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WATSONIA
JOURNAL AND PROCEEDINGS OF
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OF THE BRITISH ISLES
VOLUME 16
EDITED BY
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1986-87
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Watsonia, 16, 1-14 (1986)
Taxonomy of Elodea Michx in the British Isles
D. A. SIMPSON
School of Botany, Trinity College, Dublin 2, Ireland
ABSTRACT
Three species of Elodea are now known to occur in the British Isles: E. canadensis Michx, E. nuttallii (Planch.)
H. St John and E. callitrichoides (Rich.) Casp. The taxonomic histories of the genus and three species are
reviewed. Keys and descriptions of the species are given, together with details of their habitats and distribution.
INTRODUCTION
Elodea Michx is a New World genus, of which several species are known to be adventive in the Old
World. A recent investigation (Simpson 1983) has shown three species to be adventive in the
British Isles, namely E. canadensis Michx, E. nuttallii (Planch.) H. St John and E. callitrichoides
(Rich.) Casp. (following Cook & Urmi-K6nig’s (1984) recommendation that plants previously
referred to as E. ernstiae H. St John should now be assigned to E. callitrichoides). Simpson (1984)
outlined the introduction and spread of these species in the British Isles. This paper presents a brief
taxonomic history of the genus together with the British and Irish species, and summarizes the
taxonomic conclusions drawn from the work in the form of a key and brief systematic accounts. A
further paper in preparation will give a more detailed account of intraspecific variation in E.
canadensis and E. nuttallii. 3
The vegetative structure of Elodea is comparatively simple, consisting of a series of axillary,
branched, terete stems with narrow, sessile and minutely serrate leaves. Three decussate pairs of
leaves occur on the lowest part of the stem, above which all the leaves are in whorls. The decussate
leaves are also distinctly smaller than the whorled ones. A midrib is present in all leaves but no
other venation is apparent. Epidermal hairs are absent from both stems and leaves. A pair of
minute, entire, nodal scales (or squamulae intravaginales) are attached between the stem and leaf
base on the adaxial side of the leaf. Adventitious roots are produced at the nodes, coinciding with
the growth of a new stem. One root is produced with each stem, and root hairs occur only when the
root is in sediment. Despite its simplicity, the vegetative morphology of Elodea is, in common with
other aquatic macrophytes, highly plastic, and this has caused much of the taxonomic confusion.
Such variation appears to be brought about by a variety of environmental factors, particularly in
relation to light and temperature (Simpson 1983). The main growing season is between mid-April
and mid-September. At other times growth is very much reduced or ceases completely. The plant
overwinters by means of short, unbranched stems, or by the production of turions (hibernacula).
The latter arise in the upper leaf axils during early autumn and may remain in situ, or break free
and fall to the substrate, where they germinate the following spring.
The vegetative anatomy of Elodea reflects the plant’s morphological simplicity. The stem
consists of a single epidermal layer, a cortex consisting of parenchymatous and aerenchymatous
tissues, and a simple stele with a central protoxylem lacuna. The leaf consists of only two cell
layers. Stomata are absent and there is only a thin cuticle. Moreover there are no distinct
strengthening tissues in the stem, and only 2—3 rows of sclerenchymatous cells along the leaf
margins.
All the British and Irish species are dioecious. The female flowers are sessile and solitary in the
leaf axils, and they usually occur within the 3—5 cm of the stem apex. Each consists of a perianth
surmounting an elongated thread-like tube connected to the ovary. The latter is unilocular,
containing three to ten ovules with either basal or parietal placentation. The ovary and lower part
2 D. A. SIMPSON
of the tube are enclosed by a membranous spathe (Fig. 1). The precise nature of the flower
structure is disputed and some workers consider it to have a hypanthium (St John 1965; Lawrence
1976), whilst others suggest that it has an ovarian beak (Wylie 1904; Scannell & Webb 1976; Webb
1977; Dandy 1980). Close examination of the flower reveals that the tube is continuous with the
ovary, and that there is no separate style within the tube. This suggests that the flower is
epigynous, and that the latter interpretation is more likely to be correct.
Initially all parts of the flower are enclosed by the spathe. However the tube elongates rapidly,
carrying the perianth bud to the surface and, when above the water, the bud opens, revealing three
sepals, petals, stigmas and staminodes. The perianth is minute, being c. 3—7 mm in diameter. The
flowers are ephemeral and, once mature, they rarely survive for more than 48 hours. They seem to
be produced only when the top of the plant is within c. 0-5 m of the water surface and in still water
(Simpson 1983). Consequently the production of flowers is infrequent, and this has undoubtedly
added to the difficulties of separating the species.
In E. canadensis and E. callitrichoides, the general structure of the male flowers is similar to the
female, the main differences being that the ovary is absent, and that the staminodes and stigmas
are replaced by nine stamens. In E. nuttallii, the tube is absent and the spathe is more or less
spherical, whilst at anthesis the whole flower breaks free from the plant and floats ‘to the surface.
Pollination in all species is achieved by the pollen grains being scattered on the water surface and
drifting to the stigmas of the female flowers (St John 1965). Male flowers have only been recorded
once in the British Isles, from a population of E. canadensis in the Braid Hills near Edinburgh,
Midlothian, v.c. 83 (Douglas 1880). These plants were first found in 1879, but were not seen after
1903. I made an intensive search for male plants between 1978-1983, but without success, and it is
most likely that no male plants of the three species are present in the British Isles. It is, of course,
possible that such plants have been overlooked, but if they are present, then they must be rare.
In the absence of male flowers, reproduction is vegetatively apomictic. This is achieved by stem
breakage, the stems being extremely brittle. The detached stems quickly produce adventitious
Ficure |. Elodea, structure of flowers. A, E. nuttallii whole female flower. B, E. nuttallii, lower portion of
flower showing spathe and interior of ovary. C, E. canadensis, perianth of male flower from specimens
gathered near Edinburgh, v.c. 83 (K). Scale bars = 1 cm.
TAXONOMY OF ELODEA 3
roots, and soon become established as new plants. The effectiveness of this method as a means of
rapidly increasing a population has been amply demonstrated by E. canadensis and, to a lesser
extent, by E. nuttallii.
Determining the precise methods of spread is something of a problem which has never been
completely resolved. In canals or rivers it has probably been aided by water movement carrying
broken stem pieces. In both cases stem breakage is often increased by boat traffic, and pieces of
plant can be carried within the wake caused by the boat. However, it is difficult to identify dispersal
methods between isolated habitats, and in these cases agencies such as waterfowl could be
involved. Although there is no direct evidence to support this, incidental observations during the
present work suggest that plants can survive for several days in a humid atmosphere, and it is also
possible that material could cling to, say, birds’ feathers by the small teeth along the leaf margin. In
some cases deliberate introduction by Man is likely to have occurred.
E. canadensis has the most widespread distribution of the three species. It is native to North
America, being particularly common in the northern and eastern United States and southern
Canada. It is also adventive in most of continental Europe and Australasia, as well as parts of
Africa and Asia. E. nuttallii is also native to North America, although it is more or less confined to
the northern ard eastern United States. It is local but increasing rapidly in western continental
Europe (Wolff 1980) and has also been introduced into Japan (Kunii 1982). E. callitrichoides is
native to South America where it is common in southern Uruguay and northern Argentina. It has
been introduced to continental Europe, and is presently known at a few localities in France and
West Germany.
TAXONOMIC HISTORY OF THE BRITISH AND IRISH SPECIES
THE GENUS
Elodea was first described by Michaux (1803) in his Flora Boreali-Americana, but there were two
earlier homonyms mentioned by de Jussieu (1789) and Ventenat (1799). These were orthographic
variants of an earlier name, E/odes Adanson (1763), given to members of the Hypericaceae. Both
de Jussieu and Ventenat attributed the homonyms to Adanson; however neither homonym was
validly published, and Elodea Michx remains nomenclaturally correct.
Doubt was expressed by Babington & Planchon (1848) and later St John (1962) over the precise
authorship of the name. Both produced a range of evidence to suggest that L. C. M. Richard was
the original author. St John stated that, for many years, the staff at P believed that Flora Boreali-
Americana was written by Richard and all names in this Flora should have been accredited to him.
Further evidence of this was provided by Hooker (1842), who commented that Richard was the
anonymous author of the Flora. Richard (1814) himself made some claim to authorship by stating
that “‘je vais commencer par le description d’une plante qui appartent a un genre encore peu connu
et auquel j’ai donné le nom d’Elodea’’. St John also noted the following statement, written by
Richard, attached to the holotype of E. guyannensis Rich.: ‘“‘J’ai retrouvé ce dessin et cette
description dans une cartier d’observations fait 4a Cayenne. Je ne l’avais pas sans les yeux lorsque
Jai fait le caractére de la Flora Boreali-Americana de Michaux’. St John concluded that this
evidence was sufficient proof of Richard’s authorship. He therefore used the citation ‘Richard in
Michaux” throughout his papers. However there is, in fact, no conclusive evidence of Richard’s
authorship. The citation is not employed in Index Kewensis and, as far as is known, any other work
applicable to the genus apart from St John. Index Nominum Genericorum does mention Elodea
Rich., but as a later homonym of Elodea Michx, the former referring to Richard’s (1814)
description of the genus. As Elodea Rich. is a later homonym it is invalid. Consequently in the
absence of published evidence to the contrary, the name should be credited to Michaux alone and
not to Richard in Michaux.
Elodea was described by Michaux as monotypic, the species being E. canadensis, and he placed
it in the Linnaean Class Triandra, believing the plants to have hermaphrodite flowers. This is
interesting, because only pistillate flowers are to be found on the holotype. St John (1962)
concluded that Michaux had either examined hermaphrodite flowers, which may occur very
occasionally on this otherwise dioecious species or, more likely, he confused staminodes with
anthers. Whatever the reason, Michaux’s description of Elodea as hermaphrodite undoubtedly
4 D. A. SIMPSON
caused much of the confusion over the precise use of the name. Richard (1814) assigned a further two
hermaphrodite species, both from South America, to Elodea, and he described a new genus,
Anacharis, which contained dioecious material, also from South America. Rafinesque (1818), in a
review of Pursh’s Plants of North America, proposed the rejection of Elodea Michx on the grounds
that Elodes, which was in contemporary use for segregates of Hypericum, was similar to Elodea. He
suggested that the name Philotria Raf. should be adopted instead, but this did not gain widespread
acceptance. Nuttall (1818) described a monotypic new genus, Udora, which was applied to dioecious
material from North America. However, the description of this genus was essentially the same as
that given by Richard (1814) for Anacharis, and it appears that Nuttall was unaware of Richard’s
work (St John 1962). Nevertheless, by 1848, three generic names were in use for material covered by
the present-day concept of Elodea. Elodea sensu Michaux was hermaphrodite, Udora was both
hermaphrodite and dioecious and Anacharis was dioecious.
Babington & Planchon’s (1848) revision brought about a number of changes. The most significant
of these was the reduction of Elodea to the synonymy of a new hermaphrodite genus Apalanthe
Planch. By this time it was widely believed that Michaux had erroneously regarded his material as
hermaphrodite, and because of this, Babington & Planchon considered Elodea to be incorrect for
such material. Udora was also reduced to the synonymy of Apalanthe and Anacharis, the latter name
being maintained for dioecious material. However Caspary (1857, 1858) restored Elodea and his
interpretation of the genus included both hermaphrodite and dioecious plants. This concept rapidly
gained acceptance in both America and Europe but, in the British Isles, the debate continued,
fuelled by attempts to put a correct name to the British and Irish taxon. By the late 1870s, the
majority of British and Irish botanical opinion agreed that the floral differences were not important
enough to warrant the separation of two or more genera. This is confirmed by Index Kewensis which
shows that, by 1885, Elodea had become generally accepted for both hermaphrodite and dioecious
material.
Apart from a few minor nomenclatural changes, little further attention was paid to the genus until
Victorin’s (1931) revision, in which Elodea was again separated on flower structure. In this case he
used Anacharis for dioecious material and Philotria for hermaphrodites. The basis for the change
was his examination of a small amount of isotype material in P which did not have mature flowers.
From this, Victorin concluded that Michaux’s original description was erroneous, and consequently
he rejected Elodea. Victorin believed it important to recognize the two genera as distinct for two
reasons. Firstly, the number and arrangement of the stamens and staminodes differed between
them. Secondly, work by Santos (1923, 1924) had shown that in dioecious material corresponding
chromosomes differed in size between the sexes, whereas Victorin assumed that they were equal in
the hermaphrodite plants.
St John’s monograph (1962, 1963, 1964, 1965) again grouped both hermaphrodite and dioecious
material into the one genus. However, two subgenera were recognized, Elodea for the 14 dioecious
species and Apalanthe for the three hermaphrodite ones. Subgenus Elodea was divided into two
sections, Elodea and Natator H. St John, by the behaviour of the male flowers at maturity. In section
Elodea they remain attached to the plant, whilst in section Natator they break free and float to the
surface. St John added a total of ten new species to the seven previously recognized and accepted by
him.
THE SPECIES
E. canadensis was first described by Michaux (1803) as a hermaphrodite plant, and this was later
taken up by Richard (1814). However Babington & Planchon (1848), having determined its
dioecious nature, transferred it to Anacharis. It is of interest that their morphological description of
the species differed from that of Michaux. In particular, the leaves are described as being linear-
lanceolate and acute, whereas Michaux described them as oblong and obtuse. Their observations
were made, in fact, on two specimens of staminate plants (Drummond, K; Cleghorn, K) which had
abnormally longer, narrower leaves. Babington & Planchon also produced the first published
description of the British and Irish plant, which they named Anacharis alsinastrum Bab. They were,
however, unsure of the precise relationship between this plant and American material. Therefore
they suggested the epithet alsinastrum to “‘prevent it being confounded with the American species
and thus extending their range far beyond what may prove to be their natural limits’’. Their use of the
epithet was derived from the plant’s apparent similarity to Elatine alsinastrum L.
TAXONOMY OF ELODEA 5
Caspary’s (1857, 1858) interpretation of the species included both hermaphrodite and
dioecious plants. The hermaphrodite part was based upon Michaux’s description of the type
specimens and on his personal examination of material from Bethlehem, Pennsylvania
(Schweinitz, K); these plants are referred to E. schweinitzii (Planch.) Casp., a true hermaphro-
dite species, by St John (1962). The dioecious part was based on Babington & Planchon’s
descriptions of species assigned to Anacharis. Two further species were described: E. planchonii
Casp. and E. latifolia Casp., both of which were later recognized by St John (1965) to be E.
canadensis. E. planchonii was based on the Drummond and Cleghorn specimens mentioned
above, and Caspary considered these to be sufficiently distinct to be treated as a separate species.
The same applied to E. latifolia which had markedly broad, ovate leaves. Indeed, both
represented a wide spectrum of variation which misled Caspary. Nevertheless, they were treated
as separate species for many years until Victorin (1931) reduced E. planchonii to a variety of
Anacharis canadensis and St John (1965) recognized both to be phenotypic variants of the latter.
Moreover E. canadensis was considered to be both hermaphrodite and dioecious until St John
(1962, 1965) recognized it to be nearly always dioecious. He also recognized E. planchonii to be
based on staminate plants of this species.
The first specimens of E. nuttallii to be described were assigned to Serpicula verticillata L.f. (=
Hydrilla verticillata (L.f.) Royle) by Muhlenberg (1813) as var. 6B angustifolia. However Nuttall
(1818) made this variety synonymous with his Udora canadensis Nutt. The latter species was
described as having oblong-ovate to linear-lanceolate leaves and with staminate flowers sessile
and breaking free at anthesis. Nuttall based his description partly on Michaux’s type material and
partly on specimens collected by himself in Philadelphia (BM). The latter now represent the
holotype specimens of E. nuttallii. The behaviour of the staminate flowers, which is characteristic
of E. nuttallii, is also seen in H. verticillata, which may account for Muhlenberg describing his
plant as a variety of S. verticillata. Babington & Planchon (1848) made U. canadensis
synonymous with Anacharis nuttallii, and they suggested that British and Irish A. alsinastrum was
closely related to this species, if not the same plant. Caspary (1857, 1858) went further and
reduced A. nuttallii to the synonymy of E. canadensis.
Some years later, Small (1903) applied the name Philotria minor (Engelm.) Small to plants
with linear, acute leaves and flowers less than 3 mm in diameter, occurring in the central part of
the United States, whilst Rydberg (1906) described P. angustifolia Britton ex Rydb., which was
similar to P. minor, but with larger leaves and slightly larger flowers, and which occurred in the
central-eastern part of the United States. A third new combination was described in Britton &
Brown (1913), namely P. nuttallii Rydb. ex Britton & Brown. The name was used for plants
which were previously assigned to P. canadensis, but with more narrowly oblong and somewhat
acute leaves. The characteristic feature of all three species was the staminate flowers, which were
sessile and broke free at anthesis. St John (1920) amalgamated P. angustifolia and Elodea minor
(Small) Farw. (= P. minor) into one species, E. occidentalis (Pursh) St John. E. occidentalis was
described as having linear, flaccid leaves and a globose-apiculate staminate spathe which was c. 2
mm long. St John also recognized E. nuttallii, which differed from E. occidentalis in having
oblong-lanceolate, firm leaves and an ovate-lanceolate staminate spathe 5-6 mm long. However
he expressed doubt about the status of E. nuttallii, having seen specimens which showed the
morphology of nuttallii but the floral structure of occidentalis. St John (1962, 1965) combined the
two as E. nuttallii. The epithet, although later than occidentalis, had to be retained because
Pursh’s original use of the name was illegitimate and this invalidated its use by St John. It is of
interest that the characters used to delimit FE. nuttallii in St John (1920) are not used in his later
interpretation.
E. callitrichoidés has had a simple taxonomic history. It was first described by Richard (1814)
as the type species of the genus Anacharis. It was transferred to Elodea by Caspary (1857, 1858)
and remained in this genus as E. callitrichoides apart from a brief return to Anacharis suggested
by Victorin (1931). St John (1963) separated E. ernstiae from E. callitrichoides, and all European
material assigned to the latter was subsequently referred to this new species. However Cook &
Urmi-K6nig (1984) suggested that the characters used for separating the two species have no
diagnostic value. Therefore E. ernstiae should be referred back to E. callitrichoides, a view which
is accepted by the present author.
6 D. A. SIMPSON
MATERIALS AND METHODS
Extensive use was made of both living and herbarium material in the study. Herbarium specimens
were obtained from ABD, BM, CGE, K, LANC, OXF, P and RNG. A number of private herbaria
were also examined, including the Freshwater Biological Association (FBA) and herb R. Stokoe.
Although the work was concerned primarily with British and Irish material, specimens from
Europe and America were also looked at for comparison.
Particular emphasis was placed on the use, wherever possible, of recently gathered living
material. Such material was collected throughout the British Isles, but with a concentration in
north-western England, where a wide range of variation was noted. A full list of sites is given in
Simpson (1983), and voucher material for these collections is in LANC. Chromosome numbers
given in the species descriptions were obtained from counts which I made on British and Irish
material from 20 localities. These are also given in Simpson (1983). The counts were made from
root-tips, which were pre-treated in paradichlorbenzene for two hours and fixed in 1:3 acetic
alcohol overnight at 0°C., followed by hydrolysis in IN HC! at 60°C for eight minutes. They were
then stained in basic fuchsin for two hours and squashes prepared in either aceto-orcein or 45%
acetic acid. Ten root-tips per plant and five plants per species were examined at each locality.
Voucher material is in LANC.
(A)
(B)
Figure 2. £. canadensis. A, silhouettes of plants. B, silhouettes of leaves. C, perianth of female flower. Scale
bars = 1 cm.
TAXONOMY OF ELODEA 7.
KEY TO SPECIES
hk; Leaves linear-oblong, oblong-lanceolate, oblong, oblong-ovate or
ovate, rarely linear-lanceolate. Leaf apices broadly acute or obtuse,
rarely narrowly acute, (0-7—)0-8—2.3 mm wide c. 0-5 mm below the
nt tM op. UL ba ee oe oe ee RAS 1. E. canadensis
1 Leaves linear or linear-lanceolate. Leaf apices narrowly acute or
acuminate, 0-2—0-7(—0-8) mm wide c. 0-5 mm below the apex ......... Z
2 At least some leaves strongly recurved. Leaf lamina often strongly
twisted. Leaf margin teeth 60—90(—100) um long. Adventitious
root-tips (in living material) white or grey green. Sepals of female
flowers 16-2 San, Lome AIG. Fe Ga Pe) Za a 2. E. nuttallii
ps Leaves never strongly recurved. Leaf lamina rarely strongly twisted.
Leaf margin teeth (80—)110—140 um long. Adventitious root-tips (in
living material) red. Sepals of female flowers 3-1—4-3 mm long ....... 3. E. callitrichoides
SPECIES DESCRIPTIONS
1. ELODEA CANADENSIS Michx, Fl. Bor.-Amer., 1: 20 (1803). Udora canadensis (Michx)
Nutt., pro parte, Gen. N. Amer. Pl., 2: 242 (1818). Serpicula canadensis (Michx) Eaton, Man.
bot., 5th ed., 390 (1829). Anacharis canadensis (Michx) Planch. in Ann. Mag. Nat. Hist., ser. 2,
1: 85 (1848). Philotria canadensis (Michx) Britton in Science, 11 (2): 5 (1895). TYPE: Environs
de Montreal, Michaux (holotype: P). Fig. 2.
Anacharis alsinastrum Bab. in Ann. Mag. Nat. Hist., ser. 2, 1: 83 (1848).
Elodea latifolia Casp. in Mber. kgl. Pruss. Akad. Wiss., 46 (1857).
Elodea planchonii Casp. in Mber. kgl. Pruss. Akad. Wiss., 47 (1857). Philotria planchonii (Casp.)
Rydb. in Bull. Torrey bot. Club, 35: 462 (1908). Elodea canadensis var. planchonii (Casp.)
Farw. in Amer. Midl. Nat., 10: 203 (1927). Anacharis canadensis var. planchonii (Casp.)
Victorin in Contr. Lab. Bot. Univ. Montréal, 18: 40 (1931).
Elodea oblongifolia Casp. in Jb. wiss. Bot., 1: 467 (1858).
Philotria iowensis Wylie in Proc. Iowa Acad. Sci., 17: 82 (1910). Elodea iowensis (Wylie) Wylie in
Nat. Hist. Bull. Iowa State Univ., 6: 48 (1913).
A B Cc
aaa GN a YRS
LENS a5.
Yomuiltae \
aX Za x
i oEN Pana
ane NS oe \
Vigne a
D E F
Ficure 3. Leaf posture types. A, spreading. B, patent. C, erecto-patent. D, arcuate-deflexed. E, slightly
deflexed c. 2-4 mm from the leaf base. F, strongly recurved, with leaf bases often touching or overlapping the
stem.
8 D. A. SIMPSON
Dioecious. Lowermost leaves on stem decussate, ovate, 1-8—9-0 x 0-7—2-0 mm; median and upper
leaves in whorls of 3, linear-oblong, oblong, oblong-lanceolate, oblong-ovate, ovate, rarely
linear-lanceolate, sometimes weakly twisted, 4-5-17-0 x 1-4—-5-6 mm; leaf apices obtuse or
broadly acute, rarely narrowly acute, (0-7—)0-8—2-3 mm wide c. 0-5 mm below the apex; leaf
posture (Fig. 3) spreading, patent, erecto-patent or arcuate-deflexed, usually firm; leaf-margin
teeth 40—70(—80) um long. Female flowers with sepals oblong-elliptic, cucullate at apex, 1-9-2-8 x
0-6-1-7 mm, recurved, greenish-white, streaked with purple around apex and midrib; petals
elliptic-spathulate, 1-8—2-5 x 0-8-1-5 mm, strongly recurved, translucent, whitish; staminodes
linear, c. 1 mm long, white; stigmas strongly recurved or slightly bifid, flattened, 2-3—3-2 mm long,
sparsely papillose, the papillae (110—)120—215 um long, often purple. Male flowers similar to
female but with staminodes, stigmas and ovary absent; stamens 9, anthers bilocular. Flowering
period June-September. Adventitious root-tips white or grey-green. 2n = c. 24.
MILES 100
ee SS
|
t ¢
Pe. |
a \
» —_
’
4 A
4
FiGuURE 4. Distribution of E. canadensis in the British Isles.
TAXONOMY OF ELODEA 9
Habitat and distribution (Fig. 4). E. canadensis occurs on fine substrates at c. 0-15—4 m depth, in
unshaded, eutrophic to meso-oligotrophic water-bodies, where turbulence through water-flow or
wave action is minimal. It is, therefore, most frequently found in lowland ponds, lakes, canals,
slow-moving rivers and streams. It is also an early colonizer of new habitats, such as artificial lakes
and water-bodies recovering from pollution. EF. canadensis is widely distributed throughout most of
the British Isles, being absent mainly from upland areas in the west and north, where habitats are
generally unsuitable.
(B)
Ficure 5. E. nuttallii. A, silhouettes of plants. B, silhouettes of leaves. C, perianth of female flower. Scale bars
= 1 cm.
10 D. A. SIMPSON
2. ELODEA NUTTALLII (Planch.) H. St John in Rhodora, 22: 27—28 (1920). Anacharis nuttallii
Planch. in Ann. Mag. Nat. Hist., ser. 2, 1: 86 (1848). Philotria nuttallii (Planch.) Rydb. ex
Britton & Brown, Jllustr. Fl. n.-e. U.S., 2nd ed., 105 (1913). TYPE: Philadelphia, Nuttall
(holotype: BM). Fig. 5.
Serpicula verticillata L.f. var. B angustifolia Muhl., Cat. Pl. Amer. Sept., 84 (1813). Elodea
canadensis var. B angustifolia (Muhl.) Farw. in Amer. Midl. Nat., 10: 203 (1927).
Serpicula occidentalis Pursh, Fl. Amer. Sept., 33 (1814), nom. illegit. Elodea occidentalis (Pursh)
H. St John in Rhodora, 22: 27—29 (1920), nom. illegit.
Udora canadensis (Michx) Nutt., pro parte, Gen. N. Amer. Pl., 2: 242 (1818).
Philotria minor Small, Fl. s.-e. U.S., 47 (1903). Elodea minor (Small) Farw. in Rep. Mich. Acad.
Sci., 17: 181 (1916).
Philotria angustifolia Britton ex Rydb., Fl. Colorado, 15 (1906).
Ficure 6. Distribution of E. nuttallii in the British Isles.
TAXONOMY OF ELODEA 11
Dioecious. Lowermost leaves on stem decussate, ovate to linear-lanceolate, 2-8 x 0-5—5-0 mm;
median and upper leaves in whorls of 3—4(—5), linear or linear lanceolate, often strongly twisted,
5-5—35-0 x 0-8-3-0 mm; leaf apices acuminate or narrowly acute, 0-2—0-7(—0-8) mm wide c. 0-5
mm below the apex; leaf posture (Fig. 3) strongly recurved, spreading, patent, erecto-patent,
arcuate-deflexed, or slightly deflexed c. 2-4 mm from the leaf base, firm or flaccid; leaf-margin
teeth 60—90(-—100) um. Female flowers with sepals oblong-elliptic, ovate, cucullate at apex, 1-6—
2:5 x 0-9-1-8 mm, recurved, greenish-white streaked with purple around apex and midrib; petals
suborbicular-spathulate, 1-9-2-6 x 0-9-1-8 mm, strongly recurved, translucent, whitish; stami-
nodes linear, c. 1 mm long, white; stigmas strongly recurved, entire or slightly bifid, flattened,
2-2—3-:2 mm long, sparsely papillose, the papillae 60—100(—110) um long, sometimes purple. Male
flowers with perianth sessile within the globose spathe, the whole flower breaking free at anthesis
and floating; staminodes, stigmas and ovary absent; stamens 9, anthers bilocular. Flowering period
June—September. Adventitious root-tips white or grey-green. 2n = c.48.
Habitat and distribution (Fig. 6). E. nuttallii occurs in similar habitats to E. canadensis. Although
widely distributed in England, it is still less common than E. canadensis, but is continuing to
increase. In Wales it is known in v.cc. 35, 41, 42, 46 and 51, whilst in Scotland it has been recorded
from widely separated localities in v.cc. 72, 83 and 106. In Ireland it has been found in L. Neagh at
Drumenny, Co. Tyrone, v.c. H36, and has also been introduced into a pond at the National
Botanic Gardens, Glasnevin, Co. Dublin, v.c. H21.
(A)
(B)
Figure 7. E. callitrichoides. A, silhouettes of plants. B, silhouettes of leaves. C, perianth of female flower.
Scale bars = 1 cm.
12 D. A. SIMPSON
3. ELODEA CALLITRICHOIDES (Rich.) Casp. in Mber. kgl. Pruss. Akad. Wiss., 47—48
(1857). Anacharis callitrichoides Rich. in Mem. Inst. France, 12 (2): 7-8 (1814). TYPE:
Montivideo, Commerson (holotype: P-JU). Fig. 7.
Elodea ernstiae H. St John in Darwiniana, 12: 644 (1963).
Dioecious. Lowermost leaves decussate, ovate to linear-lanceolate, 3-0-14-0 x 0-5-1-2 mm;
median and upper leaves in whorls of 3, linear or rarely linear-lanceolate, rarely strongly twisted,
9-25 x Q-7—2-:2 mm; leaf apices acuminate rarely narrowly acute, 0-2—0-6 mm wide c. 0-5 mm
below apex; leaf posture (Fig. 3) spreading, patent, erecto-patent, arcuate-deflexed or slightly
deflexed c. 2-4 mm from the leaf base, usually flaccid; leaf-margin teeth (80—)110—140 um long.
Female flowers with sepals linear-oblong, linear-elliptic, cucullate at the apex, 3-1-4-3 x 0-9-1-6
r
gst
fy
iT
Ficure 8. Distribution of E. callitrichoides in the British Isles. Map includes the 10 km squares from which the
plant now appears to be extinct.
TAXONOMY OF ELODEA 13
mm, spreading or slightly recurved, grey-green or greenish-white, heavily streaked with purple
around the apex and midrib; petals oblong-elliptic, 2.9-3-8 x 1-0—1-8 mm, spreading or slightly
recurved, translucent, whitish; staminodes linear, 1-5—3-8 mm long, white; stigmas spreading,
deeply bifid, terete, 4.2-6-6 mm long, densely papillose, the papillae 120-215 um long, purple.
Male flowers similar to female but with staminodes, stigmas and ovary absent; stamens 9, anthers
bilocular. Flowering period October—April. Adventitious root-tips red. 2n=c.32.
Habitat and distribution (Fig. 8). E. callitrichoides has been found in canals, ponds and slow-
moving rivers. It is, however, a casual species and is currently known to occur in only two localities
in v.cc. 13 and 42. It has not been recorded from either Scotland or Ireland.
ACKNOWLEDGMENTS
I wish to thank Dr G. Halliday for guidance during the course of the work and Prof. D. A. Webb
for advice on nomenclature. Thanks are also due to the many correspondents who sent records,
and to the Biological Records Centre for preparing the distribution maps.
REFERENCES
ADANSON M. (1763). Familles des plantes. Paris.
BaBINGTON, C. C. & PLANCHON, J. E. (1848). On Anacharis alsinastrum, a supposed new British plant. Ann.
Mag. Nat. Hist., ser. 2, 1: 81-88.
Britton, N. L. (1895). Philotria canadensis. Science, 11 (2): 5.
Britton, N. L. & Brown, A. (1913). An illustrated Flora of the northern United States, Canada and the British
possessions from Newfoundland to the southern boundary of Virginia and from the Atlantic Ocean
westward to the 102nd meridian., 2nd ed. New York.
Caspary, R. (1857). Conspectus systematicus Hydrillearum. Mber. kgl. Pruss. Akad. Wiss.: 39-51.
Caspary, R. (1858). Die Hydrilleen (Anacharideen Endl.). Jahrb. wiss. Bot., 1: 377-513.
Cook, C. D. K. & Urmi-KOnic, K. (1984). Elodea ernstae back to E. callitrichoides. Watsonia, 15: 117.
Danpy, J. E. (1980). Elodea, in Tutin, T. G. et al., eds. Flora Europaea, 5: 4-5. Cambridge.
De Jussieu, A. L. (1789). Genera plantarum secundum ordines naturales disposita. Paris.
