. 422 text + 102 indexes. Middlesbrough Borough Council Leisure Services
Department, Middlesbrough. 1993. Price not stated. (no ISBN).

Cheap printing was one great advantage that the Victorians and Edwardians enjoyed over their
successors. Many a local society in those days could rise to a thick, meaty-looking annual report,
which was so impressively produced that it was virtually guaranteed eternity by being bound and
placed on library shelves. Till only very recently its modern counterpart would have had to make do
as often as not with some slight, duplicated effort which almost asked to be discarded once it had
been read. Now, at last, presentable alternatives are arriving to rescue us from that era of
deprivation and making possible once again such long-despaired-of luxuries as catalogues of
museum collections — of which the couple under review are praiseworthy examples of the new genre.

The Rubus collection at Liverpool Museum is by no means as rich or extensive as those in some of
the other national institutions, but it is large enough and sufficiently fully redetermined in the light
of the latest knowledge to justify this very useful compilation for the benefit of vice-county
Recorders and fellow-specialists in the group. 225 species from 120 collectors are represented, many
of them via the exchange clubs but a particular high proportion W. R. Linton’s, whose herbarium
was transferred from the University of Liverpool a few years back (and a pleasing vignette of whom
features appropriately on the title page). The catalogue excludes a considerable number of further
specimens that have been critically (re-)determined since it went to press, also a sizeable residue of
material, much of it unmounted, that has so far defeated current specialists.

The Dorman Museum in Middlesbrough acquired the herbarium of Miss Stovin in 1922. The
specimens of British vascular plants and charophytes, 1900 in all, fill 20 albums and are mostly
localised. The earlier ones were of her own collecting, mainly round Doncaster, but many came
later from contemporary botanists all over Britain in the years 1794-1840. The catalogue is
essentially a computer print-out of the data taken from the labels, separately indexed by species,
(present-day) counties of localities, and collectors. The work was made possible by a grant from the
Pilgrim Trust — as other local museums could usefully note.

D. E. ALLEN

Wild plants of the Phoenix Park. P. A. Reilly, with contributions by D. L. Kelly, D. M. Synnott & J.
McCullen. National Botanic Gardens, Glasnevin & The Phoenix Park, Office of Public Works,
Dublin. 1993. Price IR£7.50 (ISBN 0—7076—-0331-5); reprinted from the journal Glasra.

When our Flora of Inner Dublin was published in 1984, it was said by some that it was a pity we did
not include the National Botanic Gardens or indeed the Phoenix Park, which latter adjoins the outer
boundary set for the Inner Dublin Flora. However the habitat of this open parkland is very different
from the streets and open spaces of the inner city. Now Paddy Reilly has given us the opportunity to
discover the Phoenix Park as a site in its own right for botanical exploration.

The Phoenix Park (Pairc Fhionn Uisce — park of the pale water: nothing to do with birds or ashes)
is, according to the history given by John McCullen, the “largest enclosed recreational space (707
ha) within any capital city in Europe’’. He tells us, indeed, that this area is equivalent to the total
area of all the London parks, including Hyde Park. That alone would make it of botanical interest. It
is, in fact, a large deerpark, enclosed since 1667 and managed to a varying degree as such since 1623.
To this day, a herd of c. 500 fallow deer remain in the Park, which has been managed since 1860 by
the Office of Public Works.

The main part of the book consists of the Flora which is written by Paddy Reilly and prefaced with
a brief summary of the history, climate and soils of the Park. The soils are mostly of Limestone drift,
but are notable for their lack of permeability. Thus one might anticipate the species there being of
neutral or lime-rich soils, some confined, perhaps, to the few gravel pits and esker remnants in the
Park. Reilly then follows this with a history of the flora, having meticulously checked printed

BOOK REVIEWS 443

records and herbarium specimens. The earliest records are from Caleb Threlkeld’s Synopsis
Stirpium Hibernicarum, a Flora/herbal “of native plants such as grow in the vicinity of Dublin”,
published in 1726. This included the Phoenix Park. known to him then as the King’s Deer Park.
Already, weeds such as Cardamine hirsuta and Coronopus squamatus were present. A few
woodland/hedge species are also mentioned — Ajuga reptans, Malus sylvestris, [lex aquifolium and
Arum maculatum, though only the last two remain today. Possibly there was a greater relict
woodland flora in Threlkeld’s time than now. The history continues with similar small glimpses of
the flora, since, as Reilly states, at no time was the whole flora of the Park ever listed, which would
be invaluable now for comparison. It is to be hoped that this present Flora will serve that purpose in
years to come. In summary, he states that between 1726 and 1975, 89 species were listed for the Park
and of those, only 49 survive today. Most of those that have disappeared are wetland plants, but also
species of calcareous habitats (the exhaustion of the gravel pits in the 1880s being cited as relevant)
and, of course, of woodlands. Reilly singles out these ‘missing’ species and lists them, noting their
occurrence in the county of Dublin as a whole. Twenty-four species, especially from aquatic and
limestone grassland habitats, are now rare in the county, reflecting the decrease in appropriate
habitat countywide.

There follows a reference to strictly calcicole and calcifuge species, as these proved to be rarer,
especially the latter as acid soils only occur in one artificially created habitat supporting the
otherwise rare species Rumex acetosella. This intriguing habitat was the site used for fuel storage
during the 1939-45 war. The fuel was, of course, turf (or peat), virtually the only source of fuel in
Ireland at the time. Great banks of turf lined the roads near the American Ambassador’s residence
right into the 1950s, as testified by an interesting photograph. Such photographs are interspersed
throughout the text and help to illustrate the types of habitat referred to. This includes a very
attractive series of 20 colour photographs of landscape, habitat and individual species. This
introductory section ends with useful notes on the different habitats in the Park and associated
species as well as a summary of the features of each of the five districts delimited for the purpose of
the Flora.

The Flora per se lists some 318 species in Latin and English, with their occurrence within each of
the five districts as well as annotations on sites where necessary. This, along with the map provided,
is accessible to botanists with at least a rudimentary knowledge of field botany and is an invaluable
reference for future botanical work in the area. The chapter on bryophytes (so often evaded in
Floras) is useful especially as the authors (D. L. Kelly and D. M. Synnott) list nine habitats and the
species most commonly associated with each. The small number of hepatics (eleven) found is
presumably in part due to the comparatively dry nature of the climate. A total of 87 mosses were
found, including one new to Ireland (Bryum creberrimum). One confusing aspect of this chapter is
that for many species there is only one site listed and there is no direct reference to the botanical
districts of the main Flora.

I found it interesting to compare the species found in the Phoenix Park with those listed for Inner
Dublin. As predicted, the Park does have a greater pastoral element, with species such as Primula
veris, P. vulgaris, Veronica chamaedrys and Luzula campestris occurring commonly in the Park but
completely absent (at least in 1984) from the inner city. Less surprisingly, woodland species (e.g.
Sanicula europaea, Conopodium majus, Arum maculatum, Allium ursinum) and some aquatics
(Potamogeton spp., three Glyceria spp., Eleocharis palustris) as well as many sedges are present
only in the Park. Three species of inner Dublin surprisingly absent from the Park are Corylus
avellana, Ulmus glabra and U. procera. There is no mention of the planting of Corylus and though
many elms were planted, none seem to have seeded themselves.

The book concludes with the history of the Park by J. McCullen, which is interesting in itself, but
also gives insight into the origins of the particular flora which exists in the Park today. Particular
reference is given to tree planting which only got under way after about 1745 and has continued in
several phases of intensive planting ever since. The six appendices are an extension of this chapter,
mostly listing the species of tree planted (c. 136, including cultivars), where they occur and, in the
case of ceremonial trees by whom and when they were planted. This is of horticultural interest and
illustrates the immense diversity of trees, warranting the term ‘arboretum’ to be applied to the Park.

Thus the book is very attractively presented, remarkably free of spelling and typographical errors
— I spotted only two or three, the most misleading being the date of birth of Caleb Threlkeld, which
is listed as 1672 and 1676 in the same paragraph! It is interesting reading for any amateur of Floras

444 BOOK REVIEWS

and their history, but of course should also tempt botanists to visit the Park and discover its richness
for themselves.

M. SHEEHY SKEFFINGTON

An annotated checklist of the flowering plants and ferns of Main Argyll. G. Rothero & B. Thompson.
Pp. iv + 132. Published by The Argyll Flora Project. Glenlussa. 1994. Price. £6.00 (ISBN 0-
9522852—0-7 paperback).

It is a pleasure to welcome another first-rate local checklist to the growing number for Scotland.
Gordon Rothero and Bernard Thompson are to be congratulated on producing a good working
checklist for the plants of Main Argyll in a commendably short time — although several botanists had
done major groundwork for them. Perhaps we are at last realising that field botanists do not want to
wait for ever for the perfect work (which is unattainable anyway) nor a weighty volume which
repeats all the taxonomic detail of a national Flora. Much more useful is an easily carried
informative booklet cheap enough for revised editions to be economically possible. This latter the
authors have given us.

Main Argyll (v.c. 98) is the core of the mainland of Argyllshire less the Kintyre peninsula (v.c.
101). For the Checklist this sprawling and diverse area is divided into seven roughly natural areas.
The checklist, in family order, is clearly set out; what is known, and more importantly, what
requires more investigation, are pointed out. Obviously there is plenty to do both in investigating
critical taxa, especially subspecies, and checking up on old and doubtful records. To help with
localities there is a most useful gazeteer. The booklet is bound in a soft, damp-resistant cover and
slips easily into a large pocket.

All the reader has to do is-buy this indispensible manual, go to Argyll —all 3,700 sq. km. of it— and
help the authors with data for a second edition!

D. M. HENDERSON

Scarce plants in Britain. Compiled and edited by A. L. Stewart, D. A. Pearman & C. D. Preston.
Pp. 508, 16 tables, 49 figures, numerous maps. Joint Nature Conservation Committee, Peter-
borough. 1994. Price £34.00 (ISBN 1—873701-66-7).

Scarce plants in Britain is the result of a co-operative project begun in 1990 originally involving the
Nature Conservancy Council (N.C.C.), the Institute of Terrestrial Ecology (I.T.E.) and the
Botanical Society of the British Isles (B.S.B.I.). The Joint Nature Conservation Committee
(J.N.C.C.) took over the N.C.C.’s responsibility following the major reorganisation in 1991.

This is a most welcome and important publication. It at last draws attention to the substantial
group of plants which does not meet the fundamental criterion of national rarity (recorded from 15
or fewer 10-km squares), but which form the next tier, by definition, occurring in 16-100 10-km
squares. However as rarity is not, of course, synonymous with threat and can in many cases simply
reflect natural distribution, some nationally scarce plants are far more immediately threatened with
extinction than our classic rarities.

Franklyn Perring points out in his foreword that the nationally scarce plants were a relatively
neglected group even in the preparation of the original Atlas of the British flora (1962) and many
lowland plants, especially arable weeds and wetland species, have undergone significant declines
(Scandix pecten-veneris, Centaurea cyanus and Sium latifolium for example) and are more likely to
disappear from the British flora than some of our extremely rare denizens of remote mountains
whose populations are more stable.

The conservation importance of the nationally scarce plants is well explained in the introduction
highlighting their use in habitat evaluation and changes in their status as indicating changes in land
use or land management. The aims, methods and data acquisition are all very clearly explained and
the importance of voluntary contributions in terms of data supply and species accounts speaks for
itself. The maps are well produced but given the constant reference to the Atlas of the British flora

BOOK REVIEWS 445

(1962) throughout the book it would have been useful to have the maps the same size so that they
were compatible with the very useful overlays supplied in that work. The vice-county map (p. 11) is
larger still and lacks a superimposed grid thus making the accurate location of records (dots) near
vice-county boundaries difficult.

The species accounts are generally of a high standard and as a whole contain a wealth of useful,
interesting information. Although the format is consistent, with so many contributors the style and
the amount of biological information is a little variable and reference citation is sometimes thin or
lacking.

The chapter on habitats of scarce species in Britain is excellent and provides concise accounts of
the major habitat types including an historical background and an outline of the most important
factors which have led to their decline or modification. In the introduction to the chapter the habitat
and species approach to conservation are well explained and to some extent reconciled. However,
having pointed out that ‘“‘both are essential if effective conservation measures are to be developed
and applied”’, it is surprising that nowhere in the chapter is there any mention or reference to the
major work British plant communities edited by John Rodwell. In such an important contribution to
species conservation as Scarce plants in Britain, | feel it is a missed opportunity not to make a
stronger link or cross-reference to the most important contribution to a general understanding of
British vegetation in recent years, although two volumes of the work are cited in the references from
individual species accounts.

Finally the chapter assessing the changes in our knowledge of species from 1962-1992 (that is from
the date of the original Ad/as to the present work) makes some important points. For example, some
species (tabulated) on the current maps show an increase in records compared with those of the
Atlas. This is almost entirely due to increased recording and in some cases may mask a decline in
number of sites within a 10-km square. Where a decline is shown it is likely to be significant.

Despite these minor reservations Scarce plants in Britain is an immensely valuable and well-
produced book. It is written with clarity and conciseness and will be constantly referred to by
amateur and professional botanists and conservationists. A mark of the success of the project is that
the Conservation Agencies have already acted upon the results and several species are currently
being studied to obtain further information and also to bring records right up-to-date. The compilers
should be congratulated.

P. LusBy

Flora of Glamorgan. A. E. Wade, Q. O. N. Kay & R. G. Ellis. Pp. viii + 379 with nine colour
plates, 14 black and white figures and numerous maps. H.M.S.O., London. 1994. Price £29.95
(ISBN 0-11—3100046-9).

To the uninitiated, South Wales conjures up visions of mining valleys, factories and industrial
dereliction. This, however, is something of a calumny. There are extensive areas of industrialisation
and urbanisation but much of the botanical vice-county of Glamorgan (v.c. 41), to a greater or lesser
extent influenced by Man, supports a rich and varied flora. In habitats ranging from sand-dunes and
limestone sea-cliffs in the south to sandstone, shale and limestone Coal-measures ascending to 600
m in the north is a flora containing 80% of Welsh vascular plant species.

The Flora of Glamorgan, the first since Professor A. H. Trow’s 1906-1911 publication, was
almost a quarter of a century in gestation and is a Flora in the traditional style being basically a list of
plants together with supporting chapters appertaining to the flora. However, such a summary does a
grave injustice to this excellent work.

The introductory chapters are informative and readable and include ‘A short history of botany
and botanists in Glamorgan’ by R. G. Ellis and A. E. Wade, ‘Soils of Glamorgan’ by P. S. Wright,
‘The geological history of Glamorgan’ by M. G. Bassett and R. M. Owens and ‘Land cover of
Glamorgan, a satellite image’ by A. J. Morton. This last chapter is a little disappointing, as although
it contributes the frontispiece of the book and lists the vegetation and non-vegetation types
discerned these are difficult to identify in the colour plate and there are no outlines to assist in this.
Surprisingly there is no mention of climate in the book although this is directly relevant to remarks

446 BOOK REVIEWS

in Chapter 5, ‘The history, ecology and distribution of the flora of Glamorgan’ by Q. O. N. Kay, and
climate is important for anyone interested in plant distribution.

That chapter is a brief account of the ecology and plants of the three main areas of v.c. 41, the
Gower Peninsula, the Uplands and Border Ridges, and the Vale of Glamorgan rather than an
enumeration and description of vegetation types. As such it makes more interesting reading. Some
indication of the extent of these areas, the current (as opposed to past) influences of Man and
potential threats would have been useful.

The next three chapters cover vascular plants, bryophytes and lichens, respectively. There are
some 1,190 or so native or introduced and 550 casual vascular plant species including about 140
microspecies. Arrangement and nomenclature follows Flora Europaea (with more up-to-date
names when appropriate) and English and Welsh names are given. For each species there is
information on status, frequency, habitat and distribution in Glamorgan. For many species there
are data on floral biology often followed by a list of insect pollinators. Recording was based on 5-km
squares of the National Grid. When species occur in less than ten such squares, localities are listed.
For more frequent species maps are provided (a total of 1,019) with pre-1960 and later records
distinguished. Many specimens of critical groups (Dryopteris, Rubus, Rosa, Euphrasia, Taraxacum,
Hieracium and Carex) were checked by experts.

The chapter on bryophytes by A. R. Perry treats distribution of the 459 recorded species based on
a 10-km square survey and in the instance of rarer species, site information is provided. As with
vascular plants coverage of the county has been thorough.

The chapter on lichens is by A. Orange. Although there has been no systematic survey of these
plants, 518 lichens and 38 non-lichenised fungi are recorded on a 10-km grid square basis. This
represents little more than a compilation of available records but with so few competent recorders
represents a good start. For all species, pre-1960 and later records are distinguished; there are brief
habitat details and, for rare species, localities.

The work ends with lists of collectors and contributors, a four-figure grid reference gazetteer and
a map of the botanical vice-county of Glamorgan.

One’s first impression on glancing through this softback book is that it is a good Flora and this is
confirmed on closer examination. Quality of production is high and the colour photographs of plants
and sites are excellent. This is a first-rate addition to the long list of British local Floras.

A. J. E. SMITH

Watsonia 20: 447-452 (1995) 447

Obituaries

MARJORIE DEVEREAU
(1915—1994)

Majorie Devereau, whose sudden death occurred on 19 December 1994, was one of the two
assistant authors of the Flora of the Isle of Man, published in 1984 (though bearing a date two years
earlier). Together with her friend and principal field companion, Dr Larch Garrad, she was
responsible for the culminating impetus that carried the long-deferred work to its conclusion, taking
upon herself the onerous task of typing the greater part of the manuscript.

Born on 28 May 1915 and brought up in Douglas, one of four children of a senior official in the
Island’s Board of Education, Miss Devereau spent her entire career as a primary schoolteacher.
Though her first post was ‘across’ (as the Manx say), in Bootle, and she returned to take up further
ones in England during and after the Second World War, her heart always remained in the Isle of
Man and it was with relief that she was able to spend her final 23 years of teaching there.

After a chance meeting with Dr Garrad at an evening class in marine biology in 1965, she took up
in earnest the interest in botany that she had had from an early age, an interest which nevertheless
had to compete with amateur dramatics, gardening, garden history and learning the Manx language.
Quickly developing impressive proficiency as a field botanist, she was soon making many notable
finds, especially in Carex, a genus she was particularly proud of having mastered. C. dioica and C.
diandra were both added to the Manx list through her acumen.

Retirement in 1975 brought her more leisure and allowed her at last to attend B.S.B.I. field
meetings elsewhere in the British Isles, as well as to visit choice botanical areas on the Continent.
She also undertook most of the Isle of Man coverage for the B.S.B.I.’s Monitoring Scheme and was
compiling the records for two of its 10-km squares for the new Af¢las at the time of her death.
Retaining a sharp eye to the end, she celebrated her penultimate field season in fitting style by
detecting two new county records of Trifolium species in a stretch of ground repeatedly worked over
by botanists from John Ray onwards.

Though she formed no herbarium (her vouchers were pressed under her living-room carpet) and
was too diffident ever to publish on her own account, the size of her contribution to knowledge of
the vascular flora of her native island will fortunately be abundantly apparent to posterity through
the great number of records that stand to her name in print.

D. E. ALLEN

DOROTHY MARY GREENE
(1928—1994)

Mrs Dorothy Greene died on 31 October 1994. Although few B.S.B.I. members will have known
her, she played a crucial réle behind the scenes as database manager at the Biological Records
Centre, Monks Wood, in the 1980s. Those who use the animal and plant distribution maps and
atlases produced by B.R.C. owe a great deal to her dedicated work as a pioneer manager of
biological databases.