Douc tas, D. (1880). Notes on the Water Thyme (Anacharis alsinastrum Bab.). Hardwickes Sci. Gossip, 16:
227-229.
Hooker, W. J. (1842). Editorial note. J. Bot., Lond, 4: 432.
Kuni, H. (1982). The critical water temperature for the active growth of Elodea nuttallii (Planch.) St John.
Jap. F. Ecol. ,/327 111112.
LAwRENCE, D. K. (1976). Morphological variation of Elodea in western Massachusetts. Rhodora, 78: 739-749.
MicnHaux, A. (1803). Flora Boreali-Americana. Paris.
MUHLENBERG, G. H. E. (1813). Catalogus plantarum Americae septentrionalis. Lancaster.
NUTTALL, T. (1818). The genera of North American plants and a catalogue of the species to the year 1817.
Philadelphia.
PLANCHON, J.-E. (1849). Note additionelle—Egeria. Ann. Sci. Nat. 3° sér., Bot., 11: 79-81.
PursH, F. T. (1814). Plants of North America. London.
RAFINESQUE, C. S. (1818). Review of Pursu, F. T., Plants of North America. Am. Monthly Mag., 2: 175.
RicHarD, L. C. M. (1814). Sur les Hydrocharidées. Mem. Inst. de France, 12 (2): 1-4.
RybBERG, P. A. (1906). The flora of Colorado. Agr. Exp. Sta. Colo. Agr. Coll. Bull., 100: 15.
RybBerG, P. A. (1908). Notes on Philotria Raf. Bull. Torrey bot. Club, 35: 457-465.
St JouNn, H. (1920). The genus Elodea in New England. Rhodora, 22: 18-29.
St Joun, H. (1962). Monograph of the genus Elodea 1. Res. Stud. Wash. St. Univ., 30: 19-44.
St JoHN, H. (1963). Monograph of the genus Elodea 3. Darwiniana, 12: 639-652.
St JoHN, H. (1964). Monograph of the genus Elodea 2. Caldasia, 9: 95-113.
St JoHNn, H. (1965). Monograph of the genus Elodea 4. Rhodora, 67: 1-35, 155-181.
Santos, J. K. (1923). Differentiation among chromosomes in Elodea. Bot. Gaz., 75: 42-59.
SANTOS, J. K. (1924). Determination of sex in Elodea. Bot. Gaz., 77: 353-376.
SCANNELL, M. J. P. & Wess, D. A. (1976). The identity of the Renvyle Hydrilla. Ir. Nat. J., 18: 327-331.
Simpson, D. A. (1983). Experimental taxonomic studies of Elodea Michx in the British Isles. Ph.D. thesis,
University of Lancaster.
14 D. A. SIMPSON
Stmpson, D. A. (1984). A short history of the introduction and spread of Elodea Michx in the British Isles.
Watsonia, 15: 1-9.
SMALL, J. K. (1903). Flora of the south-eastern United States. New York.
SPRENGEL, C. P. J. (1828). Systema vegetabilium, 16th ed. Berlin.
STEUDEL, E. G. (1841). Nomenclator botanicus, 2nd ed. Stuttgart.
VENTENAT, E. P. (1799). Tableau du regne végétal selon la methode de Jussieu. Paris.
Victorin, M. (1931). L’Anacharis canadensis. Histoire et solution d’un imbroglio taxonomique. Contr. Lab.
Bot. Univ. Montréal, 18: 1—43.
WEsB, D. A. (1977). An Irish Flora, 6th ed. Dundalk.
Wo rr, P. (1980). Die Hydrilleae (Hydrocharitaceae) in Europa. Gott. Flor. Rundbr., 14: 33-56.
Wy le, R. B. (1904). The morphology of Elodea canadensis. Bot. Gaz., 37: 1-22.
(Accepted February 1985)
Watsonia, 16, 15—19 (1986) 15
Variation in the colour of the Keel petals in
Lotus corniculatus L., 2.
Clines in Yorkshire and adjacent counties
T. J. CRAWFORD
Department of Biology, The University, York, YOI 5DD
and
D. A. JONES
Unit of Genetics, The University, Hull, HU6 7RX
ABSTRACT
The distribution of the dark-keeled morph of Lotus corniculatus L. (Leguminosae) has been surveyed at 182
sites in Yorkshire and neighbouring counties. The dark morph is rare in the west but increases eastwards,
attaining high frequencies on the eastern coast. The slope of the cline is probably irregular, having steeper
phases approximately 30 km from each coast. The cline is weak south of the study area, but it is pronounced
north of the River Humber. The selective agent responsible for variation in morph frequencies has yet to be
identified.
INTRODUCTION
The cyanogenesis polymorphism of Lotus corniculatus L. (Birdsfoot Trefoil) is well known and a
number of selective forces partly responsible for maintaining the polymorphism have now been
identified (Jones 1977; Compton et al. 1983a). Less well documented, although the variation is
visually obvious, is the polymorphism for keel petal colour. Plants may bear either flowers with
completely yellow keel petals (‘light’) or flowers which have red-brown tips to the keels (‘dark’).
The intensity of dark pigmentation varies greatly between the dark-keeled plants, ranging from a
rich chocolate-brown encompassing the tip to a pale, diffuse, red pigmentation mainly confined to
the front surface of the keel tip. The latter pattern has sometimes been described as ‘pale’ but as
the distinction between pale and dark keels is somewhat subjective the two groups are together
classed as darks in the present paper.
Keel colour is determined by a pair of alleles showing tetrasomic inheritance, dark keel being
dominant to light (Hart & Wilsie 1959; Buzzell & Wilsie 1963; Ramnani & Jones 1984). Breeding
work on North American cultivars (Bubar & Miri 1965) has suggested an epistatic interaction
between loci involved in the expression of keel colour and cyanogenesis so that an excess of light-
keeled acyanogenic plants is expected. However, in spite of extensive surveys of natural
populations in Britain and continental Europe (Jones & Crawford 1977; Compton et al. 1983a;
Compton et al. 1983b), no association between cyanogenesis and keel colour has been found; nor
has breeding work on British material provided evidence for any epistatic interaction (Ramnani &
Jones 1984).
Rather surprisingly, it seems that the dark keel tip was first recorded in 1955 and was regarded as
an unusual variant in natural populations and as occurring at moderate frequencies in a minority of
cultivars (Hart & Wilsie 1959). Attention was drawn to the widespread nature of the
polymorphism, at least in western Europe, by Jones & Crawford (1977). Apart from the western,
coastal, dune region of the Netherlands, where some populations monomorphic for light keels
were found, polymorphism with varying frequencies of dark-keeled plants was almost universal. In
‘joquiAs yoed JopuN UMOYSs sI pojdwies sjueyd jo
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PETAL COLOUR POLYMORPHISM IN LOTUS 17
England and Wales there was evidence for a clinal increase in frequency of the dark-keeled morph
with latitude although there was no evidence for a similar cline in continental Europe. In Scotland,
Abbott (1981) found that western populations generally have lower frequencies of dark-keeled
plants than do those of central and eastern regions. Taking his data together with those from
England and Wales, he suggested that the dark-keeled morph was relatively rare throughout
western regions of the British Isles, but to the east the frequency increased with latitude so that it
became the commoner morph in the north-east.
In this paper we describe the status of the polymorphism in Yorkshire and adjacent counties.
Although the detailed picture remains unclear it seems that rapid clinal change occurs in this region
in both west-east and north-south directions.
METHODS
Plants were scored for keel colour at 182 locations within the area shown in Fig. 1. Because L.
corniculatus can spread by vegetative reproduction, only plants separated by at least 1 m were
scored. Although keel colour could be used to discriminate between closely adjacent plants, such a
procedure would introduce a bias against extreme frequencies. At least 100 plants were scored at
41%, and at least 50 plants at 69% of the sites.
Samples were combined within National Grid 10-km squares by the following procedure. Where
the same site had been sampled more than once, the numbers of light and dark-keeled plants were
summed over all replicate samples. The totals were then adjusted downwards to yield a single
composite sample, equal in size to the largest individual sample but with the proportions of the two
morphs reflecting those in the unadjusted totals. Data from different sites within a 10-km square
were combined by simple summation, using adjusted totals for sites sampled more than once.
RESULTS AND DISCUSSION
Fig. 1 shows the proportion of dark-keeled plants in each 10-km square for which any information
is available. The total number of plants scored, counting only the largest of replicate samples, is
shown under each pie symbol.
The difference in the frequency of dark-keeled plants between western and eastern areas is
striking, particularly in the more northern regions. For individual sites, both Red Bank (GR 34/
471.678) in the west, with 0 out of 50 plants dark, and Flamborough (GR 54/240.722) in the east,
with 99 out of 103 plants dark-keeled, represent extremes greater than any previously recorded in
the British Isles. The results so far available suggest that the slope of the west-east cline is irregular.
Dark-keeled plants remain rare for about 30 km from the western coast. There then seems to be a
relatively sudden increase in darks, followed by a more gradual increase eastwards, until about 30
km from the eastern coast the dark keel frequency again increases rapidly. Whether these changes
relate to proximity to the coast or more simply to longitude remains to be seen. Certainly it should
be possible to distinguish between the alternatives by intensive sampling in western promontories
such as North Wales, the Lake District and Galloway.
Sampling has been less intense in the southern part of the survey area, but it can be seen from
Fig. 1 that the west to east increase in dark-keeled plants becomes more marked near to, but not
necessarily coincident with, the River Humber. All available samples, including those from Jones
& Crawford (1977), south of the National Grid 400 km Northing have been combined for each
100-km square, treating replicate samples as above. Table 1 shows that the eastwards increase in
dark-keeled plants is reduced south of the River Humber. The data in Table 1, together with those
presented in Fig. 1, confirm the suggestion (Jones & Crawford 1977; Abbott 1981) that the
frequency of the dark morph, rare in the west, increases with latitude on the eastern side of the
British Isles. It is clear, however, that the increase is not continuous, but occurs rather suddenly at
a considerable distance south of the Scottish border.
Combining samples within 10-km squares clarifies the general picture at the expense of local
detail. Although there is some local heterogeneity between sites within 10-km squares, the
variation is certainly small compared with the scale of the clinal changes.
18 T. J. CRAWFORD AND D. A. JONES
TABLE 1. PERCENTAGE FREQUENCY OF DARK-KEELED LOTUS CORNICULATUS AND, IN
BRACKETS, TOTAL SAMPLE SIZE FOR NATIONAL GRID 100-KM SQUARES
Easting
Northing 1 2 3 4 5 6
3 12(538) 26(159) 47(2028) 35(392)
2 2(100) 22(32) 35(3333) 43(633) 35(120)
1 30(672) 25(64) 30(221)
0 22(50) 4(29) 16(186)
The nature of the selective forces that might maintain the polymorphism remains a mystery.
Jones & Crawford (1977) found no differences between the morphs for winter survival or for seed
production in experimental plots. In natural populations, Jones et al. (in prep.) could find no
associations between flower, pod or seed production with keel colour morph, nor with flower
damage or seed predation by phytophagous insects. They also showed that pollinating insects
failed to discriminate between the morphs in experimental populations. They suggest, however,
that the frequency of dark-keeled plants may increase slightly as the flowering season progresses.
In this context it is worth noting that the sites at the western end of the cline were in general
sampled later than those to the east. There is some evidence (Buzzell & Wilsie 1963; Jones &
Crawford 1977) that darks occur at a greater frequency in progeny raised under glass than in the
parental populations from which the seeds were collected. This possibility deserves further
attention. An explanation related to temperature has some a priori appeal because greater solar
radiation absorbed by dark-keeled flowers may influence pollen, ovule or seed development. On
the other hand, there is considerable altitudinal variation along the cline reported in this paper and
it is clear that altitude itself has little effect on the morph frequencies.
Although dark-keeled plants are common at some inland locations in Europe (Jones &
Crawford 1977), the regularly high frequencies of dark-keeled plants in north-eastern Britain are
exceptional. It seems that the light morph more usually predominates and it may be inferred that
selection commonly operates in favour of light keels. Nevertheless, populations monomorphic for
light keel colour are rare and monomorphism for dark keels has yet to be recorded. It may be that
some form of frequency-dependent selection is operating. Alien introductions, particularly on
recently seeded roadside verges, are sometimes encountered. These plants differ from native
plants in a number of respects (Jones 1977, 1982; Bonnemaison & Jones 1986), most strikingly in
their long internode lengths. Their keel colour frequencies are sometimes distinct from the
surrounding native populations and they have been excluded from this survey. Monitoring of alien
populations may reveal changes in their keel colour frequencies over time and provide important
information on the magnitude of the selective forces involved.
We would be grateful to receive information on this polymorphism from any localities. Care
should be taken to exclude Lotus uliginosus, which is monomorphic for yellow keels. We require
grid reference (as accurately as possible), date, brief description of habitat (e.g. chalk grassland,
roadside verge) including any indications that the plants may be alien, and numbers (not
proportions) of light and dark-keeled morphs. Plants should be sampled at intervals not less than
| m and, although sample sizes of at least 100 are ideal, smaller samples are still useful.
ACKNOWLEDGMENTS
We thank Hazel Beacon and Harry Vallack for drawing Fig. 1. We are also grateful to the
N.E.R.C. for supporting part of this work.
PETAL COLOUR POLYMORPHISM IN LOTUS te)
REFERENCES
AssotTrt, R. J. (1981). The keel petal colour polymorphism of Lotus corniculatus L. in Scotland. New Phytol.,
88: 549-553.
BONNEMAISON, F. & Jones, D. A. (1986). Variation in alien Lotus corniculatus L. 1. Morphological differences
between alien and native British plants. Heredity, (in press).
Busar, J. S. & Miri, R. K. (1965). Inheritance of self-incompatibility and brown keel tip in Lotus corniculatus
L. Nature, 205: 1035-1036.
BuzzELL, R. I. & WitsiE, C. P. (1963). Genetic investigation of brown keel tip colour in Lotus corniculatus L.
Crop Sci., 3: 128-130.
Compton, S. G., BEESLEY, S. G. & Jones, D. A. (1983a). On the polymorphism of cyanogenesis in Lotus
corniculatus L. TX. Selective herbivory in natural populations at Porthdafarch, Anglesey. Heredity, 51:
537-548.
Compton, S. G., NEwsomeE, D. & Jongs, D. A. (1983b). Selection for cyanogenesis in the leaves and petals of
Lotus corniculatus L. at high latitudes. Oecologia, 60: 353-358.
Hart, R. H. & Wisi, C. P. (1959). Inheritance of a flower character, brown keel tip, in Lotus corniculatus L.
Agron. J., 51: 379-380.
Jones, D. A. (1977). On the polymorphism of cyanogenesis in Lotus corniculatus L. VII. The distribution of
the cyanogenic form in Western Europe. Heredity, 39: 27-44.
Jones, D. A. (1982). Lotus corniculatus in the roadside verge. Lotus Newsletter, 13: 3-6.
Jones, D. A. & CRAWFORD, T. J. (1977). Variation in the colour of the keel petals in Lotus corniculatus L. I.
The polymorphism in Western Europe. Heredity, 39: 313-325.
Ramnanl, A. D. & Jones, D. A. (1984). Inheritance of the brown keel tip character in Lotus corniculatus L.
Bangladesh J. Bot., 13: 52-59.
(Accepted April 1985)
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Watsonia, 16, 21-23 (1986) 21
Variation in Senecio jacobaea L. (Asteraceae) in the British Isles
J. W. KADEREIT
Institut fiir systematische Botanik, Universitat Heidelberg, Germany
and
PD SEBEL
Botany School, Downing Street, Cambridge, CB2 3EA
ABSTRACT
The variation in Senecio jacobaea L. in the British Isles is described. Two subspecies, subsp. jacobaea and
subsp. dunensis (Dumort.) Kadereit & P. D. Sell, stat. nov., the former with two varieties, var. jacobaea and
var. condensatus Druce, are recognized. The nomenclature and descriptions are formally set out and a list of
herbarium specimens and published records are given for subsp. dunensis and var. condensatus.
VARIATION
The most significant variation within Senecio jacobaea L. in Britain and Ireland seems to occur in
coastal populations. The normal plant of inland grassy places is tall, with widely-spaced leaves and
a large open inflorescence of capitula which usually have ray florets and an outer row of glabrous
achenes. In some coastal localities are to be found populations of shorter plants with dense
arachnoid hairs on the involucres and peduncles, the ray florets absent or rudimentary and the
outer row of achenes hairy. In the British Isles such populations have been recorded from the
Scottish and Irish coasts and in continental Europe they occur on the Baltic or North Sea coasts of
southern Norway, southern Sweden, E. Germany, Poland, Denmark, the Friesian Islands,
Heligoland, the Netherlands and Belgium. Meijden (1976) has suggested that this taxon may be
worth the rank of subspecies and we support this.
In other British coastal localities, particularly on dunes and shingle, occur populations of plants
which have a greater tendency to produce a swollen area just below the basal leaves (described by
Harper & Wood (1957) as a crown); they are shorter with up to three stems and have a dense
inflorescence with short-liguled florets. This variant, which we have not seen from outside the
British Isles, we consider best placed as a variety of subsp. jacobaea.
Further variation which occurs in subsp. jacobaea is not at the moment considered worthy of
recognition. Occasional plants of inland populations are without ray florets. Plants with narrow
ligules have been called var. stenoglossus Brenan & Simpson (Brenan & Simpson 1949), but do not
seem to fit in an ecological niche, although they may be more frequent in the west of the British
Isles than the east. Plants from chalk grassland tend to be more slender with very finely cut leaves,
a persistent basal rosette and small capitula. A plant collected by G. Crompton on a sandy path
through machair, Daleburgh, S. Uist on 12th August 1964 (CGE) differs from any other specimen
we have seen in having a clothing of short glandular hairs.
NOMENCLATURE
Senecio jacobaea L. was first described by Linnaeus on page 870 of Species Plantarum in 1753. The
diagnosis is taken partly from the Hortus Cliffortianus and is partly new. It is reasonable to assume
22 J. W. KADEREIT AND P. D. SNELL
the specimen in the Linnaean herbarium, Savage Catalogue 996/44, was used when making the
amendments to Hortus Cliffortianus and it is designated as the lectotype.
Var. nudus Weston (1777) has been used by some authors for the coastal subspecies but there is
no evidence as to whether Weston referred to coastal or inland plants. Var. flosculosus Lam. &
DC. (1805) was said in the original publication to grow on dunes and other sandy places and is
presumably our coastal subspecies, although neither we nor Meijden (1976) have seen specimens
of it from France. S. flosculosus Jordan (1848), described without reference to the earlier
published variety, was based on plants from shady pastures near Lyon and is not referable to the
coastal populations. Var. discoideus Wimmer & Grab. (1829) was described from an inland site
near Leobschiitz (Glubczyce), Poland. S. dunensis Dumort. (1827) was described from coastal
dunes in Belgium and is thought to be the most appropriate name to take up for the maritime
subspecies.
The types of both var. condensatus Druce (1916) and var. abrotanoides J. Murr. ex Druce (1926)
are referable to the variant with condensed habit and short ligules, the former having priority.
CHROMOSOME NUMBER
Bocher & Larsen (1955) referred to the ligule size of plants collected from Ireland of which they
counted the chromosomes. Plants with long, narrow ligules from dunes near Lady’s Island Lake,
Co. Wexford, with 2n = 40 were probably referable to var. jacobaea of subsp. jacobaea. The plants
from a roadside near Skull Harbour, Co. Cork, with 2n = 40 and possibly those from dunes near
Kincasslagh, Co. Donegal, with 2n = 32 could belong to var. condensatus. Both have smaller than
usual ray florets. Both 2n = 40 and 2n = 80 have been recorded for S. jacobaea by other authors.
SENECIO JACOBAEA L., Sp. Pl., 870 (1753). Described as “Habitat in Europae pascuis”’
(Lectotype: Savage Cat. 996/44, LINN).
S. jacobaea var. nudus Weston, Bot. Univ., 2nd ed., 3: 641 (1777). Described without habitat or
locality.
(a) subsp. JACOBAEA
Stems 1-3, up to 150 cm; peduncles with more or less numerous arachnoid hairs; ray florets usually
present; outer row of achenes glabrous.
(i) var. JACOBAEA
S. jacobaea var. discoideus Wimmer & Grab., Fl. Siles. 2 (2): 153 (1829). Described from near
Leobschiitz (Glubczyce), Poland.
S. flosculosus Jordan, Cat. Jard. Dijon, 30 (1848). Described from near Lyon, France.
S. jacobaea var. stenoglossus Brenan & Simpson in Proc. R. Ir. Acad., B, 52: 69 (1949). TYPE:
Hedgebank in a lane north of Banbridge, Co. Down, 5 August 1939, J. P. M. Brenan 5730
(Holotype: K).
Stems not markedly swollen below the basal leaves, usually solitary, up to 150 cm; leaves rather
widely spaced; inflorescence often large and open; ligules 7-9 X 1-7—3-0 mm, rarely absent.
(ii) var. CONDENSATUS Druce in Rep. botl Soc. Exch. Club Br. Isl., 4: 201 (1916). TYPE:
Dunes and sea cliffs, Arbroath, Forfar, July 1915, G. C. Druce (Holotype: OXF).
S. jacobaea var. abrotanoides J. Murr ex Druce in Rep. botl Soc. Exch. Club Br. Isl. , 7: 774 (1926).
TYPE: Between Poolewe and Dundonnell, W. Ross, August 1925, G. C. Druce (Holotype:
OXF).
Stems markedly swollen below the basal leaves, 1—3, up to 40(—45) cm; internodes shorter, leaves
and inflorescence more dense; ligules 5—7(—9) 1-2 mm.
Distribution
v.c. 25: Shingle by the sea, Shingle Street, E. Suffolk, 2 August 1976, P. D. Sell 76/195a (CGE).
Shingle beach between Aldeburgh and Thorpeness, E. Suffolk, 16 September 1975, P.D.
Sell 75/115 (CGE).
v.c. 27; Old dunes, Winterton Ness, E. Norfolk, 30 July 1962, P.D. Sell 62/465 (CGE).
VARIATION IN SENECIO JACOBAEA 23
v.c. 28: Shingle between Cley Beach and Blakeney Point, W. Norfolk, 31 July 1975, P.D. Sell
75/104 (CGE).
v.c. 68: Dunes by the sea, Ross Links, north of Bamburg, Northumberland, 14 July 1982, P.D.
Sell 82/142 & C.M. Pannell (CGE).
v.c. 90: Dunes and sea cliffs, Arbroath, Forfar, July 1915, G.C. Druce (OXF).
v.c. 97: Shingle between Sallachan Point and Corran Ferry, Ardgour, Argyll, 30 June 1976, P. D.
Sell 76/169 (CGE).
v.c. 105: Mellon Charles, W. Ross, July 1926, G.C. Druce (OXF). Between Poolewe and
Dundonnell, W. Ross, Aug. 1925, G.C. Druce (OXF).
(b) subsp. DUNENSIS (Dumort.) Kadereit & P. D. Sell, stat. nov. S. dunensis Dumott., Fl. Belg.,
66 (1827). Described from dunes on the Belgian coast.
S. jacobaea var. flosculosus Lam. & DC. in Lam., Fl. Fr., 3rd ed., 4: 163 (1805). Described from
dunes and sandy places in France.
Stems solitary, up to 30 (to 60) cm; peduncles often with dense arachnoid hairs; ray florets absent
or rudimentary; outer achenes hairy.
Distribution
v.c. 108: Dunes by Farr Bay, W. Sutherland, 25 July 1959, P.D. Sell 59/164, N.D. Simpson & C.
West (CGE). Strathy Bay, south of Strathy Point, W. Sutherland, 23 July 1982, P.J.O.
Trist 107/30/82 (CGE).
v.c. 111: Shell-sand, grassy pasture at seashore, near North Jetty, Copinsay, Orkney, 5 August
1933, H.H. Johnstone 4816 (OXF).
v.c. H1: Banna sandhills, Kerry, July 1888, R. W. Scully (OXF). Fermoyle, Dingle Peninsula, N.
Kerry, August 1955, J.L. Farquharson (BM). “Abundant on most of the Kerry
sandhills, to the almost total exclusion of the rayed form’”’ (Scully 1916).
v.c. H9: “On Aran and on most of the Connemara dunes the variant without ligulate florets
predominates, but every grade of intermediate between it and the typical form is usually
found nearby” (Webb & Scannell 1983, p. 115).
v.c. H12: Rosslare, Wexford, 17 September 1926, G.C. Druce (BM). Rosslare Spit, October 1980,
observed by D.A. Webb & J.R. Akeroyd (J.R.A. pers. comm.).
v.c. H16: Dog’s Bay, Roundstone, Co. Galway, 9 September 1955, C.E. Raven (CGE).
v.c. H27: Sandhills, Mullet, W. Mayo, July 1905 (BM). Dunes south of Annagh Head, Mullet
peninsula, W. Mayo, 2 June 1979 (Curtis et al. 1981).
REFERENCES
ALLEN, D. E. (1961). Senecio jacobaea var. stenoglossus Brenan & Simpson. Proc. bot. Soc. Br. Isl., 4: 160.
BOcHER, T. W. & LarsEN, K. (1955). Chromosome studies on some European flowering plants. Bot. Tidsskr.,
52: 125-132.
BRENAN, J. P..M. & Simpson, N. D. (1949). The results of two botanical journeys in Ireland in 1938-9. Proc.
Roy. Irish Acad., B., 52: 57-84.
Curtis, T. G. F., McGoucu, H. N. & AKERoyp, J. R. (1981). The flora of the Mullet peninsula, West Mayo
(H27). Bull. Ir. biogeog. Soc., 5: 38-46.
Harper, J. L. & Woop, W. A. (1957). Senecio jacobaea L., in Biological Flora of the British Isles. J. Ecol. , 45:
617-637.
MEUDEN, R. VAN DER (1976). Het verspreidingagebied van Senecio jacobaea L. var. nudus Weston. Gorteria, 8:
57-61.
ScuLLy, R. W. (1916). Flora of County Kerry. Dublin.
Wess, D. A. & SCANNELL, M. J. P. (1983). Flora of Connemara and the Burren. Cambridge.
(Accepted May 1985)
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Watsonia, 16, 25—30 (1986) 25
Two natural hybrids in Ranunculus L. subgenus Batrachium
(DC.) A. Gray
S. D. WEBSTER
Department of Botany, The University, Whiteknights, Reading, RG6 2AS
ABSTRACT
Two natural Ranunculus hybrids are reported from southern Britain, whose identities have been elucidated by
their chromosome numbers. The hybrid recorded from S. Hants., v.c. 11, has the triploid complement (2n =
24) consistent with the parentage R. omiophyllus Ten. X R. peltatus Schrank, whilst the tetraploid complement
(2n = 32) in plants from S. Devon, v.c. 3, indicates R. baudotii Godr. X R. peltatus.
INTRODUCTION
In groups such as Ranunculus L. subgenus Batrachium (DC.) A. Gray, in which both phenotypic
plasticity and hybridization are rife, unusual or intermediate plants have sometimes been dismissed
as phenotypic variants of the ‘nearest’ species or explained away as hybrids on rather inconclusive
grounds. Although chromosome numbers are of limited taxonomic value in this group (since
several species exist at a number of ploidy levels, and several species share each chromosome
number), there are cases in which they provide important evidence in interpreting morphological
data and in resolving the identity of hybrids and intermediates (Sgrensen 1955; Cook 1966; Turala
& Wolek 1971). Two natural hybrids are reported whose chromosome numbers have proved useful
in this context. Vouchers are deposited in the herbarium of the Botany Department, University of
Reading (RNG).
THE HYBRIDS
R. OMIOPHYLLUS TEN. X PELTATUS SCHRANK FROM HOLMESLEY BOG
This plant was collected in June 1982 from Holmesley Bog in the New Forest (S. Hants., v.c. 11,
GR 41/22.01), a site which has been popular with collectors of batrachian Ranunculi over many
years and has supported at least four other members of the group including one hybrid. The most
striking features of this plant are that it has laminar leaves which are very similar to those of R.
omiophyllus, and, like this species, has lunate nectar pits, but, unlike R. omiophyllus, it also
possesses capillary leaves and bears larger flowers (Fig. 1).
In these respects the plant resembles very closely a hybrid from Copthorne Common (E. Sussex,
v.c. 14, GR 51/31.39) which was first discovered by Mr T. Hilton in 1896 and described by Groves
& Groves (1901) as R. Hiltonii (Fig. 2). Later, Williams (1926) reported the hybrid flowering at the
same site, and many specimens of it exist in various herbaria, but Prof. C. D. K. Cook was unable
to re-find it at Copthorne in 1956, and concluded that it had become extinct there. A curious
feature of the Copthorne hybrid was its high fertility. This was mentioned in the Groves’ paper,
and herbarium material studied by Cook (1966) showed nutlets with normal embryos and pollen
containing about 90% of well-formed grains which were larger than those of either parent. From
this evidence, Cook (1966) suggested that R. x hiltonii was an amphidiploid which had arisen from
R. omiophyllus X peltatus.
In contrast, the Holmesley hybrid is highly sterile (fewer than 1% of well-formed pollen grains,
and no seeds set). Mitotic root-tip squashes revealed a triploid chromosome complement (2n = 24)
for this plant (Fig. 3) — the first recorded count for this hybrid. This is consistent with the parentage
S. D. WEBSTER
26
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auioyjdoy wo (muoyiy x “Yy) snipyad x snyXydomuo -y “7 FANDIA
‘Bog Aajsowyjox wor snjpyad x snjyxkydomuo -y Jo ayanoyjis “][ FANS
HYBRIDS IN RANUNCULUS SUBGENUS BATRACHIUM 27
Ficure 3. Root-tip mitosis in R. omiophyllus x peltatus from Holmesley Bog. 2n = 24.
R. omiophyllus (2n = 32) X peltatus (2n = 16) and it also adds circumstantial evidence supporting
the hypothesis that the fertile hybrid from Copthorne was an amphidiploid (2n = 6x = 48) of the
same parentage.
The Holmesley plant was found at the margin of a New Forest stream in a marshy depression in
an area of closely cropped turf surrounded by heathland dominated by Ulex europaeus L. and
Calluna vulgaris (L.) Hull. It is regularly disturbed by humans, horses and cattle, and supports an
interesting collection of other marsh plants. Both parents were found near the site. A description
of the plant is given below.
Prostrate perennial herb. Leaves laminar or divided into capillary segments. Laminar leaves
opposite or alternate; petiole 30-120 mm long; lamina up to 19 mm long and 26 mm wide, reniform
to sub-orbicular, with lobes narrowest at their base, margin crenate. Capillary leaves alternate;
petiole 12-30 mm; lamina globose with few (15-60) segments. Segments rigid or flaccid; divergent,
fine or flattened. Intermediate leaves invariably present. Stipules ovate, much resembling those of
R. omiophyllus, adnate to petiole for half their length or less, apex obtuse. Pedicel in fruit 21-50
mm, shorter than petiole of opposed laminar leaf. Sepals up to 3 (rarely 4) mm long, spreading,
caducous, green. Petals 3—8 mm long, obovate, contiguous or not; nectar pits lunate; stamens 13-
20; carpels 25—43, glabrous or hairy. Receptacle hairy.
One of the most extraordinary features of this plant is that it overwinters in the heterophyllous
state. Plants collected in February 1984 from the same site had laminar, capillary, and intermediate
leaves.
R. BAUDOTI GODR. HYBRID FROM SLAPTON LEY
This plant was collected in June 1982 in Slapton Ley (S. Devon, v.c. 3, GR 20/82.43), a natural
lake about 3 km long and 0-5 km wide that lies parallel to the South Devon coast between Torcross
and Slapton and about 150 m from the sea. R. tripartitus DC. (“‘occasional’’), R. circinatus Sibth.
(“rare”), R. trichophyllus Chaix and R. peltatus (both “locally abundant”) and R. baudotii
(“‘occasional’’) were reported as occurring here in 1968 (Brookes & Burns 1979). The hybrid plant
was found during an attempt to re-find R. baudotii. This plant (Fig. 4) is superficially very similar to
the latter species, having short, rigid capillary leaves, laminar leaves which are deeply trifid, and
glabrous carpels. However, the petals are atypical of R. baudotii, being short, broad and rounded,
and, more importantly, the nectar pits are shortly pyriform: they have an upper border, unlike the
simple lunate nectar pit of that species.
Presumably, R. baudotii is involved as one parent; R. aquatilis L. (2n = 48) and R. peltatus (2n =
16, 32, 48) are possible candidates for the other. A natural hybrid between R. baudotii and R:
aquatilis is already known (Erikson 1905; S¢rensen 1955), and has been found in coastal regions of
England, Denmark, France and Sweden (Stace 1975). The latter hybrid is intermediate between its
parents in floral and vegetative characteristics except that the sepals are blue (as in R. baudotii),
28 S. D. WEBSTER
Figure 4. Silhouette of specimen of R. baudotii hybrid from Slapton Ley with drawing of petal showing shortly
pyriform nectar pit.
Sj
= d=
10jum
FiGURE 5. Root-tip mitosis in R. baudotii hybrid from Slapton Ley. 2n = 32.
{
HYBRIDS IN RANUNCULUS SUBGENUS BATRACHIUM 29
the carpels are hairy (as in R. aquatilis), the mature pedicels are uncurved (unlike R. baudotii)
and some leaves are morphologically intermediate between laminar and capillary. These plants are
sterile and pentaploid (2n = 40). Cook (1966) reported synthesizing a hybrid between R. baudotii
and R. peltatus. This hybrid looked and behaved like R. peltatus except that it was less vigorous,
had smaller flowers and glabrous carpels and was sterile.
The Slapton plant differs morphologically from these hybrids. Male fertility, estimated by
percentage of well-formed pollen grains, was high (90% or more) and the nutlets were well formed
and germinated readily. Mitotic root-tip squashes revealed a tetraploid chromosome complement
(2n = 32) for this plant (Fig. 5). These observations suggest that R. peltatus (2n = 32) and not R.
aquatilis is involved with R. baudotii in the parentage of this hybrid. However, this plant differs
from the synthesized hybrid of the same parentage described by Cook, and no natural hybrid of
this identity has apparently been previously reported. R. baudotii has recently been reduced to a
subspecies of R. peltatus (Cook 1983).
This plant was found on the eastern side of the Ley, in ‘bays’ in the Phragmites fringe, where the
water was about 0-5 m deep. A number of other interesting plants were found here, including
Lemna minuscula Hert., Potamogeton pectinatus L., Chara connivens Salzm. ex A. Br. and the
coenobial green alga, Hydrodictyon. A description of the plant is given below.
Erect annual herb. Leaves laminar or divided into capillary segments. Laminar leaves opposite
or alternate; petiole up to 60 mm long; lamina up to 15 mm long and 20 mm wide, reniform with
brown markings, deeply 3-lobed, lobes cuneate, margin crenate. Capillary leaves alternate, shorter
than internodes; petiole 1-10 mm long; lamina 25-35 mm long, globose to obconical. Segments
35-80, fine, usually rigid but some flaccid. Intermediate leaves occasionally present with capillary
portion of leaf proximal. Stipules triangular, adnate to petiole for 0.75 or more of their length,
apex acute. Pedicels up to 160 mm long, longer than the petiole of the opposed laminate leaf.
Sepals c.3 mm long, caducous, brown-tipped. Petals 3-5 mm long, broadly ovate. Nectar pits
shortly pyriform, with an upper border. Stamens 13-26. Carpels (25—)40-—55(-—72), glabrous.
Receptacle hairy.
DISCUSSION
Cytological data — in this case, chromosome numbers — may provide useful evidence in Ranunculus
subgenus Batrachium in the context of unusual hybrid and intermediate plants. Such evidence can
be particularly valuable since hybrids in this group are often not morphologically intermediate
between their parents, and hybrids derived from reciprocal crosses often differ. In some cases,
parental features are inherited very unequally whilst the hybrid may exhibit new features not seen
in either parent. Cytological data are needed for many other plants of unknown identity in the
group.
ACKNOWLEDGMENTS
This work was funded by the N.E.R.C. and was carried out at the Department of Botany,
University of Reading in conjunction with the Freshwater Biological Association (River
Laboratory). I am very grateful to Prof. D. M. Moore, Prof. V. H. Heywood and Mr D. F.
Westlake for their valuable advice.
REFERENCES
Brookes, B. S. & Burns, A. (1969). The natural history of Slapton Ley Nature Reserve. III. The flowering
plants and ferns. Fld Stud., 3(1): 125-157.
Cook, C. D. K. (1966). A monographic study of Ranunculus subgenus Batrachium (DC.) A. Gray. Mitt. bot.
Stsamml., Miinch., 6: 47-237.
Cook, C. D. K. (1983). One new taxon and two new combinations in Ranunculus subgenus Batrachium (DC.)
A. Gray. An. Jard. bot. Madr., 40: 473.
Erikson, J. (1905). Nagra vaxtfund fran Blekinge Batrachium baudotii X peltatus. Bot. Notiser, 1905: 319-320.
30 S. D. WEBSTER
Groves, H. & Groves, J. (1901). A new hybrid water Ranunculus. J. Bot., Lond., 39: 121-122.
SORENSEN, T. (1955). Hybriden Ranunculus baudotii x R. radians. Bot. Tidsskr. , 52: 113-124.
Stace, C. A. (1975). Hybridization and the flora of the British Isles. London.
TurRALA, K. & Wo_eEK, J. (1971). A natural tetraploid hybrid of Ranunculus subgenus Batrachium from the
Nowy Targ Basin. Acta biol. cracov, Sér. bot., 14: 157-161.
WILLIAMS, I. A. (1926). Ranunculus X hiltonii. J. Bot., Lond. , 64: 250.
(Accepted July 1985)
Watsonia, 16, 31-36 (1986) 31
The distribution of Poa angustifolia L. in Britain
C. SARGENT, O. MOUNTFORD and D. GREENE
Institute of Terrestrial Ecology, Monks Wood Experimental Station, Abbots Ripton,
Cambridgeshire, PEI7 2LS
ABSTRACT
Data collected during a survey of British Rail land included a total of 232 new 10 km square records for Poa
angustifolia L. The biology of the plant is considered in relation to the railway environment, and reasons for the
extended distribution pattern are discussed.
INTRODUCTION
Poa angustifolia L. is considered native to Britain (Clapham ef al. 1981; Hubbard 1968), although
information about its distribution is incomplete (Barling 1959; Hubbard 1968). Unless in flower the
plant is inconspicuous, and at a glance may be taken for Festuca rubra L. Its taxonomic position has
been uncertain and, in some vice-counties, it has not been recorded as a separate species or even as
a segregate of the P. pratensis L. group.
In Britain, Hubbard (1968) gave habitat information as “rough hill grassland, especially on
chalky and limestone soils, frequently growing among tufts of Bromus erectus”. Grime & Lloyd
(1973) described the species from limestone grassland, whilst Barling (1959) found the grass most
commonly with Brachypodium pinnatum (L.) Beauv., Bromus erectus Hudson and Avenula
pubescens (Hudson) Dumort. Of 135 British Floras consulted (bibliography at Monks Wood
Experimental Station), 49 included information about P. angustifolia. The habitats described are
grouped and listed in Table 1, and the number of times each habitat is mentioned is given. Some
Floras mention more than one habitat. Similarly, records held at the Biological Records Centre,
Monks Wood Experimental Station (B.R.C.) were tabulated and all those giving habitat
information are also included in Table 1. Many recorders did not include such information, and
absolute frequencies are not, therefore, calculated. Herbarium sheets held at BM and K were also
examined, and habitats recorded on these were found to correspond closely to the breakdown
given in Table 1 (most of the K records are duplicated by B.R.C., and therefore contribute to
Table 1).
Semi-natural habitats are freely-draining soils (usually calcareous) supporting coarse grassland
or woodland (often beech), sand, gravel and shingle, although several collections at K were from
wet, heavy soil. These plants tended to have at least some leaves from sterile tillers wider than 2
mm. More artificial habitats include roadside verges, walls, wasteland, docks and airports. Two
herbarium sheets labelled ‘‘London bomb-site flora’? were found. However, the most common
habitat recorded is railways.
The oldest specimen examined was collected by J. Dickson between 1789 and 1791 (BM), and
was recorded from woodland, the only habitat seen listed on herbarium sheets prior to 1850. No
locality is given for this or for a woodland collection made by Pelham in 1797 (K). The earliest
railway specimen was collected in 1897 by S. T. Dunn from near Rugby and was obtained through
the Watson Botanical Exchange Club for herb. A. B. Jackson at BM. The first roadside record,
also in herb. A. B. Jackson at BM, is dated 1910.
Amongst the Floras examined, the railway habitat is first mentioned by Hanbury & Marshall
(1899). In the Bedfordshire plant atlas (Dony 1976), the distribution map is annotated ‘railway
banks’, and elsewhere Dony (1953) suggested the grass may have been sown.
32 C. SARGENT, O. MOUNTFORD AND D. GREENE
TABLE 1. HABITATS GIVEN IN FLORAS AND ON BIOLOGICAL RECORDS CENTRE CARDS
Habitat Floras! BeRC?Z
Chalk grassland, downs, escarpments Ly if
Sand dunes, pits, warrens 4 i
Heaths (dry acid soils) 5 2
Woodland 11 9
Hedgerows, dry banks 16 S)
Railway verges 14° 28
Roadside and motorway verges 11 26
Docks, airports and wasteland 2) 9
Gardens, parks, allotments 2 7
Walls, pavements 8 19
Quarries, gravel, shingle 3 4
In shade 3 4
Near water if 3
' A total of 135 British Floras were examined (list at Monks Wood Experimental Station): Poa angustifolia was
mentioned in 49.
* All 10 km sq records for P. angustifolia held by B.R.C. were examined. Habitat information was given on
only 110 cards.
* The references listing railway verges as a habitat for this species are: Benoit & Richards 1963; Bowen 1968;
Cadbury et al. 1971; Dony 1953, 1967, 1976; Ellis 1983; Graham et al. 1972; Hanbury & Marshall 1899; Kent
1975; Messenger 1971; Roberts 1982; Roe 1981; Stearn 1975.
P. angustifolia is common in two noda of the Arrhenatheretum found along railway lines (Sargent
1984a). The relevant part of the synoptic table is reproduced (Table 2), and shows frequent
associates together with recorded abundance. The grass is common on warm, freely draining
slopes, either with Brachypodium pinnatum and Bromus erectus (mean pH 7-8), or with Festuca
rubra, Achillea millefolium L. and Plantago lanceolata L. (mean pH 7:3). In both noda,
Arrhenatherum elatius (L.) Beauv. ex J. & C. Presl, Dactylis glomerata L. and Rubus fruticosus
agg. are abundant, reflecting the character of the rough, calcicolous, railway grasslands.
It is very likely that the nodum in which B. pinnatum is constant at level V (more than 80% of
samples) is comparatively recently burnt grassland with little nutrient in the soil (Green 1983),
whilst that in which F. rubra attains the same level has not been burnt for some time. When data
collected on separate occasions from the same sites in Southern and Western Regions of British
Rail were analysed and modelled to show change, a marked increase in the area of F. rubra
grassland was found, and this was associated with changes in management: controlled sward
burning had been almost completely abandoned in these Regions with the demise of steam engines
and consequent reduction in fire hazard from engine box sparks. However, chemical weed control
of a narrow (3 m) strip adjacent to the track bed, introduced to replace systematic management of
the whole verge, has not proved entirely adequate, and it is likely that verge burning will increase
in future.
DISTRIBUTION
During a survey of British Rail land, P. angustifolia was found in a total of 316 x 10 km squares
(sites within 646 squares were surveyed). In 232 of these the plant had not previously been
recorded. These data considerably extend the known range of P. angustifolia from south-eastern
Britain into the north and west. A distribution map is shown in Fig. 1. Eight-figure grid references,
community and habitat data are held at Monks Wood.
The British Rail land survey was completed under contract with the Nature Conservancy
Council: objective data were collected from a total of 721 sites, each consisting of 100 m of track,
randomly distributed within 26 geographic strata covering the entire rural network, and from a
further 243 areas visited to document their biological interest. P. angustifolia was recorded from
430 of these locations.
DISTRIBUTION OF POA ANGUSTIFOLIA 33
TABLE 2. FREQUENT ASSOCIATES AND RECORDED ABUNDANCE OF POA ANGUSTIFOLIA
WITHIN DEFINED NODA (SARGENT 1984a) OF THE MOLINIO ARRHENATHERETEA ON
RAILWAY LAND
Nodum 1 Nodum 2
Poa angustifolia IV (1-5) II (1-2)
Arrhenatherum elatius V (1-5) IV (1-5)
Dactylis glomerata IV (1-5) III (1-4)
Poa pratensis III (1-5) II (1-3)
Rubus fruticosus III (1-5) III (1-5)
Plantago lanceolata V (1-3) Il (1-2)
Centaurea nigra III (1-5) III (1-4)
Lathyrus pratensis IT (1-4) II (1-3)
Heracleum sphondylium II (1-4) II (1-2)
Festuca rubra V (1-5)
Rumex acetosa II (1-2)
Holcus lanatus II (1-5)
Taraxacum officinale II (1-2)
Achillea millefolium IV (1-2)
Leucanthemum vulgare III (1-4) II (1-4)
Cerastium fontanum II (1-2)
Equisetum arvense IT (1-4)
Potentilla reptans III (1-5)
Vicia sativa subsp. nigra II (1-4)
Lotus corniculatus II (1-3)
Brachypodium pinnatum IV (3-5)
Bromus erectus II (1-5)
Cirsium arvense Ill (1-4
Convolvulus arvensis Ili (1-5)
Festuca arundinacea II (1-4)
Viola hirta II (1-3)
No. samples 215 54
Mean pH a3 7:8
Constancy: II = 20-40%; Il = 40-60%; IV = 60-80%; V = >80%.
Cover: 1 = <2%; 2 = 2-5%; 3 = 6-20%; 4 = 21-50%; 5 = >50%.
Nomenclature follows Tutin et al. (1964-1980).
DISCUSSION
Railway banks are comparatively warm (Suominen 1969) and are engineered to be freely draining
(Sheail 1979). The verges are untrampled and are grazed by small mammals and invertebrates
only. In the past, they have been managed by annual or more frequent mowing and burning. Only
selective herbicides, including 2,4-D and Picloram, have been used on verges, and spraying is
restricted to a strip approximately 3 m wide adjacent to the track bed. Spent ballast and cinders are
commonly tipped along slopes, and may imitate sand, gravel and shingle. Nitrogenous wastes from
trains drain into the cess and down embankments (Sargent 1984b).
Xeromorphic summer leaves (Barling 1965; Galkin & Seredin 1973; Sukhova 1974) enable P.
angustifolia to compete successfully in this environment. Although grazing response has been
variously reported (Ammar 1978; Barling 1959; Bublienko 1980; Grime & Lloyd 1973; Osichnyuk
& Panova 1978; Tkachenko et al. 1979), it is likely that the absence of heavy grazing is
advantageous, and certainly, on untrampled railway slopes, the thin, wiry rhizome (Khoang
T’yung 1975; Tuganaev & Pestereva 1976) is not at risk. Further, the rhizome may be important in
recovery from burning. Early flowering and seeding (Prokudin et al. 1982) may have favoured P.
angustifolia when verges were mown or scythed annually; the selective herbicides at present in use
34 C. SARGENT, O. MOUNTFORD AND D. GREENE
Ficure 1. Distribution map of Poa angustifolia in Great Britain. The data are plotted from 805 records
occurring within 570 10-km squares. The British Rail land survey provided 430 records covering 316 10-km
squares, 232 of these are squares in which P. angustifolia had not previously been recorded. A = British Rail
land survey; ™@ = other railway data; O = other data (for some of which no habitat information is available).
Railway data are plotted in precedence over other data.
are unlikely to produce permanent damage (Shikhotov & Kuchin 1975). This combination of
habitat and management favours the establishment of P. angustifolia on railway land.
The very marked clustering of previous records in south-eastern Britain indicates that, despite
the limitations in previous recording introduced by variable taxonomic treatment, the present
distribution pattern shows an extension of range into suitable, albeit man-made, habitats.
It is clear, however, that railway land, because access is restricted, has been under-recorded.
What is less clear is the status of P. angustifolia in Britain. Is it really a native species of downland
DISTRIBUTION OF POA ANGUSTIFOLIA és)
and beech hangers in the south-east? Or has it been introduced with grass seed and spread via
railway lines and roadside verges to other suitable habitats? The majority of previous records from
Scotland are from ports (Fig. 1), and the plant is known from docks and airports (Table 1).It may
have arrived as a cargo alien: M. McCallum Webster annotated material (K) from Selkirk
“introduced, with wool shoddy”’’. However, the earliest records, dating from the late 18th century
(before railways were built) are from woodland, and it is likely that at least part of the population is
native.
A genecological study could be most interesting. Because the species is largely apomictic
(Akerberg 1939; Love 1948) it is possible that populations from different sources have maintained
some genetic integrity, showing a preferred distribution of chromosome numbers. However, the
very wide range of numbers reported (Bolkhovskikh 1969; Edmondson 1980; Love & Léve 1961;
Majovsky et al. 1972) and the occurrence of different numbers in the same sward (collections by
Hubbard at K), indicates that the situation is complex.
The railway is often very distinct from surrounding land, especially in north-western Britain. Its
continuity has enabled a number of species more commonly associated with warm, dry, southern
habitats to extend their range: Daucus carota L. and Rosa stylosa Desv. are amongst other good
examples (Sargent 1982), and the phenomenon is certainly not unique to P. angustifolia.
ACKNOWLEDGMENTS
The authors would like to thank Christopher Preston (Biological Records Centre, Monks Wood),
and herbarium staff at the British Museum (Natural History) and Royal Botanic Gardens Kew,
where Dr T. A. Cope was particularly helpful. We are most grateful to Mrs Jean King for searching
the literature, and to Mrs June Stokes for typing the manuscript. The data were collected during a
survey of British Rail land under contract with the Nature Conservancy Council. We are grateful to
both these organizations.
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Grime, J. P. & Lioyp, P. S. (1973). An ecological atlas of grassland plants. London.
HaAnsury, F. J. & MARSHALL, E. S. (1899). Flora of Kent. London.
36 C. SARGENT, O. MOUNTFORD AND D. GREENE
Hupsarp, C. E. (1968). Grasses, 2nd ed. Harmondsworth.
Kent, D. H. (1975). The historical flora of Middlesex. London.
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Obshch. Ispyt. Prir. Otd. Biol., 80: 128-135.
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(Accepted May 1985)
Watsonia, 16, 37—42 (1986) a7
A reconsideration of the taxonomic status of Poa balfouri
Parnell (Gramineae)
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28 High Street, Balsham, Cambridge, CBI 6DJ
ABSTRACT
Poa balfourii Parnell (Gramineae) has held the doubtful status of species since Parnell described and discussed
it in 1842. Observations by the author on wild populations of Poa glauca suggest that the plant, successively
described by Parnell (1842), and discussed by Sowerby (1872) and Hubbard (1968), is a distinctive variant of
Poa glauca that may not merit formal taxonomic recognition.
INTRODUCTION
Poa balfourii was first described by Parnell (1842). He named it after Dr J. H. Balfour of Glasgow
University, who had collected the type specimens from Ben Voirlich, Mid Perth, v.c. 88, in 1842.
Parnell stated that it was closely allied to Poa nemoralis, whilst Sowerby (1872) referred to its
similarity to P. glauca Vahl but noted that it was less rigid and less glaucous, and lacked the bluish
tinge of P. glauca. Hubbard (1968) considered it to be closely related to P. glauca and very similar
to a mountain variety of P. nemoralis L., and added ‘‘a rare imperfectly known mountain grass’’.
Tutin (1952) considered the taxon only briefly and commented that it was similar to P. glauca but
differed ‘in the short truncate ligule’. P. balfourii was not recognized by Edmondson (1980).
Among the many herbarium specimens that I have seen mislabelled “P. balfourii’’, both P.
glauca and P. nemoralis are commonly found. Many 19th century herbarium sheets of P. balfourii
were annotated with alternative determinations, together with comparative notes on the similarity
of the taxon to the numerous species and varieties of montane Poa then recognized; and the
species has often been subsumed within one or other of these, e.g. P. nemoralis L. var. balfourti
(Parn.) Hook. f. and P. glauca Vahl subsp. balfourii (Parn.) Syme.
A factor complicating the variation pattern is facultative agamospermy. Poa nemoralis and P.
glauca, together with Poa palustris, form section Stenopoa and are facultatively agamospermous.
Gustafsson (1946) said of section Stenopoa that “‘morphologically they stand rather close . . . no
diploids exist and the whole complex is polyploid and apomictic’’. He noted that the variation
occurring in Sfenopoa is very wide: crosses between species frequently take place and there is a
“difficulty in correctly demarcating them’’. He added a further complication when he stated that
“members within the section certainly hybridize with one another and also with polyploids out of
other sections”. The evolution of apomictic groups is associated with polyploidy, as seen in the
multiple chromosome number recorded in P. nemoralis and P. glauca, which is some indication
that polyploidy is a source of new variation in this group. There is variation in the outline of the
panicle, the number and length of spikelets, the colour and the distribution of colour over the
whole plant and the rigidity of culm and blades. The consequent variation in an environmentally
heterogeneous area is sufficient to obscure the taxonomic distinction between P. nemoralis and P.
glauca. Ecological variants that have been assumed to be genetic variants have been recognized as
species, such as Poa montana, P. polynoda and P. balfourii itself. This paper, based on my field
observations since 1969, assesses the taxonomic status of P. balfourii.
38 Pod. OG: TRIST
THE SEARCH FOR POA BALFOURII
I started to look for Poa balfourii in 1969 in Snowdonia, to which area I returned in 1979. During
the intervening years of the 1970s I searched mica schist crags in the Killin area of Mid Perth,
v.c. 88, to follow up old records. My study was directed to areas facing north, and from north-
east to south-east, in habitats of partial shade, total exposure and varying shelter.
Throughout this period I sent many specimens of putative P. balfourii to the late Dr C. E.
Hubbard, all of which he referred to “‘the mountain variants of the polymorphic Poa nemoralis’’.
Nevertheless, he encouraged my search and begged me to find him plants of P. balfourii to bring
into cultivation. Earlier, in 1958, Hubbard (in litt.) had noted that ‘“‘P. balfourii is generally
separable from P. glauca by its lax often nodding or inclined inflorescence, but here again further
study of the two in Britain is required”. Hubbard remained in doubt and (pers. comm. 1972) said
“these mountain Poa’s are much in need of critical study in the field and the experimental
ground’’. I continued my field studies, but it was clear to me that he was never quite satisfied
with the status of P. balfourii.
VARIATION IN POA GLAUCA
Poa glauca in Britain is found both in fully exposed positions and in sheltered habitats at
altitudes of c. 670—920 m. It is essentially a grass of cliffs and rocky slopes, and is less usually
found on screes. The fully exposed habitat is seen on small ledges on vertical rock faces, in small
pockets on flat protruding rocks and occasionally on an isolated rock fallen from a cliff. These
sites, where there are often no abutting rocks on either side or above, are completely exposed to
wind and rain. P. glauca would appear to occupy a dry habitat but most of its stations occur
where there is an overhead source of water, or are on a rocky slope receiving water from above.
These sites of extreme exposure probably represent the most common habitats of P. glauca.
More sheltered habitats are found in mini-corries and rocky gullies running to the base of
cliffs, which offer varying degrees of shelter and certainly afford a less stringent environment
than the exposed cliff-face. Within these areas are small sites which are sheltered from the wind
on one or more sides, and where an additional element is shade. The sites generally face east,
north-east or south-east and experience a very limited amount of direct sunshine.
In rocky habitats, Alchemilla alpina appears to be the most common associate, with Saxifraga
aizoides, S. oppositifolia, Silene acaulis and Festuca vivipara as frequent associates; occasional
associates within a metre square include Alchemilla glabra, Cerastium alpinum, Galium boreale,
Oxyria digyna, Polygonum viviparum, Euphrasia spp., and Deschampsia caespitosa. It is unusual
to find more than three or four associated species at any one site, as the limited size of the
habitat only allows the development of a small community. The habitat is similar to that of P.
alpina L.
Hubbard (1968), in his description of P. glauca said “‘it is covered with a whitish wax” and that
this distinguished it from P. balfourii and P. nemoralis. This would seem to give taxonomic
standing to a feature modified by the environment. My observations on the occurrence of this
whitish wax is that it only occurs on P. glauca in exposed positions and that there is no trace of
this feature on P. glauca in shade and shelter. The same applies to the deep mauve colouring
that is found only on plants in exposed situations, when most of the culm, the base or up to half
the length of the blades, and the whole of the length of the sheath, is a deep mauve, and the
spikelets are a dark green with tinges of deep mauve and often with a gold or bronzed tip.
In situations where there is some adjacent rock shelter, the mauve colouring is modified in
extent and shade of colour. In fully exposed sites the rigidity of culm and blades is a feature, but
in shade and shelter the rigidity of culm and blade is absent; the panicle is loose and nodding and
the colour of the whole plant is grey-green, light green or even sometimes a dark green. The
deep mauve colour of culm, blades and sheaths is reduced to a light mauve zone c. 1 cm in length
at the base of the culm. While the spikelets are devoid of mauve colouring, they are occasionally
found with a light gold tip. As I hope to show in the following discussion, it is these shade
variants that have formerly been treated as a separate species, P. balfourii.
STATUS OF POA BALFOURII 39
RELATIONSHIP OF POA BALFOURII TO P. GLAUCA
The diagnostic characters given for P. balfourii by Parnell (1842) were as follows: ‘‘Florets slightly
webbed. Ligule prominent, obtuse. Upper leaf nearly as long as its sheath. Outer palea 5-ribbed.
Stem compressed”’.
The ‘“‘webb” of the florets (silky hairs of the lemma keels), is extremely variable in length and
density and often almost absent in P. glauca. In the shade variant of P. glauca, the webbing is
sparse as described for P. balfourti. The ligule is not always prominent but always obtuse. The
ligule of P. glauca of shade and exposure has a range of 0-8—3-0 mm but the upper limit beyond 2-5
mm is uncommon.
The relation of the length of the upper leaf to the length of the sheath in P. glauca is also very
variable. The upper leaf of the culm is frequently as long as or longer than its sheath. Parnell gave
the outer palea (lemma) as 5-ribbed, a feature which he also attributed to P. caesia (P. glauca).
With regard to the compressed stem character, it is found that the culm of the shade variant of P.
glauca may be compressed, nearly round or almost square-sided.
Sowerby (1872) described P. balfourii in detail, in much the same manner as Parnell, but also
gave no measurements. He distinguished it from P. glauca in that the plant was “‘not rigid as in P.
eu-glauca: the ligule is longer and the whole plant less glaucous and without the bluish tinge of P.
eu-glauca’’. This is a fair description of the shade variant of P. glauca.
Hubbard (1968) gave a full description of P. balfourii, supported by measurements. He
described it as ‘‘a loosely tufted perennial . . . culms erect, leaves greyish green . . . ligule blunt,
the uppermost 1-3 mm. . . panicles loose, inclined or usually nodding 4—10 cm long; spikelets 5—7
mm greyish green, glumes finely pointed”. Hubbard said it is “‘very closely related to P. glauca”’
which he said “has stiffer culms, firmer blades, stiffly erect and usually open panicles’. These
attributes apply to the P. glauca of exposed habitats. The details of Hubbard’s description of P.
balfourii perfectly matches that of P. glauca of shade and shelter, except for the extreme of the
range of measurements of both ligule and spikelet, which are uncommon.
Further evidence of the identity of P. balfourii comes from a study of the probable type
specimens. The type specimens of Poa balfourii in E, on sheets 87/83, 8 & 9 from Ben Vorlich,
1842, named P. balfourii by Parnell, have been annotated as ‘“‘no doubt one of the original
specimens’’. The specimens on both sheets have light green spikelets with gold tipped lemmas, and
in my opinion are shade variants of P. glauca.
COMPARATIVE MEASUREMENTS OF POA BALFOURII, P. GLAUCA AND P. NEMORALIS
A range of measurements of P. glauca and those attributed to P. balfourii are given in Table 1. A
comparison of the measurements of the various parts of P. glauca and P. balfourii given by
Hubbard (1968) and my own records of P. glauca of exposed and shaded positions, respectively
TABLE 1. MEASUREMENTS OF POA BALFOURII, P. GLAUCA AND P. NEMORALIS
Hubbard (1968) P. glauca*
P. balfourii P. glauca Of exposure Of shade & shelter Poa nemoralis*
Height, cm 15-40 14-40 (9—)15-—25(-34) (25—)27-—32(-40) (18—)25-30(-5S0)
Blade length, cm 8 2-8 (2-)2-8-5-0(-5-5) (3-5—)6-0-8-5(-10) (1-5—)2-5-7-0(-10)
Blade width, mm 2-3 2-4 (0-S—)1-0-—2-0(—2-6) (1-:3—)1-7—2-5(—2-8) (0-7—)1-0-—2-0(-2-3)
Upper ligule length,mm 1
Panicle length, cm 4
Spikelet length, mm 5-7 4-6 (2-5—)3-0—4-5(-6-3) (3-S5—)4-0-5-8(-6-0) (2-0—)2-5—3-0(-3-6)
Lower glume length,mm 3-4 3-4-5 (2-5—)2-8—3-8(—4-2) (2-6—)2-8-3-5(—4-0) (2-0—)2-5—3-5(-3-8)
Upper glume length,mm 3-5-5 3-4-5 (3-0—-)3-3—-4-5(—4-7) .(3-2—)3-5—4-0(—4-7) (2-5—)3-0—3-5(-3-8)
Lemma length, mm 3:5-4:5 3-4-0 (3-0—)3-5—4-0(-4-6) (3-3-)3-5—4-3(-4-6) 0-2-0-5
& 1-2-5 i225 __ 0-B-2-5(-3-0) (1-5—)3-0-10-0(—12-0)
10 2-10 (1-2-)2-0-6-0(-7-0) (4-0-)6-0-8-0(—9-5) (2-2-)3-0—-5-0(—6-0)
*Measurements made by P.J.O.T. on c. 150 plants (250 culms) from both the field and the herbarium.
40 P. J. O. TRIST
show no significant taxonomic differences. Two measurements only call for comment and these,
which have been previously mentioned, relate to P. balfourii. They are the upper range of the
ligule (3 mm) and the spikelet (7 mm). These upper limits do occur but are unusual. The range of
the lengths of the spikelets, glumes and lemmas given by Hubbard (1968) for P. glauca and P.
balfourii and those of mine for P. glauca of exposure and shade, are comparable respectively.
The plant height in all four taxa shows that plants of exposure can be up to 6 cm shorter than
those of shade. Of the length of blades, my measurements show a slightly greater length from
plants in shade compared with those of exposed sites; the width of blades shows 0-8 mm less in the
lower range in exposure.
The number of spikelets per plant of P. glauca has a range in exposed sites of (5—)10—20(-26),
and in shade of (4—)10—28(—34). The number of spikelets may have no significance and is possibly
influenced by habitat and available nutrients, but within the range of the sample we are also
probably looking at hybrid variants. The lowest branches of the panicle number 1-2 and 3 is
uncommon in both exposure and shade. The length of the branches is variable, 5—14 (-18) mm in
exposed sites dnd 7-18 (—22) mm in shade; the variation is not an unusual feature. |
The variation in the height, rigidity of culm and number of spikelets did not escape the
observation of Parnell (1842). Under P. balfourii he described a var. rigida Parn. as a “‘short stout
variety . . . with a simple panicle of few spikelets”. An 1842 specimen on sheet 74/84,2 (E) from
Ben Vorlich was recorded as Poa balfourii var. rigida. It has a ligule of 2-5 mm and is a typical,
small specimen of P. glauca, with rigid culms and blades which I have already described from
exposed sites and which is confirmed by Parnell (1842) ‘“‘found in exposed situations’. He also
described a var. extensa as a “‘tall and slender variety with a simple panicle of few spikelets’’, the P.
glauca of shade.