Dorothy Douglas was born in Coleraine, Northern Ireland, on 6 June 1928, the daughter of a
schoolteacher. After attending Coleraine High School, she entered Trinity College, Dublin,
graduating with a B.Sc. in Natural Science in 1951. At a Trinity College Bible-reading she met |
Stanley Greene, the son of a Co. Cork landowner, who she married at a teetotal wedding in 1953.
(Their presence at a Bible-reading and at any teetotal event will astonish those who knew them only
in later life!) After her marriage Dorothy initially worked as a medical technician, but she was
employed as a part-time demonstrator in Botany at the University of Birmingham after Stanley
obtained a post there in 1955. In 1959 she obtained an M.Sc. for a thesis on British bryophytes, and

448 OBITUARIES

in 1961 she began to study Antarctic bryophytes, the main research interest of her husband. She was
probably the first British woman scientist to ‘go south’ with the British Antarctic Survey, and her
visit to South Georgia in the summer of 1967-68 was one of the highlights of her lite. She retained
vivid memories of hazardous transfers from ship to ship in rough weather, of the wonderful
landscape of South Georgia and of the superb seafood available from the Antarctic waters. She was
officially taken on to the staff of B.A.S. in 1969, and transferred to the Institute of Terrestrial
Ecology’s research station at Penicuik, Midlothian, with the rest of the Birmingham team in 1974.
During this period she was the co-author of a number of papers on polar bryophytes, and she began
work on A conspectus of the mosses of Antarctica, South Georgia, the Falkland Islands and Southern
South America. This detailed checklist of 1,959 taxa, with relevant synonyms and a detailed and
meticulously compiled bibliography, was eventually published in 1986. She also set up a database of
Antarctic botanical records, based on specimens in the B.A.S. herbarium. This database, which was
fully operational by 1971, was one of the first of its kind.

In 1978 Dorothy’s marriage to Stanley broke up, and she transferred to Monks Wood. In 1979 she
became database manager at the Biological Records Centre. The ten years she spent at B.R.C. were
marked by major changes in the structure of the database. Dorothy did not write computer
programs herself, but her knowledge of the needs of biologists and the strengths and limitations of
databases allowed her to develop the B.R.C. system in collaboration with computer specialists.
When she arrived at Monks Wood only the minimum details of a record were routinely placed on
computer file. In 1981 the database was radically revised so that full details were added for new
records. Programs were developed to check the records for geographical inconsistencies, and new
software enabled maps to be plotted by computer rather than mechanically or by hand.

The first task Dorothy tackled at B.R.C. was the preparation of maps for the new edition of the
B.S.B.I. Sedges handbook. This began with a major trauma, when the first set of maps lacked many
of the significant records plotted in the Adlas of the British flora but which, it turned out, had never
been computerized. However, Dick David stepped in to work with Dorothy and revised maps were
produced. At the same time she and Jenny Moore set up a database of Charophyte records, from
which distribution maps and a catalogue of the Natural History Museum collection were published.
The days working with Jenny Moore in London provided a welcome break from the Monks Wood
routine, and the resulting publications revived interest in these neglected plants. Experience with
these data sets enabled her to tackle some large data sets built up by recording schemes which had
been set up after the successful completion of the Atlas of the British flora. She was responsible for
computerizing the records for the atlases of bryophytes, butterflies, mammals, marine algae,
myxomycetes and woodlice, and for other publications currently in press.

Dorothy brought a formidable energy and determination to her work at B.R.C. She would
regularly start work at 6.30 or 7 a.m., taking advantage of the more rapid performance of the
computer before it was slowed down by other users. Sustained by cigarettes and frequent cups of
almost black coffee, she worked from Monday to Friday and sometimes through the weekend as
well. She affected a cynical attitude to recorders and recording, and frequently expressed her
frustration at the tendency of records to ‘multiply in captivity’, as she put it, or to emerge ‘from
under the bed’ after she had been told that the database was complete. Much of her work involved
labour-intensive manipulation of computer files, using software which was less sophisticated than it
is now. She took a pride in the speed with which she could deal with mountainous piles of computer
print-out. If she was given a large job she would often put it aside, saying that she could not possibly
deal with it now, but by the next morning it would be dumped unceremoniously in the centre of your
desk with a curt note ‘Please check by lunchtime, D.’. The permanent employees at B.R.C. found
that working with her was both enjoyable and rewarding, and she was always at hand to help in a
crisis. However, sometimes she could be caustic to those who, in her opinion, were not pulling their
weight. Anyone complaining that they could not complete a task in time was likely to be advised to
cut down on the excessive number of hours they spent sleeping. Sadly, a few of those she attempted
to goad into action failed to see the human being behind the ruthless front she presented to them.
She resented anything that interfered with the job in hand, avoiding administrative chores, internal
seminars and meetings outside Monks Wood whenever she could. In the evenings she returned
home to relax with her long-haired cat, Sheba, and a stiff drink. When not working over the
weekend she cajoled Sheba into her travelling basket and drove off on Friday afternoon in her MG
_ Midget to Herefordshire, where she was restoring and modernizing a decrepit old cottage with two

OBITUARIES 449

friends. After a weekend of strenuous labour on the cottage, she would be back at work at9 a.m. on
Monday, proudly displaying her work-torn hands.

Dorothy’s last task at B.R.C. was to manage the migration of the database to ORACLE, a far
more efficient database management system than had hitherto been available to B.R.C. During the
period between her appointment in 1979 and her retirement in 1989 the database had nearly
doubled in size, from 2.5 to 4.5 million records. She retired to her Herefordshire cottage, the
modernization of which had been completed with her characteristically good taste and eye for
detail. She died of cancer, after a short illness. Her friends will cherish memories of her forthright
and vigorous personality.

J. M. Crorr & C. D. PRESTON

LEONARD ALBERT LIVERMORE
(1919—1994)

When Len Livermore died of cancer on 5 August 1994, Lancashire lost one of its best and most
independent botanists.

Len was born on 7 January 1919 at Lancaster, the son of Frederick Charles and Rachael
Livermore. He was educated at local schools and in 1935 joined Lansil Ltd of Lancaster as a
laboratory technician working on textiles. In 1939 he joined the Territorial Army and was called up
two days before the Second World War began. After brief training he was sent to France as an
artillery signaller, saw action in 1940, and was lucky enough to escape home from Dunkirk. In 1941
he transferred to the Royal Army Ordnance Corps and was sent to North Africa to destroy
dangerous captured ammunition as the Allies pushed west. He rose to the rank of Staff Sergeant
before being demobilized in 1946. He returned to Lancaster and Lansil, and obtained qualifications
as an industrial chemist at night school, ultimately gaining Fellowship of the Royal Institute of
Chemistry in 1956. He retired in 1980 as Works Chief Chemist when Courtaulds Ltd closed their
Lancaster factory.

In 1947 Len met his future wife, Pat, through a mutual friend. Together they discovered the
pleasures of fell-walking and photography in the Lake District. They were married in 1949, thus
beginning a happy marriage and a lifelong partnership. Their only child, David, was born in 1958.

Len’s interest in botany came via horticulture. He grew orchids and other tropical plants (he was a
founder member of the Bromeliad Society) and, later, alpines, with a special fondness for Primula
and Cyclamen. He won many prizes at horticultural shows in the North-West for his displays of
greenhouse plants and cut flowers. Gradually, through Pat’s interest in photographing wild flowers
using a camera Len had bought her after winning an Amateur Gardening competition, he became
interested in wild plants. They joined the B.S.B.I. in 1974, and in 1980 retirement allowed him to
take up botany full-time.

Botany was treated with the same care and dedication as explosives, chemistry and horticulture.
Len’s most important contribution to botany, Flowering plants and ferns of North Lancashire, was
jointly written with Pat. This local Flora (using the term in its best sense) was the first for the area
since Wheldon & Wilson’s Flora of West Lancashire (which covered all v.c. 60) in 1907. It was
recorded between 1976 and 1986, almost exclusively by Len and Pat. They complemented each
other, Len seeking out the more critical groups, ferns and aquatics, leaving Pat to look for the
commoner plants and the grasses, rushes and sedges.

Len would always ask permission to record on private land. He would knock on a farm door, and
spend ten minutes or more chatting about the farming and explaining what he was doing, whilst
establishing his local credentials. This exemplary approach paid off, and Len and Pat were never
once met with refusal, allowing them into places never botanized previously. Some sites off the
beaten track were absolute gems; I accompanied them on one recording trip in 1982 to the Greta
Gorge, where sunlight dappled through the alder woods on to the white sheets of large bittercress
(Cardamine amara), still the most extensive and beautiful stand I have ever seen.

The Flora was carefully planned, and Len’s determination to get proper coverage of all groups
and tetrads was influenced by an unfair Watsonia review of another local Flora. The Flora was

450 OBITUARIES

concisely presented with tetrad maps in the current fashion, each dot coloured by hand. He carefully
costed it (even down to complimentary copies, but he forgot all his family would want one too!), and
kept the price down so that people would buy it. The Flora was very well received.

Len was not daunted by critical species, and he particularly enjoyed pottering in the meadows
around Lancaster looking for Taraxacum, amassing a very respectable number of species
(Livermore & Livermore 1991b). In May, the conservatory had row after row of carefully labelled
yoghurt pots with clocking dandelion heads. He retained many critically-determined voucher
specimens for reference; the collections have now been deposited at Lancaster University (LANC)
and Liverpool Museum (LIV). Each autumn as the botanical season came to a close, Len and Pat
would switch attention to fungi, this time Pat providing the expertise and Len the support.

With the bit between his teeth Len was not one to stop, and detailed accounts of the canals, coast,
dismantled railways and the City of Lancaster were produced in rapid succession, again with Pat.
These reports were published by persuading companies to type and print them as a public relations
exercise. Len would carefully negotiate with the senior executives and then encourage the typists
often with garden produce; after many years in management in the private sector, he knew how to
get what he wanted. As the reports were not for sale, he deposited copies in libraries, and put notes
in B.S.B.I. News so that others knew they were available.

Together with Pat and other friends, including Margret Baecker, Ruth Berry, John Leedal and
Neil Robinson, he also spent many happy botanical holidays abroad in Canada, Turkey and
Europe. On occasions, with the aid of some wild horses, Margret and I even managed to drag him |
over the Lancashire border into Cumbria to look for Sorbus lancastriensis!

Len was committed to conservation. For many years he was on the Northern Regional Committee
and the Scientific and Conservation Committee of the Lancashire Naturalists’ Trust (now the
Lancashire Wildlife Trust). He had strong views on nature conservation, and had little hesitation in
letting people know when he thought they were doing nothing or doing the wrong thing. Rightly, he
felt that conservation policies should be based on sound information, and the initial objective of the
Flora was to gather systematic data to determine priorities for conservation in North Lancashire. He
felt too little attention was paid to groups like lower plants, fungi and invertebrates, which were
largely ignored by the statutory nature conservation bodies in selecting sites, simply because they
rarely collected the information. He and Pat made the point to the Nature Conservancy Council
with the outstanding list of fungi from Gait Barrows National Nature Reserve, which is unique for
its detail and coverage. Len was of the non-intervention school for site management and thought
that some sites would be better left to look after themselves.

Len was a shrewd, tough, independent northerner who knew his own mind. He preferred action
to words, and had little time for committees or people who just talked. He held forthright views on
the Civil Service, the state of the manufacturing industry and botanical referees who did not answer
letters. He would fervently argue his corner over a cup of tea and a cigarette, and would usually give
better than he got. He was always very cagey with his information, and most of us rapidly learnt not
to pass on his records to third parties; he was upset if people re-used his records without taking the
courtesy to ask him first. He taught me, a mere southerner, a lot both about botany and handling
people, and I remember with pleasure botanizing with him in the field.

When he was diagnosed as having cancer in 1992, true to form he would not give in and carried on
as normal, researching his family history and caring for his garden. He is survived by Pat and their
son David.

PUBLICATIONS OF LEN AND PAT LIVERMORE

Livermore, L. A. & Livermore, P. D. (1987). Flowering plants and ferns of North Lancashire.
Privately published, Lancaster.

Livermore, P. D. & Livermore, L. A. (1987). Fungi of Gait Barrows National Nature Reserve.
Nature Conservancy Council, Blackwell.

Livermore, L. A. & Livermore, P. D. (1989). Flowering plants, ferns and rusts of the Lancaster
Canal in the Lancaster District. Privately published, Lancaster.

Livermore, L. A. & Livermore, P. D. (1990). Coastal plants and rust fungi of the North Lancashire
coast. Privately published, Lancaster.

Livermore, L. A. & Livermore, P. D. (1990). Plants and rust fungi of the dismantled railway lines in
the Lancaster District. Privately published, Lancaster.

OBITUARIES 45]

Livermore, L. A. & Livermore, P. D. (1991a). Lancaster’s plantlife — a botanical survey. Privately
published, Lancaster.

Livermore, L. A. & Livermore, P. D. (1991b). Taraxacum flora of north Lancashire. B.S.B.I. News
57: 9-10.

THE2Ge Rich

PETER JAMES WANSTALL
(1924—1993)

Peter Wanstall died suddenly at his home in Thrandeston, Suffolk, on 29 January 1993. He had been
a member of the B.S.B.I. since 1949 and had served as a member of the Council for two terms
(1955-59, 1965-69), as the Society’s representative on the Biological Council and as a member of
the Conservation Committee. Perhaps his chief contribution, however, was as Honorary Secretary
to the Publications Committee, a post that he held from 1954 to 1968. During this period he edited
two of the Society’s conference reports, A Darwin centenary (1961) and Local Floras (1963). He was
elected an Honorary Member in 1968.

Peter was born on 21 March 1924 in Guildford, a town for which he retained a lasting attachment.
However, the family moved to Brighton when he was 12, and it was in the Sussex downland that his
love of nature was nurtured and grew. He attended Varndean School, Brighton, where he ran the
Field Club, and Queen Mary College, University of London. The College had been evacuated to
King’s College, Cambridge at the outbreak of war and this was to have a profound influence on
Peter’s educational development, for, as he once told me, the ‘Cambridge years’ were among the
most stimulating and formative of his life.

In 1946 he joined the staff of the Botany Department of Queen Mary College, where he remained
for his entire professional career, retiring in 1984. He taught courses on bryophytes, pteridophytes
and gymnosperms, plant ecology and, latterly, plant geography. In the 1960s he also conducted the
University extramural course for the Certificate in Field Biology. As a teacher, Peter came into his
own in practical classes and tutorials, where he was much more at ease than in the lecture room. His
research interests lay in the field of bryophyte ecology and, for some time, he worked on the
structure and growth of populations of two mosses, Polytrichum formosum and P. juniperinum.
Sadly, he published little, not through the lack of ability, for he possessed a good critical mind and
wrote well, but mainly, I think, through lack of self-confidence.

For more than 40 years, Peter worked to further the cause of British natural history, giving
generously of his time and expertise to the many organizations of which he was a member. With his
background and interests, it is not surprising that he became involved in the conservation movement
as it developed in the late 1950s and early 1960s. From its earliest days he played a prominent part in
the affairs of the Essex Wildlife Trust and, on moving to Thrandeston, became active in the work of
the Suffolk Wildlife Trust as Chairman of the Conservation Management Committee. Peter had an
especially close relationship with the Field Studies Council, extending over most of the first 50 years
of its existence. Apart from running courses at several of the Council’s centres, he served in a variety
of capacities but notably as a member of the Executive Committee from 1960. At the time of his
death he was also Chairman of the Publications Committee, having held the office since 1987.

The British Bryological Society, the Linnean Society of London and the Suffolk Naturalists’
Society were other organizations to benefit from Peter’s membership. Joining the British
Bryological Society in 1946, he served, inter alia, as referee for the moss families Andreaeaceae,
Polytrichaceae and Buxbaumiaceae, as a member of the Council for four terms between 1952 and
1984, as Vice-President (1986-87) and as President (1988-89). In 1983 he performed an outstanding
service for the Society as principal organizer of the Diamond Jubilee Meeting held at Bedford
College, London and Box Hill. He was a staunch supporter of the Linnean Society, having been
elected a Fellow in 1953. Serving as a member of Council (1975-80) and as a Vice-President (1976—
77), he also played a large part in the organization of the Society’s joint meeting with the British
Bryological Society in 1987. On a social level he was Honorary Secretary-Treasurer of the Linnean
Dining Club for eleven years, standing down in 1989. After retiring, he quickly became involved in

452 OBITUARIES

the activities of the Suffolk Naturalists’ Society, as a Council member (1984-89) and as Botany
Section Editor of Suffolk natural history, the Society’s Transactions.

Peter greatly enjoyed foreign travel, having a keen interest in other cultures, both modern and
ancient. Two countries in particular held a special place in his affection — France and Nigeria. France
he came to love through his many visits to Provence and the eastern Pyrenees with student groups
and, more recently, as leader of courses for the Field Studies Council. His first experience of Nigeria
was in 1966 when he spent three months teaching at the University of Ife (Ibadan Campus) under
the auspices of the ‘VISTA’ scheme. He made two shorter visits in 1971 and 1976-77 as an external
examiner at the University of Ibadan. It was during the first of these that he and I travelled widely
together, collecting and sightseeing in the north-east of the country. Our journey, of almost 5000
km, was not without its share of difficult and trying situations, but Peter’s good humour and calm,
positive approach to problems invariably helped us through. His bryophyte collections from 1966
and 1971 are now housed in the Cryptogamic Herbarium of the Natural History Museum, London
(BM).

Lepidoptera were another passion dating back to his school-days. He had a considerable
knowledge of the group and always included a moth trap with the equipment taken on field courses
to France. I can see him now, deftly wielding his butterfly net wherever we stopped to collect on our
travels through Nigeria.

Peter Wanstall’s enthusiasm, sociability and courtesy never failed him. He was a wonderful
companion in the field, a wise counsellor and a generous and loyal friend. At his funeral, the church
was filled to capacity, a striking testimony to the affection and esteem in which he was held.

To his wife, Jane, and children, Sally, Ian and Giles, we extend our sincere condolences in their
great loss. .

A. J. HARRINGTON

SS

B.S.B.I. PUBLICATIONS

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 and R. W. David. 1982. 268 pages, with a line drawing and distribution map
for every species. Paperback. ISBN 0 901158 05 4.

Ne. 2 UMBELLIFERS OF THE BRITISH ISLES
T. G. Tutin. 1980. 197 pages, with a line drawing for each species. Paperback. ISBN 0 901158 02 X. Out
of print: new edition in preparation.

No. 3 DOCKS AND KNOTWEEDS OF THE BRITISH ISLES
J. E. Lousley and D. H. Kent. 1981. 205 pages, with many line drawings of British native and alien taxa.
Paperback. ISBN 0 901158 04 6. Out of print: new edition in preparation.

No. 4 WILLOWS AND POPLARS OF GREAT BRITAIN AND IRELAND
R. D. Meikle. 1984. 198 pages, with 63 line drawings of all species, subspecies, varieties and hybrids.
Paperback. ISBN 0 901158 07 0.

No.5 CHAROPHYTES OF GREAT BRITAIN AND IRELAND
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No. 7 ROSES OF GREAT BRITAIN AND IRELAND
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LIST OF VASCULAR PLANTS OF THE BRITISH ISLES
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ATLAS OF THE BRITISH FLORA
F. H. Perring and S. M. Walters, 1990. Reprint of 3rd ed, 1982. 468 pages. Distribution maps of over
1700 species including updated maps for 321 Red Data Book species. New bibliography of updated
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ALIEN PLANTS OF THE BRITISH ISLES
E. J. Clement and M. C. Foster, 1994. 590 pages. Lists 3586 species found growing wild and includes
frequency of occurrence, areas of origin, means of introduction, references to descriptions and
illustrations and location of herbarium material. Excludes grasses, to come in Vol. 2. Paperback. ISBN
0 901158 23 2.

SCARCE PLANTS IN BRITAIN
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which occur in 16-100 10-km squares with maps showing records before and after 1970 and text giving
details of habitat and associated species. Of vital importance to all conservationists. ISBN 1 873701 667.

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OF THE

BRITISH ISLES

BSB. ..BeS. BL)

The B.S.B.I. was founded in 1836 and has a membership of
2,/00. It is the major source of information on the status and

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a panel of referees available to members to name problem
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Details of membership and any other information about the
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c/o Department of Botany,

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WATSONIA

THE JOURNAL OF
THE BOTANICAL SOCIETY
OF THE BRITISH ISLES

VOLUME 20 SMITHSON TA
MAR 0.9 1999
C/BRARIES

EDITED BY

J.R. AKEROYD, J. R. EDMONDSON, R. R. MILL,

E. C. NELSON, C. D. PRESTON anp B. S. RUSHTON

1994-95

PRINTED FOR THE SOCIETY BY
Eaton Press Limited, Wallasey, Merseyside

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

a — = —— =—- > - — —— _ _ — ———_—- —-

DATES OF PUBLICATION

Part 1, pp. 1-88, 31st March 1994

Part 2, pp. 89-184, 6th September 1994
Part 3, pp. 185-328, 17th February 1995
Part 4, pp. 329-452, 30th August 1995

p. 11, line I] up.

p. 78, line 17 up.
ps5, line 2.

p. 131, caption.

p. 136, line 11 up.

pe 182, line 20:
p.205, header.
[De ALD JVI tore

p. 300, line 20 up.

p. 377, line 2 up.

p. 388, line 18 up.

p27: line 21.

Errata

For:
For:
For:
For:
For:
For:
For:
For:
For:
For:
For:

For

gerardi read: geradii

littorale read: litorale

16 May read: 15 May
sahlinanum read: sahlinianum
marshalli read: marshallii —
platyphyllus read: platyphyllos
185-194 read: 205-227
sylvatica read: silvatica
Anisandra read: Anisantha
alpinim read: alpinum
Hippophaé read: Hippophae
: 1 amarckii read: lamarckii

INDEX
WATSONIA VOLUME 20 (1994-95)

Prepared by C. R. Boon

Acacia 174

Acaena inermis (v.c. 44) 426; novae-zelandiae
(v.c. 46) 292; ovalifolia (v.c. 77) 426

Acanthus 439

Acer negundo 341; palmatum ‘Senkaki’ 309; pla-
tanoides ‘Crimson King’ 309

Achillea ptarmica 10

Aconitum X cammarum (v.c. 77) 286; napellus x
variegatum (v.c. 77) 286

Aceorus 185; calamus 10

Adiantum 164; capillus-veneris 382, 383

Adler, W., Oswald, K. & Fischer, R., with M. A.
Fischer (ed.) — Exkursionsflora von Oster-
reich (Bk Rev.) 303-304

Aethusa cynapium subsp. agrestis (v.c. 4) 429

Agrimonia procera (v.c. 28, 104) 292, (v.c. 50)
426

Agropyron maritimum 385, 386; repens 385

Agrostis capillaris 235, 394; castellana (v.c. 3, 58)
300; scabra (v.c. 83) 300; stolonifera 8, 35,
234, 235, 388, 392

Ajuga reptans 443

Akeroyd, J. R. — Obit. of Brian Thomas Styles
(1934-1993) 173-175

Akeroyd, J. R. & Beckett, G. — Petrorhagia
prolifera (L.) P. W. Ball & Heywood (Cary-
ophyllaceae), an overlooked native species

in eastern England 405—407

_ Akeroyd, J. R., with T. G. Tutin et al. (eds) &

M. E. Newton — Flora Europaea, 2nd

edition, vol. 1: Psilotaceae to Platanaceae

| (Bk Rev.) 81-82

_ Alchemilla 303; mollis (v.c. 40, 48) 426; tytthantha
(v.c. 77) 292

Alisma lanceolatum 35; plantago-aquatica 35, 235

Allaby, M. (ed.) — The concise Oxford dictionary
of botany (Bk Rev.) 73

| Allan, B. & Woods, P. J. B. — Wild orchids of

Scotland (Bk Rev.) 73-74

. Allen, D. E. — Additional sets of continental

| Rubus exsiccatae in British herbaria 409

| - Allen, D. E. — Four new Anglo-Sarnian species of

) i} Rubus L. (Rosaceae) 143-151

| Allen, D. E. — Rubus aghadergensis D. E. Allen

| (Rosaceae): a new name for an Irish bramble

61

Allen, D. E. — Rubus percrispus D. E. Allen &
R. D. Randall (Rosaceae) in Dorset (v.c. 9)
407-408

Allen, D. E. — Typification of Rubus pullifolius
W. C. R. Watson (Rosaceae) 408

Allen, D. E. — Rev. of A catalogue of the Herbar-
tum of the British Flora collected by Margaret
Stovin (1756-1846) (by M. Simmons) 442

Allen, D. E. — Rev. of Catalogue of the Brambles
of Britain and Ireland in the herbarium of
Liverpool Museum (LIV) (by M. Palmer
(comp.), with J. Edmondson (ed.)) 442

Allen, D. E. — Rev. of The Wiltshire Flora (ed.
B. Gillam) 158-160

Allium schoenoprasum (v.c. 46) 436; scorodopra-
sum (v.c. 98) 301, (v.c. 58) 436; subhirsutum
(v.c. 17) 301; ursinum 443; vineale (v.c. 104)
301

Alnus Miller in Ireland, Variation and hybridis-
ation of, 67—70; cordata (v.c. 58) 420; gluti-
nosa 10, 67-69, 235; incana 67, 69, 70, (v.c.
77) 287, (v.c. 26, 77) 420; x pubescens 69, 70

Alopecurus aequalis 11; x brachystylus (v.c. 4, 77)
435; bulbosus 8; geniculatus 35, 234, 235;
geniculatus x bulbosus (v.c. 44) 435; x plettkei
(vic; 44) 435; pratensis 11, (v.c. 102) 435;
pratensis x geniculatus (v.c. 4, 77) 435

Althaea officinalis 9, 63

Alyssum saxatile (v.c. 42) 422

Amelanchier lamarckii (v.c. 77) 427

Ammi majus (v.c. 38) 294, (v.c. 40) 429

Ammophila arenaria 207, 214, 388

Amsinckia micrantha (v.c. 40, 59) 430

Anabasis 440

Anagallis arvensis subsp. arvensis 63; minima
(v.c. 104) 289, (v.c. 40) 423; tenella 10, 12, 63

Anaphalis margaritacea (v.c. 12, 40) 432

Andromeda polifolia 9, 12

Anemone nemorosa (v.c. 102) 420

Angelica sylvestris 11

Anisantha diandra (v.c. 38) 300; sterilis (v.c. 73)
435

Annual General Meeting (1993) (Rpt) 85-87,
(1993) (Rpt) 181-182, (1994) (Rpt) 327-328

Antennaria 330; dioica 11, 172

Anthriscus caucalis (v.c. 58) 429

453

454 INDEX TO WATSONIA VOLUME 20

Anthyllis vulneraria subsp. carpatica (v.c. 25) 293,
subsp. vulneraria 63

Apera spica-venti (v.c. 13) 300

Apium graveolens 10; inundatum 10, 35, 53, (v.c.
78) 429; nodiflorum 63, 235

Aquilegia stuartii 276; vulgaris 63

Arabis caucasica (v.c. 39) 422; hirsuta (v.c. 102)
422; scabra 383

Arctostaphylos alpinus (v.c. 104) 289

Arenaria balearica (v.c. 40) 420; laricifolia 418;
serpyllifolia subsp. leptoclados (v.c. 81) 287

Armeria 239, 253; maritima 78, 252, 253, 392-394

Artemisia absinthum 63, (v.c. 81) 433; stelleriana
(v.c. 75) 433

Arum maculatum 443

Aruncus dioicus (v.c. 47) 290

Arundo Stricta 56

Asarum europaeum 304

Asparagus horridus 305; stipularis 305

Asplenium marinum (v.c. 81) 285; trichomanes
subsp. trichomanes (v.c. 50) 286; tricho-
manes-ramosum 164; viride 164, 303

Aster laevis (v.c. 69) 432; laevis x novae-belgii
(v.c. 46) 298, (v.c. 73) 433; lanceolatus (v.c.
69) 433; tripolium 11, 63, 392, 394; x versico-
lor (v.c. 46) 298, (v.c. 73) 433

Astilbe rivularis (v.c. 98) 290

Astragalus danicus (v.c. 108) 293, (v.c. 108) 428

Athyrium 164; filix-femina 78; flexile (v.c. 92) 286

Atriplex x gustafssoniana (v.c. 46) 287; hortensis
(v.c. 38) 420; littoralis (v.c. 35, 46) 287;
longipes (v.c. 46) 420; portulacoides (v.c. 46)
287, 388; praecox (v.c. 104) 287; prostrata
(v.c. 81) 287; prostrata x longipes (v.c. 46)
287

Atropa belladonna (v.c. 75) 295

Avena fatua 63, (v.c. 79) 300; sterilis subsp.
ludoviciana (v.c 40) 435

Azolla filiculoides 8, 13, 28, (v.c. 46) 286, 294

Baecker, M. — Rev. of The Flora of Ditchley: wild
flowers of an Oxfordshire estate (by A. J.
Dunn) 158

Bailey, J. P., with R. E. Dempsey & R. J. Gornall
— Contributions to a cytological catalogue of
the British and Irish flora, 4 63-66

Baker, A. J. M., Proctor, J. & Reeves, R. D.
(eds) — The vegetation of ultramafic (serpen-
tine) soils (Bk Rev.) 74-75

Baldellia ranunculoides 10, 35

Barbarea intermedia (v.c. 78) 422; stricta 8; vul-
garis (v.c. 100) 421

Baumann, H., with W. T. Stearn & E. R. Stearn
(trans.) — Greek wild flowers and plant lore in
ancient Greece (Bk Rev.) 439

Baxter, T. — The eternal yew (Bk Rev.) 75-76

Beckett, E. — Illustrated Flora of Mallorca (Bk
Rev.) 304-305 -

Beckett, G., with J. R. Akeroyd — Petrorhagia

prolifera (L.) P. W. Ball & Heywood (Car-
yophyllaceae), an overlooked native species
in eastern England 405-407

Bellis perennis 214

Berberis darwinii (v.c. 50, 77) 420

Berula erecta 11

Beta vulgaris subsp. maritima 63

Betula 311; alba 311

Bidens cernua 10, 235; frondosa (v.c. 29) 433;
tripartita 11, 35, 235, 433

Binggeli, P. — Rev. of The release of genetically-
engineered organisms (eds B. Shorrocks &
D. Coates) 164-165

Birkinshaw, C. R. — Aspects of the ecology and
conservation of Damasonium alisma Miller
in Western Europe 33-39

Bittrich, M., with K. Kubitzski & J. G. Rohweder
(eds) — The families and genera of vascular
plants. Vol 2: flowering plants, dicotyledons,
Magnoliid, Hamamelid and Caryophyllid
families (Bk Rev.) 162-163

Blackstonia perfoliata (v.c. 28) 294

Blamey, M. & Grey-Wilson, C. — Mediterranean
wild flowers (Bk Rev.) 157 :

Blechnum spicant 11

Block, M., with M. J. Bunting & D. Briggs — The
Cambridge British Flora (1914-1920) 195—
204

Blunt, W. & Stearn, W. T. — The art of botanical
illustration (Bk Rev.) 305-307

Blysmus compressus 9, 11; rufus 11

Book Reviews 73-83, 157-166, 303-316, 439-446

Botrychium 164; lunaria 9, 12, 164

Bowen, H. J. M. — Obit. of Ronald D’Oyley
Good (1896-1992) 169-171

Brachypodium pinnatum (v.c. 79) 435; sylvaticum
Srey

Brassica juncea (v.c. 41) 289, (v.c. 25) 423; nigra
64, 382

Bridson, G. D. R. — Rev. of The art of botanical
illustration (by W. Blunt & W. T. Stearn)
305-307

Briggs, D., with M. J. Bunting & M. Block — The
Cambridge British Flora (1914-1920) 195-
204

Briggs, M. — Annual General Meeting (1993)
(Rpt) 85-87, (1994) (Rpt) 327-328

Briggs, M. — Obit. of Edgar Dunstan Wiggins
(1911-1993) 179-180

Briggs, M. — Rev. of Marianne North at Kew
Gardens (by L. Ponsonby) 79-80

Briggs, M. — Rev. of Mediterranean wild flowers
(by M. Blamey & C. Grey-Wilson) 157

Briza media 10

Bromopsis inermis (v.c. 69) 300; ramosa 64

Bromus commutatus 9; hordeaceus (v.c. 102) 435;
interruptus 64

Brummitt, R. K. — Vascular plant families and
genera. A listing of the genera of vascular

INDEX TO WATSONIA VOLUME 20

plants of the world according to their families,
as recognised in the Kew Herbarium, with an
analysis of relationships of the flowering plant
families according to eight systems of classift-
cation (Bk Rev.) 439-440

Brunnera macrophylla (v.c. 64) 295

Bryonia dioica 64

Bryum creberrimum 443

Bull, A. — Eric Edees, Norfolk and Norfolk
brambles 319-320

Bull, A. L., with H. E. Weber — Rubus edeesii H.
E. Webber & A. L. Bull (Rosaceae), a new
bramble species from Britain and Germany
345-349

Bunting, M. J., Briggs, D. & Block, M. — The
Cambridge British Flora (1914-1920) 195-
204

Bupleurum falcatum L. (Apiaceae) in the British
flora. The status of, 115-117, subsp. cernuum
116, subsp. falcatum 115-117; tenuissimum
11

Burbidge, R. B. — Rev. of Wild orchids of
Scotland (by B. Allan & P. J. B. Woods) 73-
74

Burns, A. , with D. McClintock — Obit. of Kath-
leen Margaret Hollick (1913-1993) 322

Burtt, B. L. — Rev. of Phylogeny and classifi-
cation of the Orchid family (by R. L.
Dressler) 76-77

Butler, J. K., with P. J. O. Trist — Puccinellia
distans (Jacq.) Parl. subsp. borealis (O.
Holmb.) W. E. Hughes (Poaceae) in main-
land Scotland and the Outer Isles 391-396

Butomus umbellatus 10, 53

Buxus harlandii 309; microphylla 309; sempervir-
ens 64

! Calamagrostis 51, 52, 54-58, 397, 399-403; canes-

cens 11, (Wigg.) Roth and their hybrids in S.
E. Yorks, v.c. 61, northern England. Cala-
magrostis stricta (Timm) Koeler, 51-60,
(illus.) 53, (v.c. 24) 300, (Wigg.) Roth and
their hybrid populations in S. E. Yorks, v.c.
61, northern England. A graphical analysis
of the characters of Calamagrostis stricta
(Timm) Koeler, 397-404; canescens x stricta
51, 52, (illus). 53, 57-59, 397, 398, 400, 403;
neglecta 51; purpurea 58; stricta (Timm)
Koeler, C. canescens (Wigg.) Roth and their
hybrids in S. E. Yorks., v.c. 61, northern
England 51-60, (illus.) 53, (Timm) Koeler,
C. canescens (Wigg.) Roth and their hybrid
populations in S. E. Yorks., v.c. 61, northern
England. A graphical analysis of the charac-
ters of, 397-404

| Walla palustris (v.c. 59) 434

} Callitriche 35, 260; brutia 234, 235; hamulata 11;

| platycarpa 8, (v.c. 75) 431; stagnalis 235;

truncata 10

455

Calluna vulgaris 11, 291

Caltha palustris 10

Calystegia x lucana (v.c. 73) 295; sepium subsp.
roseata (v.c. 45) 295; sepium x silvatica (v.c.
73) 295; silvatica (v.c. 80, 108) 430; sold-
anella 214

Campanula glomerata 382; persicifolia (v.c. 46)
297; portenschlagiana (v.c. 49) 431

Cannabis 82; sativa 82

Cardamine amara 449; flexuosa 235; hirsuta 443;
pratensis 235; raphanifolia (v.c. 79) 289

Carduncellus pinnatus 304

Carex 321, 446, 447; acuta 8; acutiformis 312;
appropinquata 53; arenaria 388; x beckman-
nii (v.c. 98) 299; curta 10; demissa 312;
diandra 10, (v.c. 78) 299, 447; dioica 12, 447;
distans 11, 392; disticha 10, (v.c. 93) 299;
divisa 10; divulsa subsp. divulsa (v.c. 58) 434;
echinata 10; elata 9, 11, 53, (v.c. 36) 299;
elongata 10, 312; extensa 11; flava 321; hosti-
ana 10; laevigata 11; limosa 11; magellanica
(v.c. 46) 299; muricata 321, subsp. lampro-
carpa (v.c. 89) 299; nigra 11; ornithopoda
(v.c. 64) 299; otrubae (v.c. 104) 299; palles-
cens 10, (v.c. 83) 299; panicea 11; paniculata
11, 53, (v.c. 86) 299; paniculata x diandra
(v.c. 98) 299; pilulifera 7; pseudocyperus 11,
(v.c. 46) 299; pulicaris 9, 10; punctata (v.c. 3)
434; remota 11, 235, (v.c. 102) 434; rostrata
11, 53, 434; spicata (v.c. 74) 299; vesicaria 9,
53, 235, (v.c. 108) 434; viridula subsp. brach-
yrrhyncha 10, subsp. viridula 9, (v.c. 40) 434;
vulpina 11

Catabrosa aquatica 10, (v.c. 78) 299, (v.c. 78) 435

Ceanothus 309

Centaurea cyanus 444; montana (v.c. 50) 297;
nigra (v.c. 75) 432; scabiosa (v.c. 81) 297, 382

Centaurium erythraea (v.c. 80) 294; tenuiflorum
170

Cephalanthera rubra 172

Cerastium 214; arvense 182; diffusum 214; font-
anum subsp. holosteoides (v.c. 73) 287;
pumilum (v.c. 49) 421; semidecandrum 214,
(v.c. 79) 287

Ceratocapnos claviculata 182, (v.c. 104) 286

Ceratochloa cathartica (v.c. 11) 300, (v.c. 44) 435;
marginata (v.c. 22) 300

Ceratophyllum demersum 11, (v.c. 99) 286; sub-
mersum 11

Chaenorhinum minus (v.c. 93) 296

Chaerophyllum hirsutum (v.c. 77) 429

Chamerion angustifolium 8, 13

Chapman, S. B. & Rose, R. J. — Changes in the
distribution of Erica ciliaris L. and E. x
watsonii Benth. in Dorset, 1963-1987 89-95

Chara 17, Red data books of Britain and Ireland:
Stoneworts (Bk Rev.) 80-81; aspera 417;
globularis 8, 417, subsp. globularis 10, var.
annulata 8, var. virgata 8; hispida subsp.