From Table 1 we can also compare the measurements of this shade variant of P. glauca (P.
balfourii) with those of P. nemoralis from the mountain. The height range of the two taxa is similar
and relates to local environment. For the length of the culm blades, the upper range of
measurements is similar. In the lower scale, P. nemoralis appears smaller but in this sample it is
influenced by small specimens of P. nemoralis var. coarctata; the same may be said of the width of
the blades. In the range of panicle lengths, P. nemoralis shows a wider variation and the small
panicles of the sample of the var. coarctata have influenced the lowest length of 1-5 cm; both the
size and outline of the panicle of both taxa are extremely variable in relation to environment. In
the lengths of the spikelets of both taxa the range of each is almost identical, but in P. nemoralis it
is slightly smaller. In general, P. glauca produces fewer spikelets than P. nemoralis and some of the
spikelets of the lower branches of the latter are often depauperate. A comparison of the length of
the glumes and lemma shows little more than a 0-5 mm difference. The significant difference in the
morphology of the two taxa lies in the shape and length of the ligule: in P. glauca the ligule is 0-8-
2:5(—3-0) mm and obtuse and in P. nemoralis it is 0-2—0-5 mm and truncate. In addition, the shape
of the glumes in the latter is long-pointed and lanceolate and in the former pointed and ovate-
elliptic, but these shapes are variable.
RELATIONSHIP TO POA NEMORALIS
Hubbard (1968) echoed Parnell’s protologue by stating that ‘in its form of growth, P. balfourii is
very similar to the mountain variety of P. nemoralis with rather large spikelets, differing mainly in
its longer conspicuous ligules”. This ‘‘mountain variety” similarity leads us to consider the
montane variants of P. nemoralis.
Many varieties of P. nemoralis have been described and in the present context I shall only refer
to some which I consider to be recognizable. Two varieties are more or less exclusively lowland,
var. vulgaris Gaud. of woodlands and var. angustifolia Parn. of shade and hedgerows, and need not
be considered. Var. glaucantha (Gaud.) Reichenb. and var. coarctata Gaud. are both found on
mountain crags, but also in the lowlands, the former in woodland clearings and the latter on walls.
[I have also noted var. montana Gaud. in herbaria and I have attempted to match specimens of this
variety with plants from the mountains. It is very variable in colour, in the outline of the panicle
and number of spikelets, which have a size-range of 3-3—6-0 mm. The ligule is 0-5 mm long. In fact,
all of these varieties of P. nemoralis have a common diagnostic factor in the shape and length of the
STATUS OF POA BALFOURII 41
ligule, which is truncate, often with a pinched point in the centre facing the leaf, and is 0-2—0-5 mm
long — the two hallmarks of P. nemoralis.
Hubbard (pers. comm. 1972) said that ‘‘a longer liguled P. nemoralis has been called var.
montana or var. parnellii”’ and I believe his reference was to Poa montana Parn. (P. nemoralis var.
montana Koch) which Parnell (1842) described with a “conspicuous obtuse ligule’. An 1887
specimen (CGE) has a 3 mm ligule; it was gathered on Creag an Lochain, Killin, labelled Poa
montana by C. C. Babington, and referred to Poa balfourii by Arthur Bennett. This long-liguled
Poa montana, referred to P. balfourii and confused with a mountain variety of P. nemoralis, is a
variant of P. glauca.
A collection by E. S. Marshall labelled P. balfourii from Garbh Beinn, no. 1649 (CGE), has
been annotated by Hackel: ‘“This is so near to what in our Alps we call P. nemoralis var. montana
that I can find no marked difference’. On the same sheet, Hackel also referred to Marshall’s 1896
collection from Aonach Beg, no. 1/648, and named P. balfourii var. montana, which he said “‘only
differs from no. 1649 by its longer ligule . . . so that probably no clear line can be drawn between
P. balfourii and the alpine forms of P. nemoralis’’. Hackel’s note continues, “‘. . . and shall often
meet with specimens which must remain doubtful, or in which it will depend more upon the skill of
the observer than upon real positive characters’’. This appears to be an indirect reference but an
undisclosed admission that “the skill of the observer” should include observations on the habitat.
There is a further problematic variant which I have seen on a number of occasions in N. Wales
and in Scotland. In general appearance it is more like P. nemoralis than P. glauca: it is lax, with its
spikelets generally nodding, and does not exhibit the rigidity of leaf and culm of P. glauca growing
in exposed sites. Its glumes are long and narrow as in P. nemoralis and unlike the ovate-elliptic
glume of P. glauca. The ligules are 0-5—0-8 mm and obtuse. Are these variants of P. glauca x P.
nemoralis or variants of one or other taxon? These plants are surely those which Hubbard often
quoted to me as “the mountain variant of the polymorphic P. nemoralis with slightly longer
ligules’’. That mountain variants of both P. nemoralis and P. glauca exist cannot be disputed, but
variants with ligules >0-5 mm cannot be accepted as P. nemoralis and are variants of P. glauca.
TAXONOMIC CONCLUSIONS
Poa glauca and Poa nemoralis are morphologically similar, and the taxa frequently present
problems of separation. In P. glauca the glumes are long-pointed and ovate-elliptic, and in P.
nemoralis generally lanceolate and sometimes almost ovate, and variants are found where the
differences are blurred. There are therefore occasions when the distinction between the two taxa
can only be resolved on the shape and length of the ligule.
Field observations on the ecology of Poa glauca have shown that plants are found in a range of
situations from extreme exposure to shade and shelter, which induces a variety of phenotypes. The
slender variant of green-coloured P. glauca, with no covering of whitish wax and without any deep
mauve coloration, usually found as a single-culmed plant in the shade and shelter of rocks, is that
montane Poa variant to which the name Poa balfourii Parnell has been given. It is considered
therefore that P. balfourii has been incorrectly treated at the level of species, and that it is merely
an infraspecific variant of P. glauca. Whether its particular morphology is a genotypic or a
phenotypic response to its shade environment is unknown.
ACKNOWLEDGMENTS
I thank the Curator of the Botany School Herbarium, University of Cambridge (CGE), for his
permission to cite records, and the Curator of the Herbarium of the Royal Botanic Gardens,
Edinburgh (E), for the loan of, and permission to cite from, the collections of Poa balfourii made
by R. Parnell and others. I am grateful to Dr J. R. Edmondson for his opinion and guidance and to
Dr S. M. Walters for reading the MS. and offering much valued advice.
42 P: J. OCTRSE
REFERENCES
Epmonpson, J. R. (1980). Poa, in Tutin, T. G. et al., eds. Flora Europaea, 5: 159-167. Cambridge.
Gustarsson, A. (1946). Apomixis in higher plants. Part 1. The mechanism of apomixis, pp. 111, 252. Lund.
HuBBARD, C. E. (1968). Grasses, pp. 179, 181. Harmondsworth.
PARNELL, R. (1842). The grasses of Scotland, p. 145. Edinburgh.
SowErby, J. (1872). English Botany, 11: 121. London.
Tutin, T. G. (1952). Poa in CLapHaM, A. R., TuTIn, T. G., & WaArRBuRG, E. F. Flora of the British Isles, pp.
1435-1443. Cambridge.
(Accepted June 1985)
Watsonia, 16, 43-54 (1986) 43
The distribution, ecology, history and status of Gastridium
ventricosum (Gouan) Schinz & Thell. in the British Isles
P.O; TRIST
28 High Street, Balsham, Cambridge, CB1 6DJ
ABSTRACT
Gastridium ventricosum (Gouan) Schinz & Thell. (Nit-grass) is a native of south-western England and occurs as
a casual in the Channel Islands. From herbaria and manuscript records, it appears that it has been recorded
from 34 vice-counties in Britain over the past 200 years. Few records have been reported since the publication
of the Atlas of the British Flora (Perring & Walters 1962). At the request of the Nature Conservancy Council, a
field survey of its current status was carried out in July and August 1980, 1981 and 1982. Habitat descriptions,
plant associates, estimated populations, soil textures and conservation status were recorded.
INTRODUCTION
Perring & Walters (1962) give dots for records of Gastridium ventricosum in 107 10 km squares for
pre-1930 and 18 post-1930 records for England and Wales and two in the Channel Islands. With
three exceptions, all records were south of the Wash and 98% were south of the line between the
River Severn and the River Blackwater. Hubbard (1968) records that it is “‘probably native in short
grassland on limestone and chalk: elsewhere a weed of arable land, occasionally locally abundant
in corn fields on light or heavy soils’. Perring & Farrell (1977) report “‘At one time it was recorded
from about 28 vice-counties extending as far north as Yorkshire, though it was undoubtedly only
casual in many inland stations. Since 1960 it has only been reported from six localities in Cornwall,
Dorset, Somerset, Hampshire, Gloucestershire and Glamorgan, and from Guernsey and Sark”’.
The current native sites of G. ventricosum are found in short virgin grassland on limestone or
soils derived from limestone. It can no longer be regarded as a weed of arable land. As such, in
1980 it was only known in a small arable area of S. Hants., v.c. 11 (Trist 1983). Considering its
isolation in the native sites, it is unlikely that we shall see many more casual plants.
METHODS
Inquiries on the location of known sites were made from members and county recorders of the
B.S.B.I. Old records in herbaria were examined, together with numerous county Floras and other
manuscripts. Many arable sites which have been recorded in the past 50 years in southern England
have been recently surveyed. The last known records from vice-counties have been listed. Soil
samples were taken within and without the area of 2m? quadrats to a depth of c. 10 cm. For pH
determination, dry soil was treated with a soil indicator containing bromothymol blue and methyl
red. The method does not match the accuracy of the use of the glass electrode but it does give an
indication of a pH range, which is given in the text.
DISTRIBUTION
Perring & Walters (1962) show that G. ventricosum formerly had a wide distribution over the
southern half of Britain. The greater majority of these records were referable to arable colonists
and casuals. The author has only been able to trace 22 records since 1950, of which 50% were
casuals or colonists.
44 P. J. O. TRIST
FicureE 1. Distribution of Gastridium ventricosum in the British Isles, based on the author’s 1980-82 survey.
@ native sites; circles denote arable colonists and casuals.
The field surveys of 1980—82 have revealed two facets in the history of the distribution of this
grass. Firstly, its former status as an arable colonist is almost lost and will inevitably disappear.
Secondly, there are a number of native sites on coastal downland, which have previously only been
recorded on one or two occasions and some which have not been previously recorded. The records
of 1982, compared with those of 1960, are predominantly native.
Recent fieldwork reveals that G. ventricosum occurs in only nine vice-counties, in 11 10-km
squares with 36 sites in 24 1-km squares. In southern Wales there are five sites in the Gower
Peninsula and one near Cardiff. In England there is one site in North Somerset, three in South
Somerset, one in South Devon, ten in Dorset, two in West Gloucestershire and one in the Isle of
Wight. In South Hampshire, there are five sites of arable colonists. In the Channel Islands there
STATUS OF GASTRIDIUM VENTRICOSUM 45
TABLE 1. GASTRIDIUM VENTRICOSUM LOCATIONS
Distance
Vice- Grid from sea_ Altitude
County Location (10 km) Aspect Slope km m Habitat*
3 South Devon 20/9.5. SE a 0-10 60 G
5 South Somerset A 31/0.4. S 20 0-5 60 G
South Somerset B 31/0.4. SE 20 0-8 45 G
South Somerset C 31/0.4. S 20 0-8 30 G
6 North Somerset 31/4.2. SSW 20 20-0 50 G
9 Dorset A 40/0.7. S 10 0-25 60 G
Dorset B 40/0.7. S 10 0-25 60 G
Dorset C 40/0.7. SSW 40-60 0-19 30 G
Dorset D 40/0.7. S 30-55 0-19 30 G
Dorset E 40/0.7. S 25-30 0-25 83 G
Dorset F 40/0.7. S 30 0-25 170 G
Dorset G 40/0.7. S 10 0-03 100 G
Dorset H 40/0.7. S 5) 0-20 100 G
Dorset I 40/0.7. S 30 0-20 100 G
Dorset J 40/0.7. S 20 0-20 100 G
10 Isle of Wight 40/9.7. S 0-003 5 G
34 Clifton A 31/57. SSE 15-20 8-0 100 '®
Clifton B 3L/9.7. SSW 35-40 8-0 30 Cc
41 Gower A 21/5.8. SSE 30 0-95 50 G
Gower B 21/4.8. SSW 30-35 0-35 30 G
Gower C 21/4.8. WSW 20-25 0-09 35 G
Gower D 21/4.8. ESE 20 0-10 35 G
Gower E 21/4.8. S 15 0-85 40 G
Nr Bridgend 21/9.6. S 35-40 0-40 40 G
11 Lymington A 40/3.9. Level arable 0-05 <e) A
Lymington B 40/3.9. ae) 0-50 <3 A
Lymington C 40/3.9. o 0-50 <3 A
Lymington D 40/3.9. sa 0-50 <3 A
Lymington E 40/3.9. ee 1-80 <3 A
S Sark
La Collenette WV/4.7. S 0-92 98 R
By School WV/4.7. S 0-47 100 R
By Chapel WV/4.7. N 0-92 104 R
Mill Lane WV/4.7. Ww 0-14 94 R
La Coupée WV/4.7. S 0-80 98 R
Near Mill WV/4.7. WwW 0-70 102 R
Guernsey
S 1-00 15 R
Le Douit, St Peter’s WV/3.7.
* Habitat key: G, Grassland; A, Arable; C, Cliffs; R, Ruderal.
are six sites in Sark and one in Guernsey: these have not been seen by the author but the sites, soil
and plant associates indicate that the plants are casuals.
Fig. 1 shows the distribution of G. ventricosum in the British Isles plotted from the 1980-82
survey by the author. Table 1 gives the location, aspect and slope of the habitats and the distance
from, and the height above, the sea.
PHENOLOGY
G. ventricosum is an annual grass with a life cycle spanning twelve months. It sets seed in
September—October and germination takes place in six to eight days. Within 14 days of
germination, the single-leaf seedling is 1-1-5 cm long. In the first winter of this work, it was
46 P.O SERGE
TABLE 2. GASTRIDIUM VENTRICOSUM: SIZE OF PLANTS IN DIFFERENT HABITATS
Native plants Arable colonists
Downland Rabbit grazed In wheat
Height, cm 5-50 1-5-9-5 37-97
Spike length, cm Zoo 0-5-2:-7 3-5-16-5
Blade length, cm 0-5-6:5 0-3-2:-7 4-5-20-0
Ligule length, mm 0-5-3-2 0-5-1-6 2:0-— 4-2
recognized that seedling development would be subject to climatic conditions of habitat. From
November to April 1980-81, a night frost was recorded on 58 occasions at Balsham, Cambridge, of
which 21 were in February. In the autumn of 1980, seeds were sown in pots at Balsham and, during
the above period, most of the seedlings in pots were either directly killed or unseated by frost: this
climate would not have been experienced in a native coastal habitat.
In cultivation, the single leaf growth remained static until late April and remained slow until late
in the spring. By mid-May, the characteristic leaf curl developed and in the following three weeks
into June there was a gradual acceleration of tillering up to early July, when the spikes broke the
sheath. The long period of low temperatures, when a range of 1—10°C of frost was recorded, would
have retarded leaf development until late April.
From year to year it appears that spike exsertion may take place any time from mid-June to
mid-July. D. E. Coombe (in litt. 1980) reported anthesis from plants in Dorset on 29th June 1952
and, following a mild winter at Balsham in 1983-84, I recorded first spike exsertion on 19th June.
Coombe (in litt. 1984) reported spike exsertion in the Avon Gorge on 6th July and, from my survey
of 1980-82, I found exsertion from the first to the third week of July. The variation in date of spike
exsertion is at least partly due to variation in the date of germination, which is influenced by
climatic conditions. Seed sown at Balsham on 24th March 1981 produced plants which exserted
spikes after 23rd September. The late period of growth of this grass is marked. In the Gower
Peninsula in August 1980, fresh green spikes had developed on new branches of a culm where the
primary spike had already gone to seed.
The culm has a final height range according to habitat, and detailed measurements are given in
Table 2. In an open association of grasses and herbs, the height varies between 5 and 40 cm; and
where an open sward has the occasional low gorse bushes or hawthorn on the perimeter of the
G. ventricosum colony, the several influences discussed in the notes on native habitats will increase
the height to c.50 cm. The height of the arable colonist plants varies between 37 and 97 cm and is
often about twice that of the average downland plant and is related to both height and density of
the wheat crop; and no doubt there is also some growth response from fertilizers applied to the
wheat. Culms are frequently branched even among the smallest plants, but rarely so in the tall
arable plants.
ECOLOGY
SOILS AND GEOLOGY OF THE NATIVE SITES
The soils to be described were taken from a depth of 10-15 cm. The Somerset sites are found on
isolated deposits derived from the Rhaetic Limestone which is a junction-bed between the Lower
Lias and the Keuper Marl; they contain inter-bedded limestone beds and clay or shale. At one, the
soil is a chocolate-brown silty clay loam with a low clay fraction. The soil aggregates are up to 2-5
cm across and break readily into fine granules and contain small rounded fragments of hard
limestone and red-brown shale. At another, the soil is similar and is a grey-brown silty clay loam
with a low clay fraction. The aggregates are up to 1-3 cm across, breaking readily into fine granules
and contain fragments of limestone and grey shale. At both sites, the soils contain fibrous roots and
are stable to water.
At the Dorset sites the soil is a brown silty clay loam with a low clay fraction and overlies the
STATUS OF GASTRIDIUM VENTRICOSUM 47
Purbeck Limestone. Much of the area has in the past been shallow quarried and outcrops of rock
and small stones are much in evidence on the surface. It was difficult to uncover a profile and the
interpretation of a halt to an auger may be a large stone or flat stone platform. Much loose rock is
encountered and soil depth may vary from 10—60 cm. On a higher part of this area, soil depth is
very shallow and most augerings indicate c.10 cm over hard limestones. The silty clay loam samples
comprised mainly small aggregates of 1-5 mm of angular fragments with partly rounded edges and
angular blocks of 1—2 cm across, together with flat limestone flakes of 3-10 mm. These soils wet
slowly, are stable to water and are free-draining.
On the cliffs of the Avon Gorge and on the Gower coast, the soils overlie Carboniferous
Limestone. At the former sites, the soil is a red-brown sandy loam with granules up to 2 cm across.
At the latter, the texture is more or less similar but three of the sites have more fine sand and one a
little more clay. The aggregates break easily, are stoneless and contain many fibrous roots; they
wet easily and are stable to water.
The soil descriptions clearly have much similarity and are considered to be from native sites of
G. ventricosum. No soil at any site showed any evidence of wetness or indication of impeded
drainage. The aggregates have a good granular structure, break readily and are friable, making a
good open medium for root growth. All sites are on slopes where rainfall has a steady run-off and,
with good drainage, less calcium is leached or being made soluble.
The soil samples taken from each site show that, with two exceptions, the pH range of 6-5—7-0 is
common to all sites: the exceptions being 7-8 at the Clifton A site and 8-3 at Clifton B (Lovatt
1981). The soil is naturally well drained on all of the native sites and is shallow, c.10 cm, and in the
absence of rainfall, soon becomes dry. The breakdown of plant remains is mainly by insects and
under dry conditions they become the main agents in preventing acidity. Woodlice were recovered
from the Dorset and Somerset samples. Trist (1983) has recorded variation in soil pH tolerance by
G. ventricosum in arable from 5-8—6-5 and as low as 5-0—5-8 in a local condition in an old
stackyard.
THE NATIVE HABITAT
Many records in county Floras erroneously give the impression that G. ventricosum is native in
arable and woodland margin habitats. In the literature dating back to the 19th century there is
scarcely any mention of our native sites or descriptions of our native habitats. Trow (1911),
recording for Glamorgan, had no knowledge of native plants in the county which had been
collected by Groves, 1903 in BM, OXF and NMW,, and Riddelsdell, 1907 in BM. This latter record
and that of Druce, Monknash Cwm, 1929, together with White, Clifton, 1912, all refer to sites as
native but give no specific description of the native habitat. It appears that only four of the 22
native sites now known were recorded prior to c.1912 and the only native sites which had any
attention were in the Avon Gorge.
There are features of habitat which are common to all native sites of G. ventricosum in England
and Wales. There is an open sward of short calcareous grasses and herbs. The soil has good drainage
properties and the texture only varies slightly from a silty clay loam to a sandy loam. The depth is
shallow and frequently overlies rock. The surface, often littered with loose stones, has rock exposure
and is open to the wind which contributes to erosion. The sites are on slopes, some of which are
steep, or often on the brow of the slope. Such areas have well defined transverse tracks or sections of
exposed bands of rock, adjacent to which the ground is bare through climatic and rabbit erosion. The
slopes are in general facing south and normally in close proximity to the sea or a tidal estuary.
At most of the Dorset sites, the rabbit is the conservationist and, in its absence, there would be
considerable change in the open grassland conditions. G. ventricosum occurs in small patches of
broken ground between stones and surface rocks where the turf is heavily grazed and eroded by
rabbit scratching. It only grows to a height of 1-8 cm between patches of open, non-aggressive
grasses. On one Dorset site, a grass enclosure presented a varied habitat: small gorse was dispersed
and there was little broken ground. Three small sites of G. ventricosum were found where the soil
depth was c.10 cm, while over most of the enclosure there was a depth variation up to 60 cm. Gorse
was at each site and had been periodically controlled by fire. This retards the gorse in the first year
and allows an open area for the autumn germination of G. ventricosum which later attains a height
of 15-20 cm; but where germination has taken place below the gorse, the seedlings have to
compete with the gorse recovery and the light factor gives growth up to 47 cm.
48 P. J. O. TRIST
In the Gower Peninsula, a site on a slope overlooking the sea had a shallow soil with rock
exposure. The calcareous sward had scattered small gorse which had been fired. In the open sward,
G. ventricosum of 12-18 cm high had no competition with its associate grasses but, within gorse
patches, its height doubled to 18-38 cm.
The taller plants of G. ventricosum within gorse areas may, in part, be attributed to the light
factor but in these coastal gorse habitats the height may also be influenced by gorse management.
The habitat is frost free but exposed to wind and sea spray. In the micro-climate between gorse
cover, there is less exposure to wind and variation in temperature. Where gorse is periodically fired
there would be an increase in available phosphorus from the gorse ash in the top few centimetres of
soil and a release of nitrogen from the breakdown of nodules on the gorse roots. This small
addition of P and N could make some contribution to the growth height of G. ventricosum found
within areas of low gorse, which at 10 sites have been noted to occupy 25—65% of ground cover in
2m.
At the Somerset sites, the slopes have much broken ground largely due to weathering. The
broken ground provides the open requirement of G. ventricosum, in spite of Sanguisorba minor
making some ground cover and Hypericum perforatum creating low shade and competition. On the
margin of the colonies of this grass there were hawthorn, blackthorn and dog-rose, 1-5—2 m high
(1980). At a metre distant from the scrub, G. ventricosum was 15 cm high and, under scrub shade,
it had grown out to the light to 43 cm high. Similar growth has been recorded on wheat headlands;
where this grass is in competition and deprived of light it will grow to 97 cm (Trist 1983). There is a
big difference between the 55 cm height of gorse in the Gower and the 2 m height-of hawthorn at
the Somerset sites, but the deprivation of light has a similar effect on the growth of G. ventricosum.
If scrub is not controlled, there is a gradual loss of habitat caused by the advancing growth of
parent scrub and seedlings. While G. ventricosum can withstand some light deprivation,
responding by increased growth height, colonies adjacent to expanding scrub move their ground to
more open sites. At one of the Somerset sites it is likely that an extension of the G. ventricosum
area has occurred over a number of years, during which time the scrub has increased its growth
following the abandonment of livestock grazing. Recorders in the past have noted fluctuations in
site populations and positions. While the climatic influence on seed production is of great
significance, this is only one facet. The survival of populations is dependent on seed germination in
an open habitat and this latter is more important. In 1981, R. G. B. Roe (in litt.) reported ‘‘there
was a good crop this year again in the same general area but not specifically in the same places as in
1980”.
WOODLAND HABITATS
Where G. ventricosum has been recorded from woodland margins and rides, it can only be
considered as a short-lived casual introduction. This is supported by herbarium sheet annotation.
As examples we have “‘borders of a wood, Twine Hills, near Wells’’, 1883, sheets in CGE and BM;
and ‘‘a clearing in woodland, near Buck’s Green, Sussex’”’, 1941, sheets in BM and K. In the
former, the wood border record is giving the impression of a habitat but the site no doubt refers to
an arable headland bordering a wood. In the latter, it is known that there are arable lands on either
side of the wood. There seems little doubt that the grass was introduced on boots or on wheels
from the arable land. The only record for Yorkshire is from a ride in Brocodale Wood, Wentvale,
1937, recorded by W. A. Sledge who considered it a gamekeeper’s introduction.
THE ARABLE HABITAT
The movement of G. ventricosum seed from its native habitat and its establishment as a colonist in
arable cultivation is discussed by Trist (1983). All of the features of the native habitat are absent
from the arable habitat.
Although this grass as an arable colonist is now only known in South Hampshire, it was formerly
reported from many counties some 50-150 km from the sea. In arable land it survives under very
different conditions and there is no preference for slope or aspect. It has been found in a wide
range of soils from light gravel sands to heavy clay, which in turn have poor to satisfactory
drainage. Its tolerance to a lower pH has been recorded. Its arable weed associates are naturally
different from those of its native grassland. Probably the most significant native habitat factors are
the maximum light requirement and an absence of plant competition; under cereals both of these
STATUS OF GASTRIDIUM VENTRICOSUM 49
factors are restricted, especially during the early phases of establishment. The author has surveyed
a large number of cereal crops from Dorset to Surrey where G. ventricosum was formerly reported:
in all fields, the crop density made it impossible for this grass to survive and no plants were seen.
SALT TOLERANCE
A feature of the habitat of G. ventricosum is proximity to the sea. In the majority of sites, this has
been found to be between 30 and 100 m, although one Somerset site is 500 m and another 20 km
from a tidal estuary. Many of the cliff sites are subject to sea spray and there is evidence that this
grass has a tolerance to a low degree of salinity. To the east of Lymington, plants were found in
arable land which had recently been flooded with sea water and had good growth (Trist 1983). This
area may be the reference given by Johns (1893) under Gastridium lendigerum ‘‘growing in fields
near the sea occasionally overflowed in the south of England’’. Other saline habitat records include
“sea shore near Brighton” J. Hardy, 1840 (in MANCH) and Hanbury & Marshall (1899) record
“edge of low marshy ground above the sea shore’’, Eastwear Bay, Kent; these sites could have
been subject to high tides. Swete (1854) records in the tidal Avon Gorge, “‘side of the river below
Cook’s Folly’’; White (1887) records that it was refound on the “‘bank of Avon” by W. E. Green
prior to 1882.
THE PLANT ASSOCIATES OF G. VENTRICOSUM
The associated species set out in Table 3 do not include species occurring less than four times. The
total species recorded at 22 native sites was 110. The records from the Rhaetic Limestone show
numbers of less than half those recorded from the Purbeck and Carboniferous Limestone, but it is
known that the species on the former were under-recorded. The species list reflects a dry, well
drained, calcareous soil which is common to all sites. Sanguisorba minor occurred in 18 sites,
Thymus praecox subsp. arcticus in 13 and Koeleria macrantha and Pilosella officinarum in 11 sites.
Where G. ventricosum can be presumed to be native, the site is generally species rich. Some of the
associated flora is rare and is exemplified by Althea hirsuta, first found in 1875 near one of the
Somerset sites and refound in 1950 in a similar habitat of “open stony pasture’ (Roe 1981).
Further examples of other interesting species are found in Ophrys sphegodes, Linum bienne and
Gentianella anglica in Dorset; Bromus madritensis, Potentilla tabernaemontani and Rubia peregrina
in the Avon Gorge; and the last two species which also occur with Scilla verna in the Gower. The
associated flora of the arable colonist G. ventricosum included Agrostis gigantea, Cirsium arvense,
Polygonum aviculare and Veronica persica, a collection of arable weeds which reflects the alien
habitat.
BRIEF HISTORY OF RECORDS
The earliest known record of G. ventricosum in Britain is given by Ray (1688), ‘“‘near Tunbridge
Wells, Kent’’, found by a Mr Doody who assisted Ray with his Synopsis and Historia Plantarum.
There is no account of the habitat but, 200 years later, Wolley-Dod (1937) recorded “‘locally plentiful
in cornfields between Tunbridge Wells and Eridge Rocks, 1883”. In 1726, Dillenius found at Norton
St Philip ‘in a field sown with trefoil’, the first record for North Somerset. There are no further early
records until we have the first from North Devon at Woodbury and Cornwood, 1784 (Martin &
Fraser 1939); the first record in the Avon Gorge is by Dyer, 1789 ‘“‘near the New Hotwell, Clifton”
(Shiercliff 1789) and Mansel-Pleydell (1895) gives the first record for Dorset as 1799.
Since about 1850 up to modern times there have been very few records described from native
habitats, but there has been much recorded error in describing arable land and woodland as native
habitats. Perhaps with the exception of the Avon Gorge, there has been no continuity of records in
the 34 vice-counties in which G. ventricosum has been recorded; and only two to three records per
county are known for 14 vice-counties, which were widely dispersed from Herefordshire to
Yorkshire. The distribution illustrates the movement of the cereal colonist by seed corn
transportation. The recognized seasonal fluctuation in the germination of this grass would also lead
to the abandonment of search for records. Over the period 1850-1950 there were considerable
fluctuations in the national acreage of cereals and, where G. ventricosum had become established in
arable land, there would be frequent loss of the habitat by land going down to grass. In the past 30
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STATUS OF GASTRIDIUM VENTRICOSUM 4
TABLE 4. GASTRIDIUM VENTRICOSUM: LAST KNOWN RECORDS IN VICE-COUNTIES,
1823-1979
Vice-county Location Source Habitat Year
Pembs. 45 ? J. L. Knapp, BRIST arable 1824
E. Norfolk 27 ‘Gillingham & Cley Smith (1824) arable 1823
Hunts. 31 Monks Wood, Abbots Ripton A. Fryer, OXF arable 1882
Caerns. 49 Aber A. Ley, BM waste 1886
E.. Kent 15S Staplehurst E. S. Marshall, BM, BRIST, OXF arable 1893
Merioneth 48 Aberdovey D. A. Jones, in MS Flora of waste 1907
Merioneth, NNW
Selkirks. 79 Galashiels I. M. Hayward, OXF waste 1916
Scilly 1b Old Town, St Mary’s H. Downes, in Thurston & Vigurs roadverge 1922
(1922)
E. Cornwall 2 Nr. St Austell railway W. Tresidder, in Thurston waste 1928
station (1929)
Surrey 17 By the Way & Arun Canal J. E. Lousley, RNG arable 1936
S.W. Yorks. 63 Brocodale Woods, Wentvale W. A. Sledge, herb. W.A.S. woodland 1937
N. Essex 19 Gosfield C. E. Hubbard, K, OXF, herb. arable 1939
E. C. Wallace
Herts. 20 Near Cowheath Wood, S. Phelp, in Dony (1967) arable 1946
Bayford
N. Devon 4 Northam Burrows Wright (1949) waste 1948
S. Essex 18 West of Hadleigh Castle E. C. Wallace, K, OXF, herb E.C.W. grassland 1950
W. Suffolk 26 Lakenheath J. E. Lousley, BM waste 1952
W. Kent 16 Chattenden Wood F. Rose, MNE woodland 1954
W. Sussex 13. Between Crawley & Horsham B. Welch, BM arable 1954
Herefs. 36 Eywas Harold Common F. M. Day, K waste be)
Worcs. 37 Evesham C. W. Bannister, BM waste 1958
Mons. 35 Near Raglan E. K. Horwood, B.R.C. field record arable 1958
Warks. 38 West of Alcester M. Clarke & J. Kiernan, WAR arable 1960
E. Suffolk 25 Clapper Farm, East Bergholt P. H. Raven & J. F. M. Cannon, BM grassland 1961
Beds. 30 Ampthill H. J. M. Bowen, K waste 1962
W. Cornwail 1 Trelissick Gardens, B. E. M. Garratt, no voucher, wall 1979
King Harry Margetts & David (1981)
years there have been very few reports of G. ventricosum but an understanding of the native site
has been developing while the arable colonist has been disappearing. Apart from the last
remaining arable colonist stronghold in the Lymington area, where this grass has been known for
over 100 years, there has been only one recent report of an arable colonist: ‘‘arable between
Preston and Osmington, Dorset’, B. Marcan, 1972 in BM. This record from arable land was
adjacent to a cliff path where H. J. M. Bowen recorded a native site in 1955. The improved
technology of cereal production has by its crop density made it impossible for this grass to continue
to thrive in the cereal habitat (Trist 1983). The modern study of the native habitat has led to
increased observation within areas where sites have previously been recorded. While there are still
relatively few known sites, this grass can be found with large populations.
The last known records of G. ventricosum from 25 vice-counties over the period 1823-1979 are
given in Table 4.
CLIMATIC INFLUENCE ON PLANT POPULATIONS
The average annual rainfall at the rain gauge stations nearest the G. ventricosum sites is given in
Table 5 and shows a significant variation in its range of 752-1178 mm. From records of height
growth and plant populations recorded at the native sites, there is no indication that either of these
factors are influenced by total annual rainfall.
Lovatt (1981) records that the Avon Gorge populations have mainly been found in unshaded,
52 P.3: OMERBE
TABLE 5. ANNUAL RAINFALL AVERAGES FOR 1941-1970 TAKEN FROM THE NEAREST
APPROPRIATE RAIN GAUGE STATION TO THE GASTRIDIUM VENTRICOSUM SITES AND
ADJUSTED FOR ALTITUDE DIFFERENCE
Location Rain Gauge Station
——_—— > Average
Site Altitude Site Altitude Annual Rainfall
m m mm
N. Somerset 50 The Grange, Somerton 32 752
S. Somerset 60 Chapel Cleeve 27 793
Avon Gorge, W. Gloucs. 30-100 Oakfield Road, Clifton 61 851
Dorset 50-100 Panorama, Swanage 37 906
S. Devon 54 Rea Barn Road, Brixham 50 950
Gower Peninsula, Glam. 30-50 Penmaen 87 1178
south-facing rocky habitats, prone to desiccation. The ‘rocky habitat’ is common to most of the
known native sites and, as the grass has continued to survive in this habitat, it is some indication of
its ability to withstand dry conditions. This is further exemplified by Lovatt when he comments that
long droughts and high temperatures, causing moisture deficit, kill out competitors to the
advantage of G. ventricosum: the resulting bare ground provides an opportunity for the survival of
more plants in the following season. However, where this grass is associated with scrub, a dry year
may have some affect on other grasses and herbs but not very much effect on scrub seedlings and
the increase in G. ventricosum plants in the following year, encouraged by bare ground, may be
temporary. In the end, the advance of scrub defeats this grass, not by deprivation of moisture but
by crowding out and excessive shade, in fact by the loss of essential open habitat.
Lovatt in the same paper says “‘that lack of rain is the most important weather factor but long
periods of sun and high temperature are also required’’. The question of amounts of rain and
periods of sunshine has to be related to the seasonal demands in the development of the plant. It is
the relative absence of rain in a hot dry summer which gives the right conditions for good seed
production and some amount of rain in the autumn to encourage germination.
If the climatic conditions for seed production can be met, then the extreme exposure of native
cliff habitats, which are frost free, can be tolerated, and if rabbits are present, the loss of some
seedlings by grazing or disturbance is compensated by grazing control of other plants and seed
burial.
If a grazing site is abandoned, such as is the case at one Somerset site, and runs to scrub, long grass
and herbs, the chance of existing populations surviving in later years will not only depend on climatic
influence but on grass and scrub control by man and animal. At the same site in 1980, there was a
colony of 18 G. ventricosum plants spread over 30m? on a steep slope with broken ground and with
2m high thorn scrub creating a fair amount of shade. At an adjacent site 35 m along the slope where
the grass was much shorter, the herbs smaller and there was no scrub, there was a colony of 70 plants
in 0-25 m*. This discussion follows the conditions required for the maintenance of a population, give
or take the annual differences caused by ‘climatic influence’: a good or poor seed year. It does not
explain the very wide annual variations in populations which occur.
Of 22 native sites, all but three are less than 1 km from the sea and this affinity appears as a
strong feature in the habitat requirement of this grass. Two of these three sites are in the Avon
Gorge, and both are 8 km from the sea. One of the Somerset sites that probably represents the
eroded remains of a cliff, is now some 20 km from the sea. It is of interest to consider that in
Britain, G. ventricosum is probably at its northern limit in the south-west and prefers a mild coastal
habitat. By contrast, in early July 1983, the author found this grass at Le Chateau Bonaguil in Lot-
et-Garonne which is c. 100 km from the tidal estuary of the R. Garonne. This surprise is explained
if we turn to Rouy (1913) who records a wide distribution ‘‘dans une grande partie de la France’.
Table 6 gives population estimates of G. ventricosum at 24 native sites in Britain. The causes of
the variation in populations from site to site are manifold.
STATUS OF GASTRIDIUM VENTRICOSUM 53
TABLE 6. POPULATION ESTIMATES AT 24 NATIVE SITES OF GASTRIDIUM VENTRICOSUM IN
BRITAIN
Estimated
number of
Locality Ey plants Year
Clifton B 34 4 1980
Dorset F 9 15 1981
Dorset G 9 20 1981
Is. of Wight 10 20 1982
Gower D 41 20 1980
Gower E 41 25 1980
Clifton A 34 ZS 1980
Dorset A 9 25 1980
Dorset J 9 30 1980
S. Somerset A 5 45 1980
Dorset I 9 65 1980
Gower C 4] 80 1980
S. Somerset B 5 100 1982
S. Somerset C 5 100 1982
Dorset B 9 100 1981
Nr. Bridgend 41 100 1982
Dorset H 9 110 1980
N. Somerset 6 150 1980
Gower B 41 155 1980
Gower A 4] 330 1980
Dorset D 9 730 1980
Dorset C 9 870 1980
Dorset E 9 1000+ 1981
S. Devon 3 1000+ 1983
CONSERVATION
The conservation status of G. ventricosum is generally satisfactory. Most sites are off the beaten
track, often in areas of rough walking, and have some difficulty of access, which would deter the
average rambler. It can therefore be said that public pressure is not a problem.
The Dorset sites are wardened and the rabbit is allowed to survive and act to advantage as a
grazier. Sites with encroaching hawthorn and those with gorse do present a threat to habitats, but
some control is being exercised. At the Somerset sites, in the absence of livestock grazing, animal
droppings and inorganic fertilizers, the phosphorus status is lowered and unchecked hawthorn and
dog rose is encouraged. However, at one site, some action has been taken recently by the County
Trust for Nature Conservation and plans are in hand to control scrub at another.
Gorse is a frequent associate of G. ventricosum but, on the coastal habitats of the Gower, there
has been a time-honoured custom of burning the gorse annually or at least every 2-3 years. An
additional control is the fact that sites are on shallow soils over rock, so that growth height and
total cover is checked. Also in the Gower, there is control of reversion to rough grazing by sheep.
On one of the Dorset sites, a large ungrazed enclosure has three colonies of G. ventricosum
associated with gorse. The latter is to be controlled and cattle grazing re-introduced.
The small colony on a cliff face in the Isle of Wight is probably made up of colonists from a
native site further back on the slopes, not yet found. The site is in danger of wind and wave erosion
and no action can be taken.
54 P. J. OUERISE
ACKNOWLEDGMENTS
I am much indebted to a large number of people who have given assistance with information and
help in the field. In particular I thank C. W. Bannister, Dr H. J. M. Bowen, Dr D. E. Coombe,
Mrs P. J. Copson, G. Ellis, T. Evans, F. Fulcher, J. H. Hemsley, P. Johnston, Dr Q. O.N. Kay, J.
G. Keylock, Dr C. M. Lovatt, Mrs B. Marcan, L. Margetts, D. McClintock, Mrs M. H. Marsden,
R. J. H. Murray, E. G. Philp, M. Porter, Capt. R. G. B. Roe, Dr F. Rose, Mrs P. Ryan, B.
Shepard, E. Spackman, Dr W. A. Sledge, Dr C. A. Stace, Mrs F. Le Sueur, Mrs S. E. Thomson,
M. A. Turner, J. Venner, E. C. Wallace and T. C. E. Wells. I am also indebted to Dr N. H. Pizer
for his help in the examination of soils and pH determinations; to C. D. Preston for the
preparation of the distribution map; to the Curators of various herbaria cited in this paper for
permission to cite records and to others who have provided herbaria and personal records; to the
World Wildlife Fund for their interest and kind assistance with a grant for the 1980 survey. The
annual rainfall averages were supplied by courtesy of the Director General of the Meteorological
Office who is thanked. Finally I am grateful to Dr D. E. Coombe who read the draft manuscript
and who offered much valuable advice and guidance.
REFERENCES
Dony, J. G. (1967). Flora of Hertfordshire, p. 111. Luton.
Druce, G. C. (1929). Plant notes. Rep. botl Soc. Exch. Club Br. Isl., 8: 765. S
Hansury, F. J. & MarsHALt, E. S. (1899). Flora of Kent, p. 394. London.
HUuBBARD, C. E. (1968). Grasses, 2nd ed., p. 313. Harmondsworth.
Jouns, C. A. (1893). Flowers of the field, Appendix, p. 16. London.
Lovatr, C. M. (1981). The history, ecology and status of Gastridium ventricosum (Gouan) Schinz & Thell. in
the Avon Gorge, Bristol. Watsonia, 13: 287-298.
MANSEL-PLEYDELL, J. C. (1895). Flora of Dorset, p. 303. Dorchester.
Marcetrts, L. J. & Davip, R. W. (1981). A revision of the Cornish Flora, p. 316. Redruth.
Martin, W. K. & Fraser, G. T., eds (1939). Flora of Devon, p. 691. Arbroath.
PERRING, F. H. & WALTERS, S. M. (1962). Atlas of the British Flora, p. 400. London.
PERRING, F. H. & FARRELL, L. (1977). British red data books, 1. Vascular plants, p. 78. Lincoln.
Ray, J. (1688). Fasciculus Stirpium Britannicarum, pp. 11-12. London.
Rog, R. G. B. (1981). The flora of Somerset. Taunton.
Rouy, G. (1913). Flore de France, 14, p. 74. Paris.
SHIERCLIFF, E. (1789). The Bristol and Hotwell Guide, p. 70. Bristol.
SmiTH, J. E. (1824). The English Flora, p. 88. London.
SwETE, E. H. (1854). Flora Bristoliensis, p. 88. London.
TuurstTon, E. (1929). Note on the Cornish flora. J. R. Inst. Cornwall, 23: 74-86.
TuuRSTON, E. & Vicurs, C. C. (1922). A supplement to F. Hamilton Davey’s Flora of Cornwall. J. R. Inst.
Cornwall, 21: 1-172.
Trist, P. J. O. (1983). The past and present status of Gastridium ventricosum (Gouan) Schinz & Thell. as an
arable colonist in Britain. Watsonia, 14: 257-261.
Trow, A. H. (1911). Flora of Glamorgan, p. 175. Cardiff.
White, J. W. (1912). Bristol Flora, p. 649. Bristol.
Wo Ley-Dop, A. H. (1937). Flora of Sussex, p. 503. Hastings.
WriGut, F. R. E. (1949). Plant notes. Trans. Devon Assoc., 81: 59.
(Accepted November 1984)
Watsonia, 16, 55—63 (1986) 55
Observations on the morphology and fertility of Juncus x
surrejanus Druce ex Stace & Lambinon in north-western Wales
T. H. BLACKSTOCK
Nature Conservancy Council, Penrhos Road, Bangor, Gwynedd, LL57 2LQ
and
R. H. ROBERTS
51 Belmont Road, Bangor, Gwynedd
ABSTRACT
The range of morphological variation of hybrid plants from a number of different populations of Juncus x
surrejanus Druce ex Stace & Lambinon (Juncaceae) in north-western Wales is described. The variability of the
hybrid is compared with that of its parent species, J. acutiflorus Ehrh. ex Hoffm. and J. articulatus L., growing
in the same localities. The production of viable (i.e. germinable) seed by this hybrid, as found in many of the J.
X surrejanus populations investigated, is reported for the first time from British plants, which had previously
been described as seed-sterile.
INTRODUCTION
The hybrid Juncus acutiflorus Ehrh. ex Hoffm. X J. articulatus L. was first found by W. H. Beeby
in Surrey in 1882 (Beeby 1884). Specimens sent to F. Buchenau for his opinion were tentatively
identified as ‘J. lamprocarpus, Ehrh., var. macrocephalo affinis”, but with the further comment
that they were ‘‘more likely a hybrid between lamprocarpus and acutiflorus.’”’ Beeby, however,
having observed the plant during a second season, was impressed by its complete failure to produce
seed and its intermediacy in some respects between the two species. Consequently, in the following
year, he sent a further gathering of specimens to Buchenau (Beeby 1885) and at the same time
drew his attention to the constant sterility of the plant. To this Buchenau replied that “from the
form of the perianth, and from your observations, probably = J. acutiflorus x lamprocarpus.” This
hybrid was listed as “x Juncus Surrejana Dr.” by Druce (1929) but the name Juncus X surrejanus
Druce ex Stace & Lambinon has only recently been validated (Lambinon & Stace 1983).
Cytological evidence was obtained by Timm & Clapham (1940), who showed that while J.
acutiflorus had a chromosome number of 2n = 40 and J. articulatus 2n = 80, the plants they had
identified as the hybrid had 2n = 60 and a study of meiosis in these plants confirmed their hybrid
origin. Further chromosome counts of 2n = 60 for this hybrid have been obtained by Zandee
(1981) from various localities in western Europe.
Although Timm & Clapham found that J. x surrejanus was very common in the Oxford area,
subsequent records of it in the British Isles have been very uneven. A recent map of its distribution
in Wales (Ellis 1983), for example, shows a preponderance of records in Merioneth, v.c. 48, where
it has been recognized in many places by Benoit & Richards (1961), but only a very thin sprinkling
of records elsewhere. This is possibly because identification of the hybrid is not always as
straightforward as the published descriptions suggest. Whilst Timm & Clapham, like Beeby,
emphasized the complete sterility of the hybrid plants they had studied, Chapple (1948) found that
though, in general, capsules are not formed, when they are, “‘an occasional unformed seed will be
found’’. Chapple’s use of the word “unformed” was unfortunate: no doubt what he meant was
““poorly-developed”’.
56 T. H. BLACKSTOCK AND R. H. ROBERTS
Subsequent descriptions of J. x surrejanus have either overlooked or ignored Chapple’s
observations and most of them follow Timm & Clapham (1940) in emphasizing the complete
absence of seed production (Richards 1962; Stace 1975). Nilsson & Snogerup (1972), on the other
hand, state that, rarely, one or a few seeds are formed in some capsules, and their illustration of
the hybrid shows a moderately well-developed capsule.
Our own interest in this hybrid arose from our independent observations that plants collected
from a number of localities in north-western Wales, which appeared from their morphology to be
this hybrid, occasionally produced plump, normal-looking seeds. Furthermore, the extreme
variation found amongst the hybrid plants was much greater than one is led to expect from
published descriptions. Consequently, in 1983 a more extended search was made for this hybrid to
enable its morphological variability to be investigated more thoroughly, and sufficient material was
gathered late in the year in order to estimate the frequency with which seeds occur and to test their
viability. It was also possible to locate plants in two localities in Anglesey and Caernarvonshire
where Zandee (1981) had previously reported hybrid material confirmed by him cytologically.
TABLE 1. COMPARATIVE CHARACTERS FOR J. X SURREJANUS AND ITS PARENT SPECIES**
Rhizome system
Flowering stem
*Length of flowering
stem, cm
*Number of leaves per
stem
Appearance of leaves
*Length of second leaf
from stem apex, cm
Flattening of leaves
Inflorescence branches
Peduncles of flower
heads
*Number of heads of
flowers per
inflorescence
*Number of flowers per
head
*Length of perianth
segments, mm
Length of inner
perianth segment
relative to outer
Tips of outer perianth
segments
Length of ripe capsule
relative to perianth
Colour of ripe capsule
J. articulatus
Subcaespitose or shortly
creeping
Decumbent, flexible
35-8 (s.d. 15-4), 14-58
3-7 (s.d. 1-2), 2-8
Ribbed, dull, soft and
curved
8-8 (s.d. 3-4), 3-0-15-3
Strongly compressed
Generally few;
ascending at a narrow
angle to the vertical
Long
18-2 (s.d. 11-9), 3-55
6-4 (s.d. 2-6), 2-15
2-6 (s.d. 0-2), 2-3-3:3
Equal
Straight
Longer (up to 3-5 mm)
and protruding
Black and shining
J. acutiflorus
Far creeping
Erect, stiff
71-7 (s.d. 16-4), 27-110
2:7 (s.d. 0-6), 2-4
Smooth, shining, stiff
and straight
28-7 (s.d. 6:2),
18-0—41-2
Subterete
Many; basal long,
ascending almost
vertically; shorter
branches widely
spreading
Short
50-1 (s.d. 24-9), 11-102
8-3 (s.d. 3-3), 2-20
2-1 (s.d.0-2), 1-5=2°5
Greater
Curved outwards
Longer (up to 3-0 mm)
and protruding
Brown
* Figures given represent sample mean (standard deviation), range.
** Data for all characters are based on plants collected from populations in north-western Wales.
J. X surrejanus
Far creeping
Intermediate
68-5 (s.d. 22-0), 20-126
3:8 (s.d. 0-8), 2-6
Smooth, somewhat shining,
moderately flexible and
often curved
14-0 (s.d. 4-7), 6-2—27:8
Subterete or somewhat
compressed
Variable in number; long
branches ascending almost
vertically; shorter branches
widely spreading
Variable; mostly
intermediate
24-9 (s.d. 15-9), 3-65
11-7 (s.d. 5-6), 2-34
2:7 (s.d. 0-3), 2:0-3-4
Generally equal,
occasionally slightly longer
Often straight, sometimes
slightly curved outwards
Very little longer and
scarcely protruding
Greenish-brown
MORPHOLOGY AND FERTILITY OF JUNCUS x SURREJANUS 57
MATERIAL AND METHODS
Altogether, material was gathered from 22 plants of the hybrid from 15 separate localities in
north-western Wales; material of the parent species was also collected from each locality when
available. The hybrid was found in a variety of different habitats, including a range of lowland mire
types which had generally been partially drained and where J. X surrejanus at times occurred in
considerable abundance.
The search for seeds was made by opening undehisced capsules, and all the seeds obtained,
regardless of their quality, were placed immediately on damp blotting-paper in Petri dishes.
At the same time a count was made of the number of flowers per head and data were later
collected for culm stature, number of leaves per stem, length of the second leaf from the stem apex
and number of heads per inflorescence. A sample of five or more perianth segments from dissected
flowers was measured under the low power of the microscope, inner and outer segments being
measured separately. Capsules were also measured when present. Similar morphological data were
collected from material of the parent species for comparison.
Living material of one hybrid plant from Cors Geirch was brought into cultivation and actively-
growing root-tips were used to obtain a somatic chromosome count. Root-tips were fixed in acetic
alcohol and subsequently hydrolyzed in 0-1M HCl prior to being stained and squashed in aceto-
orcein.
J. articulatus
J. acutiflorus
J. X surrejanus
Leaf Number
Core re rereeerereressns
er rr |
eee ee
Settee e sees
.
reer ee ee ee |
37-39 40-42
1-3. 4-6 = =©7-9 10-12 13-15 16-18 19-21 22-24 25-27 28-30 31-33 34-36
Leaf Length (cm)
FicureE1. Histograms of leaf lengths of J. x surrejanus and its parent species for samples from populations in
north-western Wales. In all cases measurements were made on the second leaf from the stem apex.
58 T. H. BLACKSTOCK AND R. H. ROBERTS
RESULTS
MORPHOLOGY
Morphologically the hybrid shows a very wide range of variation in almost all characters; data for
the hybrid and both parent species are summarized in Table 1. The mean stature (69 cm) is similar
to that of J. acutiflorus (72 cm) but the range of variation (20-126 cm) is greater than in that species
(27-110 cm). The flowering stems are stouter than those of J. articulatus, but are usually flexible
and never as stiffly erect as those of J. acutiflorus. The habit of the hybrid plants is generally
intermediate between that of the parents, most of those observed being partly decumbent and only
rarely prostrate.
The leaves of the hybrid are mostly intermediate in appearance between those of the parents,
often somewhat curved, smooth and shining and never as straight and stiff as those of J.
acutiflorus, nor as dull and strongly compressed as those of J. articulatus. The number of leaves per
stem varies front2 to 6, compared with 2 to 4in J. acutiflorus and 2 to 8 in J. articulatus. Although
there is overlap with its parent species in leaf length (Fig. 1), the leaves of the hybrid tend to be
intermediate (mean 14-0 cm) between the much shorter leaves of J. articulatus (mean 8-8 cm) and
the much longer leaves of J. acutiflorus (mean 28-7 cm). These mean values agree remarkably
closely with those given by Timm & Clapham (1940) for plants from the Oxford district: J.
articulatus, 9-3 cm; J. X surrejanus, 13-8 cm; and J. acutiflorus, 28-0 cm.
The inflorescence in general resembles more closely that of J. articulatus, both in its longer (but
more spreading) branches and in the number of heads per inflorescence (range 3-65). The number
of flowers per head is often much greater (range 2-34) than that reported by Timm & Clapham
(1940) (range 3-6), and the heads themselves are often enlarged by continuous flowering, so that
there can be a larger number of flowers per head than in either J. acutiflorus or J. articulatus (Fig.
2). This was also remarked on by Timm & Clapham (1940) and by Nilsson & Snogerup (1972).
eee ee reeee
60 pe reer ee ee eee eee J. articulatus
Sheer iaee J. acutiflorus
SS Xx suirejands
50 eee
!
I
40 !
” |
Tb I
© I
2 ee
=
mao
—_
rab) |
2 1
E
3 1
FA 20 |
!
!
|
Ema PS EAL ee
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10
i |
! [6 PE Saseste crises
I
I ;
ee cas eo tee eceeeees ——
1-3 4-6 7-9 10-12 13-15 16-18 19-21 22-24 25-27 28-30 31-33 34-36
No. of flowers per head
Ficure 2. Histograms of number of flowers per head in inflorescences of J. X surrejanus and its parent species
for samples from populations in north-western Wales.
MORPHOLOGY AND FERTILITY OF JUNCUS x SURREJANUS 39
J. articulatus
J. acutiflorus
J. < surrejanus
Number of segments
1:3-1-4 1-5-1-6 1-7-1-8 1-9-2-:0 2-1-2-2 23-24 2.5-2-6 2.7-2:8 2-9-3-0 3-1-3-2 3-3-3-4
Length of perianth segments (mm)
FicureE 3. Histograms of perianth segment lengths of J. X surrejanus and its parent species for samples from
populations in north-western Wales. Measurements of both inner and outer perianth segments are included.
A feature of the hybrid plants which becomes more noticeable later in the season (mid-October
to November), and which can give some of them a striking and attractive appearance, is the ability
to proliferate both from the nodes of the stem and from the axes of the flower heads. Such plants
were also noticed by Timm & Clapham (1940) who were undoubtedly correct in suggesting that
plantlets arising from proliferation constitute an effective method of vegetative reproduction.
Flower size in the hybrid and its parent species is very variable, as can be seen in Fig. 3. The
mean length of perianth segments in different hybrid plants ranged from 2-2 mm -— similar to that in
many plants of J. acutiflorus — to 3-3 mm, which is larger than that in any of the plants of J.
articulatus studied. The size of the flowers varies a good deal even among those of a single head,
e.g. 2-4 mm-—3-4 mm in the Plas Bog hybrid, and the mean for the aggregate of all samples of
perianth segment length in the hybrid (2-7 mm) is greater than the mean in either J. acutiflorus (2-1
mm) or J. articulatus (2-6 mm). The inner perianth segments are longer than the outer in J.
acutiflorus, whereas in the hybrid they are equal in length, or the inner may be very slightly longer.
The frequency with which capsules are developed in different plants of the hybrid varies from
20% to almost 100% of the flowers. The capsules are more or less intermediate in shape between
those of the parents and their colour is greenish-brown in contrast to the shining black capsules of
J. articulatus and the brown capsules of J. acutiflorus; they are also shorter in relation to the
perianth so that only the beaks of the capsules protrude beyond the perianths at maturity. This is
also true of capsules in which seeds have developed, and only rarely do these show a better
development than the empty ones. It is possible that the capsules swell in response to pollination
even though fertilization does not take place. |
60 T. H. BLACKSTOCK AND R. H. ROBERTS
Timm & Clapham (1940) found that examination of pollen in aceto-carmine was not helpful in
distinguishing the hybrid as about 80% of the tetrads appeared normal. Lambinon & Stace (1983),
on the other hand, state that the pollen of the hybrid is largely sterile. The pollen of hybrid plants
in north-western Wales was found to be highly variable in both quantity and quality. Anthers and
pollen from freshly-gathered material of J. X surrejanus as well as its parent species were stained in
lignin pink. In contrast to J. acutiflorus and J. articulatus, where abundant and evenly-formed
pollen tetrads are produced, the anthers of many hybrid plants were either empty or contained
only very few grains. In others there was a full complement of pollen, but closer examination
showed that often a high proportion of the grains had malformed tetrads with one or more of the
units either smal! and misshapen or completely lacking. Most anthers also contained some
completely empty grains, and in a few instances a small supernumerary unit was observed
amongst the other four parts of the tetrad. Only occasional plants had a relatively high proportion
(up to c. 60%) of normal-looking pollen.
SEED PRODUCTION
Out of 22 plants of the hybrid examined only two failed to yield any seed at all. In the other 20 the
proportion of flowers which produced seed varied from 0-3% to 20%. The quality of the seed
varied from completely empty testae through thin, partly-developed seeds to some which were
plump and normai-looking; these last were as large as the seeds of J. articulatus and occasionally
larger (up to 0-75 mm long). The quality of the seed also varied from one flower to another on the
same plant and even within the same capsule, when more than 1 seed was present. In the great
majority of plants (16), seeds were found to occur singly within the capsules, only rarely were two
or three found together and only in one plant were there more, as can be seen from Table 2, which
also shows the number of flowers examined, the number of seeds found and the number of these
which germinated.
TABLE 2. SEED PRODUCTION AND GERMINATION IN J. xX SURREJANUS
Max. no.
No. flowers No. seeds seeds No. seeds
Plant no. Locality (v.c.) examined found capsule ' germinated
1 Cwm Nantcol (48) 85 1 1 0
2 Bethesda (49) 92 6 1 2
3 Cors Geirch (49) 44 3) 1 2
4 Hirdre Uchaf (49) 10 2 2 0
5 Nant Ffrancon, 1 (49) 530 24 1 9
6 Nant Ffrancon,2 (49) 380 36 1 |
7 Pant Glas, 1 (49) 84 28 8 8
8 Pant Glas, 2 (49) 59 3 1 0
9 Llyn Ystumllyn (49) 85 0 — —
10 Craig Wen, 1 (52) 63 0 — =
11 Craig, Wen, 2 (52) 127 l 1 0
12 Gwenfro Isaf, 1 (52) 45 7 3 0
13 Gwenfro Isaf, 2 (52) 38 3 il 0
14 Llanbedrgoch (52) 390 1 1 1
15 Lianeilian, 1 (52) 82 10 3 3
16 Llaneilian, 2 (52) 32 6 1 0
17 Malltraeth Marsh (52) 51 1 1 0
18 Mynydd Bodafon (52) 67 4 1 3
19 Newborough
Warren, | (52) 88 1 1
20 Newborough
Warren, 2 (52) 270 5 l 2
21 Plas Bog, | (52) 170 14 5
pa Plas Bog, 2 (52) 90 l l 0
TOTAL 2882 157 39
MORPHOLOGY AND FERTILITY OF JUNCUS x SURREJANUS 61
SEED GERMINATION
The first seed germinated five weeks after sowing; others germinated at intervals from 25th
November 1983 to 12th April 1984. As soon as they had germinated the seedlings were placed on
sterilized compost. In a few instances the seedlings failed to develop chlorophyll or showed some
other abnormality, made no further growth and eventually died. Out of 39 seeds which
germinated, eight have failed, but the other 31, from eleven separate plants, are growing (some of
them quite vigorously) and it is worth noting that, of these, two are from the Cors Geirch plant
whose identity was confirmed cytologically (2n = 60).
DISCUSSION
Observations on the plants from different populations of J. x surrejanus have revealed that this
hybrid exhibits considerable morphological variation in north-western Wales. Although there is
generally a broad overlap with its parent species, the hybrid shows a degree of intermediacy in
some characters, e.g. growth habit, mean number of leaves per stem, mean leaf length and
inflorescence branch angles. In other respects, however, J. X surrejanus tends more closely to
approach one or other of its parents, especially J. articulatus, which it resembles, for instance, in
the inner and outer perianth segments of each flower being more or less equal in length and in the
frequent occurrence of vegetative proliferation. This wide morphological overlap can result in
considerable difficulties in the recognition of J. X surrejanus in the field. In our experience,
identification can only be tentative until mature inflorescences of all three taxa are available for
comparison in a particular locality. At that stage the ripe capsules of both J. acutiflorus and J.
articulatus protrude well beyond the perianth segments whereas, when capsules are formed in the
hybrid, they hardly exceed the perianths. The length of the perianth segments in at least some
flowers is generally greater than in either of the parent species and, late in the year, when
continued flowering has often taken place, the combination of large flowers with non- protruding
or undeveloped capsules and large number of flowers per head gives the hybrid a particularly
distinctive appearance, as may be seen from the silhouettes in Fig. 4.
The morphological variability of J. x surrejanus may be in part due to the range of environmental
conditions in which it and its parent species grow. The plants included in this investigation came from
a variety of lowland habitats, including damp depressions in fixed-dune grassland, partially drained
valley mires and flood-plain alluvial marshes, amongst which there is a considerable range of soil
conditions, water-table behaviour and vegetation structure. Although the majority of localities
reported for J. X surrejanus are in the lowlands, there is a specimen of this hybrid in UCNW collected
at c. 1200 feet [370 m] on Snowdon by S. M. Walters in 1950. The extreme variability of J. articulatus
is well known, and some of the characteristics of different variants of this species are associated with
certain ecological conditions (Clapham 1949), so that the morphology of hybrid progeny in a
particular locality may be influenced by an interaction of environmental and genetical factors.
All the previously published descriptions of J. X surrejanus in the British literature emphasize
the feature of complete seed-sterility for this hybrid. In marked contrast, many of the hybrid plants
examined from populations in north-western Wales have been found to be partially fertile and
produce germinable seed, albeit in usually very small quantity. Although there is no conclusive
evidence that either backcrosses or F, hybrids have become established in the wild, the possibility
that hybrid complexes might occur should be borne in mind in future population studies of these
jointed rushes. It is also not inconceivable that some of the morphological variation of J. x
surrejanus described here and elsewhere could be accounted for by the inclusion of individuals of
subsequent generations, derived at least partly from F, plants. Nilsson & Snogerup (1972) have
suggested that the plants referred to by Timm & Clapham (1940) as “‘large 80” might have resulted
from backcrossing of J. X surrejanus to J. articulatus. These “‘large 80” plants had a mixture of
characters of both J. acutiflorus and J. articulatus, but were seed-fertile with a chromosome
number of 2” = 80, which led Timm & Clapham (1940) to postulate a number of other possibilities
regarding their taxonomic status, and the situation clearly requires further cytological and
experimental investigation.
T. H. BLACKSTOCK AND R. H. ROBERTS
62
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MORPHOLOGY AND FERTILITY OF JUNCUS x SURREJANUS 63
ACKNOWLEDGMENTS
We would like to thank Dr N. Blackstock for preparing root-tip squashes, M. Hesketh for
producing the silhouettes, Dr C. A. Stace for kindly examining some of our specimens and Dr M.
Zandee for providing details of localities in north-western Wales. Our thanks are also due to the
North Wales Naturalists’ Trust and various other landowners for allowing us access to sites.
REFERENCES
Beesy, W. H. (1884). J. acutiflorus, var. macrocephalus. Rep. botl Soc. Exch. Club Br. Isl., 1: 79.
Beesy, W. H. (1885). Juncus lamprocarpus X acutiflorus. Rep. botl Soc. Exch. Club Br. Isl., 1: 96.
Benoit, P. M. & RicHarps, M. (1963). A contribution to a Flora of Merioneth, 2nd ed., p. 49. Haverfordwest.