456 INDEX TO WATSONIA VOLUME 20

hispida 9; penduculata 11; vulgaris 10, 11, 34,
35, subsp. crassicaulis 11, subsp. longibract-
eata 10, var. papillata 10, var. vulgaris 10
Chater, A. O. — Rev. of Sedges and their allies in
Dorset. An atlas of the distribution of the
family Cyperaceae in Dorset (by D. A. Pear-
man) 312-313
Chater, A. O. — Rev. of The fern guide. A field
guide to the ferns, clubmosses, quillworts and
horestails of the British Isles (by J. Merry-
weather & M. Hill) 78-79
Chater, “A. ©. & Rich? h*€:"G-— Rorippa
islandica (Oeder ex Murray) Borbas (Brassi-
caceae) in Wales 229-238
Chelidonium majus (v.c. 81) 286, 418
Chenopodium capitatum (v.c. 13) 287; cheno-
podioides 11; ficifolium (v.c. 46) 420; rubrum
235, (v.c. 46) 287; vulvaria 64
Cherrill, A. J. & Lane, A. M. — Bracken (Pterid-
ium aquilinum (L.) Kuhn) infestation of
rough grazing land in the catchment of the
River Tyne, northern England 105-114
Chionodoxa forbesii (v.c. 77) 436
Church, J. M., with N. F. Stewart — Red data
books of Britain and Ireland: Stoneworts (Bk
Rev.) 80-81
Cicuta virosa 12
Circaea x intermedia (v.c. 50) 429; lutetiana x
alpina (v.c. 50) 429
Cirsium acaule 382; dissectum 10; eriophorum
382; heterophyllum 64; heterophyllum x
palustre (v.c. 78) 432; palustre 10, 64; vulgare
64; x wankelii (v.c. 78) 432
Cistus clusii 304
Cladium mariscus 9, 10
Cladonia rangiformis 214, 223
Claytonia perfoliata 214; sibirica 236
Cleave, A. — Field guide to trees of Britain,
Europe and North America (Bk Rev.) 307-—
308
Clematis vitalba 381, 382
Climacoptera 440
Coates, D., with B. Shorrocks (eds) — The release
of genetically-engineered organisms (Bk
Rev.) 164-165
Cochlearia anglica 394; danica (v.c. 81) 289, (v.c.
42) 423; officinalis (v.c. 81) 289
Coeloglossum viride 10, (v.c. 46) 301, (v.c. 3) 437
Coincya monensis subsp. recurvata (v.c. 58) 289
Colchicum autumnale 9, 10, 18, (v.c. 41) 301,
(v.c. 49) 436
Colutea buhsei 309; multiflora 309
Comes, H. P. — Senecio vulgaris L. subsp. dentic-
ulatus (O. F. Muell.) P: Dy Sell and S.
vulgaris subsp. vulgaris var. vulgaris on
Jersey (Channel Islands) 205-227
Conolly, A. P. — Obit. of Irene Mary Vaughan
(1889-1993) 175-179 ;
Conopodium majus 64, 443

Conyza sumatrensis (v.c. 11, 13, 14) 433

Coombes, A. J. — Rev. of Index of garden plants
(by M. Griffiths) 308-309

Cornus sanguinea 382

Coronopus squamatus (v.c. 77) 289, (v.c. 81) 423,
443

Corrigiola litoralis 35

Corydalis 309

Corylus avellana 443

Corynephorus canescens 388

Cotoneaster — Great Orme Berry — renamed. The
native British, 61-63, (key) 62-63, 309;
section Cotoneaster series Cotoneaster 62;
affinis (v.c. 3, 46) 292; bullatus (v.c. 25, 26,
47, 75) 293, (v.c. 50, 69) 427; cambricus
J. Fryer & B. Hylm6, nom. et stat. nov. 62, .
309; canescens 62; cinnabarinus 62; conspi-
cuus (v.c. 41, 49) 292; dammeri (v.c. 58, 77)
427; dammeri x conspicuus (v.c. 41) 292;
dielsianus (v.c. 47) 293; divaricatus (v.c. 58)
427; franchetii (v.c. 26) 428; frigidus (v.c. 25,
44) 427; frigidus x salicifolius (v.c. 50) 427;
hjelmqvistii (v.c. 41) 293, (v.c. 47, 49) 427;
horizontalis (v.c. 47, 73, 75) 293; integerri-
mus 62, var. anglicus 62; integrifolius (v.c.
77) 292; intermedius 62; juranus 62; lacteus
(v.c. 47) 427; obtusisepalus 62, 63; rehderi
(v.c. 46, 49) 293; salicifolius (v.c. 41, 47) 292,
(v.c. 49) 427; scandinavicus 62; simonsii (v.c.
108) 427; sternianus (v.c. 4, 29, 41, 44) 293,
(v.c. 35, 44) 428; x suecicus (v.c. 41) 292;
villosulus (v.c. 77) 427; x watereri (v.c. 50)
427

Crackles, F. E. — A graphical analysis of the
characters of Calamagrostis stricta (Timm)
Koeler, C. canescens (Wigg.) Roth and their
hybrid populations in S. E. Yorks., v.c. 61,
northern England 397-404

Crackles, F. E. — Calamagrostis stricta (Timm)
Koeler, C. canescens (Wigg.) Roth and their
hybrids in S. E. Yorks., v.c. 61, northern
England 51-60

Crambe maritima (v.c. 75) 289

Crassula helmsii (v.c. 35, 49, 91, 104) 289, (v.c.
26, 40) 424; tillaea (v.c. 13) 289, (v.c. 19) 424

Crataegus 102, 330, 341; section Crus-galli 341;
laciniata (v.c. 57) 428; laevigata 102, 182; x
media 102; monogyna Jacq. (Hawthorn) at
an upland site. A comparison of the growth
and morphology of native and commercially,
obtained continental European, 97-103, 182,
214, 381, 382, subsp. nordica 102; persimilis:
(v.c. 35) 428

Crepis paludosa 64, (v.c 102) 432

Crinodendron hookerianum 309

Crocosmia x crocosmiiflora (v.c 81) 436; panic-
ulata (v.c. 46) 436; pottsii x aurea (v.c. 81)
436

Crocus 321; vernus (v.c. 46) 436

INDEX TO WATSONIA VOLUME 20

Croft, J. M. & Preston, C. D. — Obit. of Dorothy
Mary Greene (1928-1994) 447-449

Cronk, Q. C. B., Marner, S. K. & Primavesi, A.
L. — Lectotypification of Rosa rothschildii
Druce (Rosaceae) 409-410

Cronk, Q. C. B. & Primavesi, A. L. — Further
Druce Rosa taxa (Rosaceae) 411-412

Cruciata laevipes (v.c. 93) 431

Cullen, J. — Rev. of Stearn’s dictionary of plant
names for gardeners (by W. T. Stearn) 80

Cuscuta epithymum (v.c. 59) 430

Cyclamen coum (v.c. 14) 423

Cynodon dactylon 35

Cynoglossum germanicum 172

Cynosurus cristatus 388

Cyperus longus 9, 12

Cyrtomium falcatum (v.c. 11) 286

Cystopteris 164

Cytisus scoparius 430, subsp. maritimus 296; stria-
tus (v.c. 35) 428

Dactylis glomerata 214, 382

X Dactylodenia st-quintinii (v.c. 104) 301

Dactylorhiza 263, 264, 266, 268, 272; fuchsii 263—
266; incarnata subsp. coccinea 266, subsp.
incarnata 10, 266, subsp. pulchella 265, 266,
(v.c. 2) 437; incarnata x purpurella (v.c. 81)
301; lapponica (v.c. 104) 301; x latirella (v.c.
81) 301; maculata 10, 265, 266; majalis subsp.
praetermissa 263-266, subsp. purpurella 266,
subsp. traunsteineri 263—267 , 270, 271, subsp.
traunsteineri var. bowmanii M. N. Jenk., var.
nov. 263, (illus.) 267, 271, 272, subsp. traun-
steineri var. eborensis 271, subsp. traunstein-
eri var. francis-drucei 271, subsp. traunstein-
eri var. traunsteineri 263, 271; praetermissa
10; traunsteineri 263

_ Damasonium alisma Miller in Western Europe.
Aspects of the ecology and conservation of,
33-39, (illus.) 36; bourgaei 33

Danthonia decumbens 11

Darmera peltata (v.c. 98) 424

Daucus carota 382, subsp. carota 64

David, Richard William (‘Dick’) (1912-1993)
(Obit.) 320-321

Davies, P., with R. Gibbons — Field guide to wild
flowers of Britain and Northern Europe (Bk
Rev.) 307-308

} Davis, Peter Hadland (1918-1992) (Obit.) 167—

169

1 Davis, Peter, Recollections of, 168-169

it Dawson, H. J. & Ingrouille, M. J. — A biometric

survey of Limonium vulgare Miller and L.

humile Miller in the British Isles 239-254

§ De Langhe, J.-E., with J. Lambinon, L. Delvo-
salle & J. Duvigneaud — Nouvelle flore de la
Belgique, du nord de la France et des régions
voisines (Bk Rev.) 77-78

Delvosalle, L., with J. Lambinon, J.-E. De

457

Langhe & J. Duvigneaud — Nouvelle flore de
la Belgique, du nord de la France et des
régions voisines (Bk Rev.) 77-78

Dempsey, R. E., Gornall, R. J. & Bailey, J. P. —
Contributions to a cytological catalogue of
the British and Irish flora, 4 63-66

Deschampsia cespitosa 11, subsp. parviflora (v.c.
35) 300

Desmond, R., assisted by C. Ellwood — Diction-
ary of British and Irish botanists and horticul-
turists, including plant collectors, flower
painters and garden designers (Bk Rev.) 207

Devereau, Marjorie (1915-1994) (Obit.) 447

Deyeuxia neglecta 51

Dianthus armeria 170; prolifer 405

Diascia 309

Dicentra 309

Dickinson, N. M. — Rev. of The vegetation of
ultramafic (serpentine) soils (eds A. J. M.
Baker, J. Proctor & R. D. Reeves) 74-75

Digitalis 167; davisii 167

Digitaria ischmaeum (v.c. 3) 300

Diphasiastrum alpinum 164, (v.c. 83) 285; com-
planatum 164

Diphazium sibiricum 164

Dipsacus fullonum 64, (v.c. 91) 432

Doronicum plantagineum (v.c. 40) 433

Dowlen, C. M. & Ho, T. N. — Gentianella ciliata
(L.) Borkh. in Wiltshire (v.c. 8) 279

Draba aizoides 383; muralis (v.c. 79) 289

Dressler, R. L. — Phylogeny and classification of
the Orchid family (Bk Rev.) 76-77

Drimia maritima 304

Drosera intermedia 10; longifolia 12; rotundifolia
9, 12, 64

Druce in Northamptonshire.
Address, 1994 185-194

Dryopteris 446; aemula (v.c. 41, 73) 286; carthus-
tana 11; carthusiana x dilatata (v.c. 50) 286;
carthusiana x expansa 182; cristata 9, 11; x
deweveri 182, (v.c. 50) 286; oreades 78

Dudley-Smith, S. — Sonia Holland: in memoriam
173

Dudman, A. A. & Richards, A. J. — Seven new
species of Taraxacum Wigg. (Asteraceae),
native to the British Isles 119-132

Dunn, A. J. — The Flora of Ditchley: wild flowers
of an Oxfordshire estate (Bk Rev.) 158

Duvigneaud, J., with J. Lambinon, J.-E. De
Langhe & L. Delvosalle — Nouvelle flore de la
Belgique, du nord de la France et des régions .
voisines (Bk Rev.) 77-78

Presidential

Echinochloa crus-galli (v.c. 46) 300

Echium vulgare 64

Edees, Eric, contribution to British batology 317—
SS

Edees, Eric, Norfolk and Norfolk brambles 319-
320

458

Edees, Eric Smoothey (1907-1993) (Obit.) 317-
320

Editorial. A new direction for Watsonia 1

Edmondson, J. R. — Rev. of Exkursionsflora von
Osterreich (by W. Adler, K. Oswald & R.
Fischer, with M. A. Fischer (ed.)) 303-304

Edmondson, J. R. — Rev. of Nouvelle flore de la

Belgique, du nord de la France et des régions
voisines (by J. Lambinon, J.-E. De Langhe,
L. Delvosalle & J. Duvigneud) 77-78

Edmondson, J. R. — Rev. of The families and
genera of vascular plants (ed. K. Kubitzski).
Vol. 2: flowering plants, dicotyledons, Mag-
noliid, Hamamelid and Caryophyllid fami-
lies (eds K. Kubitzski, J. G. Rohweder & M.
Bittrich) 162-163

Edmondson, J. (ed.) with M. Palmer (comp.) —
Catalogue of the Brambles of Britain and
Ireland in the herbarium of Liverpool
Museum (LIV) (Bk Rev.) 442

Elatine alsinastrum 35; hexandra 35, (v.c. 96) 288,
(v.c. 42) 421

Eleocharis acicularis 12, 35; austriaca (v.c. 59)
298; multicaulis 11, (v.c. 35) 434; palustris 27,
35, 235, 312, 443, subsp. palustris 312, subsp.
vulgaris 313; quinqueflora 10; uniglumis 8

Eleogiton fluitans 11

Ellis, R. G., with A. E. Wade & Q. O. N. Kay -
Flora of Glamorgan (Bk Rev.) 445-446

Elodea callitrichoides 8; canadensis 8, 29; nuttallii
8, 13, 29, (v.c. 61, 73) 298, (v.c. 40) 433

Elymus caninus 64; repens subsp. arenosus 385,
386, var. maritimum 385

Elytrigia 386; atherica 387, 388; x oliveri (v.c. 3)
300; repens (L.) Desv. ex Nevski subsp.
arenosa (Spenner) A. Léve (Poaceae) in
north-western Europe 385-390, (map) 389,
subsp. repens 64, 385, 387, var. maritima 386;
repens x atherica (v.c. 3) 300

Empetrum nigrum 9, 11, 291, subsp. nigrum (v.c.
52) 423

Entyloma ficariae 48

Epilobium brunnescens (v.c. 12, 102) 429; cilia-
tum (v.c. 102) 428; komarovianum (v.c. 2,
98) 294; montanum x ciliatum 35, (v.c. 104)
294, (v.c. 47) 428; obscurum 35; roseum (v.c.
73) 294; tetragonum subsp. tetragonum 9

Epipactis helleborine 176, (v.c. 72) 301, 316;
leptochila 170; palustris 10, (v.c. 2) 301;
phyllanthes 316

Equisetum arvense 235; arvense x palustre (v.c.
101) 285; x dycei (v.c. 104) 285; fluviatile x
arvense (v.c. 24) 285; fluviatile x palustre (v.c.
104) 285; hyemale (v.c. 78) 285, (v.c. 2) 419;
x litorale 78, (v.c. 24) 285; palustre 235; x
rothmaleri (v.c. 101) 285; sylvaticum 315:
telmateia 164; variegatum 164

Eragrostis pectinacea 35

Eranthis hyemalis (v.c. 46) 420

INDEX TO WATSONIA VOLUME 20

Erica 91; ciliaris L. and E. x watsonii Benth. in
Dorset, 1963-1987. Changes in the dis-
tribution of, 89-95, (map) 90, 91; cinerea 10;
mackaiana 275, 276; mackaiana x tetralix
275; mackayi 276; x praegeri 277; ‘Stuartii’
276; stuartii 276; x stuartii: the authorship
reconsidered 275-278, ‘Charles Stuart’ 276,
‘Stuartil’ 276; tetralix 10, 64, 89, 91, 93, 94,
275, 276, subsp. stuartii 276, (illus.) 277; x
watsonii Benth. in Dorset, 1963-1987.
Changes in the distribution of Erica ciliaris
L. and, 89-95

Erigeron acer (v.c. 75) 433; annuus (v.c. 77) 433;
karvinskianus (v.c. 50) 298, (v.c. 42, 44) 433

Erinus alpinus (v.c. 77) 431

Eriophorum angustifolium 10, 182; vaginatum 10

Erodium maritimum 64; moschatum (v.c. 46) 429

Erophila glabrescens (v.c. 46, 73) 289, (v.c. 12,
38, 43, 52, 77, 98) 422-423; majuscula (v.c.
38) 422

Erucastrum gallicum (v.c. 49) 423

Eryngium maritimum 214

Escallonia 312; macrantha (v.c. 46) 289

Esfandiaria 440

Euonymus europaeus 64

Eupatorium cannabinum 12

Euphorbia esula x waldsteinii (v.c. 3) 294; lathyris
64; portlandica 64, 211, 214; x pseudovirgata
(v.c. 3) 294

Euphrasia 320, 446; anglica 170; anglica x nemor-
osa (v.c. 44) 296; arctica 11; arctica subsp.
borealis x confusa (v.c. 44, 46) 296; arctica
subsp. borealis x scottica (v.c. 44) 296; con-
fusa x tetraquetra (v.c. 2) 296; foulaensis 392;
heslop-harrisonii (v.c. 104) 296; nemorosa x
confusa (v.c. 25, 47) 431; rostkoviana subsp.
rostkoviana x confusa (v.c. 44) 296

Evans, P. R., with A. T. Jones— A comparison of
the growth and morphology of native and

commercially obtained continental Euro- |

pean Crataegus monogyna Jacq. (Hawthorn)
at an upland site 97-103

Falcaria vulgaris (v.c. 14) 429
Fallopia x bohemica (v.c. 3, 69) 287, (v.c. 38) 421;

convolvulus (v.c. 102) 421; japonica x sachal- |

inensis (v.c. 3, 69) 287, (v.c. 38) 421
Festuca arenaria 412; arundinacea 8; brevipila
(v.c. 12) 435; heterophylla (v.c. 78) 434;

huonii (v.c. 3) 299; longifolia (v.c. 3) 435; .
ovina subsp. ophioliticola (v.c. 77) 434; rubra ©
235, 382, 388, L. new to England. Two «
subspecies of, 412, subsp. arctica 412, subsp. ~
commutata (v.c. 35) 434, subsp. juncea 392, —

394, 412, subsp. megastachys (v.c. 25) 299,
subsp. scotica 412; rubra x Vulpia fasciculata
(v.c. 69) 435; vivipara (v.c. 83) 299

X Festulpia hubbardii (v.c. 69) 435

Ficaria bulbifera 43, 46; calthifolia 43, 45, subsp. |

a

-- ———

INDEX TO WATSONIA VOLUME 20

grandiflora 46; calthifolius 43; grandiflora 43,
44, 46; intermedia 43, 45; nudicaulis 45;
pumila 43, 45; ranunculoides 43, 45, subsp.
grandiflora 46, var. divergens 43, 45; step-
porum 47; transsilvanica 43, 45; verna 43, 45,
subsp. bulbifera 43, 46, subsp. calthifolia 45,
subsp. grandiflora 46, 47, subsp. pumila 45,
var. aurea 43

Field, M. H. — The status of Bupleurum falcatum
L. (Apiaceae) in the British flora 115-117

Filago arvensis 157; minima (v.c. 46) 298

Filipendula ulmaria 12, 27, 235

Fischer, M. A. (ed.), with W. Adler, K. Oswald
& R. Fischer — Exkursionsflora von Oster-
reich (Bk Rev.) 303-304

Fischer, R., with W. Alder, K. Oswald & M. A.
Fischer (ed.) — Exkursionsflora von Oster-
reich (Bk Rev.) 303-304

Foeniculum vulgare (v.c. 73) 429

Forsythia x intermedia (v.c. 35) 296; suspensa x
viridissima (v.c. 35) 296

Fowler, B. R. — Obit. of Eric Smoothey Edees
(1907-1993) 317

Frangula alnus 10, 311

Frankenia laevis (v.c. 3) 288

Fraxinus excelsior 382

Fritillaria meleagris (v.c. 58) 301

Fryer, J. & Hylmo, B. — The native British
Cotoneaster — Great Orme Berry — renamed
61-63

Fucus 214

Fumaria 201, 320; bastardii (v.c. 79) 286; capreo-
lata (v.c. 81) 420; densiflora (v.c. 61) 286;
muralis 214; purpurea (v.c. 100) 420

Gagea durieui 305; iberica 305; lutea 64, 173;
nevadensis 305

Galeopsis angustifolia (v.c. 40) 430; bifida (v.c.
102) 430; segetum 11

Galium mollugo (v.c. 79) 297, (v.c. 78) 431,
subsp. erectum (v.c. 50) 297; palustre 235,
subsp. elongatum (v.c. 46) 431; saxatile 264,
sterneri (v.c. 81) 297; uliginosum 10; verum
214

Gaudinia fragilis (v.c. 11) 300, 388

Gaultheria mucronata (v.c. 58) 423; shallon (v.c.
46) 423

Genista anglica 9, 11; monspessulana (v.c. 5) 294;
tinctoria 9

Gentiana pneumonanthe 11; verna 166

Gentianella 279; amarella subsp. amarella (v.c. 1)
294; campestris (v.c. 50) 294; ciliata 160, (L.)
Borkh. in Wiltshire (v.c. 8) 279

Gentianopsis 279; ciliata 279

Geranium 214; endressii x veriscolor (v.c. 77) 294;
himalayense (v.c. 77) 429; lucidum 64; molle
214; x oxonianum (v.c. 77) 294; pratense 185;
psilostemon (v.c. 77) 429

459

Geum macrophyllum (v.c. 46) 292; rivale 11;
urbanum (v.c. 102) 426

Gibbons, R. & Davies, P. — Field guide to wild
flowers of Britain and Northern Europe (Bk
Rev.) 307-308

Gibbs, P. E. — Rev. of Algarve plants and land-
scape (by D. J. Mabberley & P. J. Placito)
163-164

Gillam, B. (ed.) — The Wiltshire Flora (Bk Rev.)
158-160

Gladiolus communis subsp. byzantinus (v.c. 17)
436

Glaucium flavum 64

Glaux maritima 388

Glyceria 443; declinata 8, 35, 235; distans 391, var.
prostrata 391, 392; fluitans 34, 35, 234, 235;
maxima 12, 15, (v.c. 81) 299

Gnaphalium luteo-album 64; uliginosum 11, 35,
234, 235

Good, Ronald d’Oyley (1896-1992) (Obit.) 169-
TA

Goodyera repens (v.c. 81) 301

Gornall, R. J., with R. E. Dempsey & J. P. Bailey
— Contributions to a cytological catalogue of
the British and Irish flora, 4 63-66

Gornall, R. J., with C. A. Stace, J. Squirrel &
Y. Shi— Chromosome numbers in Hieracium
L. section Alpina (Fries) F. N. Williams 367—
377

Graham, G. G. & Primavesi, A. L. — Roses of
Great Britain and Ireland: BSBI handbook
No. 7 (Bk Rev.) 160-161

Greene, Dorothy Mary (1928-1994) (Obit.) 447—
449

Greenwood, B. D. — Rev. of Flora of Flintshire
(by G. Wynne) 315-316

Grey-Wilson, C., with M. Blamey — Mediterra-
nean wild flowers (Bk Rev.) 157

Griffiths, M. — Index of garden plants (Bk Rev.)
308-309

Groenlandia 326; densa 10, (v.c. 104) 298

Gunnera 420; tinctoria (v.c. 75, 99) 294

Gymnadenia conopsea 11, subsp. borealis (v.c.
46) 437, subsp. conopsea 11, subsp. densi-
flora 11, (v.c. 46) 437; conopsea x Dactylo-
rhiza fuchsii (v.c. 104) 301

Gymnocarpium robertianum (v.c. 46) 286

Hackney, P. — Rev. of Red data books of Britain
and Ireland: Stoneworts (by N. F. Stewart &
J. M. Church) 80-81

Halliday, G. — Two subspecies of Festuca rubra L.
new to England 412

Haloxylon 440

Hammada 440

Hammarbya paludosa 11, (v.c. 2) 436

Hampton, M. & Kay, Q. O. N. — Sorbus domes-
tica L., new to Wales and the British Isles
379-384

460

Harrington, A. J.— Obit. of Peter James Wanstall
(1924-1993) 451-452

Harrison, P. — Recollections of Peter Davis 168—
169

Hartzell, H. Jr — The yew tree. A thousand
whispers. Biography of a species (Bk Rev.)
75-76

Hebe salicifolia (v.c. 50) 296, (v.c. 73) 431

Hedera colchica (v.c. 77, 98) 429, var. dentata
429; helix 314, 381, 382

Hedge, I. C. — Rev. of Vascular plant families and
genera. A listing of the genera of vascular
plants of the world according to their families,
as recognised in the Kew Herbarium, with an
analysis of relationships of the flowering plant
families according to eight systems of classifi-
cation (by R. K. Brummitt) 439-440 |

Hejny, S. & Slavik, B. (eds) — Kvétena Ceske
Republiky (Flora of the Czech Republic).
Vol. 1. Huperziaceae — Urticaceae (Bk Rev.)
Te

Helianthemum nummularium 382; salicifolium
304

Helichrysum italicum 309

Henderson, D. M. — Rev. of An annotated check-
list of the flowering plants and ferns of Main
Argyll (by G. Rothero & B. Thomson) 444

Heracleum mantegazzianum (v.c. 50) 429; sphon-
dylium subsp. sphondylium 64

Herminium monorchis 172

Hermodactylus tuberosus (v.c. 6) 301

Heywood, V. H. — Obit. of Peter Hadland Davis
(1918-1992) 167-168

Hieracium 282, 285, 323, 351, 367-369, 376, 419,
446; of the section Alpestria (Fries) F. N.
Williams. A new Shetland, 282-284; L.
section Alpina (Fries) F. N. Williams. Chro-
mosome numbers in, 367-377, Eleven new
British species of, 351-365; subgen. Hiera-
clum 367; subgen. Pilosella 367; alpinum
367-370, 373, 374, 376, 377; attenuatifolium
282, 283; australius 283; backhousei 352, 370;
caledonicum (v.c. 91) 298, (v.c. 94) 432;
calenduliflorum 356, 367, 369, 370, 374-377;
calvum P. D. Sell & D. J. Tennant, sp. nov.
351, 352, 370; celsum 375; completum P. D.
Sell & C. West, sp. nov. 351, 353, 354, 370;
curvatum 354; dicella (v.c. 75) 298; duriceps
(v.c. 80) 297; eximium 354, 356, 358, 367,
369,370. 375,376) S79: fereximiune 354.
forma tenellum (Backh.) P. D. Sell & C.
West, stat. nov. 351, 354, 370, var. tenellum
354; eximium/calenduliflorum intermediate
370, 375; eximium/memorabile intermediate
371, 375; exotericum (v.c. 91) 297; globosi-
florum (v.c. 104) 298, 352, 363, 367, 371;
graniticola 352, 371, 376; grovesii 359, 371,

376; hanburyi 352, 354, 361, 368, 369, 371;

375, 376, forma atraticeps (Pugsley) P. D.

INDEX TO WATSONIA VOLUME 20

Sell & D. J. Tennant, stat. nov. 351, 354, 371,
f. hanburyi 354, forma pusillum P. D. Sell &
D. J. Tennant, forma nov. 351, 354, 371, var.
atraticeps 354, var. humile 354; hanburyi/
tenuifrons intermediate 371, 374, 375; holo-
sericeum (v.c. 104) 298, 367, 371, 375, 376;
insigne 354, 371, forma celsum P. D. Sell &
D. J. Tennant, forma nov. 351, 354, 367, 371,
375, f. insigne 354; kennethii P. D. Sell &
D. J. Tennant, sp. nov. 351, 355, 360, 372;
larigense 352, 372, 375; leptodon P. D. Sell &
D. J. Tennant, sp. nov. 351, 356, 358, 369,
372, 375; leyi 198; macrocarpum 367, 369,
372-376; marginatum 357, 372, forma chae-
tocephalum P. D. Sell & C. West, forma nov.
351, 357, 372, f. marginatum 357; memora-
bile 369, 372; milesii P. D. Sell & C. West, sp.
nov. 351, 352, 357, 372; mundum P. S. Sell &
C. West, sp. nov. 351, 355, 358, 360, 372;
murorum 376; nigrescens 368, 369, 376; nota-
bile 356, 372; optimum P. D. Sell & C. West,
sp. nov. 351, 359, 369; pensum P. D. Sell &
C. West, sp. nov. 351, 355, 360, 372; persci-
tum P. D. Sell & C. West, sp. nov. 351, 354,
355, 358, 360, 361, 373; probum P. D. Sell &
C. West, sp. nov. 351, 353, 362, 373; pseudo-
curvatum 354, 373; pseudopetiolatum 373,
375; pulmonarioides 376; salticola (v.c. 99)
297; severiceps (v.c. 98) 432; spenceanum W.
Scott & R. C. Palmer, sp. nov. 282, 283;
subglobosum P. D. Sell & C. West, sp. nov.
351, 355, 358, 360, 363, 373; subgracilentipes
367, 373; subhirtum (v.c. 75) 297; subumbel-
latiforme (v.c. 91) 297; tenuifrons 357, 367,
369, 373-376; tenuifrons/memorabile inter-
mediate 373, 375; umbellatum 368, 376,
subsp. bichlorophyllum 368, subsp. umbella-
tum 368; vagum (v.c. 75) 297; vennicontium
(v.c. 104) 298; villosum 376

Hill, M. O., Preston, C. D. & Smith, A. J. E.
(eds) — Atlas of the bryophytes of Britain and
Ireland. Volume 1: Liverworts (Hepaticae
and Anthocerotae) (Bk Rev.) 161-162

Hill, M. O., Preston, C. D. & Smith, A. J. E.
(eds) — Atlas of the bryophytes of Britain and
Ireland. Volume 2: Mosses (except Diplolep-
ideae) (Bk Rev.) 161-162

Hill, M., with J. Merryweather — The fern guide.
A field guide to the ferns, clubmosses, quill-
worts and horsetails of the British Isles (Bk
Rev.) 78-79

Himantoglossum hircinum 170

Hippophae rhamnoides 64, (v.c. 77) 294, 388

Hippuris vulgaris 10, 53

Ho, T. N., with C. M. Dowlen — Gentianella
ciliata (L.) Borkh. in Wiltshire (v.c. 8) 279

Holcus lanatus 234

Holland, Sonia C. (1912-1993) (Obit.) 171-173

Holland, Sonia: in memoriam 173

INDEX TO WATSONIA VOLUME 20 461

Hollick, Kathleen Margaret (1913-1993) (Obit.)
322

Holodiscus discolor (v.c. 73) 424

Holyoak, D. T. — Rumex frutescens Thouars x R.
obtusifolius L. (Polygonaceae), a previously
undescribed hybrid dock, and new records of
R. x wrightii Lousley in West Cornwall (v.c.
1) 412-415

Honkenya peploides 64, 211, 214, 223

Hordeum jubatum (v.c. 75) 435; marinum 11, 64;
murinum subsp. murinum 64; vulgare 99

Hottonia palustris 9, 10, 53, (v.c. 77) 423

Humulus lupulus 64

Hunnybun, Edward Walter (1848-1918) 196-197

Huperzia selago 11

Hyacinthoides non-scripta 64, 154, 155; non-
scripta x hispanica (v.c. 49) 436

Hydrocharis morsus-ranae 10

Hydrocotyle vulgaris 10

Hylm6, B, with J. Fryer. — The native British
Cotoneaster — Great Orme Berry — renamed
61-63

Hypericum calycinum (v.c. 77) 421; x desetangsii
(v.c. 98) 422, nothosubsp. desetangsii (v.c.
24, 44) 288, nothovar. carinthiacum 422;
elodes 9, 12; hirsutum (v.c. 2) 422; humifu-
sum 35; maculatum 11; perforatum x macula-
tum subsp. obtusiusculum (v.c. 24, 44) 288;
perforatum x maculatum (v.c. 98) 422; tetrap-
terum 12

Iberis amara 64

Ilex x altaclerensis 309; aquifolium 443

Impatiens capensis (v.c. 40) 429; glandulifera 8,
13, 236; noli-tangere (v.c. 46) 294; parviflora
(v.c. 93) 294, (v.c. 91) 429

Ingram, G. I. C., with D. E. Wilson — Dis-
tribution of Lady Orchid (Orchis purpurea
Hudson) in an East Kent wood 154-156

Ingrouille, M. J., with H. J. Dawson — A bio-
metric survey of Limonium vulgare Miller
and L. humile Miller in the British Isles 239-
254

Inula conyzae 382

Tris foetidissima 382, (v.c. 73) 436; pseudacorus
1S 3

Isoetes 78; echinospora 78; lacustris 78

Isolepis cernua 312; setacea 11, 312

Jenkinson, M. N. — A new variety of Narrow-
leaved Marsh-orchid in ee Hampshire
(v.c. 11) 263-273

Jones, A. T. & Evans, P. R. — A comparison of
the growth and morphology of native and
commercially obtained continental Euro-
pean Crataegus monogyna Jacq. (Hawthorn)
at an upland site 97-103

Juncus 214; acutiflorus 12; ambiguus, 12, (v.c. 81)
434; articulatus 34, 35, 235; articulatus x

acutiflorus (v.c. 79) 434; bufonius 12, 235,
392; bulbosus 10, 35; compressus (v.c. 41)
434; conglomeratus 12; x diffusus 12, (v.c.
46) 434; effusus 9, 10, 35, 153, 154, 234, 235,
f. spiralis 153, var. spiralis J. McNab in the
Inner Hebrides 153-154, var. suberectus 153;
effusus x inflexus (v.c. 46) 434; foliosus (v.c.
73, 86) 298; gerardii 11, 394; kochii 235;
maritimus 299; squarrosus 10, 12; subnodulo-
sus 11, (v.c. 68) 434; x surrejanus (v.c. 79)
434; tenageia, cf. 35; tenuis (v.c. 79) 298

Kay, Q. O. N., with M. Hampton — Sorbus
domestica L., new to Wales and the British
Isles 379-384

Kay, QO. O.N., with A. E. Wade & R. G. Ellis —
Flora of Glamorgan (Bk Rev.) 445-446

Kent, D. H. — Rev. of Dictionary of British and
Trish botanists and horticulturists, including
plant collectors, flower painters and garden
designers (by R. Desmond with C. Ellwood)
307

Knees, S. G. — Rev. of The genera of the Umbelli-
ferae (by M. G. Pimenov & M. V. Leonov)
157-158

Kniphofia praecox (v.c. 49) 436

Koeleria macrantha 223, (v.c. 79) 300; vallesiana
383

Kohlrauschia 405; prolifera 405

Kraft, T. & Nybom, H.— DNA fingerprinting and
biometry can solve some _ taxonomic
problems in apomictic blackberries (Rubus
subgen. Rubus) 329-343

Kubitzski, K. (ed.) — The families and genera of
vascular plants (Bk Rev.) 162-163

Kubitzski, K., Rohweder, J. G. & Bittrich, M.
(eds) — The families and genera of vascular
plants. Vol. 2: flowering plants, dicotyle-
dons, Magnoliid, Hamamelid and Caryo-
phyllid families (Bk Rev.) 162-163

Laburnum anagyroides (v.c. 49) 428

Lactuca saligna 11; serriola (v.c. 43, 51) 297;
virosa (v.c. 5) 297

Lagurus ovatus 210, 214

Lambinon, J., De Langhe, J.-E., Delvosalle, L.
& Duvigneaud, J. — Nouvelle flore de la
Belgique, du nord de la France et des régions
voisines (Bk Rev.) 77-78

Lamiastrum galeobdolon subsp. argentatum (v.c.
25) 295, (v.c. 57, 80) 430

Lamium amplexicaule (v.c. 102) 430; hybridum
(v.c. 100, 108) 430; purpureum 157

Lane, A. M., with iS J. Cherrill — Bracken
(Pteridium aquilinum (L.) Kuhn) infestation
of rough grazing land in the catchment of the
River Tyne, northern England 105-114

Lantana montevidensis 102

Larix kaempferi (v.c. 77) 419

462 INDEX TO WATSONIA VOLUME 20

Lathraea clandestina (v.c. 48) 296, (v.c. 40) 431

Lathyrus aphaca 64; grandiflorus (v.c. 73) 293,
(v.c. 12) 428; japonicus (v.c. 11, 91) 293, 388;
nissolia (v.c. 44) 293, (v.c. 40) 428; palustris
10

Lavatera arborea (v.c. 50, 51) 288; cretica (v.c. 3)
288; olbia (v.c. 35) 422

Lavin, J. C. & Wilmore, G. T. D. (eds) — The
West Yorkshire plant atlas (Bk Rev.) 441

Leersia oryzoides 8

Lemna gibba 8, 13; minor 35, 235, (v.c. 108) 298,
434; minuta (v.c. 38) 298, (v.c. 40, 42) 434;
trisulca (v.c. 78) 434

Leonov, M. V., with M. G. Pimenov — The genera
of the Umbelliferae (Bk Rev.) 157-158

Leontodon saxatilis (v.c. 81) 432

Lepidium heterophyllum (v.c. 26) 423; latifolium
11, (v.c. 40) 423; ruderale (v.c. 26, 40) 423

Leucanthemum lacustre x maximum (v.c. 26) 433;
x superbum (v.c. 26) 433; vulgare 64

Leucojum aestivum 8, (v.c. 79) 436

Leycesteria formosa (v.c. 50) 432

Leymus arenarius (v.c. 104) 300, 394

Ligustrum vulgare 382

Limonium 162, 240-244, 246, 248, 252; binervo-
sum 369; britannicum subsp. celticum var.
celticum 65, subsp. celticum var. pharense 65;
humile Miller in the British Isles. A bio-
metric survey of Limonium vulgare Miller
and, 239-254; vulgare Miller and L. humile
Miller in the British Isles. A biometric survey
of, 239-254

Limosella aquatica 11, 35, (v.c. 73) 431

Linaria vulgaris 65

Lindernia dubia 35

Linum usitatissimum (v.c. 50) 429

Listera ovata 265

Lithospermum officinale (v.c. 46) 295; purpuro-
caeruleum 382

Littorella uniflora 12, 417

Livermore, Leonard Albert (1919-1994) (Obit.)
449-451

Lobelia dortmanna 65; urens 170

Lonicera involucrata (v.c. 77) 432; japonica (v.c.
12) 432; periclymenum 65; pileata (v.c. 77)
432; xylosteum (v.c. 75) 297, (v.c. 77) 432

Lopez, G. — Rev. of Illustrated Flora of Mallorca
(by E. Beckett) 304-305

Lophocolea bidentata 161

Lotus 439; corniculatus 65, 99; glaber 10; pedun-
culatus 12

Lusby, P. — Rev. of Scarce plants in Britain (eds
A. L. Stewart, D. A. Pearman & C. D.
Preston) 444-445

Luzula campestris 443; pallidula 12; sylvatica 65

Lychnis coronaria (v.c. 46) 287, (v.c. 12, 46, fe)
421; flos-cuculi 9, 10

Lycopodiella inundata 11, (v.c. 70) 419

Lycopodium 164; annotinum (v.c. 108) 419; clav-

atum 11, (v.c. 58) 419

Lycopus europaeus 35, (v.c. 104) 295

Lysimachia nummularia 235; thyrsiflora 11; vul-
garis 11, 53

Lythrum borysthenicum 34, 35; hyssopifolia 35,
(v.c. 35) 428; portula 35, 235; salicaria 9, 11,
532235

Mabberley, D. J. & Placito, P. J. — Algarve plants
and landscape (Bk Rev.) 163-164

Malcolmia maritima (v.c. 49) 289

Malus sylvestris 443

Malva moschata 65; pusilla (v.c. 4) 288, (v.c. 40)
422

Marner S. K., with Q. C. B. Cronk & A. L.
Primavesi — Lectotypification of Rosa roths-
childii Druce (Rosaceae) 409-410

Matricaria discoidea 235, 394; recutita (v.c. 108)
433

Matthiola incana 65, (v.c. 46) 289

McClintock, D. & Burns, A. — Obit. of Kathleen ©
Margaret Hollick (1913-1993) 322

McMillan, N.F. — Rev. of Supplement to Flora of
Cheshire (by A. Newton) 79

Meconopsis 324

Medicago arabica (v.c. 83) 293; lupulina 65

Meikle, R. D. — Obit. of Sir George Taylor (1904—
1993) 324-325

Melica nutans 65

Mentha 78; aquatica x spicata (v.c. 102) 430;
X piperita (v.c. 102) 430; pulegium 10, 35;
x smithiana 12; spicata (v.c. 102) 431; spicata
x longifolia (v.c. 3, 81) 431; spicata x suaveo-
lens (v.c. 24) 295, (v.c. 47) 431; suaveolens
(v.c. 83) 295; x verticillata 11; x villosa (v.c.
24) 295, (v.c. 47) 431, var. alopecuroides 295;
x villosonervata (v.c. 3, 81) 431

Menyanthes trifoliata 10

Merryweather, J. & Hill, M. — The fern guide. A
field guide to the ferns, clubmosses, quillworts
and horsetails of the British Isles (Bk Rev.)
78-79