CHAPPLE, J. E. G. (1948). Juncus acutiflorus (Ehrh.) Hoffm. x J. articulatus L. Rep. botl Soc. Exch. Club Br.
isk, 3: 370-371
CiapHaM, A. R. (1949). Taxonomic problems in Galium and Juncus, in WiLmotr, A. J., ed. British flowering
plants and modern systematic methods, pp. 72-74. Arbroath.
Druce, G. C. (1929). Notes on the second edition of the “British plant list’”’. Rep. botl Soc. Exch. Club Br. Isl.,
8: 867-877.
E.uis, R. G. (1983). Flowering plants of Wales, p. 266. Caerphilly.
LAMBINON, J. & Stace, C. A. (1983). Note nomenclaturale sur deux Juncus hybrides: J. x kern-reichgeltii
Jansen & Wachter ex Reichgelt et J. x surrejanus Druce ex Stace & Lambinon. Bull. Jard. bot. Nat. Belg.,
53: 507-S08.
Nitsson, O. & SNoGERUP, S. (1972). Drawings of Scandinavian plants 75-80. Bot. Notiser, 125: 203-211.
RICHARDS, P. W. (1962). Juncus, in CLAPHAM, A. R., TuTin, T. G. & WarBurRG, E. F. Flora of the British Isles,
2nd ed., p. 992. Cambridge.
Stace, C. A. (1975). Juncus acutiflorus Ehrh. ex Hoffm. x J. articulatus L., in StacE, C. A., ed. Hybridization
and the flora of the British Isles, pp. 465-466. London.
Timm, E. W. & CrLapHaM, A. R. (1940). Jointed rushes of the Oxford district. New Phytol., 39: 1-11.
ZANDEE, M. (1981). Studies in the Juncus articulatus L. - J. acutiflorus Ehrh. — J. anceps Laharpe — J. alpinus
Vill. aggregate, 1. A cytotaxonomic survey. Proc. K. ned. Akad. Wet. ser. C, 84: 243-254.
(Accepted November 1984)
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PLATE 1. Facsimile of page 1 of the list sent to Lord Bute. The last five pages of the list were written by Hope
but the first four are in an unknown hand. It is perhaps significant that the only two puzzling names are on page
1. See discussions of Veronica hederaceus and Valeriana pyrenaica.
Perhaps these names are the
misunderstandings or carelessness of a transcriber and they remained uncorrected by Hope. See also the
discussion of Deschampsia setacea.
Watsonia,- 16, 65-74 (1986) 65
James Robertson and the botany of Bute, 1768
J. H. DICKSON
Botany Department, The University, Glasgow, G12 8QQ
ABSTRACT
In 1768, during a visit of 18 days to the Isle of Bute, James Robertson recorded 445 plant species, comprising
357 angiosperms, one gymnosperm, 22 pteridophytes, 26 bryophytes, 16 lichens, 22 algae and one fungus. This
long list demonstrates the high standard of botanical recording in mid-18th century Scotland. Excluding about
12 species that were certainly or probably misidentified, 35 species of vascular plant have no present records for
the island. These include several geographical and/or ecologically noteworthy plants such as Anacamptis
pyramidalis, Apera spica-venti, Calamagrostis canescens, Coeloglossum viride, Cuscuta europaea, Deschampsia
setacea, Polygonum viviparum and Vaccinium vitis-idaea. In the manuscript, “‘[Valeriana] peregrina” may be a
transcriber’s error for Veronica peregrina.
INTRODUCTION
Employed at first as a gardener at the Royal Botanic Garden, Edinburgh, James Robertson
became a highly assiduous, field-working associate of John Hope, Professor of Botany in the
University of Edinburgh and King’s Botanist (Fletcher & Brown 1970). According to Horn (1966)
Robertson made the first recorded ascents of many Highland peaks. He was a skilled botanical
artist (Lightfoot 1777, plate 28; Robertson 1768). After his Scottish journeys, in the 1770s he
botanized in St Helena, Cape Province, India and China. Though Robertson has never been
entirely forgotten (Fletcher 1959; Duncan 1980; Mackechnie 1958; Slack 1958; Slack & Dickson
1959), little has been written about him and he has sometimes been ignored or unmentioned by
historians of botany.
Lodged in the library of the Marquess of Bute at Mount Stuart, Isle of Bute, are documents
which reveal the efficiency and skilfulness of botanical recording in mid-18th century Scotland.
They were written in 1768, four years before the visit by Lightfoot who published the first Flora of
Scotland (1778 but dated 1777). There is a two-page letter, with the date 12th August 1768, which
introduces a list of plants headed Plantae in Insula Bute nascentes (Plate 1). On eight pages of
double columns and one page with a single column are 436 binomials mostly of vascular plants but
also of bryophytes, lichens and algae. In addition there is a six-page document headed Remarks
made by James Robertson on the Island of Bute 1768. These documents had been sent by John
Hope to the third Earl of Bute, politician and devoted botanist. According to Hope’s letter to Lord
Bute, Robertson prepared a Catalogue of Exotic Trees and a List of the four first classes of Animals
but these appear to have been lost.
| The early history of botanical exploration in Scotland has been summarized by Fletcher (1959)
_ and Balfour (1979) and is mentioned by Fletcher & Brown (1970); in none of these works is there
any reference to the island of Bute. Botanical observations for Scottish islands prior to the mid-
18th century are not unknown but are mostly insubstantial: Perring (1953) wrote concerning
Martin Martin in the Hebrides, Sibbald (1710) concerning islands in the Firth of Forth and Raven
(1950) concerning Ray on the Bass Rock. Wallace’s list for Orkney (1700), however, includes over
250 names of vascular plants. The long list for Bute may well be the earliest extant compilation for
a Scottish island made by professional Linnaean botanists. It appears to be the earliest thorough
listing for any part of the west of Scotland.
Robertson travelled very widely in Scotland (Hope 1769; Robertson 1768). He botanized in the
Highlands at least as early as 1766, and from 1767 to 1771 made very extensive tours each spring to
66 J. H. DICKSON
autumn in the service of the Commissioners on the Forfeited Estates (Smith 1982). He kept
journals of his tours (Anonymous 1966; Mitchell 1897). In 1768 he was instructed to examine
‘“‘marine plants” (Foulis 1788) and was “. . . employed . . . in search of native plants on the sea
coast and western isles of Scotland’’, according to an editorial note in Robertson (1768). Leaving
Edinburgh on 10th May, apart from Bute, he visited Lanarkshire, Ayrshire, Wigtownshire,
Arran, Mull and Skye. From the Remarks we know that Robertson sailed on 17th June 1768 to
Kilchattan, Isle of Bute, from Brodick, Isle of Arran, where two years earlier he had been
awaited as an adviser in “. . . Farming Including planting and wood Nursery” (Anonymous
1982). On 4th July, Robertson left Bute from the north end, crossed to Colintraive and made for
Inverary (Foulis 1788). In September 1768 Robertson collected Eriocaulon on Skye (Hope 1769),
where he stayed until 8th October.
Though Hope’s British herbarium is lost, there is a notebook, kept in the Royal Botanic
Garden (Anonymous 1907; Balfour 1900, 1901), which lists the specimens under the heading A
Catalogue of British Plants in Dr. Hope’s Hortus Siccus, 1768. Despite the heading, the
Catalogue gives dates as late as the 1780s. The Catalogue mentions Bute 45 times, including nine
references to vascular plants not in the list sent to Lord Bute. There is nothing to indicate that
these additional records were made by anyone other than Robertson or at any time other than
1768. Therefore the 1768 recording of the Bute flora totals 445 species of all classes.
48 species not now known from Bute are in the list. About 12 are erroneous, several must be
treated with caution, while others, for the most part distinctive species, are phyloceo aia
notable for the Clyde area or west of Scotland in general.
The nomenclature and taxonomic arrangement of the list is that of Hudson (1762) who
adopted the Linnaean system. A copy of Hudson’s book with Hope’s book-plate and a few
annotations is kept in the library of the Botany Department of the University of Glasgow, where
there is also a second copy of Hudson signed by Hope on the title pape: These may very well be
the books used by Robertson and Hope in drawing up the list.
Diagonal crosses are placed before 31 names in the list (Plate 1). These indicate that the plants
were not known to Hope as growing in the vicinity of Edinburgh. Six names in the list are
followed by interrogation marks. Very brief comments follow four names. A pointing hand
indicates ‘‘Sisymbrium moniense’’. These are all the annotations that Hope made to the list. Only
two names of vascular plants in the list are puzzling.
It is clear from the Catalogue that Hope’s herbarium included specimens collected by
Robertson from Bute. In the absence of the specimens there can be no modern revision of the
records.
Though many of the place names mentioned in the Remarks derive from the southern half of
the island (Fig. 1), Robertson travelled widely on the island and may have visited Inchmarnock,
the small island off the western coast. He saw and commented on the varied geology of the island
which is bisected by the Highland boundary Fault and includes base-rich rocks (Hill 1979). He
visited the southernmost parts of the island but it is not explicitly stated that he climbed the
highest ground, Windy Hill, in the north, reaching 280 m. It would seem likely that such a
thorough and energetic worker did so. However, he did not record the extensive stands of
Cladium mariscus (L.) Pohl, a rarity in the Clyde area (Ballantyne 1897), from Bull Loch in the
north.
The annotations to the list, the puzzling names, the erroneous and doubtful records, as well as
the phytogeographically and ecologically notable species, are discussed below. The binomials are
reproduced as written in the list except that the ligatures of a and e are separated and the long s
as used in the 18th century is shortened (Tables 1 & 2). Modern synonyms of the Linnaean and
Hudsonian binomials (Dandy in Stearn 1973) are given in brackets and follow Clapham, Tutin &
Warburg (1981).
Though there have been several publications on the botany of Bute in recent years (Dickson
1981, 1983; Dickson & Boyd 1982; Kerr 1980; Mackechnie 1971; Marshall 1980), they are all
short contributions. However, some 535 species of vascular plants have been recorded for the
island since the last published listing by Ballantyne (1911). In making comparisons between the
list and the modern flora I have used data compiled by the Biological Records Centre (1981), and
recording from my field work in 1981 to 1985. Botanically minded members of the Buteshire
Natural History Society have been helpful.
a en
JAMES ROBERTSON AND BUTE BOTANY
To Inveraray
Colintraive
Bogany Point
Loch Greenon Rothsay ° ( Rothesay)
| (Greenan Loch) f, 3
Ascok (Ascog)
Mountstewart
Inchmarnock (Mountst uart)
e \ Kerry Lamonde
(Kerrylamont) _
Brough-hag °®
Scatpsie eet
Bay (Bruchag)
Kicating
Bay ( Kilchattan)
Kilhaton
OiKmily- whe hes Old Chapel
Omiles 1 2 (St Blane’s Church)
Dunaguile
(Dunagoil)
67
Figure 1. Map of the Isle of Bute. Place names underlined are those as written in the Remarks. Where
different, modern spellings are shown in brackets. Freshwater lochs are shown in black.
68 J. H. DICKSON
TABLE 1. SPECIES NOW ABSENT FROM BUTE
Manuscript binomials are followed by their modern equivalents in brackets. A: No records since Robertson; B:
No records since Ballantyne (1911); C: No records since Lee (1933); some of Lee’s records may refer to the
19th century.
1. Weeds and garden escapes.
Aethusa cynapium (A. cynapium L.) Fool’s Parsley
Agrostema githago (A. githago L.) Corncockle
Agrostis spicaventi (Apera spica-venti (L.) Beauv.) Silky Apera
Anthemis nobilis (Chamaemelum nobile (L.)All.) Chamomile
Brassica campestris (B. rapa L. ssp. sylvestris (L.) Janchen) Wild Turnip
Bromus secalinus (B. secalinus L.) Rye Brome
Cheiranthus cheiri (C. cheiri L.) Wallflower
xCuscuta Europaea (C. europaea L.) Large Dodder
xHelleborus viridis (H. viridis L.) Green Hellebore
[Lolium] temulentum (L. temulentum L.) Darnel
Scleranthus annuus (S. annuus L.) Knawel
[Trifolium] agrarium (*T. campestre Schreb.) Hop Trefoil
2. Plants of unimproved pasture, wetlands, heaths, rough ground and the coast.
Aira setacea (Deschampsia setacea (Huds.) Hack.) Bog Hair-grass
x[Arundo] calamagrostis (Calamagrostis canescens (Weber) Roth) Purple Smalireed
[Carex] inflata (+C. vesicaria L.) Bladder Sedge
Carex pilulifera (C. pilulifera L.) Pill-headed Sedge
[Cerastium| Semidecandrum (C. semidecandrum L.) Little Mouse-ear Chickweed
Filago germanica (F. vulgaris Lam.) Common Cudweed
Fumaria claviculata (Corydalis claviculata (L.) DC.) White Climbing Fumitory
[Lycopodium] selago (Huperzia selago) Fir Clubmoss
xOphioglossum vulgat. (O. vulgatum L.) Adder’s Tongue
[Orchis| pyramidalis (Anacamptis pyramidalis (L.) Rich.) Pyramidal Orchid
Orobus tuberosus (Lathyrus montanus Bernh.) Bitter Vetch
xPoa loliacea (Desmazeria marina (L.) Druce) Darnel Poa
xPolygonum viviparum (P. viviparum L.) Alpine Bistort
x[Satyrium| albidum (Pseudorchis albida (L.) A. & O. Léve) Small White Orchid
xSatyrium viride (Coeloglossum viride (L.) Hartman) Frog Orchid
Saxifraga granulata (S. granulata L.) Meadow Saxifrage
xVaccinium vitis-idaea (V. vitis-idaea L.) Cowberry
[Vicia] lathyroides (V. lathyroides L.) Spring Vetch
[Vicia] sylvatica (V. sylvatica L.) Wood Vetch
3. Aquatic plants
Myriophyllum spicat (M. spicatum L.) Spiked Water-milfoil
[Potamogeton|] lucens (P. lucens L.) Shining Pondweed
Sparganium natans (S. minimum Walbr.) Small Bur-reed
Zostera marina (Z. marina L.) Eel-grass
4. Certainly or probably misidentified plants. (Not recorded since Robertson’s visit).
xAcrostichum ilvense (*Woodsia alpina (Bolton) S. F. Gray) Alpine Woodsia
Bunium bulbocastanum (B. bulbocastanum L.) Great Pignut
[Carex] distans (C. distans L.) Distant Sedge
[Carex] divisa (C. divisa L.) Divided Sedge
[ Equisetum| fluviatile (* E. telmateia Ehrh.) Great Horsetail
Juncus acutus (J. acutus L.) Sharp Rush
| Potamogeton| compressum (P. compressus L.) Grass-wrack Pondweed
xPotentilla argentea (P. argentea L.) Hoary Cinquefoil
xSedum sexangulare (S. sexangulare L.) Insipid Stonecrop
Sium latifolium (S. latifolium L.) Water Parsnip
Thymus serpyllum (T. serpyllum L.) Thyme
Ulmus campestris (U. procera Salisb.) English Elm
“See Dandy in Stearn (1973). tSee Jermy et al. 1982.
Dr Prrrrrwarrnory
> rrrPrrmrrrrrUmrrUrrY>y
Dr >>
EE
JAMES ROBERTSON AND BUTE BOTANY 69
TABLE 2. ANNOTATIONS AND PUZZLING NAMES
Manuscript binomials are followed by their modern equivalents in brackets.
Cardamine hirsuta? (*C. flexuosa With.) Wood Bitter-cress
[Carex] vesicaria? (+C. rostrata Stokes) Beaked Sedge
Draba verna? (Erophila verna (L.)) Spring Whitlow Grass
Populus nigra is it indigenous (P. nigra L.) Black Poplar
Prunus cerasus Huds. Geen=ang. (P. cerasus L.) Sour Cherry
Sambucus nigra is it indigen: (S. nigra L.) Elder
[| Sisymbrium] moniense (Rhychosinapis monensis (L.) Dandy ex Clapham) Isle of Man Cabbage
x[Valeriana] peregrina? (Veronica peregrina?) American Speedwell?
Veronica hederaceus (Veronica hederifolia?) Ivy-leaved Speedwell?
*See Dandy in Stearn (1973). ¢See Jermy et al. (1982).
SPECIES NOW ABSENT FROM BUTE
More than 320 of the vascular plants recorded by Robertson are widespread, mostly abundant
species still growing on the island. The following are also still present on Bute but are more or less
rare in west-central Scotland and on the Clyde isles: Baldellia ranunculoides, Bidens tripartita,
Chaerophyllum temulentum, Conium maculatum, Glaucium flavum, Mertensia maritima,
Osmunda regalis, Parietaria judaica and Saxifraga aizoides. However, most interest centres on the
species which have disappeared from Bute since the 18th century.
Of the 47 species listed in Table 1 the great majority have not been recorded since Robertson’s
visit. Weeds of arable farming constitute an important group (category 1). Agrostemma is an
outstanding example of a bad weed rendered all but extinct in Britain by improved agricultural
techniques. According to Hennedy (1865: 23) it was ‘“. . . plentiful on Bute . . .”. Lee’s record
(1933) may refer to the 19th century. The species of Cuscuta are all very rare in Scotland now and
C. europaea is not considered native. The genus can hardly be in error but could the species have
been C. epilinum Weihe? In the Remarks Robertson observed that the inhabitants ‘‘. . . sow oats,
barley and some peas, a good deal of flax, and some hemp...” (my italics). Robertson’s
discovery of Apera, the only one for the island, may well be the first Scottish record.
The introduced and commonly ruderal Chenopodium bonus-henricus, recorded by Robertson
and last seen in 1945, is another plant which has much declined, like the arable weeds. It may have
escaped from cultivation, as probably had Tanacetum. Such as assessment applies with even
greater force to Helleborus, Chamaemelum and Cheiranthus, all well-known garden plants and
none native in Scotland. Robertson’s specimen of Cheiranthus came from Rothesay where perhaps
it grew on the walls of the ancient castle, but the species is not there now.
Category 2 (Table 1) covers a diversity of species including three plants often growing at
altitudes far higher than those reached on Bute. However there need be no doubting Robertson’s
discoveries of Huperzia selago, Polygonum viviparum and Vaccinium vitis-idaea on grounds of his
unfamiliarity with the species. In 1767 he had recorded Polygonum viviparum on hills in the
Highlands and, for Ben Wyvis, he lists Vaccinium myrtillus, V. uliginosum and V. vitis-idaea. That
Robertson knew his mountain plants is further made clear in a letter Hope wrote to Joseph Banks
in February 1767. None of these three species is restricted to high altitudes. Huperzia was last
reported for Bute by Ballantyne (1911). Still growing near sea-level in Kintyre (Cunningham &
Kenneth 1979), it could well have inhabited moorland on Bute, especially in the north of the island
where it may even linger. If it has disappeared, possibly grazing or fire was the cause (Page 1982).
Polygonum viviparum has been recorded from the hills of northern Arran and also from three of
__ the low hills of Knapdale (Cunningham & Kenneth 1979) which reach 550 m, 270 m higher than the
Bute hills. However, the cliffy ground at the northern end of the island might harbour this species
and also Vaccinium vitis-idaea which in Kintyre grows ‘“‘on most hills from c. 500 ft. up”
_ (Cunningham & Kenneth 1979). Fruitless hunts for these three species were made in 1982 and 1983
at the northern end of Bute, though some ground remains unsearched.
Not least because Robertson is known to have drawn four orchids, there are firm reasons for
believing he had a good knowledge of these plants. His record of Anacamptis remains the only one
70 J. H. DICKSON
for the Clyde area, though there is a recent record for south-western Kintyre (Cunningham &
Graham 1979). The scarcity of this calcicolous orchid in Scotland was already realized by Lightfoot
(1777) who stated “In dry pastures, but very rare, as near Kiloran in the island of Colonsa’’.
Disturbed by holidaymakers and grazing animals, the very small areas of shell sand which remain on
the western side of Bute and on Inchmarnock were searched in 1984 with negative results. Similarly,
neither Coeloglossum nor Pseudorchis has been refound but they could still be discovered, both
being inconspicuous, especially Coeloglossum. Both Deschampsia setacea and Calamagrostis need
comment. If correct, this is the earliest Scottish record of D. setacea which, though scattered over
Britain, is absent from all of southern Scotland and there are no Clyde area records (Perring &
Walters 1976). Unknown now on Bute, the genus Calamagrostis is virtually absent from the Clyde
area; the most likely species is C. epigejos, known from Kintyre (Cunningham & Kenneth 1979).
If in small stands in tall growing vegetation, another readily overlooked species is Ophioglossum
vulgatum. However, if it has disappeared from Bute, this may be part of a marked decline over the
last 300 years (Page 1982). Similarly, Corydalis claviculata may linger; perhaps referring to a record
made 30 years earlier, Lee (1933) gave the species as growing at Scalpsie. That plants known to
Robertson, but with no published records since his visit, can still be refound is shown by the
discoveries of Carex pallescens and C. pulicaris in 1982 (Dickson & Boyd 1982) and, more
surprisingly, of Populus tremula in 1984. However, aspen has only been encountered in the south-
east of Inchmarnock where there is a small stand of fairly well-grown trees. On the mainland of Bute
aspen may linger vegetatively as a small shrub in rocky ground or on a cliff as it does on Ailsa Craig
and elsewhere round the Scottish coasts. .
If only because of the state of knowledge of both taxonomy and topographical botany in the mid-
18th century, it is inevitable that doubts arise about some of the entries comprising the long list of 380
vascular plants. However there are at least twelve species which deserve detailed consideration or
can be freely accepted as mistakenly identified (Category 4, Table 1). On chorological and ecological
grounds Woodsia alpina is an impossible record. Young Cystopteris fragilis ,a species absent from the
list but occurring now on Bute, may have been the source of error (Jermy et al. 1978). Robertson also
found Conopodium majus (Gouan) Loret and not Bunium bulbocastanum; the two are superficially
similar (Tutin 1980). Carex distans may possibly still occur on Bute. However, the then undescribed
C. binervis Sm. could have been confused with C. distans, as claimed by Smith (1800) with reference
to Lightfoot. C. binervis is such an abundant plant of moorland on Bute that Robertson must have
seen it. Carex divisais unlikely to be correct; the only two modern records of C. divisain Scotland are
possibly introductions (Jermy et al. 1982). Confusion with another species is the probable explana-
tion but which species is far from clear.
In all likelihood Robertson saw the common plant now called Equisetum fluviatile L. and not E.
telmateia. E. fluviatile is readily found at Greenan Loch, where Robertson botanized, as it is at other
lakes on the island. Robertson’s Juncus acutus was certainly J. maritimus. The only 20th century
Scottish records for Potamogeton compressus were made in Angus where the species is now extinct
(Ingram & Noltie 1980). The record for Bute, likely to be an error, was perhaps based on P.
obtusifolius Mert. & Koch, well-known on the island now. Potentilla argentea is not a plant of the
Clyde area at present. It is absent from the western seaboard of Scotland except for Dumfriesshire.
A misidentification seems unlikely. The Catalogue uses both the names Argentea and Argentina, the
second followed by “‘ad vias passim” (Anonymous 1907: 169). This clearly refers to P. anserina L., as
it does in Hudson (1762). Perhaps there was a slip of the pen, the common P. anserina being
intended. However, that being the case the ‘‘X”’ is hard to explain. Hope must have known P.
anserina near Edinburgh.
The occurrence of Sedum sexangulare is extremely unlikely. It resembles S. acre, which Robertson
found. Possibly the source of confusion may have been the then undescribed Sedum anglicum which
Robertson must also have seen. It was described for the first time by Hudson (1778). Sium latifolium
is a very rare Scottish plant with only two localities, both on the eastern coast. Perhaps Robertson
saw Berula erecta (Huds.) Colville, not now known on Bute, or more likely, Apium nodiflorum (L.)
Lag., known now on the island. Robertson would have seen Thymus praecox subsp. arcticus
(Durand) Jalas and not 7. serpyllum, in Britain only found in the Breckland of East Anglia.
Ulmus procera occurs only as a planted tree in Scotland and, because there had already been
extensive tree planting on Bute long before 1708, Robertson may have seen that species. More
likely, however, he saw the common, native Ulmus glabra Huds. In his 1767 journal Robertson uses
JAMES ROBERTSON AND BUTE BOTANY a
only the name U. procera; it is highly improbable that he did not see U. glabra during his extensive
travels.
ANNOTATIONS AND PUZZLING NAMES
The few species listed with qualifications and the two puzzling names are given in Table 2. The
interrogation marks placed against Cardamine hirsuta and Draba verna are difficult to explain
unless Robertson found only poor specimens. Difficulty in separating Carex rostrata from
C. vesicaria may have made Robertson cautious and hence the interrogation mark.
The questioning of Populus nigra, if that is what Robertson saw, was well justified. Lightfoot
(1777) was also doubtful. No modern botanist has claimed that Black Poplar is indigenous in
Scotland. Perhaps also Robertson saw Sambucus only in the vicinity of buildings. Otherwise it is
difficult to see why the native status is questioned. That such a thought occurred to him gains
credance from the entry in Hope’s Calendarium Florae (Anonymous 1907; 127) which states “Aug.
3. Sambucus niger. On a rock in a den, north of Aughtermughty, therefore a native’.
‘Prunus cerasus Huds. Geen. = ang.” is a doubly noteworthy entry. It is the only one with a
vernacular name, apart from one alga, and the only one referring to Hudson by name. The latter
part I take to mean that the name in English is Gean, a name normally applied to Prunus avium
(L.) L., as is done in the Catalogue. Probably P. avium was the plant seen by Robertson. The
garden cherries, now seldom if ever seen wild in western Scotland, could well have been grown at
Mount Stuart and seen by Robertson there. However, the list avowedly consists of indigenous
plants.
Robertson, who had already found Rhynchosinapis monensis on the Ayrshire coast in 1766,
found this British endemic on Bute where it was also seen by Lightfoot (1777: 353) “‘. . . about a
mile to the south of Mount Stewart . . .”’. It is discussed by Ray (1724) and listed in both editions
of Linnaeus (1754, 1759) but is omitted from Hudson (1762) as Hope’s use of the hand symbol
indicates.
The entry ‘“‘X [Valeriana] peregrina?” is one of the most intriguing in the list. It is not a formal
binomial used by any other author. Perhaps the interrogation mark implies that “‘peregrina’”’ was
being used informally, meaning literally foreign; the plant was unfamiliar and hence not considered
indigenous. If so, this would be the only example of such a usage in the list. Moreover, elsewhere in
the list the expression “‘is it indigenous” is written. What species of Valeriana could it have been?
Valeriana peregrina purpurea albave of Bauhin, called Valeriana cornucopiae by Linnaeus, is the
modern Fedia cornucopiae (L.) Gaertner. This is an annual, arable weed of the Mediterranean
region. Not unlike a Valerianella but with conspicuous flowers, it has never been claimed as part of
the British flora. If this is what was intended, why was the Linnaean binomial not used? Perhaps
Valeriana pyrenaica should be given consideration. ‘This robust species is locally well established in
Scotland now and is known from Bute. Hope did not have a specimen but he knew the species
(Anonymous 1907: 149). If the plant Robertson found was V. pyrenaica, why was the epithet
peregrina used at all? Moreover why was the cross used? Could it be that Veronica peregrina was
intended? The close proximity of Valeriana and Veronica in the list makes this a tempting conclusion
(Plate 1). Veronica peregrina is very well established in the old walled garden at Kames (Dickson
1981). From the Remarks it seems likely that Robertson visited the castle there (Fig. 1), though the
walled garden was not constructed until 1786 (Ross 1880). Veronica peregrina is not in the Catalogue.
Therefore Hope and his associates may only have known it from the few lines in Species Plantarum
and from the figure and description in Morison (1680) where it is called Veronica annua alba
polygonifolio. That could account for both the cross and the interrogation mark. If ‘“Valeriana
peregrina’ should be read as Veronica peregrina this was the first Scottish record and one of the first
in the British Isles, long before those records made in Ireland (Bangerter 1964). Whatever the genus,
the binomial is not in Hudson. Hope should have placed a pointing hand symbol against the entry, as
was done in the list for Rhynchosinapis monensis and for 23 taxa in the Catalogue.
No 18th century author used the name Veronica hederaceus. Probably V. hederifolia L. was the
species intended. That is the name used in the Catalogue (Anonymous 1907: 148). The species is
now known to be markedly eastern in Scotland. It has not been recorded from Bute since
Ballantyne (1911).
72 J. H. DICKSON
SPECIES MISSING FROM THE LIST
There are some noteworthy absentees from the list. By these are meant distinct species, recognized
in the mid-18th century, which in all likelihood were just as common then as now. Allium ursinum
L., Drosera rotundifolia L., Hydrocotyle vulgaris L., Hypochaeris radicata L. and Rumex
obtusifolius L. are five which spring to mind. Though 30 taxa of grasses are listed, Robertson
seemingly missed Brachypodium sylvaticum (Hudson) Beauv., Deschampsia cespitosa (L.)
Beauv., Phalaris arundinacea L. and Poa trivialis L. All these species were in Hope’s herbarium.
However, Festuca rubra, though a Linnaean species and discussed by Hudson, is not in the
Catalogue and so perhaps was not known to, or misunderstood by, Robertson and Hope and this
omission can be explained. It is especially surprising that the common, conspicuous Deschampsia
cespitosa is missing from the list. Could the rare Deschampsia setacea, discussed above, be a
thoughtless transcriber’s error for D. cespitosa, unchecked by Hope? See Plate 1. Another striking
absentee is the sedge Carex nigra (L.) Reichard, which is conspicuous on Bute. Then known as
Carex cespitosa L., this species can scarcely have escaped notice by Robertson who recorded no
less than 17 sedges; the modern list totals only 23. However, none of the names used by Robertson
can be applied to C. nigra. It is hard to account for some of these omissions except to say that
nobody can find everything especially on a short visit or, perhaps, some species though recognized
or even collected never found their way on to the list finally sent to Lord Bute.
CONCLUSION
In the letter, Hope praises the list as “numerous” considering the brevity of Robertson’s visit.
Even had Robertson stayed much longer on the island this is a claim with which one can readily
agree. Robertson and Hope compiled a list of 380 species of vascular plants. It is clear that this
represents a very substantial part of the flora that there was to find. I
The disappearances of species recorded in 1768 and now extinct on Bute can for the most part be
readily explained. The many changes in agricultural practices, such as the efficient cleaning of
seeds, improvement in grassland management, drainage, fire and reclaiming of marginal land, may
well have been the major agents. The misidentified species represent only a few percent of the total
380 vascular plants and are readily understandable with regard to the state of knowledge in the mid
18th century. The likely sources of these mistaken records are widespread species familiar in the
west of Scotland.
The list in its length and detail and even in the annotations, sparse though they are,
demonstrates the assiduousness, efficiency and perceptiveness of the partnership of Robertson and
Hope. Like Hope, Robertson published very little and in consequence has suffered the undeserved
fate of being little known to British field botanists.
ACKNOWLEDGMENTS
I am most grateful to the Marquess of Bute for permission to study the documents. His lordship’s
librarians, Miss C. Armet and Mr A. Hunter were very helpful, as were Dr W. M. Boyd and Miss
Dorothy K. Marshall, honorary president of the Buteshire Natural History Society. My greatest
thanks extend to Mrs Anne Buchanan who first drew my attention to the list. Professor D. M.
Henderson and Mr D. McKean of the Royal Botanic Garden, Edinburgh, made some useful
comments.
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Proc. Soc. Antiq. Scot., 32: 11-19.
Morison, R. (1680). Plantarum historiae universalis Oxoniense. Oxford.
Pace, C. N. (1982). The ferns of Britain and Ireland. Cambridge.
PERRING, F. H. (1953). A 17th century contribution to the Scottish flora. Watsonia, 3: 36-40.
PERRING, F. H. & WALTERS, S. M. (1976). Atlas of the British flora, 2nd ed. Wakefield.
Raven, C. (1950). John Ray. Cambridge.
Ray, J. (1724). Synopsis methodica stirpium Britannicarum, 3rd ed. London.
RoBeErTSON, J. (1767). Journal. MS 2507. National Library of Scotland, Edinburgh.
RoBerTson, J. (1768). An account, and a print, of a new species of Astragalus, a plant discovered by James
Robertson in 1767. Scots Mag., 344.
Ross, W. (1880). Blain’s history of Bute. Rothesay.
SIBBALD, R. (1710). The history ancient and modern of the Sheriffdoms of Fife and Kinross; with descriptions of
both. Edinburgh.