Messenger, Kenneth Guy (1920-1993) (Obit.)
322-323

Mibora minima 210, 211, 214, (v.c. 9) 300

Microbiota 73

Milium effusum (v.c. 81) 299, (v.c. 79) 434

Mill, R. R. — Rev. of Origin and geography of
cultivated plants (by N. I. Vavilov, trans. D.
Léve) 82-83

Milne- Redhead, E. — Obit. of Sonia C. Holland
(1912-1993) 171-173

Mimulus 13; guttatus 8; moschatus 11

Minuartia 418; verna (v.c. 41) 287, 418

Moenchia erecta (v.c. 44) 287, (v.c. 46) 421

Molinia caerulea 10, 70, 182, 296, 436, subsp.
arundinacea (v.c. 44) 300, (v.c. 44) 435

Montia fontana 10, 182

Morin, N. R. — Rev. of Index Kewensis on CD-

INDEX TO WATSONIA VOLUME 20

ROM (Royal Botanic Gardens, Kew) 310—
311

Moss, Charles Edward (1870-1930) 195-196

Mountford, J. O. — Floristic change in English
grazing marshes: the impact of 150 years of
drainage and land-use change 3-24

Mountford, J. O. — Problems in reconstructing
floristic change: interpreting the sources for
English grazing marshes 25-31

Muscari armeniacum (v.c. 83) 301, (v.c. 49) 436;
neglectum (v.c. 83) 301

Myosotis laxa 235; ramosissima 210, 211, 214,
22Ga@y-e> 73) 295; scorpioides 12,28), 235;
scorpioides x laxa (v.c. 46) 295; secunda 12,
28; x suzae (v.c. 46) 295

Myosoton aquaticum (v.c. 49) 421

Myosurus minimus 8, (v.c. 3) 286

Myrica gale 9, 10

Myriophyllum alterniflorum 10, 35, 417; aquati-
cum (v.c. 46) 294, (v.c. 35, 40) 428; spicatum
8, 13, 53, 417; verticillatum 10, 53

Narcissus poeticus subsp. poeticus (v.c. 49) 301;
pseudonarcissus 10, subsp. obvallaris (v.c.
46) 436

Nardus stricta 11

Narthecium ossifragum 9, 10, 65

Nasturtium palustre 232

Nelson, E. C. — Erica x stuartii: the authorship
reconsidered 275-278

Nelson, E. C. — Rev. of The concise Oxford
dictionary of botany (ed. M. Allaby) 73

Nelson, E. C. — Rev. of The eternal yew (by T.
Baxter) 75-76

Nelson, E. C. — Rev. of The yew tree. A thousand

whispers. Biography of a species (by H.

| Hartzell Jr) 75-76

' Nelson, E. C. — Rev. of Wild and garden plants

| (by S. M. Walters) 313-314

Nelson, E. C. & Walsh, W. F. — Trees of Ireland,

| native and naturalised (Bk Rev.) 311-312

Nepeta x faassenii (v.c. 46) 295; racemosa x
nepetella (v.c. 46) 295

Newton, A. — Eric Edees’ contribution to British
batology 317-319

Newton, A. — Supplement to Flora of Cheshire
(Bk Rev.) 79

Newton, A. & Porter, M. — New species of Rubus
L. (Rosaceae) from Wales and the Welsh
Marches 133-141

| Newton, M. E., with T. G. Tutin et al. (eds) & J.

: R. Akeroyd — Flora Europaea, 2nd edition,

vol. 1: Psilotaceae to Platanaceae (Bk Rev.)

81-82

_ Nitella 17; mucronata 8; translucens 12

_ Notes 61-71, 153-156, 279-284, 405-418

Nuphar lutea 10, 43, 260

_ Nybom, H., with T. Kraft - DNA fingerprinting

j and biometry can solve some taxonomic

|

463

problems in apomictic blackberries (Rubus
subgen. Rubus) 329-343

Nymphaea alba 10, 53

Nymphoides peltata 8, 13, (v.c. 68) 295

Obituaries 167-180, 317-326, 447-452

Oenanthe aquatica 35; crocata 12, 235, 264; fistu-
losa 53; lachenalii 11; silaifolia 12

Oenothera glazioviana x biennis (v.c. 46) 429;
stricta 214

@Ollgaard, B. — Scandinavian ferns (Bk Rev.) 164

Ononis repens 382, 388; spinosa 11

Onopordum acanthium 65

Ophioglossum vulgatum 10, 63, 65, 182, (v.c. 73)
285

Ophrys apifera 165

Orchis militaris 190; morio 9, 10, 182, 264, 265,
314; purpurea Hudson) in an East Kent
wood. Distribution of Lady Orchid(, 154—
156; ustulata 11, (v.c. 41) 301

Origanum vulgare 382

Ornithopus perpusillus 65, 182

Orobanche crenata (v.c. 18) 431; rapum-genistae
(v.c. 4) 296

Osmunda crispa 418; regalis 9, 10, 314

Oswald, K., with W. Adler & R. Fischer & M.A.
Fischer (ed.) — Exkursionsflora von Oster-
reich (Bk Rev.) 303-304

Oxalis debilis (v.c. 49) 294; stricta (v.c. 50) 294

Palmer, M. (comp.), with J. Edmondson (ed.) —
Catalogue of the Brambles of Britain and
Ireland in the herbarium of Liverpool
Museum (LIV) (Bk Rev.) 442

Palmer, R. C. & Scott, W. — A forgotten Shetland
dandelion 279-281

Palmer, R. C., with W. Scott — A new Shetland
Hieracium of the section Alpestria (Fries)
F. N. Williams 282-284

Papaver atlanticum (v.c. 58) 420; dubium subsp.
lecogqii (v.c. 46) 286; hybridum (v.c. 5) 286;
orientale (v.c. 58) 420

Parapholis strigosa 65

Parentucellia viscosa (v.c. 5, 64) 296, (v.c. 46) 431

Parietaria judaica 382

Parnassia palustris 9, 12, var. condensata 78

Parnell, J. — Variation and hybridisation of Alnus
Miller in Ireland 67—70

Paspalum distichum 34, 35

Pearman, D. A. — Sedges and their allies in
Dorset. An atlas of the distribution of the .
family Cyperaceae in Dorset (Bk Rev.) 312-
313

Pearman, D. A. — Rev. of Flora of the Christ-
church area (by F. Woodhead) 314-315
Pearman, D. A., with A. L. Stewart & C. D.

Preston (eds) — Scarce plants in Britain (Bk
Rev.) 444-445
Pedicularis palustris 10; sylvatica 10, 182, L.

464

subsp. hibernica D. A. Webb (Scrophularia-
ceae) new to Wales 70-71, (v.c. 41, 46) 296,
subsp. sylvatica 70, 71, 296

Perring, F. — Presidential Address, 1994. Druce in
Northamptonshire 185-194

Perring, F. H. — Rev. of The West Yorkshire plant
atlas (eds J. C. Lavin & G. T. D. Wilmore)
44]

Persicaria amphibia 235; amplexicaulis (v.c. 77)
287; bistorta 9; campanulata (v.c. 12) 421;
hydropiper 12, 35, 234, 235; lapathifolia 65;
laxiflora 10; maculosa 34, 35, 235; minor 10;
vivipara (v.c. 73) 287

Petasites alba (v.c. 46) 298, (v.c. 50) 433; hybridus
11

Petrorhagia sect. Kohlrauschia 405; nanteuilii
405, 406; prolifera (L.) P. W. Ball & Hey-
wood (Caryophyllaceae), an overlooked
native species in eastern England 405-407;
velutina 405

Petroselinum segetum 8

Peucedanum 158; officinale 158; ostruthium 11;
palustre 10

Phacelia tanacetifolia (v.c. 50) 430

Phalaris arundinacea 65, 235, 260; paradoxa (v.c.
14) 300

Philadelphus coronaria (v.c. 38) 423; coronaria x
microphyllus x pubescens (v.c. 77) 424; x
virginalis (v.c. 77) 424

Phleum arenarium 210, 211, 214, 223, 388;
phleoides 65

Phragmites australis 8, 52, 53, 260, 286, 312

Picris echioides (v.c. 98) 297

Pieris 309; japonica ‘Pygmaea’ 309

Pilosella 367-369, 376; caespitosa subsp. colli-
niformis (v.c. 77) 297; officinarum 382

Pimenov, M. G. & Leonov, M. V. — The genera of
the Umbelliferae (Bk Rev.) 157-158

Pimpinella saxifraga 382

Pinguicula lusitanica 11; vulgaris 12

Pinus 174; pinaster 214; radiata 311

Placito, P. J., with D. J. Mabberley — Algarve
plants and landscape (Bk Rev.) 163-164

Plant Records 285-301, 419-437

Plantago coronopus 392, 394; lanceolata 382;
major 35, 235, subsp. intermedia (v.c. 25, 46)
295; maritima 214, 392, 394; media (v.c. 77)
431

Platanthera bifolia 10, (v.c. 79) 301

Poa angustifolia (v.c. 81) 299; annua 235; infirma
(v.c. 9) 299; nemoralis (v.c. 102) 435; retrof-
lexa x borealis 392; trivialis 234

Podosphaera clandestina 98

Polygonum arenastrum (v.c. 102) 421; aviculare
174; boreale (v.c. 73) 421; oxyspermum
subsp. rail (v.c. 102) 421; ; rurivagum “ c. 68)
287

Polypodium 78; cambricum (v.c. 98) ‘419:. Xx
mantoniae (v.c. 28) 285; vulgare 314; vulgare

INDEX TO WATSONIA VOLUME 20

x interjectum (v.c. 28) 285

Polypogon monspeliensis 65, (v.c. 12) 435

Polystichum 164; aculeatum (v.c. 2) 419; x bick-
nellii (v.c. 46) 419; setiferum (v.c. 91) 419;
setiferum x aculeatum (v.c. 46) 419

Polytrichum formosum 451; juniperinum 451

Ponsonby, L. — Marianne North at Kew Gardens ©
(Bk Rev.) 79-80

Pontederia cordata (v.c. 59) 301

Populus alba 8, 13; alba x tremula (v.c. 77) 422;
x canadensis 311; x canescens (v.c. 77) 422;
nigra 172, subsp. betulifolia (v.c. 46) 422;
tremula var. glabra 198; tremuloides 341;
trichocarpa (v.c. 58, 77) 422

Porter, M., with A. Newton — New species of
Rubus L. (Rosaceae) from Wales and the
Welsh Marches 133-141 |

Potamogeton 260, 316, 324, Sir George Taylor’s
studies of the genus, 325-326, 417, 443;
alpinus 10, 12, 53, 325; berchtoldii 8, 13, 28; x
cognatus 326, Asch. & Graebn. at Loch
Borralie, West Sutherland (v.c. 108), Scot-
land 415-417; coloratus 8; compressus 11;
crispus 35; drucei 255; filiformis 236, 417;
filiformis x pectinatus 326; x fluitans 255, 257-
259; x franconicus 28; friesii 12, 53; gramin-
eus 10, 417; x griffithii 325; x lanceolatifolius
326; x lintonii 11; lucens 10, 53, 255; lucens x
natans 261; macvicarii 325; natans 9, 35, 53,
255-261, 417; natans L. x P. nodosus Poir.)
in Dorset, new to the British Isles. Potamo-
geton x Schreberi G. Fisch. (, 255-262; natans
x gramineus 255, 257; natans x lucens 255,
257; x nitens 417; nodosus 255-261; noltei
261; obtusifolius (v.c. 46) 298; x olivaceus
325; panormitanus 28; pectinatus 8, 13, 28,
260, 326, 417; perfoliatus 10, 17, 416, 417;
perfoliatus x praelongus 415; polygonifolius
10, 326; praelongus 12, (v.c. 29) 298, 416,
417; praelongus x cognatus 415; pusillus 8,
17, 28, 325, var. pseudotrichoides 28, var.
tenuissimus 28; x salicifolius 11; x schreberi
G. Fisch. (P. natans L. x P. nodosus Poir.) in
Dorset, new to the British Isles 255-262;
sparganiifolius 259; x sparganiifolius 11, 255,
257, 258; x sudermanicus 325; x suecicus 326;
trichoides 9, 28, (v.c. 3, 61) 298

Potentilla anglica (v.c. 93) 292; anglica x reptans
(v.c. 46) 292; anserina 235, 394; argentea 65;
crantzii subsp. serpentini 303; erecta 264,
subsp. erecta 291, subsp. strictissima (v.c. 3,
4,5, 46) 291; x mixta (v.c. 46) 292; norvesiaa
(v. c. 80) 291; palustris 10, 65 3

Prance, G. T. — Rev. of Flora Europaea, 2nd
dino: vol. 1: Psilotaceae to Platanaceae
(ed. T. G. Tutin et al., assisted by J. R.
Ackeroyd & M. E. Newton) 81-82

Presidential Address, 1994 185-194

Press, J. R. — Rev. of Field guide to trees of

INDEX TO WATSONIA VOLUME 20

Britain, Europe and North America (by
A. Cleave) 307-308

Press, J. R.— Rev. of Field guide to wild flowers of
Britain and Northern Europe (by R. Gibbons
& P. Davies) 307-308

Preston, C. D. — Juncus effusus var. spiralis
J. McNab in the Inner Hebrides 153-154

Preston, C. D. — Potamogeton x cognatus Asch. &
Graebn. at Loch Borralie, West Sutherland
(v.c. 108), Scotland 415-417

Preston, C. D. — Potamogeton x_ schreberi
G. Fisch. (P. natans L. x P. nodosus Poir.) in
Dorset, new to the British Isles 255-262

Preston, C. D. — Sir George Taylor’s studies of
the genus Potamogeton 325-326

Preston, C. D., with J. M. Croft — Obit. of
Dorothy Mary Greene (1928-1994) 447-449

Preston, C. D., with M. O. Hill & A. J. E. Smith
(eds) — Atlas of the bryophytes of Britain and
Ireland. Volume 1: Liverworts (Hepaticae
and Anthrocerotae) (Bk Rev.) 161-162

Preston, C. D., with M. O. Hill & A. J. E. Smith
(eds) — Atlas of the bryophytes of Britain and
Ireland. Volume 2: Mosses (except Diplole-
pideae) (Bk Rev.) 161-162

Preston, C. D., with A. L. Stewart & D. A.
Pearman (eds) — Scarce plants in Britain (Bk
Rev.) 444-445

Primavesi, A. L. — Obit. of Kenneth Guy Mess-
enger (1920-1993) 322-323

Primavesi, A. L., with Q. C. B. Cronk — Further
Druce Rosa taxa (Rosaceae) 411-412

Primavesi, A. L., with Q. C. B. Cronk & S. K.
Marner — Lectotypification of Rosa rothschil-
dii Druce (Rosaceae) 409-410

Primavesi, A. L., with G. G. Graham — Roses of
Great Britain and Ireland: BSBI handbook
No. 7 (Bk Rev.) 160-161

» Primula japonica (v.c. 77) 423; x polyantha (v.c.

73) 423; veris 9, 10, 443; vulgaris 65, 443;
vulgaris x veris (v.c. 73) 423

Proctor, J., with A. J. M. Baker & R. D. Reeves
(eds) — The vegetation of ultramafic (serpen-
tine) soils (Bk Rev.) 74-75

Prunus cerasus (v.c. 50) 292; lusitanica (v.c. 49)
292; padus 11; spinosa 381, 382

Pteridium aquilinum (L.) Kuhn) infestation of
rough grazing land in the catchment of the
River Tyne, northern England. Bracken(,

105-114, 264, 265

| Pteris cretica (v.c. 5, 29) 285

_ Puccinellia 391, 394, 395; borealis 392; capillaris
SO 3925 distans 8, 13, 392, (v.c. 26) 435,
(Jacq.) Parl. subsp. borealis (O. Holmb.) W.
E. Hughes (Poaceae) in mainland Scotland
and the Outer Isles 391-396, (illus.) 393,
subsp. distans 391, var. prostrata 392; fascicu-
lata 11; maritima 11, 392, 394, 395; x mixta
395

465

Pulicaria dysenterica 65; vulgaris 35
Pulmonaria ‘Mawson’s Blue’ (v.c. 39) 430
X Pyracomeles 309

Pyrola minor 11; rotundifolia 316

Pyrus pyraster 311

Quercus humilis 305; ilex 214, 381-383; lusitanica
305; petraea 176; robur 214, 382

Radiola linoides 10

Ranunculus 43, 48; aquatilis 12; auricomus (v.c.
108) 286; baudotii 9; bulbosus 214; calthifo-
lius 45, 46; circinatus 9, 53; ficaria L. sensu
lato 41-50, (key) 44-45, f. luxurians 44, 46,
subsp. bulbifer 43, 46, subsp. bulbilifer 41,
43, 45, 47-49, subsp. calthifolius 41, 44, 45,
47, 48, subsp. chrysocephalus 41, 46-49,
subsp. fertilis 45, subsp. ficaria 41, 43, 45, 47—
49, subsp. ficartiformis 41, 44, 46-48, subsp.
grandiflora 46, subsp. grandiflorus 44, subsp.
nudicaulis 45, var. aurantiacus 43, 45, 49,
var. bulbifer 43, 45, 46, var. calthaefolia 44,
var. calthifolia 45, var. calthifolius 43, 46,
var. divergens 46, var. fertilis 43, 45, var.
ficaria 46, var. incumbens 45, var. sinuata 46,
var. sinuatus 43; ficariaefolia 44; ficariaefor-
mis 44; ficartiformis 43, 44, 46; flammula 10,
26, 35, 235, subsp. flammula 65; fluitans 9,
11; grandiflorus 43; hederaceus 10; lingua 9,
10; monspeliacus 304; nudicaulis 43; omio-
phyllus 11; ophioglossifolius 169, 171; pelta-
tus 10, 35; penicillatus 10, subsp. pseudoflui-
tans (v.c. 46) 420, subsp. pseudofluitans var.
pseudofluitans 420; praecox 43; repens 35,
234, 235; reptans 237; sardous 11, (v.c. 38,
50, 75) 286; sceleratus 65, 235, (v.c. 102) 420;
trichophyllus 8; tripartitus 11, (v.c. 41) 420

Raphanus raphanistrum 159, (v.c. 102) 423

Rapistrum rugosum (v.c. 50) 423

Reeves, R. D., with A. J. M. Baker & J. Proctor
(eds) — The vegetation of ultramafic (serpen-
tine) soils (Bk Rev.) 74-75

Reports 85-87, 181-183, 327-328

Rhamnus catharticus 79, 311

Rhinanthus angustifolius 12; minor 9, 10

Rhododendron 309, 312; aiolosalpinx 80; ponti-
cum 102

Rhus hirta (v.c. 50) 294

Rhynchospora alba 10; fusca 9, 11

Ribes alpinum (v.c. 58, 73) 424; rubrum 11

Rich, T. C. G. — Pedicularis sylvatica L. subsp. -
hibernica D. A. Webb (Scrophulariaceae)
new to Wales 70-71

Rich, T. C. G. — Obit. of Leonard Albert Liver-
more (1919-1994) 449-451

Rich, T. C. G. — Rev. of Kvétena Ceske Republiky
(Flora of “the**Czech” Republic). Vol. 1.
Huperziaceae — Urticaceae (eds S. Hejny &
B. Slavik) 77

466

Rich, T. C. G., with A. O. Chater — Rorippa
islandica (Oeder ex Murray) Borbas (Brassi-
caceae) in Wales 229-238

Richards, A. J., with A. A. Dudman— Seven new
species of Taraxacum Wigg. (Asteraceae),
native to the British Isles 119-132

Richardson, P. M. — Rev. of Flora of Northumb-
erland (by G. A. Swan) 165-166

Riley, P., with D. L. Kelly, D. M. Synnott &
J. McCullen — Wild plants of the Phoenix
Park (Bk Rev.) 442-444

Rohweder, J. G., with K. Kubitzski & M. Bittrich
(eds) — The families and genera of vascular
plants. Vol. 2: flowering plants, dicotyle-
dons, Magnoliid, Hamamelid and Caryo-
phyllid familes (Bk Rev.) 162-163

Rorippa 229, 233; amphibia 9; x anceps 8, (v.c.
61) 289; austriaca 236; islandica 35, (Oeder
ex Murray) Borbas (Brassicaceae) in Wales
229-238, (map) 230, (v.c. 44, 46) 289, (v.c.
41) 422; microphylla 8; palustris 229, 231,
232, 234-237; pyrenaica 35; x sterilis 8, 13;
sylvestris 9, 232, (v.c. 91) 422; sylvestris x
amphibia (v.c. 61) 289

Rosa 177, 196, 309, 323, taxa (Rosaceae). Further
Druce, 411-412, 446; group Transitoriae 426;
agrestis (v.c. 11) 427; x andegavensis (v.c. 17)
292; arvensis 411, var. suberecta 411; borreri
410; caesia 161, subsp. glauca (v.c. 3, 108)
427; canina 161, 382, 410, 411, (v.c. 102, 108)
426, var. latebrosa 411; canina x caesia subsp.
glauca (v.c. 1) 427; canina x obtusifolia (v.c.
25) 292; canina x rubiginosa 409-411; canina
x sherardii 409, 410; caryophyllacea 410;
x drucei 411; x dumalis 160, 161, (v.c. 1) 427;
x dumetorum (v.c. 25) 292; eglanteria var.
corstorphinae 411; eglanteria x gallica 411;
glauca (v.c. 17) 292; x hibernica (v.c. 49) 426;
x involuta (v.c. 46) 427; micrantha (v.c. 26)
292, 410, (v.c. 68) 427; mollis (v.c. 75) 292;
mollissima f. alba 411; x nitidula 409, 411;
obtusifolia 177, 409, var. rothschildii 410;
pimpinellifolia 214; pimpinellifolia x canina
(v.c. 49) 426; pimpinellifolia x mollis (v.c. 98)
426; rothschildii Druce (Rosaceae). Lectoty-
pification of, 409-410; x rothschildii 409, 410;
rubiginosa 160, 409-411, (v.c. 67, 73) 427,
f. corstorphinae 411; rubiginosa x pimpinelli-
folia 411; rugosa 79, (v.c. 80) 426; x sabinii
(v.c. 98) 426; sherardii 409, 427; x sherardii
pimpinellifolia (v.c. 46) 427; stylosa x canina
(v.c. 17) 292; tomentella 409, var. rothschildii
410; tomentosa (v.c. 68) 427, f. alba 411;
verticillacantha 410

Rose, R. J., with S. B. Chapman — Changes in the
distribution of Erica ciliaris L. and E. x
watsonii Benth. in Dorset, 1963-1987 89-95

Rothero, G. & Thomson, B. — An annotated
checklist of the flowering plants and ferns of

INDEX TO WATSONIA VOLUME 20

Main Argyll (Bk Rev.) 444

Royal Botanic Gardens, Kew — Index of garden
plants (Bk Rev.) 310-311

Rubia peregrina 382

Rubus 61, 79, 160, 285, 303, 317-319, 323, 329,
330, 332-336, 340, 341, 345, 407, 408, 419,
442, 446, exisiccatae in British herbaria.
Additional sets of continental, 409, L. (Rosa-
ceae). Four new Anglo-Sarnian species of,
143-151, L. (Rosaceae) from Wales and the
Welsh Marches. New species of, 133-141;
subgen. Rubus). DNA fingerprinting and
biometry can solve some _ taxonomic
problems in apomictic blackberries (, 329-
343; sect. Corylifolii 133, 136, 144, 145; sect.
Hystrices 133; sect. Rubus series Sylvatici
345, 347, 348; sect. Vestiti 146; series
Micantes 150; series Sylvatici 61, 148; aden-
anthoides (v.c. 79, 99) 425; adscitus (v.c. 9)
291, (v.c. 44) 425; aequalidens (v.c. 3) 291,
(v.c. 9) 425; aghadergensis D. E. Allen, nom. —
et stat. nov. 61, D. E. Allen (Rosaceae): a
new name for an Irish bramble 61; amplifica-
tus (v.c. 80) 290; anglosaxonicus 150, 407;
anglosaxonicus x echinatus 407; arcticus L.
(Rosaceae). An early Scottish record of, 418;
ariconiensis A. Newton & M. Porter, sp.
nov. 134, (map) 135, 137-139, (v.c. 44) 426;
armeniacus (v.c. 70, 75) 291, (v.c. 52) 425;
arrhenii 160; arrheniiformis (v.c. 9, 28) 290;
aspericaulis 407; atrebatum (v.c. 9) 425; bal-
fourianus 137, 139; boraeanus 146; boudiccae
(v.c. 9) 290, 319, (v.c. 53) 424; boulayi 143;
britannicus 139, 140, (v.c. 12) 426; broensis
340; caesarius D. E. Allen, sp. nov. 145, 146,
148, 150; caesius 139, 145, (v.c. 67, 68) 426;
calvatus 348; calvus 348; cardiophyllus (v.c.
2) 290; chrysoxylon 147; confinis 340; corb-
ieri 143, 146; couchii Rilstone & D. E. Allen,
sp. nov. 246, 147, 148, 150; crudelis 148;
daltrii 317; dasyphylloides 145; dasyphyllus
140, 145, 348; dentatifolius 150; diversifolius
136-139; diversus (v.c. 12) 425; drejeri 345;
dumetorum 136, var. britannicus 139, var.
diversifolius 139, var. ferox 144, 145; ebuden-
sis (v.c. 104) 290; echinatoides 150, (v.c. 79)
425; edeesii H. E. Weber & A. L. Bull, sp.
nov. 345, H. E. Weber & A. L. Bull (Rosa-
ceae), a new bramble species from Britain
and Germany 345-349, (illus.) 346, (illus.)
347; elegantispinosus (v.c. 69) 290, (v.c. 13)
425; elongatispinus 148; errabundus (v.c. 2)
290, (v.c. 64, 79) 424; eupectus 134; flexuosus
(v.c. 46) 425; formidabilis (v.c. 11) 425;
fruticosus 143, 214; fuscicortex (v.c. 50) 291;
fuscoater 144; fuscus 329-336, 339-341; gra-
bowskii 341; gratus 348; griffithianus 150;
hartmanii 329-336, 339-341; hebridensis
(v.c. 94) 426; heterobelus 150, (v.c. 3) 291;

INDEX TO WATSONIA VOLUME 20 467

hirtifolius 147; hirtus 140; hylocharis 140,
348; imbricatus 146; incurvatiformis (v.c. 49,
50) 425; infestus 329-334, 337, 338, 340;
insectifolius (v.c. 2) 291; insularis 329-334,
336, 337, 340, 342, subsp. confinis 331, 340,
subsp. insularis 340; intensior 139, 140;
iodnephes 139; iscanus A. Newton & M.
Porter, sp. nov. 133, 134, (map) 135; lacinia-
tus (v.c. 75) 290, (v.c. 42, 44) 424; latifolius
(v.c. 104) 291; leucandrus 408; leucanthemus
147; leyanus (v.c. 26) 425; lindebergii (v.c.
79) 425; lindleianus 61, 348, var. latifolius 61;
longithyrsiger (v.c. 9) 291; macrophylloides
147, 348; majusculus 408; malvernicus (v.c.
3) 291; marshallii var. semiglaber 136; mela-
nodermis (v.c. 44) 425; menkei 339, var.
pseudopallidus 339; merlinii 140; micans 150,
407, (v.c. 35) 425; milfordensis (v.c. 11) 290,
(v.c. 11) 425; mitigatus 339; mucronatoides
348; muenteri 329-334, 339-341; nemoralis
348; nemorosus 137, 139, (v.c. 46) 426; nes-
sensis 341; neumanii 340; norvicensis 319;
obvallatus 148; odoratus x parviflorus (v.c.
13) 424; ordovicum (v.c. 50) 291; oxyanchus
408; pallidus 329-336, 339, 340; pedemonta-
nus 341; percrispus D. E. Allen & R. D.
Randall, sp. nov. 148, 150, D. E. Allen & R.
D. Randall (Rosaceae) in Dorset (v.c. 9)
407-408; perdigitatus (v.c. 3) 290; phaeocar-
pus 148, (v.c. 9) 291; pictorum (map) 135,
139, 140, (v.c. 42) 426; pistoris (v.c. 9) 290;
platyacanthus 348; plicatus (v.c. 104) 290,
348; polyanthemus 329-334, 338, 340, 348;
pseudopallidus 329-336, 339, 340; pulcherri-
mus 340; pullifolius W. C. R. Watson (Rosa-
ceae). Typification of, 408; pyramidalis (v.c.
104) 290, 348; radula 146, 407; raduloides
150, (v.c. 70) 291, 407; rhamnifolius 61;
rossensis (v.c. 2) 291, (v.c. 1) 425; rubriflorus
145; rudis 150, (v.c. 28) 291, 348; scabripes
(v.c. 17) 291; scabrosus 145; scheutzii 329-
334, 339-341; sciocharis (v.c. 14) 424; segon-
tii A. Newton & M. Porter, sp. nov. 140;
septentrionalis 329-334, 336, 337, 340;
silurum (v.c. 69) 290; silvaticus 348; sprenge-
lii (v.c. 44) 425; stenopetalus 160; subiner-
moides (v.c. 104) 290; taeniarum 330, 338,
340; tenuiarmatus (map) 135, 137, 139; thyr-
sigeriformis 143; transmarinus D. E. Allen,
sp. nov. 143, 145, 150; triangularis 139; trich-
odes (v.c. 10) 425; tuberculatus 136, 137, 139,
144, (v.c. 77) 291, 345, 348; ulmifolius 146,
319, 381, 382; vagensis (map) 135, A.
Newton & M. Porter, sp. nov. 137-139, (v.c.
42) 426; vestitus 137, 146, 348; vigorosus 348;
winteri (v.c. 3) 291, (v.c. 10) 425; wirralensis
(v.c. 64) 425

Rudbeckia 80

Rumex acetosella 443, subsp. acetosella 182; con-

glomeratus 413, 415; conglomeratus x obtusi-
folius subsp. obtusifolius 414; x cornubiensis
D. T. Holyoak, hybr. nov. 413, (illus.) 414;
crispus 65, 234, 388, 415, subsp. crispus 413;
crispus x obtusifolius 413; cristatus (v.c. 12)
421; x dufftii (v.c. 46) 288; frutescens 412,
413, (illus.) 414, 415; frutescens x conglomer-
atus 413; frutescens Thouars x R. obtusifolius
L. (Polygonaceae), a previously undescribed
hybrid dock, and new records of R. x wrightii
Lousley in West Cornwall (v.c. 1) 412-415;
frutescens x obtusifolius subsp. obtusifolius
413; hydrolapathum 9, 11; longifolius (v.c.
81, 100) 421; longifolius x crispus (v.c. 79)
288; maritimus 7, 10, 23, (v.c. 81) 288;
obtusifolius 235, 413, 415, subsp. obtusifolius
413, (illus.) 414; palustris 10; patientia (v.c.
12) 421; x pratensis 413; x propinquus (v.c.
79) 288; pulcher 413; sanguineus 413; sangui-
neus X obtusifolius (v.c. 46) 288; x wrightii
Lousley in West Cornwall (v.c. 1). Rumex
frutescens Thouars x R. obtusifolius L. (Poly-
gonaceae), a previously undescribed hybrid
dock, and new records of, 412-415

Ruppia cirrhosa 11, 18; maritima 11

Rushton, B. S. — A new direction for Watsonia
(Editorial) 1

Sagina boydii 198; maritima 392, 394, (v.c. 81)
421; nodosa 10, 53; procumbens 235; sub-
ulata (v.c. 42) 421

Sagittaria sagittifolia 10, 53, 260

Salicornia 79, ramosissima 11 |

Salix 13, 28, 197, 198, 201, 296; x ambigua (v.c.
80) 422; aurita 9,10, 295; aurita x repens (Vv.c.
80) 422; caprea subsp. sphacelata (v.c. 78)
288, var. caprea 201, var. genuina 201, var.
sphacelata 201; caprea x myrsinifolia (v.c.
104) 288; cinerea 235; cinerea x repens (v.c.
11) 288; daphnoides (v.c. 77) 288; x forbyana
(v.c. 77) 288; fragilis 173; x friesiana (v.c. 69)
288; x latifolia (v.c. 104) 288; myrsinifolia x
phylicifolia (v.c. 93) 288; purpurea 12; pur-
purea x viminalis x cinerea (v.c. 77) 288;
repens 10; x sericans (v.c. 75) 288; x subseri-
cea (v.c. 11) 288; x taylorii 325; x tetrapla
(v.c. 93) 288; triandra 8; viminalis 8; vimin-
alis x caprea (v.c. 75) 288; viminalis x repens
(v.c. 69) 288

Salsola 440; kali (v.c. 104) 287

Salvia pratensis (v.c. 13) 295

Sambucus canadensis (v.c. 77) 431, var. acutiloba
431; nigra 382; racemosus (v.c. 6) 297

Samolus valerandi 53

Sanford, M. N. — Rev. of Roses of Great Britain
and Ireland: BSBI handbook No. 7 (by
G. G. Graham & A. L. Primavesi) 160-161

Sanguisorba minor 382, subsp. muricata (v.c. 26,
35, 77) 426; officinalis 9, 11

468

Sanicula europaea 443

Santolina chamaecyparissus (v.c. 77) 433

Saponaria officinalis 65

Sarracenia purpurea (v.c. 98) 288

Sasaella ramosa (v.c. 46) 434

Saxifraga granulata 11; hypnoides (v.c. 50) 424;
rotundifolia (v.c. 77) 424; tridactylites 210,
211, 214, 223, 226; umbrosa x spathularis
(v.c. 50) 290; x urbium (v.c. 50) 290

Scabiosa columbaria subsp. pratensis 78

Scandix pecten-veneris 444

Scheuchzeria palustris 11

Schoenoplectus lacustris 11, 53, 260; tabernae-
montani 9, (v.c. 81) 299

Schoenus nigricans 10, 12, (v.c. 48) 299

Scilla autumnalis 65; siberica (v.c. 49) 436

Scirpus sylvaticus 11

Scleranthus annuus 182, (v.c. 99) 421

Sclerochloa distans 391

Scott, W. & Palmer, R. C. — A new Shetland
Hieracium of the section Alpestria (Fries)
F. N. Williams 282-284

Scott, W., with R. C. Palmer — A forgotten
Shetland dandelion 279-281

Scrophularia auriculata 12, (v.c. 104) 296

Scutellaria galericulata 12, 65;.minor 12

Sedum dasyphyllum (v.c. 50) 290; spectabile (v.c.
25) 424; spurium (v.c. 58) 424; stoloniferum
(v.c. 4) 290

Selaginella selaginoides 11

Sell, P. D. — Ranunculus ficaria L. sensu lato 41-
50

Sell, P. D., West, C. & Tennant, D. J. — Eleven
new British species of Hieracium L. section
Alpina (Fries) F. N. Williams 351-365

Senecio x albescens (v.c. 59, 75) 298; aquaticus 10;
cambrensis 307; cinerea x jacobaea (v.c. 59,
75) 298; erucifolius 382, (v.c. 48) 433; jacob-
aea 382; squalidus 205, (v.c. 91, 98) 433;
squalidus x viscosus (v.c. 47) 433; subnebro-
densis (v.c. 47) 433; sylvaticus 211; vulgaris
205, 206, 209-214, 220, 222, 223, 225, 388, L.
subsp. denticulatus (O. F. Muell.) P. D. Sell
and S. vulgaris subsp. vulgaris var. vulgaris
on Jersey (Channel Islands) 205-227, subsp.
vulgaris var. hibernicus 205, subsp. vulgaris
var. vulgaris on Jersey (Channel Islands).
Senecio vulgaris L. subsp. denticulatus (O. F.
Muell.) P. D. Sell and, 205-227

Septoria ficariae 48

Seriphidium maritimum 65

Serratula tinctoria 65, 182

Sheehy Skeffington, M. — Rev. of Wild plants of
the Phoenix Park (by P. A. Riley, with D. L.
Kelly, D. M. Synnott & J. McCullen) 442-

; 444

Shi, Y., with C. A. Stace, R. J. Gornall & J.

Squirrel — Chromosome numbers in Hiera- .

cium L. section Alpina (Fries) F. N. Williams

INDEX TO WATSONIA VOLUME 20

367-377

Shorrocks, B. & Coates, D. (eds) — The release of
genetically-engineered organisms (Bk Rev.)
164-165

Silene conica (v.c. 40) 421; x hampeana (v.c. 73)
421; latifolia subsp. alba x dioica (v.c. 73)
421; noctiflora 65; nutans 214; uniflora 388

Simmons, M. — A catalogue of the Herbarium of
the British Flora collected by Margaret Stovin
(1756-1846) (Bk Rev.) 442

Sinacalia tangutica (v.c. 40) 433

Sinapis arvensis 159

Sison amomum 12, 65

Sisymbrium orientale (v.c. 99) 288; terrestre 232

Sisyrinchium bermudiana (v.c. 12) 436

Sium latifolium 10, 27, 53, 444 f

Slavik, B., with S. Hejny (eds) — Kvétena Ceske
Republiky (Flora of the Czech Republic).
Vol. 1. Huperziaceae — Urticaceae (Bk Rev.)
77 i

Smith, A. J. E. — Rev. of Flora of Glamorgan (by —
A. E. Wade, Q. O. N. Kay & R. G. Ellis)
445446

Smith, A. J. E., with M. O. Hill & C. D. Preston
(eds) — Atlas of the bryophytes of Britain and
Ireland. Volume 1: Liverworts (Hepaticae
and Anthocerotae) (Bk Rev.) 161-162

Smith, A. J. E., with M. O. Hill & C. D. Preston
(eds) — Atlas of the bryophytes of Britain and
Ireland. Volume 2: Mosses (except Diplole-
pideae) (Bk Rev.) 161-162

Smyrnium olusatrum (v.c. 57) 429

Solanum dulcamara 34, 35; nigrum (v.c. 102) 430

Soleirolia soleirolii (v.c. 26, 77) 420

Solidago gigantea subsp. serotina (v.c. 50, 81) 432

Sonchus arvensis 65

Sorbaria tomentosa (v.c. 29) 290

Sorbus 379, 382, 383; aucuparia 379-381, 383;
domestica 178, L., new to Wales and the
British Isles 379-384; intermedia 311, (v.c.
40) 427; lancastriensis 450; torminalis 182,
(v.c. 41) 292

Southworth, E. C. — Rev. of Greek wild flowers
and plant lore in ancient Greece (by H.
Baumann, with W. T. Stearn and E. R.
Stearn (trans.)) 439

Sparganium angustifolium 11; emersum 260; erec-
tum 34, 35, 235, 260; minimum 170; natans
10, 12, 170, (v.c. 5) 300

Spartium junceum (v.c. 50) 428

Spergularia marina 388, 392, 394

Sphagnum 182, 266, 288, 295

Spiraea x pseudosalicifolia (v.c. 50) 424; x rosalba
(v.c. 50) 424; salicifolia x alba (v.c. 50) 424;
salicifolia x douglasii (v.c. 50) 424

Spiranthes romanzoffiana (v.c. 110) 436; spiralis
265

Spirodela polyrhiza 10, (v.c. 46) 434

Squirrel, J., with C. A. Stace, R. J. Gornall & Y.

INDEX TO WATSONIA VOLUME 20

Shi— Chromosome numbers in Hieracium L.
section Alpina (Fries) F. N. Williams 367—
377

Stace, C. A., Gornall, R. J., Squirrel, J. & Shi, Y.
— Chromosome numbers in Hieracium L.
section Alpina (Fries) F. N. Williams 367—

| Sd

Stachys x ambigua 10, (v.c. 102) 430; arvensis

(v.c. 81) 430; byzantina (v.c. 35, 77) 430;

germanica 158; officinalis 65, (v.c. 77) 430;

palustris 10; sylvatica x palustris (v.c. 102)

| 430

- Stark, G. — Rev. of Scandinavian ferns (by B.