SLACK, A. (1958). A limestone flora on Ben Sgulaird. Glas. Nat., 18: 56-57.
Stack, A. & Dickson, J. H. (1959). A further note on the limestone flora of Ben Sgulaird. Glas. Nat., 18:
106-108.
SMITH, A. (1982). Jacobite estates of the forty-five. Edinburgh.
SmiTH, J. E. (1800). Descriptions of five new British species of Carex. Trans. Linn. Soc. Lond., 5: 264-273.
STEARN, W. T. (1973). Facsimilies with an introduction. John Ray. Synopsis Methodica Stirpium Britanni-
carum. Editio Tertia 1724. Carl Linnaeus. Flora Anglica 1754 and 1759. London.
74 J. H. DICKSON
TutTin, T. G. (1980). Umbellifers of the British Isles. London.
WALLACE, J. (1700). An account of the Islands of Orkney. London.
Witson, J. (1855). Wilson’s guide to Rothesay and the Island of Bute. Rothesay.
(Accepted April 1985)
TT eee ee eS ee ———
Watsonia, 16, 75-87 (1986) 75
Short Notes
A RELIABLE METHOD FOR DISTINGUISHING BETWEEN BETULA PENDULA AND
B. PUBESCENS
The difference in leaf shape between B. pendula Roth and B. pubescens Ehrh. is usually used to
distinguish between these species. The difference normally cited is the shape of the leaf apex
(acuminate in B. pendula but acute to subacute in B. pubescens). Many field botanists recognize
the considerable degree of variation within these species (particularly in B. pubescens) and the
consequent difficulty in identifying some specimens. Accordingly, there was felt to be a need to
develop a method of identification which took this variability into account. Consequently, a
quantitative method of expressing leaf-shape differences was developed following on the work of
Nokes (1979).
Three characters are measured as follows:
1. Leaf tooth factor (LTF). A ruler is placed between the tips of the teeth at the ends of the third
and fourth lateral nerves. The number of teeth projecting beyond this line is subtracted from the
total number of teeth between the nerves (excluding the teeth at the nerve endings). This character
was modified from the leaf tooth factor of Nokes (1979). In Fig. 1, LTF has a value of 3.
2. The distance, in millimetres, from the petiole to the first tooth on the leaf base (DFT), (Fig. 1).
3. Leaf tip width (LTW). The shortest width, in millimetres, of the leaf apex one quarter of the
distance between the apex and the leaf base (Fig. 1).
To identify a tree, these three characters are measured on five short-shoot leaves from the lower
FiGurE 1. Measurement of leaf characters.
76 SHORT NOTES
25
20
Number 15
of
Trees. Vip
Discriminant score
FiGureE 2. Frequency distribution of discriminant scores in the test sample of 104 trees. Plants with discriminant
scores less than zero are B. pubescens (56 chromosomes) and those with scores greater than zero are B. pendula
(28 chromosomes) with the exception of those plants marked with a (MM) which are misclassified.
crown and averaged. The average for each character is multiplied by a-constant and
combined in the following discriminant function:
(12 x Average LTF) + (2 X Average DFT) — (2 X Average LTW) — 23
If the solution is greater than zero, the tree is likely to be B. pendula; if it is less than zero, the
tree is likely to be B. pubescens. This method gives a correct rate of classification of 93% (tested
against chromosome number). This was based on a sample of 104 trees which were collected from
14 self-sown populations in England and Scotland. Most of these populations contained both
species. Trees which have low values for the solution (between —5 and 5) are less likely to be
correctly identified than those with higher numerical values. Of the sample tested, 15% had scores
in this range, although only seven trees (7% of the sample) were incorrectly identified. In this
sample, only B. pubescens trees were misclassified. Fig. 2 shows the distribution of discriminant
scores for the test sample.
Another method has been developed which allows a higher degree of certainty of identification
(97%). This requires considerably more calculation and will be dealt with in a later paper.
ACKNOWLEDGMENTS
The authors would like to thank Dr D. C. B. Nokes for the loan of material and for chromosome
number data. We are grateful to Dr C. A. Stace for suggesting this note. The financial support of
the Science and Engineering Research Council is gratefully acknowledged.
REFERENCE
Nokes, D. C. B. (1979). Biosystematic studies of Betula pendula Roth and B. pubescens Ehrh. in Great Britain.
Ph. D. thesis, CNAA, The Polytechnic, Wolverhampton.
M. D. ATKINSON & A. N. CODLING
School of Applied Sciences, The Polytechnic, Wolverhampton, WV1 ILY
MORE ABOUT SEA- AND LAKEBALLS
The publication, earlier this year, of my Presidential Address on ‘Seaballs and Lakeballs’ (Cannon
1985), in which I attempted to bring together information on naturally occurring plant fibre balls
and related phenomena has, as is so often the case, brought to light further information of great
SHORT NOTES 77
interest that I should have liked to incorporate in the original paper. This short note may stimulate
further interest in this unusual aspect of field botany.
Dr Elizabeth McClintock, at the Herbarium, University of California, Berkeley, has kindly
drawn my attention to three further kinds of fibre balls (McClintock 1977). The first kind results
from waste bark fibres from redwood timber extraction and processing on the northern California
coast near Fort Bragg. Although some of the bark is itself used commercially, spillages occur into
the ocean, where the fibres may become matted and twisted together to form balls which are
subsequently cast up on the shore in great numbers. British botanists will probably be less familiar
with the bark of the redwood (Sequoia sempervirens (Lamb.) Endl.) than with that of the related
giant sequoia (Sequoiadendron giganteum (Lindley) Buchholz) from the Californian Sierras, which
grows much better in our climate than the coastal redwood, and is widely planted in parks and
large gardens. However, the general similarity of the very thick fibrous barks will enable readers to
appreciate how seaballs from this markedly different source material can occur.
Dr McClintock also describes lakeballs formed from Ruppia maritima L. in Little Borax Lake,
Lake County, California. Perhaps British botanists should bear in mind the possibility of finding
fibre balls from this species, although the habitats of Ruppia known to me do not appear to be very
conducive to ball formation. The Ruppia balls shown in the photograph in her paper look
extraordinarily like Posidonia balls, but can, apparently, vary in size “from a baseball to a small
watermelon’’. She also refers to a report by H. D. Thoreau as long ago as 1854 of lakeballs in
Massachusetts. These were later stated by Ganong (1905) to be composed of fibres from
Eriocaulon sp. He also refers to a lake in Idaho where species of Ceratophyllum, Chara, Nitella,
Zannichellia and Najas are said to have been implicated in ball formation. Not many English or
Welsh botanists have the opportunity to see Eriocaulon, but perhaps Scottish and Irish members
will note this rather esoteric possibility.
Mr Peter Foss has kindly informed me of the occurrence of the ‘mystery’ Chater, Walters and
Webb seaballs (now known to be composed of Ammophila root fibres) in additional Irish localities
to that on the Dingle Peninsula where they were originally discovered (Cannon 1985). Perhaps
they may yet be found in suitable British localities, where marram occurs abundantly in dunes
above an extensive sandy beach which slopes gently into the sea in sheltered conditions? Mr Foss
hopes to publish a short paper on the new Irish localities in due course.
On a recent holiday visit to the south of France, my wife and I visited the island of Port-Cros
which, as one of the Isles d’Hyéres, has been protected as the Parc National de Port-Cros. In
addition to the land vegetation, the regulations also provide for an extensive marine reserve
stretching out for a considerable distance from the shoreline. The reserve includes, in one of the
more sheltered areas, a substantial bed of Posidonia and, to our intense amazement, in an
otherwise unprepossessing beach cafe, there was offered for sale an excellent publication about the
biology of Posidonia and its environment (Boudouresque & Meinesz 1982). It provides a complete
account of the biology of the species and its ecology, rightly emphasizing the importance of the
extensive Posidonia meadows as habitats for numerous marine animals, so contributing to the
productivity of the Mediterranean and consequently of great relevance to long-term human
economic interests. The need for conservation of Posidonia beds is rightly emphasized, as they are
threatened today in many places, through pollution and the development of marinas and other
coastal facilities.
REFERENCES
BoupDourESQugE, C. F. & MEINESZ, A. (1982). Decouverte de l’herbier de posidonie. Cahier no 4. Parc National
de Port-Cros, Hyeres.
Cannon, J. F. M. (1985). Seaballs and lakeballs — an old Mediterranean theme with a new Irish variation.
Watsonia, 15: 177-181.
Ganon, W. F. (1905). On balls of vegetable matter from sandy shores. Rhodora, 7: 41-47.
McCiintock, E. (1977). Sea balls and lake balls. Fremontia, 5: 23-24.
J. F. M. CANNON
Botany Department, British Museum (Natural History), London, SW7 5BD
78 SHORT NOTES |
DACTYLORHIZA MAJALIS (REICHB.) P. F. HUNT & SUMMERHAYES SUBSP.
CAMBRENSIS (R. H. ROBERTS) R. H. ROBERTS IN S.E. YORKSHIRE
On 29th June 1963, a population of marsh orchids with narrow leaves, some heavily marked, was
found by the author on a marsh by the River Hull, near Wansford, 5 km S.E. of Driffield, GR 54/
0.5, S.E. Yorks., v.c. 61. Two, representative, fresh specimens were sent to Kew and the late V. S.
Summerhayes suggested that the plants might be Dactylorhiza traunsteineri (Saut.) Sod, but that
he could not be certain without seeing the population and this he was never able to do.
In fact, the plants had features which were not correct for D. traunsteineri, notably the flowering
spikes were not lax and the lowest leaf was not the narrowest. In November 1981, R. H. Roberts
identified the taxon from colour transparencies, of both whole plants and close-ups of
inflorescences, depicting two typical members of the population. Roberts (pers. comm.) stated
that the plants undoubtedly belong to D. majalis (Reichb.) P. F. Hunt & Summerhayes subsp.
cambrensis (R. H. Roberts) R. H. Roberts. In 1961, this subspecies was known only from coastal
marshes on Anglesey, v.c. 52, and at Ynyslas in Cards., v.c. 46, but was found a year later in
Caerns., v.c. 49, and in 1979 at two localities in Merioneth, v.c. 48.
Associated species at the S.E. Yorks. site were Equisetum palustre, Caltha palustris, Ranunculus
flammula, Cardamine pratensis, Lychnis flos-cuculi, Parnassia palustris, Mentha aquatica,
Valeriana dioica, Succisa pratensis, Dactylorhiza fuchsii, D. incarnata subsp. incarnata, D.
incarnata subsp. pulchella, Eleocharis palustris, E. uniglumis, Carex hostiana, C. viridula subsp.
brachyrhyncha, C. panicea, C. flacca, C. diandra, C. disticha and C. dioica. &
A biometric study of the S.E. Yorks. population was carried out in 1964 and again in 1983
following the procedure adopted by Roberts (1961a). Table 1 gives sample data for the 1964 and
1983 S.E. Yorks. samples and those given by Roberts (1961b).
My data for the S.E. Yorks. D. mayjalis subsp. cambrensis population studied in 1964, including
mean measurements for various structures (Table 1), match well those for the two populations first
studied by Roberts (1961b) in Wales. Such minor differences as occur between the Yorkshire and
Welsh populations may be the result of long isolation.
In 1979, Ettlinger & Roberts (1980) found populations of D. majalis subsp. cambrensis in two
localities in Merioneth. The Merioneth populations, studied by Bateman & Denholm (1983) and
on which their description of the taxon was based, differ from the S.E. Yorks. plants and those
studied by Roberts (1961b) in a number of respects. Flower colour, shape of labella and labella
markings, as depicted in photographs exhibited by Bateman & Denholm (1982) at the B.S.B.I.
Exhibition Meeting in 1981, are markedly different from those of S.E. Yorks. plants, and R. H.
Roberts (in litt.) confirmed that on the whole the flowers of the Merioneth plants are deep
TABLE 1. COMPARISON OF THE S.E. YORKS. DACTYLORHIZA MAJALIS SUBSP. CAMBRENSIS
POPULATION WITH THOSE IN ANGLESEY AND CARDS.
Figures quoted are means with standard errors; N = sample size.
S,E; Yorks. Sib. MOrks. Cards. Anglesey
(1964) (1983)
Character N = 30 N = 10 N = 85? Nj 920
Height, mm 259 Fl 344+ 193 280453 ZISEST
Total no. of leaves 6-2+0-2 6-8+0-3 6:6+0-1 6-8+0-1
No. of non- 1-5+0-2 1-6+0-2 1-9+0-1 2:0+0-1
sheathing leaves
Leaf length, mm 129-0+4-7 128-0+7:2 124:0+2-4 123-0+2:°6
Leaf width, mm 16-:0+0-8 19-0+0-9 17-040-4 18-0+0-5
Leaf index (length/width) 8-3+0-3 6:8+0-3 7°3£0-2 6-8+0-2
Labellum length, mm 7:-9+0-2 7:5£0:3 8-4+0-1 7-940:1
Labellum width, mm 9-5+0:2 9-5+0-4 10-4+0-1 10-6+0-1
Spur length, mm 7:3+0:2 7-0+£0-3 7:9+0>1 8:-7+0-1
Spur width, mm 2:3+0:1 2°1£0°1 3-3+0:1 3:2+0-1
a) N = 86 for labellum and spur dimensions.
b) N = 118 for labellum and spur dimensions.
SHORT NOTES 79
magenta and have a diamond-shaped labellum with the labellum pattern concentrated in the
central area. The typical flower colour of the S.E. Yorks. plants is identical with that of the original
populations studied by Roberts. The leaf index of the Merioneth population sample is 5-3
(Bateman & Denholm 1983), within the range of that for D. purpurella, i.e. 3-1—-6-1 (Roberts
1961a).
Conditions at the S.E. Yorks. site changed later in the 1960s, believed to be the result of nearby
road construction possibly affecting the flow of spring water. Menyanthes trifoliata, previously
confined to a corner of the marsh remote from the D. majalis subsp. cambrensis population, spread
over the whole area; all Dactylorhiza populations were greatly reduced in numbers of plants, and
flowering spikes of the D. majalis subsp. cambrensis colony became scarce and appeared
spasmodically.
On 20th June 1982, only five plants of this taxon, only one having leaf markings, were found;
these compared well with the 1964 population sample except for average longest leaf length. The
data were as follows: average height 262 mm, average leaf number 6-2, average number of non-
sheathing leaves 2:2, average longest leaf length 105 mm, maximum width of longest leaf 14 mm,
average number of flowers in the spike 31-8.
In early July, 1983, ten plants were noted, eight with typical leaf markings and all the bracts and
upper stem suffused with pigment. The sample data for this population are given in Table 1.
Examination of the data, including notes on vegetative, inflorescence and floral features, revealed
that this population resembled that studied in 1964 in most respects. The most marked differences
were in the average height of the plants, the length of flowering spike (average 54 mm), and a
tendency for a slightly wider leaf, although the leaf index of 6-8 is the same as that for the Anglesey
colony.
Four of the plants examined in 1983 were in tall vegetation which may have affected height, but
this cannot fully account for the greater average height. A possible explanation of these observed
changes in the tetraploid D. majalis subsp. cambrensis population as revealed by the 1983 studies is
that there has been past hybridization, most probably with the diploid D.fuchsii, and subsequent
backcrossing from the F; to D. majalis subsp. cambrensis so that the population has become
slightly introgressed. This theory is in line with the findings of Lord & Richards (1977) who
demonstrated that triploid dactylorchid hybrids are by no means totally sterile, but may frequently
cross among themselves, or backcross to the parents. A photograph of a plant, taken on the S.E.
Yorks. site on 29th June 1963, has been examined by R. H. Roberts and tentatively identified as
the hybrid D. fuchsii x D. majalis subsp. cambrensis, being similar to plants of this hybrid found in
Wales. Also, a plant on the same site with intermediate characters and exceptional vigour, noted in
1982, was thought to be this hybrid and excluded from the population sample. The changed site
conditions may have favoured the survival of hybrid derivatives.
ACKNOWLEDGMENT
I wish to thank Mr. R. H. Roberts for determining Dactylorhiza majalis subsp. cambrensis from
photographs, for much valuable comment on my observations and for kindly reading the
manuscript and making suggestions.
REFERENCES
BATEMAN, R. M. & DENHOLM, I. (1982). The tetraploid marsh-orchids of the British Isles — a study in critical
group taxonomy. Watsonia, 14: 226.
BATEMAN, R. M. & DENHOLM, I. (1983). A reappraisal of the British and Irish dactylorchids, 1. The tetraploid
marsh-orchids. Watsonia, 14: 347-376.
ETTLinGER, D. T. & Roserts, R. H. (1980). Plant record. Watsonia, 13: 146.
Lorp, R. M. & Ricnarps, A. J. (1977). A hybrid swarm between the diploid Dactylorhiza fuchsii (Druce) So6
and the tetraploid D. purpurella (T. & T. A. Steph.) So6 in Durham. Watsonia, 11: 205-210.
Roserts, R. H. (1961a). Studies on Welsh Orchids, 1. The variation of Dactylorchis purpurella (T. & T. A.
Steph.) Vermeul. in North Wales. Watsonia, 5: 23-36.
80 SHORT NOTES
Roserts, R. H. (1961b). Studies on Welsh Orchids, 11. The occurrence of Dactylorchis majalis (Reichb.)
Vermeul. in Wales. Watsonia, 5: 37-42.
F. E. CRACKLES
143 Holmgarth Drive, Bellfield Ave., Hull, HU8 9DX
THE REDISCOVERY OF CAREX MARITIMA GUNN. ON THE FAIRWAYS AT
ST ANDREWS LINKS, FIFE
Carex maritima Gunn. is a rare sedge of sandy coasts in north-eastern Britain. During the present
century it has been lost from many of its former strongholds, especially in eastern Scotland where,
from E. Ross southwards, David (1982) listed the plant as still extant at only two of the 28 sites for
which he had old records.
In Fife, v.c. 85, C. maritima used to occur at three localities: at aenbannié Links, 37/4.0, where
it was recorded about a century ago (Howie 1884); at Tentsmuir, 37/4.2, where it was last seen in
the late 1960s (M. Benstead pers. comm.) but is now almost certainly extinct; and at St Andrews
Links, 37/5.1, where until summer 1984 it had not been recorded since 1911 (specimen in STA).C.
maritima was first found on St Andrews Links by Maughan (specimen in E), and was recorded by
Howie (1884) as occurring “‘on the old road that crossed the Swilkin Burn at St Andrews’.
In 1984, the sedge was first discovered on 20th June in a damp hollow in the middle of the third
fairway of the Jubilee Golf Course. This seemed a most unlikely habitat, yet since then I have
crawled across all the St Andrews fairways and — to my utter astonishment — have now found it at
14 sites on the Jubilee Course, 14 on the New Course, six on the Old Course (including three on
the eighteenth fairway!), and one on the putting green by the Swilkin Burn.
All the sites are on low-lying fairways that flood intermittently in winter (usually November —
April), and despite much searching I failed to find the sedge on any of the uncut roughs. Some of
the colonies are very large, with three sites on the Jubilee Course and five on the New Course each
holding more than 1,000 plants. Three sites probably have over 100,000 plants each, and it occurs
in these at extraordinarily high densities, with sample quadrats producing counts of 500—1,000
plants/m‘.
Apart from Carex maritima, the fairway ‘slacks’ are floristically of little interest, although
Isolepis setacea occurs sparingly on the Jubilee Course and Blysmus rufus is co-dominant with C.
maritima at two sites on the New Course. The sedge is generally found in damp grassland in which
Agrostis stolonifera, Carex flacca, Poa pratensis and Festuca rubra are abundant, together with a
few other ‘wet slack’ species such as Juncus bufonius, Sagina procumbens, Glyceria declinata,
Carex ovalis and Leontodon autumnalis.
Perhaps the most surprising aspect of this plant’s rediscovery has been to find it flourishing on
some of the most intensively managed parts of the golf links. Management of the fairways includes
regular cutting to keep the sward to a height of 2—3 cm, aeration of the turf by ‘slitting’ it in early
winter, applications of organic fertilizers in spring, and (in some years) spraying with the broad-
leaved weedkiller 2,4-D. This management may in fact favour C. maritima, as it keeps the turf
fairly open and helps to restrain taller-growing species that might otherwise eliminate it from the
sward. Indeed, on the fairways this sedge is probably at a competitive advantage — it can flower
and set seed below the height of the cutting blades, is unaffected by 2,4-D, and may be encouraged
to spread vegetatively by having its rhizomes repeatedly cut by the tractor-mounted ‘slitter’ used to
aerate the turf. It also seems able to colonize divot holes and turf cuttings quickly, and can
establish itself in reseeded areas before the grass sward becomes closed. It is surprising to note that
some of the largest colonies (those on the first fairway of the New Course) are in turf that was layed
as recently as 1974. The possibility cannot be ruled out that C. maritima was already in the turf at
the time it was layed; however, I could find no evidence of the sedge in the area from which the turf
was taken, and the habitat there does not appear at all suitable.
It is remarkable that C. maritima has remained undetected at St Andrews for so long. However,
it must be admitted that this sedge is easily overlooked, and at St Andrews it is found in places that
most field botanists would tend to leave well alone. Certainly its continued occurrence at this site is
of considerable interest, not only because of its unusual habitat, but also because it appears to be
flourishing here in greater quantity than at any of its other surviving British stations.
SHORT NOTES 81
It would be most interesting to know whether C. maritima is still to be found on other Scottish
sand dune systems where the wet slacks are now being managed as golfing fairways.
ACKNOWLEDGMENTS
I wish to thank the staff of the St Andrews Links Trust, and in particular W. Woods, for permission
to visit the site and for information about how the fairways are managed. I am grateful to G. H.
Ballantyne for details of old records, and P. K. Kinnear and H. J. Noltie for assistance with the
field survey. I also thank Mrs D. Longley for her help in preparing this note.
REFERENCES
Davip, R. W. (1982). The distribution of Carex maritima Gunn. in Britain. Watsonia, 14: 178-180.
Howie, C. (1884). Flora of Fife. Unpubl. MS., St Andrews.
S. J. LEAcH
Nature Conservancy Council, Rathcluan House, Carslogie Road, Cupar, Fife, KY1I5 4JA
FILAGO PYRAMIDATA L. REFOUND IN WEST SUSSEX
Filago pyramidara, last recorded in W. Sussex, v.c. 13, in 1905, and known from only 14 localities
in England since 1960 (Perring & Farrell 1983), was refound by H. W. M. in Halnaker Chalk-pit in
1983 and 1984.
In August 1983, one dry, shrivelled plant was found and at that time misidentified as F. vulgaris
Lam., and in September of that year a further 15 seedlings were discovered. During July and
August 1984, some 250 plants were found; E. C. Wallace and R. C. Stern suggested that these
could be F. pyramidata L., a suggestion later confirmed by F. Rose and C. Jeffrey (B.S.B.I.
Compositae Referee). By early September many hundreds of plants (possibly up to 1000) had been
discovered in the chalk-pit. Of these, six were erect and approximately 17-5 cm tall, several were
branched and semi-prostrate, and the remaining many hundreds, growing on bare, compacted
chalk with the uncommon moss Seligeria calcarea, were only 1-25 cm or less, high. In the
surrounding patches of very short, rabbit-grazed turf were several species of flowering plant
including Euphrasia nemoralis, Gentianella amarella, Centaurium erythraea and Agrostis stolo-
nifera, all very dwarfed and only 2-5 cm high or less, and also the moss Homalothecium lutescens.
In BM there is a specimen of F. pyramidata labelled ‘“‘Halnaker Hill, 8 August 1891” that was
collected by H. L. F. Guermonprez. This led us to the Guermonprez collection in the Portsmouth
City Museums (PMH), in which a herbarium sheet is labelled “‘Halnaker Hill, Sussex (in chalk-
pit)” and also “8 August 1891”. H. L. F. Guermonprez specialized in painting the British flora,
and among 2000 paintings in the collection at PMH is one of F. pyramidata painted in 1891 at
Halnaker; the sharply five-angled capitula are clearly seen in the painting, just as they are in the
growing plants at Halnaker — a key character which cannot be observed in pressed specimens.
Guermonprez’s working copy of Flora of Sussex (Arnold 1907) shows, in pencilled marginal notes,
a drawing of the leaf shape (also diagnostic) and refers again to this locality.
Could the warm weather of 1983 possibly have influenced the number of seedlings which
appeared in 1984? They may not be in such abundance every year, but their small size,
inconspicuous habit and the fact that they flower only late in the season, leads us to suggest that the
plants could well have been present but overlooked at this locality since 1891.
REFERENCES
_ Arnon, F. H. (1907). Flora of Sussex, 2nd ed. London.
82 SHORT NOTES
PERRING, F. H. & FARRELL, L. (1983). British red data books: 1. Vascular plants, 2nd ed. London.
H. W. MaTcCHAM & M. BriccGs
21 Temple Bar, Strettingham, Chichester, W. Sussex, PO18 OLB
CHROMOSOME NUMBERS OF IRISH PLANTS, 1.
This paper is the first of a projected series in which I shall provide chromosome counts, using
material of known wild origin, of native Irish species. In practice this means that the series will
cover virtually the entire Irish flora as very few previous counts of Irish material exist. Expediency
dictates that the counts cannot be done following a systematic scheme and therefore several
families will be.covered by each contribution.
Because of the wide range of material to be surveyed it is not possible to provide more than an
outline of the methods used. Generally, for meiotic counts flower buds are fixed, rapidly, in
Carnoy’s solution (6:1:3, absolute ethanol: glacial acetic acid: chloroform) or Dyer’s modification
of it (Dyer 1963). Mitotic counts are made from root material, pre-treated by placement in a cold
room for 24—48 hours or more, or rarely pre-treated with 0-2% or 0-5% colchicine for either two or
four hours. Roots are subsequently fixed, as above, and may then be hydrolyzed in SN HCl, at
room temperature, for up to one hour. Materials are stained using, preferably, the lacto-propionic
orcein method of Dyer (1963) or, more rarely, aceto-carmine. Vouchers are deposited in TCD.
Crambe maritima L. = 30' Co. Clare, v.c. H9: Portcowrugh, Inishmore, Aran
Islands; sea-shore. 1972, D. A. Webb.
Euphorbia portlandica L. n = 20 Co. Wicklow, v.c. H20: Mizen Head; sand-dunes.
1982, P. Wyse Jackson.
Hypericum androsaemum L. n = 20 Co. Roscommon, v.c. H25: shore of Lough Ree;
woodland. 1982, P. Wyse Jackson.
Hypericum canadense L. i W. Galway, v.c. H16: shore of Lough Mask; wet
flush. 1981, P. Wyse Jackson.
Hypericum tetrapterum Fries n=8 W. Galway, v.c. H16: shore of Lough Mask; wet,
peaty flush. 1981, P. Wyse Jackson.
Jasione montana L. n=6 Co. Wexford, v.c. H12: Carnsore Point; stone wall.
var. latifolia Pugsley 1982, J. A. N. Parnell.
Ranunculus bulbosus L. n= 8 W. Galway, v.c. H16: Roundstone; grassland. 1982,
var. dunensis Druce W. Bradley.
Samolus valerandi L. n= 13 Co. Wicklow, v.c. H20: Arklow; damp cliff-base.
1968, D. A. Webb.
Spergularia rupicola Lebel n=c.18° Co. Dublin, v.c. H21: Dun Laoghaire; harbour wall.
ex Le Jolis 1969, D. A. Webb.
Thymus praecox Opiz ahi Co. Clare, v.c. H9: Poulsallagh; dune grassland.
subsp. arcticus (E. Durand) 1979, P. Wyse Jackson.
Jalas
Notes:
' The 2n = 30 cytodeme has not yet been detected in Ireland.
° Confirms the only previous count of 2n = 16 made on material from the same region by D. M.
Moore (Webb & Halliday 1973).
*n = 19 was also recorded, but only from two cells; the extra chromosome may be an accessory.
* The only cytotype so far recorded in Ireland. ,
ACKNOWLEDGMENTS
I should like to thank Miss Y. Bannon for technical assistance and the staff of the Trinity College
Botanic Gardens for keeping specimens of the species in cultivation.
— —EE
SHORT NOTES 83
REFERENCES
Dyer, A. F. (1963). The use of lacto-propionic orcein in rapid squash methods for chromosome preparations.
Stain Tech., 38: 85—90.
Wess, D. A. & Hatuipay, G. (1973). The distribution, habitat and status of Hypericum canadense L. in
Ireland. Watsonia, 9: 333-344.
J. A. N. PARNELL
School of Botany, Trinity College, Dublin 2, Ireland
AN ADDITIONAL CRITERION FOR ASSESSING NATIVE STATUS
Eight criteria which can be used in assessing whether a species is a native member of the British
flora have been discussed by Webb (1985). This note outlines a further criterion which has been of
use in investigating this problem: the relationship of the species in question to oligophagous
insects.
In only a few cases (so far) has entomological evidence been invoked in assessing native status.
Rhynchosinapis wrightii is the host of two beetles of very restricted distribution. F. R. Elliston
Wright used this as evidence for its native status before it was described as a species endemic to
Lundy (Wright 1933; Lucas & Synge 1978).
Coombe (1956) showed that the presence of oligophagous insects on /mpatiens noli-tangere can
be used to distinguish sites where the species is probably native (in N. Wales and NW. England)
from sites in southern England where it is a garden escape. J. parviflora, known to be introduced,
is remarkably free from associated insects except where it grows in the vicinity of /. noli-tangere at
sites where the latter is native.
Entomological evidence must be used with the same caution as Webb’s criteria. Introduced
species can attract oligophagous insects from related native species, as in the case of /mpatiens
parviflora. Other aliens are associated with insects that were presumably introduced with them. I
have found that Carpobrotus edulis on the Lizard peninsula, W. Cornwall, frequently bears a scale
insect which Mrs Linda Huddleston kindly identified as Pulvinariella mesembryanthemi (Vallot).
This was originally described from France and only later discovered on native ‘mesems’ in South
Africa. Similarly the weevil Stenopelmus rufinasus Gyll. is associated with the introduced fern
Azolla filiculoides in Britain (Janson 1921; Flint 1979). It may be significant that both Azolla and
Carpobrotus were probably introduced as living plants rather than as seed or spores.
REFERENCES
CoomsE, D. E. (1956). Notes on some British plants seen in Austria. Veroff. geobot. Inst., Zurich, 35: 128-137.
Fut, J. H. (1979). Azolla filiculoides Lam. and its weevil, Stenopelmus rufinasus Gyll., in Yorkshire.
Naturalist, Hull, 104: 112.
Janson, O. E. (1921). Stenopelmus rufinasus Gyll., an addition to the list of British Coleoptera. Entomologist’s
mon. Mag., 57: 225-6.
Lucas, G. & SynGE, H. (1978). The IUCN Plant Red Data Book. Kew.
Wess, D. A. (1985). What are the criteria for presuming native status? Watsonia, 15: 231-236.
Wricut, F. R. E. (1933). Contribution to the flora of Lundy Island. J. Bot., Lond., 71, suppl.
C.7D. PRESTON
Institute of Terrestrial Ecology, Monks Wood Experimental Station, Abbots Ripton, Huntingdon,
PEI 25S:
THE DISTRIBUTION OF SORBUS LANCASTRIENSIS E. F. WARBURG
Warburg (1957) published the description of a new species of whitebeam, Sorbus lancastriensis,
from Humphrey Head, Westmorland, v.c. 69 (now Furness, v.c. 69b) and gave its distribution as
“apparently restricted to several places on Carboniferous Limestone round Morecambe Bay in
84 SHORT NOTES
Lancashire and Westmorland’’. Between 1982 and 1984 much of the limestone in this area was
visited by the authors, and this note details the jocations and approximate sizes of all the
populations seen of this rare, endemic taxon.
Sorbus lancastriensis is often confused with two other species of Sorbus which also occur in
north-western England, and with which it frequently grows: S. aria (L.) Crantz and S. rupicola
(Syme) Hedlund. There are three reasons for this confusion. Firstly, it is essential to examine the
correct leaves. Ideally, the three broadest leaves of a rosette from the short, lateral spurs of
branches should be compared, as these are the most constant and typical for each species. Shade
leaves, or leaves from the present year’s growth, should be avoided as these are highly variable.