@Ollgaard) 164

_ Stearn, W. T. — Stearn’s dictionary of plant names

| for gardeners (Bk Rev.) 80

Stearn, W. T., with W. Blunt — The art of

botanical illustration (Bk Rev.) 305-307

_ Stellaria holostea 65, (v.c. 102) 420; media 235;

palustris 10; uliginosa 10, 35

_ Stewart, A. L., Pearman, D. A. & Preston, C. D.

| (eds) — Scarce plants in Britain (Bk Rev.)

444-445

_ Stewart, N. F. & Church, J. M. — Red data books
of Britain and Ireland: Stoneworts (Bk Rev.)
80-81

Stratiotes aloides 9, 12

Styles, Brian Thomas (1934-1993) (Obit.) 173-
175

Suaeda maritima 11; vera 65

Succisa pratensis 10, 65

Swan, G. A. — Flora of Northumberland (Bk
Rev.) 165-166

_ Symphoricarpos x chenaultii (v.c. 44) 432; orbicu-
laris x microphyllus (v.c. 44) 432

Symphytum bulbosum (v.c. 6) 295; caucasicum
(v.c. 26) 430; grandiflorum (v.c. 47, 73) 295;
officinale 9; orientale (v.c. 50) 295; tubero-
sum (v.c. 47) 295; (v.c. 46) 430

Tamarix gallica 388

_ Tanacetum parthenium 66

| Taraxacum 79, 121, 160, 165, 235, 279, 280, 323,
| 330, 341, 446, 450, Wigg. (Asteraceae),
native to the British Isles. Seven new species
of, 119-132; sect. Celtica 121; sect. Erythro-
sperma 124; sect. Hamata 132; sect. Naevosa
125; sect. Obliqua 125; sect. Ruderalia 123,
128, 130; sect. Spectabilia 279, 280; sect.
Vulgaria 128; aequilobum 121, 123, 128;
agaurum 125; amplum 128; argutum 121;
atactum 132; breconense C. C. Haw., sp. nov.
119, (illus.) 120, 121; “British atactum’’119,
132; “British laetum” 119, 125; crispulum
128; discretum 125; edmondsonianum
H. @Ollg., sp. nov. 121, (illus.) 122, 123;
euryphyllum 126; faeroense 280; geirhildae
(Beeby) R. C. Palmer & W. Scott, comb. et
stat. nov. 280, 281; haworthianum A. A.

469

Dudman & A. J. Richards, sp. nov. 119, 123,
(illus.) 124, 125; hirsutissimum C. C. Haw.,
sp. nov. 119, 125, (illus.) 126; laetiforme 124,
125; laetum 124, 125; latens H. Ollig., sp. nov.
127, (illus.) 127, 128, 129; nigridentatum
T. Edm., sp. nov. 119, 129, (illus.) 129, 130;
ornatum 280; ostenfeldii 121; palustre 11;
pannulatiforme 130; pannulatum 130; platy-
glossum 125; sahlinianum A. A. Dudman &
A. J. Richards, sp. nov. 119, 130, (illus.) 131,
132; spectabile 280, subsp. geirhildae 280,
(illus) 281; vastisectum 128

Taxus 75, baccata 75, 382, ‘Fastigiata’ 76; brevifo-
lia 75 |

Taylor, Sir George (1904-1993) (Obit.) 324-326

Taylor, Sir George, studies of the genus Potamo-
geton 325-326

Tayloriophyton 325

Teesdalia nudicaulis 182, (v.c. 81) 289

Tellima grandiflora (v.c. 79) 424

Tennant, D. J., with P. D. Sell & C. West —
Eleven new British species of Hieracium L.
section Alpina (Fries) F. N. Williams 351—
365

Teucrium balearicum 305; botrys 66; marum 305;
scorodonia 382; subspinosum 305

Thalictrum flavum 11, (v.c. 45) 286; minus (v.c.
78) 420

Thelypteris palustris 9, 10, (v.c. 41) 285, (v.c. 57,
69) 419

Thlaspi 440; arvense 66, 159; caerulescens 78

Thomson, B., with G. Rothero — An annotated
checklist of the flowering plants and ferns of
Main Argyll (Bk Rev.) 444

Tilia cordata 182, 348; platyphyllos 182, (v.c. 50)
288

Tolmiea menziesii (v.c. 46) 424

Tolypella 17; intricata 12; nidifica 9; prolifera 12

Torilis nodosa 66

Tortula ruraliformis 223

Tragopogon porrifolius (v.c. 35) 432; pratensis
subsp. minor 66

Trichomanes 164

Trichophorum cespitosum 12

Trientalis europaea 66

Trifolium 447; campestre 66; fragiferum subsp.
fragiferum 66; glomeratum 66, (v.c. 5) 294;
incarnatum 66, subsp. incarnatum (v.c. 40)
428, subsp. molinerii 66; occidentale 214;
ochroleucon 66; ornithopodioides (v.c. 12)
428; repens 235; scabrum 66, 388; squamo-
sum 66; striatum 66; subterraneum (v.c. 39)
428; suffocatum 388

Triglochin maritimum 11; palustre 11

Tripleurospermum maritimum 392, 394

Trisetum flavescens (v.c. 102) 435

Trist, P. J. O. — Elytrigia repens (L.) Desv. ex
Nevski subsp. arenosa (Spenner) A. Léve
(Poaceae) in north-western Europe 385-390

470

Trist, P. J. O. & Butler, J. K. — Puccinellia distans
(Jacq.) Parl. subsp. borealis (O. Holmb.) W.
E. Hughes (Poaceae) in mainland Scotland
and the Outer Isles 391-396

Triticum maritimum 385, 386; repens var. y 386,
var. arenosum 386, var. maritimum 385, 386

Tropaeolum speciosum (v.c. 75) 294

Tunica prolifera 405

Tutin, T. G. et al. (eds) — Flora Europaea, 2nd
edition, vol. 1: Psilotaceae to Platanaceae
(Bk Rev.) 81-82

Typha angustifolia (v.c. 35) 301; x glauca (v.c. 35)
436; latifolia 34, 35, 235, (v.c. 102) 436;
latifolia x angustifolia (v.c. 35) 436

Ulex europaeus 214; minor (v.c. 98) 428

Ulmus 323, 382; glabra 443; procera 433

Uriginea maritima 304

Urmi, E. — Rev. of Atlas of the bryophytes of
Britain and Ireland. Volume 1: Liverworts
(Hepaticae and Anthocerotae) (eds M. O.
Hill, C. D. Preston & A. J. E. Smith) 161-
162 |

Urmi, E. — Rev. of Atlas of the bryophytes of
Britain and Ireland. Volume 2: Mosses
(except Diplolepideae) (eds M. O. Hill,
C. D. Preston & A. J. E. Smith) 161-162

Uromyces dactylidis 48; ficariae 48

Urtica dioica 235; urens 66

Utricularia australis 11; intermedia 12; minor 9,
10; vulgaris 10, 53

Vaccinium myrtillus 9; oxycoccos 9, 11, 12

Valeriana dioica 9, 11; officinalis 9, 11

Valerianella carinata (v.c. 50) 297, (v.c. 27) 432;
eriocarpa (v.c. 11) 297; locusta (v.c. 96) 297

Vaughan, Irene Mary (1889-1993) (Obit.) 175-
179

Vavilov, N. I. — Origin and geography of culti-
vated plants (Bk Rev.) 82-83

Verbascum x godronii (v.c. 26) 431; pulverulen-
tum 431; thapsus x pulverulentum (v.c. 26)
431; virgatum 66, (v.c. 35) 296

Veronica anagallis-aquatica 11, 28; beccabunga
11, 235; catenata 8, 28; chamaedrys 66, 443;
filiformis (v.c. 104) 296; hederifolia (v.c. 102)
431, subsp. hederifolia 66; x lackschewitzii
28; polita 66; scutellata 10, 35, (v.c. 102) 431;
serpyllifolia subsp. serpyllifolia 66

Viburnum lantana 382; opulus 12; tinus (v.c. 50)
297, (v.c. 12, 50) 432

Vicia herbacea (v.c. 81) 295; hirsuta 66; lathyr-
oides (v.c. 79) 293; lutea (v.c. 58) 293; minor
295; sativa 214, subsp. sativa (v.c. 12, 46, 81)
428; sylvatica 66; tetrasperma 66

- Viola canina 11; cornuta (v.c. 44) 422; kitaibeliana

INDEX TO WATSONIA VOLUME 20

210, 211, 214, 223; palustris 11, 182, subsp.
Juressi (v.c. 46) 422; persicifolia 9, 11

Vulpia bromoides 388; ciliata subsp. ambigua
(v.c. 18) 435; myuros (v.c. 98) 435

Wade, A. E., Kay, Q. O. N. & Ellis, R. G. -
Flora of Glamorgan (Bk Rev.) 445-446

Wahlenbergia hederacea 11

Walsh, W. F., with E. C. Nelson — Trees of
Ireland, native and naturalised (Bk Rev.)
311-312

Walters, S. M. — Obit. of Richard William
(‘Dick’) David (1912-1993) 320-321

Walters, S. M. — Wild and garden plants (Bk
Rev.) 313-314

Wanstall, Peter James (1924-1993) (Obit.) 451-
452

Webb, D. A. — Rev. of Irish trees and shrubs (by
P. Wyse Jackson) 311-312 i

Webb, D. A. — Rev. of Trees of Ireland, native _ |
and naturalised (by E. C. Nelson & W. F. |}
Walsh) 311-312

Weber, H. E. & Bull, A. L. — Rubus edeesii H. E.
Weber & A. L. Bull (Rosaceae), a new
bramble species from Britain and Germany
345-349

Welch, D. E. — An early Scottish record of Rubus
arcticus L. (Rosaceae) 418

West, C., with P. D. Sell & D. J. Tennant —
Eleven new British species of Hieracium L.
section Alpina (Fries) F. N. Williams 351-
365

Weston, I..— Annual General Meeting (1993)
(Rpt) 181-182

Weston, I. -— Central Lincolnshire nature
reserves, N. Lincs (v.c. 54), 16 May 1993 (Fld
Mtg Rpt) 182-183

Wiggins, Edgar Dunstan (1911-1933) (Obit.)
179-180

Wilmore, G. T. D., with J. C. Lavin (eds) — The
West Yorkshire plant atlas (Bk Rev.) 441

Wilson D. E. & Ingram, G. I. C. — Distribution of
Lady Orchid (Orchis purpurea Hudson) in an
East Kent wood 154-156

Wolffia arrhiza 9

Woodhead, F. — Flora of the Christchurch area
(Bk Rev.) 314-315

Woods, P. J. B., with B. Allen - Wild orchids of
Scotland (Bk Rev.) 73-74

Wynne, G. — Flora of Flintshire (Bk Rev.) 315-
316

Wyse Jackson, P. — Irish trees and shrubs (Bk
Rev.) 311-312

Zannichellia palustris 11

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CONTENTS

EpiToriAL A new direction for Watsonia

Mountrorp, J. O. Floristic change in English grazing Sri oe the sayarls
of 150 years of drainage and land-use change

MountTrorp, J. O. Problems in econstucting floristic change infereretine
the sources for English grazing marshes

BIRKINSHAW, C. R. Aspects of the ecology and conservation of Dyes
nium alisma Miller in Western Europe

SELL, P. D. Ranunculus ficaria L. sensu lato

CRACKLES, F. E. Calamagrostis stricta (Timm) Koes
(Wigg.) Roth and their hybrids in S.E. Yorks.,
England

NoTES

Allen, D. E. Rubus aghadergensis D. E. Allen (Rosaceae): a new name for
an Irish bramble

Payer, J. & ae B. The native British Coerener™ Great ome Bory
—renamed .

Dempsey, R. ae Gone R. fe & Teri Me P. Gonmbnten: to a
cytological ciples of the British and Irish flora, 4

Parnell, J. Variation and hybridisation of A/nus Miller in Ireland

Rich, T. C. G. Pedicularis sylvatica L. subsp. hibernica D. A. Webb
(Scrophulariaceae) new to Wales

Book REVIEWS ...

REPORT

Annual General Meeting, 15 May 1993

CHAPMAN, S. B. & RosE, R. J. Changes in the aincapation of see at
_L. and E. x watsonii Benth. in Dorset, 1963-1987

JONES, A. T. & Evans, P. R. A comparison of the growth and mennolopy
of native and commercially obtained continental European Crataegus
monogyna Jacq. (Hawthorn) at an upland site re

CHERRILL, A. J. & Lang, A. M. Bracken (Pteridium aquilinum (L. ) Kent)
infestation of rough grazing land in the catchment of the River Tyne,
northern England

FIELD, M. H. The status of ppRiiahane farm Tks (Easteaeene) in the
British flora

DupMan, A. A. & RICHARDS, A. Uf goven new species cat Tunmedeusn Wace
(Asteraceae), native to the British Isles

Newton, A. & Porter, M. New species of Rubus L. Russ Son
Wales and the Welsh Marches . es

ALLEN, D. E. Four new Anglo-Sarnian species of Ritbias ib. (Rovere)

NOTES

Preston, C. D. Juncus effusus var. spiralis J. McNab in the Inner Hebrides

Wilson, D. E. & Ingram, G. I. C. Distribution of Lady Orchid (Orchis
purpurea Hudson) in an East Kent wood

Book REVIEWS ...

OBITUARIES

REPORT

PERRING, F. Breodential Waar 1994: eres in Nowarmpronehice

C. canescens
v.c. 61, northern

Vv

PAGES

51-60

61

61-63

63-66
67-70

70-71
73-83

85-87

89-95

97-103

105-114

115-117

119-132

133-141
143-151

153-154

154-156
157-166
167-180
181-183
185-194

INDEX TO WATSONIA VOLUME 20

BunrtTING, M. J., Briccs, D. & BLockx, M. The Cambridge British Flora
(1914-1920)

Comes, H. P. Senecio eae pips Ten irounin (OC; F. Mie ) oe D.
Sell and S. vulgaris subsp. vulgaris var. vulgaris on Jersey (Channel
Islands)

CHATER, A. O. & Rien, T. C. G. Ronee sealoenalran (Cede em Marcen
Borbas (Brassicaceae) in Wales

Dawson, H. J. & INGROUILLE, M. J. A eemeie: survey oi Done
vulgare Miller and L. humile Miller in the British Isles

Preston, C. D. Potamogeton x schreberi G. Fisch. (P. natans L. x P.
nodosus Poir.) in Dorset, new to the British Isles

JENKINSON, M. N. A new variety of Narrow-leaved Marsh- atc in HS
Hampshire (v.c. 11)

NELSON, E. C. Erica x stuartii: the ASN reooneider a

NOTES

Dowlen, C. M. & Ho, T. N. Gentianella ciliata (L.) Borkh. in Wiltshire
(v.c. 8) ce

Palmer, Re @2de Scott, W. Acormaticn Shetland dandenen

Scott, W. & Palmer, R. C. A new Shetland Hieracium of the section
Alpestria (Fries) F. N. Williams

PLANT RECORDS

Book REVIEWS ..

OBITUARIES

REPORT

Annual General Meeting, 14 May 1994

Kraft, T. & Nypom, H. DNA fingerprinting and pememne can Aikes some
taxonomic st alae in apomictic blackberries (Rubus subgen.
Rubus) a)

WEBER, H. E. & (Enm. we ie fe ieee iz E nWebee & A. ne Bull
(Rosaceae), a new bramble species from Britain and Germany
SELL, P. D., WeEsT, C. & TENNANT, D. J. Eleven new British species of

Hieracium L. section Alpina (Fries) F. N. Williams ..

STACE, C. A., GORNALL, R. J., SQUIRRELL, J. & Sm, Y: Guisnastins
numbers in Hieracium L. section Alpina (Fries) F. N. Williams
Hampton, M. & Kay, Q. O. N. Sorbus domestica L., new to Wales and the

British Isles

Trist, P. J. O. Elytrigia repens a ) mea! ex Neveu iis arenosa
(Spenner) A. Love (Poaceae) in north-western Europe ee

Trist, P. J. O. & BuTLer, J. K. Puccinellia distans (Jacq.) Parl. Sega
borealis (O. Holmb.) W. E. Hughes (Poaceae) in mainland Scotland
and the Outer Isles

CRACKLES, F. E. A graphical acdnGe at ie eure of Calarnngrases
stricta (Timm) Koeler, C. canescens (Wigg.) Roth and their hybrid
populations in S. E. Yorks., v.c. 61, northern England

Notes

Akeroyd, J. R. & Beckett, G. Petrorhagia prolifera (L.) P. W. Ball &
Heywood (Caryophyllaceae), an overlooked native species in eastern
England

vi

195-204

205-227
229-238
239-254
259-262
263-273
275-278

279
279-281
282-284
285-301
303-316
317-326

327-328

329-343

345-349

351-365

367-377

379-384

385-390

391-396

397-404

405-407

A Te

aS

INDEX TO WATSONIA VOLUME 20

Allen, D. E. Rubus percrispus D. E. Allen & R. D. Randall (Rosaceae) in
Dorset (v.c. 9) t.

Allen, D. E. Typification of Rubus enifolius W. Cc. R. water n (Rose
ceae)

milen, D: E. Neat tere sets Ge eoninentall Rubee exsiccatae in Emiish
herbaria

Cronk, Q. C. B. ine S. K. & Syncs A. io Lectotypitcation ah
Rosa rouisensde Druce (Rosaceae)

Cronk, Q. C. B. & Primavesi, A. L. Further Druce as taxa (Resexe)

Halliday, G. Two subspecies of Festuca rubra L. new to England

Holyoak, D. T. Rumex frutescens Thouars x R. obtusifolius L. (Polygona-
ceae), a previously undescribed hybrid dock, and new records of R. x
wrightii Lousley in West Cornwall (v.c. 1) ah

Preston, C. D. Potamogeton x cognatus Asch. & Graebn. at Loch Borralie:
West Sutherland (v.c. 108), Scotland as

Welch, D. An early Scottish record of Rubus arcticus L. (Ressce25)

PLANT RECORDS

Book REVIEWS ...

OBITUARIES

Vil

407—408

408

409

409-410
411-412
412

412-415

415-417

418
419-437
439-446
447-452

ae

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Pe merabetiienys
sO ues xs Smaens ete
Ws Nikon mabeseus)

SOCIO

Gane
eH 60 werieg

bes
Da ate te Une tas

PRN cae eens
= cy

ARE NRG PLEA EH

MERE et
SY genitske Hs

we HE waieay CNeuyee

aegis
Pe eat

oo ew

cesta

oie

Wawa ens jwitea
Piet -ne

boty
aie dy enous

Wea IE
me Nt

pelos wes

eee
ete iE Atha ort TEN sipaeap aie Ue

se ae

Lie lvte A et a

wey
CUP AY pened pt.

ee ing et sre Me ae ae,
ihe iy

Vanfw

Nea a

Ca lad

AMY Sir
steve Nm

Werte,

pe Sm spaces
tetage je a(t:

my arate jteerar ra
he aa sn Ob eee pi ter
sgrniiteg tapi oh bent Pr ae a

ftp see petee ned epee w rivet o4
SSP RAE veo Wrepaie dpitte inyeecpondrisiant seer

eit of

Ae sprosweneete ay Ap ris beans
wit ne a Erpircy ye pee nk en

~*

Caen f eho Spiga dr cemjenl
mrestensr ge inte shite eis yeep

AAA Standing UNNI aly
besermusaeeiany

EO RR roar
bern eke

petes

oat reRejenheeion PEON enmen may
riere sy:

? datsir= Sm Ayreon
4PM emcee ley rm
nip dewyae S enes SS rte
Soares ier
wy e ap easel