Secondly, S. aria is very variable in leaf shape and size, and two specimens growing together may
be superficially quite different, prompting identification of two taxa. Thirdly, the characters used in
the key by Warburg (1952) to distinguish S. lancastriensis from S. rupicola are poor. The fruit and
leaf toothing characters are useless, and there is too much overlap in the leaf venation character to
allow diagnosis On its own. The more recent key by Game (1981) is much better and includes all the
species of Sorbus in v.cc. 66—70. The species are distinguished here according to the following key,
which is based on material of all three species from north-western England. A combination of
characters must be used. Sorbus lancastriensis is very close to S. rupicola and the relationship
between the taxa requires clarification.
jn Leaves ovate, rhombic to elliptic, rarely obovate, the base truncate to
broadly cuneate (angle with central vein 45° or more), rarely cordate,
toothed nearly to base, with (10-5)11 or more pairs of veins. Fruit
length greater than or equal to width .......... £6. SESE AP S. aria
1. Leaves obovate, oblanceolate or oblong, cuneate at base (angle with
central vein less than or equal to 42(47)°), + entire in lowest quarter,
with 10(11) or fewer pairs of veins. Fruit width greater than or equal to
lengths. 220) 0 RUA LE Ee, Meee eaten Z
jz Leaves erect, (1-7) 1-8—2-5 times as long as wide, with (5)6—9(10) pairs of
WEIS DT). VN AT dee SE ER EEO eee Re ae S. rupicola
a Leaves spreading, (1-3)1-4—1-75(1-9) times as long as wide, with (6-5)8—
LOGL) pairs:Of VEINS: FCSN A ee Seen S. lancastriensis
Sorbus lancastriensis is very characteristic of the open limestone screes, rocks, crags, scars and
cliffs within 30 km of Morecambe Bay, usually in small populations. It reaches the northern and
southern limits of the local outcropping limestone at Cunswick Scar and Warton Crag respectively.
The only locality not on limestone is at Roughholme Point where it occurs on a conglomerate
drumlin. It grows from sea-level at Arnside to about 180 m at Cunswick Scar. It prefers open scrub
and may persist in developing woodland but does not tolerate shade. It flowers and fruits freely in
most localities and regenerates readily in the absence of grazing. The localities are listed below; full
lists detailing 6-figure grid references are held by the Biological Records Centre, the vice-county
recorders, the Nature Conservancy Council (N.W. Region) and the local Naturalists’ Trusts. The
names of localities used are those from the Ordnance Survey 1:10,000 series or 1:25,000
‘Pathfinder’ series.
West Lancashire, v.c. 60 (all 34/4.7):
Eaves Wood. Two sites: Castlebarrow (above Elmslack), around the pepperpot and in the woods
below, c. 20 shrubs, the colony extending into Middlebarrow Plain (v.c. 69); and National Trust
Nature Trail, c. 15 shrubs mixed with S. aria, extending down to Waterslack.
Jenny Brown’s Point. A largely inaccessible population on Jack Scout sea-cliffs, mixed with S.
rupicola, about 30 plants in total.
Scout Wood. Five or six shrubs on the scarp edge in woodland.
Silverdale Cove. Sea-cliffs to north and south of the Cove, c. 50 plants.
Warton Crag. Four sites: Beacon Breast, 10-15 shrubs; above Scar Close, five shrubs; Three
Brothers Allotment, one shrub on limestone pavement with three S. aria plants; south of Crag
Foot, nine plants on rocky outcrop above the road, with S. rupicola.
Westmorland, v.c. 69:
eine
SHORT NOTES 85
Arnside (34/4.7). Generally distributed around Arnside, Arnside Knott and Far Arnside, usually
in small populations and frequently with S. aria. There are some populations along the Kent
Estuary (Arnside—Grubbins Wood) which have some puzzling plants intermediate with S.
rupicola, and it is not always possible to name an individual specimen. Arnside Knott, at least
five shrubs; Copridding Heath, at least three shrubs; Arnside Park, a few trees scattered in
woodland; Far Arnside, one plant in hedge by road, and then round the coast on cliffs to
Arnside, c. 200 plants, with occasional S. rupicola.
Meathop (34/4.7). In the quarry and on sea-cliffs at the mouth of the River Winster, about ten
shrubs mixed with S. aria.
Middlebarrow Wood (34/4.7). Limestone Pavement at the top of the wood.
Brigsteer (34/4.8). Three places: Burnbarrow Scar, many shrubs along the cliff edge; Crag Millot,
about 15 shrubs in pioneer scrub; Windy Howe, one plant on roadside.
Scout Scar (34/4.8). Abundant (probably in excess of 200 plants) along the edge of the cliff with
scattered shrubs in the woods and on the scree below.
Whitbarrow Scar (34/4.8). Locally abundant on cliffs from Raven’s Lodge clockwise around the
scar to Broad Oak, with occasional plants in the wood below. Probably in excess of 1000 plants,
the largest population.
Yewbarrow (34/4.8) (A. Game). Abundant in open woodland with smaller shrubs on cliff and
screes, probably 100—200 plants.
Cunswick Scar (34/4.9). Abundant along the open middle section of the scar, with one plant of S.
aria.
Furness, v.c. 69b:
Humphrey Head (34/3.7). Abundant along the sea-cliffs and scar on the west side, over 200 shrubs
mixed with a much smaller number of S. rupicola plants. One shrub also found at Roughholme
Point.
Old Park Wood (34/3.7). On cliffs on west side, c.25 plants.
Reake Hill, Leven Estuary (34/3.7). 15—20 small trees on western side of hill.
Roudsea Wood N.N.R. (34/3.8). 1 large shrub in old quarry (N.C.C. and others).
There are also records in Ratcliffe (1977) which need verification or correction. The record for
Gait Barrows N.N.R. (v.c. 60) is probably an error for S. aria. The record for Hutton Roof Crags
and Farleton Knott (v.c. 69) has not been refound. The record of S$. rupicola for Scout and
Cunswick Scars, where the species has been searched for carefully without success, is probably an
error for S. lancastriensis. There is also a National Trust record for Brigsteer Park (v.c. 69) which
has not been refound.
The authors would be pleased to hear of any further localities. Other sites in which the plant
should be looked for are about Grange, and the limestone on the west side of the Leven Estuary.
ACKNOWLEDGMENTS
The authors wish to thank Mr D. Earl and Mr & Mrs L. Livermore for their excellent company in
the field and with whom many of the localities were found or verified.
REFERENCES
Gam), A. (1981). Sorbus, in WiccincTon, M. J. & GraHaM, G. G., eds. Guide to the identification of some of
the more difficult vascular plant species. N.C.C. England Field Unit Occasional Paper No. 1, pp. 57-58.
Banbury.
— Rartcurre, D., ed. (1977). A nature conservation review, vol. 2. Cambridge.
_ Warsurs, E. F. (1952). Sorbus, in CLAPHAM, A. R., TuTInN, T. G. & WarRBuRG, E. F. Flora of the British Isles,
pp. 423-437. Cambridge.
_ Warsurc, E. F. (1957). Some new names in the British flora. Watsonia, 4: 45—46.
T. C. G. Rico & M. BAECKER
Botany Department, The University, Leicester, LE] 7RH
86 SHORT NOTES
FURTHER DISCOVERIES OF HIERACIUM BACKHOUSEI F. J. HANB.
Hieracium backhousei F. J. Hanb., which is included in section Alpina, is one of the rarest British
hawkweeds, and is endemic to Scotland. It was discovered by James Backhouse Jnr in 1886 at
Dubh Loch, near Braemar, S. Aberdeen, v.c. 92, where a small population still occurs. No definite
further localities had been reported until I found a small number of plants in 1976 on Derry
Cairngorm, in Glen Derry, S. Aberdeen, which were subsequently confirmed as this species, and
later in greater numbers locally in two places in Glen South Esk, Angus, v.c. 90, in 1981. Solitary
plants, which also should possibly be referred to this species, were noted by me in Coire Etchachan
and near Loch Etchachan in S. Aberdeen, and at the head of Glen Fee, Angus.
All the old records for H. backhousei, except for the Dubh Loch records, but including the one
from Glen Derry (Linton 1905; Pugsley 1948), appear to refer to H. memorabile (Sell & West
1967, ‘19G8)s- 465
At Dubh Loch, the population occurs in crevices and in pockets of coarse-grained granite by
small waterfalls, very close to the running water. This habitat is matched exactly in both Glen
Derry and in Glen South Esk, where the populations occur on slab rock or large boulders of
granite, on islands or projections into the stream, again in the vicinity of small waterfalls. In Glen
South Esk and at Dubh Loch the species is accompanied by another hawkweed from section
Alpina, H. calenduliflorum Backh. Additionally, H. hanburyi Pugsl. and occasionally H. atraticeps
(Pugsl.) P. D. Sell & C. West and H. pseudocurvatum auct. occur nearby. In the localities
described, all of these hawkweeds generally occur somewhat further away from the running water
than H. backhousei. It has also been noted that some of the pockets of rock where H. backhousei
grows remain temporarily filled with water after heavy rain; and it would seem that the species
requires more moisture around the roots than other species in section Alpina, as it does not stray
onto adjacent ground further away from water, although the surface rock is normally dry as is the
case with all species in section Alpina. In cultivation, however, it is surprisingly one of the easiest
species to grow and it withstands summer drought conditions as well as other species in section
Alpina.
H. backhouset is distinguished by its large capitulum, often single, from 30 mm to over 50 mm in
diameter, or occasionally with two or more heads. The basal leaves are very rigid, coriaceous,
drab, non-glossy green, being mainly elliptical and irregularly serrate-dentate with narrow-acute
teeth. The lowest cauline leaf is often very similar to the innermost basal leaf, with a few narrow
teeth which are frequently spinulose, and has some stellate hairs on the lower surface. The
majority of leaf surfaces, except at the margins, have very few or no simple, eglandular hairs
present. The involucre is often robust with broadish bracts (1-0—1-6 mm) which are abruptly acute
to sub-acute, with only few obvious, very short, glandular hairs. The ligules are a bright, lightish
yellow, with several very short simple eglandular hairs at the apices, and very few to several on the
backs, and have yellowish styles.
The number of plants in Glen South Esk greatly exceeds the number at Dubh Loch; the former
site, in v.c. 90, is therefore now the main station known for the species. It is surprising that H.
backhousei has not been noted in Angus previously, although it might have been confused with H.
tenuifrons P. D. Sell & C. West, and could occur in the described habitat elsewhere in the eastern
Highlands. Specimens have been placed in CGE and in herb. D.J.T.
ACKNOWLEDGMENT
I should like to thank Mr P. D. Sell for advice on preparation of this manuscript and confirmation
of the identities of specimens.
REFERENCES
LinTON, W. R. (1905). An account of the British Hieracia, pp. 16-17. London.
PuGs.ey, H. W. (1948). A prodromus of the British Hieracia. J. Linn. Soc. (Bot.), 54: 48-49.
|
|
|
SHORT NOTES 87
SELL, P. D. & West, C. (1967). Taxonomic and nomenclatural notes on the British flora. Watsonia, 6: 304—
305.
SELL, P. D. & West, C. (1968). Hieracium, in PERRING, F. H., ed. Critical supplement to the atlas of the British
flora, p. 78. London.
D. J. TENNANT
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Watsonia, 16, 89-103 (1986) 89
Book Reviews
The pollen loads of the Honeybee. A guide to their identification by colour and form. Dorothy
Hodges. Pp. 51 & 14, with 10 colour plates, 30 black & white plates and 7 text figures.
International Bee Research Association, London. 1984. Price £26.50 (ISBN 0-—86098-140-1).
This is the second facsimile edition of a remarkable book that was first published in 1952 and was
designed to provide a practical manual for the «dentification of pollen loads gathered by honey
bees. As the authoress explains, the colour of a pollen load is indicative of the kind of piant from
which the pollen originated. The strength of the book lies in the colour charts showing the colour of
pollen loads collected from 120 different plant species. For most species three different pollen
loads are illustrated, so that the natural colour variations, greater in some species than others, are
indicated. The colour charts are arranged in six full-page plates in the order of flowering, to assist
in the identification of pollen loads during the different seasons of the year. A further five plates of
colour paintings show the processes by which bees collect and pack pollen loads and the colours of
artificially produced pollen loads. There are also 30 pages of line drawings depicting the pollen
grains of the most important sources of food for honeybees as seen in the light microscope. These
drawings are reminiscent of those in the classic textbook on pollen by Wodehouse (1935). This is
largely because they depict intact pollen grains which have not been subjected to the treatment
with strong acids prevalent in palynological laboratories. As such they may serve as a practical
means of identifying some of the more distinctive types of pollen with a relatively modest
microscope.
The book has been revised to include a chapter, previously published elsewhere, on the methods
of melissopalynology. This is undoubtedly a useful addition for those wishing to pursue the subject
in a practical way but would have been better placed after the plates, since the stark style contrasts
dramatically with the much more readable text of the book itself. A more useful revision would
have been to have updated the literature references given. The classic textbooks concerned with
the identification of pollen are mentioned, but most have passed through several editions more
recent than those listed. Perhaps, more importantly, several new books could then have been
included which provide keys and illustrations for identification. Recent works by Moore & Webb
(1978) and Sawyer (1981), for example, contain keys and micrographs and are both modestly
priced.
It may be hoped that such information will be included in a future edition, since there will be a
continuing, and perhaps even a growing, demand for this unique book from bee keepers,
pollination biologists and palynologists.
REFERENCES
Moorg, P. D. & Wess, J. A. (1978). An illustrated guide to pollen analysis. London.
Sawy_eR, R. (1981). Pollen identification for beekeepers. Cardiff.
WobEHOUSE, R. P. (1935). Pollen grains [etc]. New York & London.
S. BLACKMORE
The Longman illustrated dictionary of botany. A. Sugden. Pp. 192, with 300 coloured diagrams.
| Longman & York Press, Harlow, Essex & Beirut. 1984. Price £3.95 (ISBN 0—582-—55696-1).
This reference book intended for amateur botanists and students of botany stands up to the
_ publishers’ claim that it defines clearly and simply both terminology and concepts used in the study
90 BOOK REVIEWS
of plant life and plant sciences. More than 1200 definitions are written in non-technical language
and illustrated by 300 diagrams, most of which are in good colour with a few in black & white. It
does not have the alphabetical arrangement customarily associated with dictionaries; but once the
system is grasped in which the words are grouped in related topics as used in study courses, a
comprehensive (alphabetical) index then leads to the section required.
There is a wide coverage of subjects relating to plants, some 23 headings including: flower
biology and anatomy, development from seed or spore to mature plant, evolution and spread of a
species and adaptation to habitat, habitat definitions, basic chemistry of plants, and a review of the
plant kingdom. Appendix one is ‘Understanding botanical words’ and Appendix two is ‘SI Units’.
The suggestion for the B.S.B.I. to review this book came from an amateur member, who
recommended it; a Museum assistant scientific officer, taking Botany ‘O’ Level, evaluated the
Dictionary as being useful for his studies; a senior taxonomist, checking for accuracy, reported this
good in general; but found that the definitions of cyme and raceme do not distinguish between the
position of the oldest flower, which makes the inflorescence determinate (cyme) or indeterminate
(raceme); also the term strophiole is not included (although the page on seed structure does
include definitions of hilum, micropyle and raphe). However these test words are possibly severe,
as in general this clear, compact and handy reference book will doubtless be very useful for those
for whom it is mainly intended: amateurs, sixth formers and first year undergraduate students.
M. Briccs
“
Leaves. G. T. Prance & K. Sandved. Pp xii + 244, with 310 colour photographs and 46 black &
white illustrations. Thames & Hudson, London. 1985. Price £22.50 (ISBN 0-—500-—54104-3).
This lavish book is the result of co-operation between representatives. of two famous botanical
institutions in America: the text, accurate, as would be expected, is by G. T. Prance, Senior Vice-
President for Science at the New York Botanic Garden; the spectacular photographs are by Kjell
Sandved, of the National Museum of Natural History, Smithsonian Institution, Washington, D.C.
It is an ‘eye-opener book’ — exploring leaf form and function from a world-wide selection and from
many angles, including shape, arrangement and colour — with much information that will be new to
general readers, and with stunning photographs to illustrate the sheer beauty and diversity of
leaves. There are sections on leaf margins, venation, hairs, prickles and poisons; dramatic sections
on modified leaves, leaves as propagules, on carnivorous and on succulent leaves; also on those
which carry spores, those patterned by leaf-miners and other predators, or those on and in which
insects are sheltered; on leaves useful to man; and finally on fossil leaves. The last chapter gives
brief advice on leaf collecting and making a herbarium, with appendices on cleaning, staining and
leaf prints, and finally philately.
Throughout, the beauty of leaves, of magnified leaf parts and the arrangement of the leaves on a
plant as well as of the whole leaf, is the main theme of the illustrations. The book is expensive; but
if you have a small coffee table waiting for a book through which one can browse and obtain much
visual pleasure and unexpected facets of knowledge on a very functional plant organ that is
widespread in the natural world, this would be the book for you.
M. BriccGs
Guide to standard Floras of the world. D. G. Frodin. Pp. 619. Cambridge University Press,
Cambridge. 1984. Price £95.00 (ISBN 0-—521-—23688-6).
This remarkable book provides a critical guide to the published work that is available for the
identification of flowering plants and ferns throughout the world. The term ‘standard floras’ is used
in the sense coined by V. H. Heywood in connection with the Flora Europaea project, i.e. the best
works currently available for those seeking information about the plants of any country or similar
area. The Guide is authoritative, well arranged and clearly the fruit of a prodigous amount of hard
and, one suspects, often unrewarding labour. It takes the form of a select bibliography with
BOOK REVIEWS 9]
annotations in which the works are assessed, almost invariably as a result of the author’s own
careful examination of the contents. Those who know the author will not be surprised to find that
this is no mere juggernaut of bibliographic drudgery, but rather is spiced with informed comment
and some idiosyncratic touches that enliven the text. In short, in addition to fulfilling its primary
function as a practical guide to prospective Flora users, it is a book that anyone with an interest in
floristics or botanical literature can browse in for many hours with considerable enjoyment.
The data are very firmly structured under nine major divisions covering the major continental
regions, after which follow sections on the various countries and other territorial aggregations.
Chapter headings are accompanied by delightful quotations that reflect the aspirations,
achievements and failures of botanical authors. The treatment is exhaustive and highly detailed.
Thus, under the British Isles, we learn that the latest work covering Scotland as a country is W. J.
Hooker’s Flora Scotica of 1821, and there is even an entry for Rockall (for which no vascular plants
have been recorded). British botanists should note that coverage at the British county Flora level is
not provided, a possibility obviously precluded by the world-wide objectives of the book. The
geographical breakdown is preceded by three extremely interesting chapters that survey the range
of publications that are available, the progress and prospects of Flora writing since 1939, and
general considerations on the style of Floras. All those engaged in Flora writing or contemplating
initiating projects of this kind should study these essays with care. If more thought were given to
what Floras are actually ‘for’ and how perceived needs can best be met in the real world, botanical
libraries would have fewer sad, rusting hulks of major, but over-ambitious works, some of which
expired with a bang of recriminations in a glare of publicity, while others subsided in obscurity with
scarcely a whimper.
In short, this is a major, highly practical work of reference that should be in all libraries with any
pretence towards coverage of botanical taxonomy and related topics. The tone of this review has
been deservedly enthusiastic but, sadly, two rather serious reservations must be made. The first is
that the cut-off date for entries is 1980, while the publication date was late in 1984. Obviously, it is
highly desirable that works of reference should be as up-to-date as possible and, while no doubt
difficulties existed, such as the author’s present location at the University of Papua New Guinea,
four years in press is grossly excessive, even for a publication as complex as the Guide. The book is
beautifully designed and produced and is worthy of the best traditions of the Press, but can it
possibly justify the staggeringly high price of £95.00? Basic working tools of this kind are especially
valuable in the developing areas of the world and in the less well-endowed libraries. One cannot
but fear that the cost will preclude its use in many situations where it would be of the greatest
value, while the major and more affluent libraries will be held to ransom, and individual purchases
will be limited to the very well-heeled. As I have said, this is a very nicely produced book, but it is
in no way a luxury publication. Surely it could have been produced more cheaply so that this,
coupled with the prospects of wider sales, could have brought the price down? Otherwise, it is
difficult to interpret the pricing policy other than as a cynical exploitation of the major scholarly
libraries as a captive market, and this is an attitude that one is loath to attribute to such a very
distinguished publishing house as C.U.P.
J. F. M. CANNON
Flowers of the Himalaya. O. Polunin & A. Stainton. Pp. xxx + 580, with 128 colour plates, 74
pages of line illustrations, an illustrated glossary and 2 maps (end papers). Oxford University
Press, Oxford. 1985. Price £29.50 (ISBN 0-—19-—217623-4).
The increasing number of B.S.B.I. members who visit the Himalaya on tours should find this
long-awaited book of interest, and indeed for many it will be required reading. It is the first
popular guide to the flora of the Himalaya, from Kashmir and Ladakh to the Nepal-Sikkim border
in the east. Polunin and Stainton have collaborated well in producing a layman’s account which
retains botanical accuracy.
It is based upon the Enumeration of the flowering plants of Nepal, prepared by the British
Museum (Natural History) in recent years, and so adopts current taxonomic treatment and
nomenclature. Inclusion of a few more synonyms would have been helpful, especially to Indian
92 BOOK REVIEWS
readers who still follow Hooker’s The Flora of British India, though this is hardly within the scope
of a ‘popular’ guide. Some 1500 species are described, found mostly above 1200 m in the upper
valleys, and in the hills and mountainous regions up to 5500 m. Naturally this is not a Flora, as only
a selection of the estimated 9000 species supported by the area covered, can be represented. The
690 colour photographs are generally of a high standard and compare with Polunin’s European
guides, though inevitably a number are disappointing. Ann Farrer’s 315 line drawings add much to
this work, being accurate and providing an attractive illustration of a wide variety of families; the
inclusion of individual flowers or fruit in detail is to be commended. There is an informative
account of where to go plant-hunting in the Himalaya, and at what seasons of the year. Brief
descriptions of the influence of climate, geology, soils, altitude and man on the flora are given. A
clear glossary of terms will aid the beginner.
The book appears remarkably clear of errors in both original content and preparation. Some of
the upper altitudinal limits are incorrect, but this is a reflection of scanty and sometimes inaccurate
entries on herbarium sheets.
Oxford University Press have settled on an acceptable price on this occasion (cf. Flowers of
Greece and the Balkans), but publishers should note that this is very much the upper limit to which
even the enthusiast is prepared to stretch.
The authors are to be congratulated on an excellent book which will prove to be a standard
reference for decades to come, as it succeeds in covering the common and attractive species
encountered by a visitor. A. Stainton is currently preparing a supplementary volume, which wiill fill
in many gaps and give a better representation of the subtropical flora of the region.
C. A. CHADWELL
Wild flowers of the Yorkshire wolds. Sylvia M. Arnold. Pp. 85, with 14 black & white photographs
and 6 text-figures. Hutton Press Ltd., Beverley, East Yorkshire. 1985. Price (soft cover) £2.95
(ISBN 0—907033-—25-3).
The Yorkshire wolds is an attractive area with a rich flora about which little has been written. This
small book contains chapters on the various habitats with short lists of species for each. Under the
heading ‘Facts and Fiction’ are given interesting items of information for selected species,
including their former medicinal and other uses. There is also a chapter on conservation.
The selection of species to be mentioned in the book appears to be arbitrary. Some rarities are
included, whilst species characteristic of wolds habitats and widely distributed receive no mention,
notably Picris hieracioides, Galeopsis angustifolia and Koeleria cristata. A few grasses are included,
but only one sedge and one rush, so that some of the most interesting species of spring-fed marshes
are omitted.
There are minor criticisms and some queries: the Butterfly Orchid that is rare in wolds woods is
Platanthera chlorantha and not P. bifolia. If Polygala calcarea has been found on the wolds,
extending its known range by sixty miles or more, as recorder I would welcome information;
likewise, if there are authentic records for Dactylorhiza praetermissa X D. purpurella, I should like
to know of them. And does Veronica filiformis really occur on field margins?
This attractive, well produced book, nevertheless makes interesting reading and is a useful guide
to the habitats and wild flowers of the wolds, particularly for the beginner and visitor.
F. E. CRACKLES
How to make a wildlife garden. J. C. Baines. Pp. 192, with 70 colour photographs and 10 line
drawings. Elm Tree Books, London. 1985. Price £8.95 (ISBN 0-—241-11448-9).
This apparently simple book does more than fulfill its avowed purpose, it introduces a number of
important ecological ideas to the general reader and makes a useful practical handbook for
botanist-gardeners. Chris Baines’ breezy and colloquial style is surprisingly adaptable; directions
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BOOK REVIEWS 93
and explanations are notably clear, and narrative parts are lively and expressive, rising even to
eloquent rhetoric on occasion.
Now that our flora and fauna have to contend with rural as well as urban wastes, it is all the more
important to promote the idea of wildlife gardens, particularly for some meadow plants and for
creatures such as newts and frogs, and butterflies. Many misconceptions are demolished in the
course of this book — that a garden for wildlife is necessarily unkempt and scruffy, for instance, or
that simply scattering a packet of wildflower seed over a waste patch will ensure a wildflower
garden. Dispensing firmly with the idea that all wild plants behave like invasive weeds, the author
explores the preferences of a wide range of species and brings all his skills as a landscape architect
into his suggestions for a wild garden.
He shows how even a small garden can contain several habitats, attractive to plants and animals
(including human ones). His ‘cottage garden service station’, close to the house, includes many
nectar-rich plants and feeding places for winter birds. A meadow patch, however small, is good for
plants, insects and small mammals; and Chris Baines suggests that after the late summer cut, one
gives a garden party to achieve the trampling effect of grazing animals — the only way to get Yellow
Rattle (Rhinanthus minor) to self-seed, he avows. A pond is not only desirable but also
practicable, if one follows the useful instructions outlined here for making a pleasantly naturalistic
one (no crazy paving edges) with grassy slopes to it and a marshy area. There is a useful tip about
drip-filling, borrowed from Arab horticulturalists. The woodland edge habitat contains Primroses,
Red Campion, Bluebells and Foxgloves as well as some more unusual plants. (From my own
experience I would sound a cautionary note about planting Oxlips in proximity to Primroses, for
though you get some remarkable hybrids, the Oxlips seem to disappear after a few years). It is
good to find that a vegetable patch has a place in this wildlife garden, reinforcing the idea that, in a
balanced environment, food plants suffer no more depredations than in any other garden —
perhaps fewer.
There are sensible notes about the provenance of bought wildflower seed, on collecting seeds
and cuttings, and on growing wild plants, and a useful appendix of societies and nurseries.
Throughout the book there is an awareness of the wild garden as a part of a wider system. Larger
mammals may come to feed (Chris Baines saw a muntjac one morning in his garden, not many
miles from Birmingham city centre), and a place to feed and roost could be a matter of life or death
for winter birds, or a vital link in the life cycle of a butterfly or moth. While its specific concern 1s to
encourage people to be active on their own patch, this book looks also to the natural history over
the garden wall, and further abroad.
F. GREENOAK
Index Filicum — Supplementum quintum pro annis 1961-1975. F. M. Jarrett, with T. A. Bence, J.
W. Grimes, B. S. Parris & J. L. M. Pinner. Pp. 245. Clarendon Press, Oxford. 1985. Price £25.00
(ISBN 0-19-—854579-7).
Indices that give the original place of publication of plant names are essential tools of trade of the
taxonomist. The most familiar of these to flowering plant botanists is Index Kewensis, which was
launched in January 1882 by the then Director of Kew, Sir Joseph Hooker, who employed B.
Daydon Jackson to start work on the project. The work, initially published as two volumes (1893,
1895), with subsequent Supplements published at five- to ten-year intervals, listed all generic and
_ specific names of flowering plants from those of Linnaeus onwards together with their places of
publication. Thus /ndex Kewensis set the style for other lists of plant names and stimulated the
Danish pteridologist Carl Christensen to compile a similar list of fern names (Index Filicum; 1905—
_ 06) and later Supplements (1913, 1917 and 1934). The fourth Supplement, covering the years 1934
to 1960, was prepared by a committee of the I.A.P.T. and published in 1965 (Regnum Vegetabile
mol. 37):
The book under review is the Fifth Supplement and is again the work of the staff at Kew under
the direction of Dr Frances Jarrett, who had maintained at Kew a running index of all fern names
published since January, 1960. What is more, she wisely included in her index all pteridophytes
(Lycopodiopsida, Equisetopsida, Psilotopsida); in this supplement, and for the first time, the
94 BOOK REVIEWS
names of fern allies are included. They had hitherto been published separately, at different
times, and by different authors. Infraspecific names are not listed except where they provide the
basionym for a new name. All entries are cross-referenced to their basionym or the name from
which they derive, whilst each basionym is cross-referenced to the name which is derived from it.
Like the Fourth Supplement, it is not a nomenclator making taxonomic decisions on synonymy,
as did the original work. Preparing a work of this nature for the press is tedious and time-
consuming, but this book has been carefully and consistently edited. The compilers have included
all papers that have arrived at Kew, and we hope and believe that very few containing new taxa
have been missed. Any papers missed in Supplement Five will be included in Supplement Six.
If such works are to be effective they must be the result of an ongoing project, as indeed this
now is. Like so many long-term and on-going data files, it lends itself to electronic retrieval/
printing techniques. Publication is already ten years behind, and we must hope that more resour-
ces can be released for this important work so that the sixth Supplement does not take so long to
edit and publish.
A. C. JERMY
Med-Checklist: a critical inventory of vascular plants of the circum-Mediterranean countries, 1
Pteridophyta (ed. 2), Gymnosperms, Dicotyledons (Acanthaceae — Cneoraceae). W. Greuter, H.
M. Burdet & G. Long (eds). Pp. c + 330. Conservatoire et Jardin botaniques, Geneva &
Secretariat Med-Checklist, Botanischer Garten & Botanisches Museum, Berlin-Dahlem. 1984.
Price Sw Fr 98.00 (ISBN 2—8277-—0151-0; 2—8279—0004-1).
The preface to this book (in both English and French) explains that the Med-Checklist scheme
was set up in 1978 under the auspices of the European Science Foundation and placed under the
scientific authority of the Organization for the Phyto-Taxonomic Investigation of the Mediterra-
nean Area (OPTIMA). Most of the compilation is carried out in three centres: Berlin, Geneva
and Montpellier, with data derived from a network of regional and national advisors.
Families, genera, species and subspecies are the only formal taxonomic ranks recognized,
although the ‘aggregate’ (informal grouping for convenience) is used. All countries bordering the
Mediterranean Sea, together with Portugal, Bulgaria, the Crimea and Jordan, are included. This
area is divided into 27 territories, of which the European coincide with those of Flora Europaea
except for Malta, which is recognized separately from Sicily. The others are the East Aegean
Islands (all the Greek Islands not covered by Flora Europaea, but included in Flora of Turkey),
Algeria, Asiatic Turkey (Anatolia), Cyprus, Egypt, Israel and Jordan (combined), Libya, Leba-
non and Syria (combined), Morocco, Sinai and Tunisia.
The checklist gives taxa in alphabetical order, and assigns each a complex unique reference
number. Authorities and places of publication are given (including the synonyms). The countries
are listed across the landscaped page in sequence around the Mediterranean approximately
clockwise, starting with Portugal and ending with Morocco. Occurrence of the taxa is marked by
a ‘+’ sign, in the appropriate column(s). The work ends with a list of basic Floras, additional
references and an index of scientific names.
This work will obviously be an essential work of reference for many botanists, not only for
those interested in the Mediterranean; for it covers the whole territory of the countries consi-
dered, e.g. all France, not just the Mediterranean part. It is backed up by a series of notulae
published at intervals in the periodical Willdenowia. These contain many new records and an
alarming number of new names and combinations (many sadly lacking any explanation or discus-
sion). Some of the nomenclatural changes are alarming, and the genus Quidproquo, a nomen
novum with the species Q. confusum for Raphanus aucheri, is surely the way to give taxonomy a
bad reputation.
It is unfortunate that the work is retrograde in that it lumps together all the recent splits from
Lycopodium which do seem to have gained acceptance, On the other hand A. J. Scott’s radical
splitting of the Chenopodiaceae is recognized, including therefore the genus Blitum for the red-
fruited Chenopodium species. There is a considerable number of unnecessary deviations from
Flora Europaea, which cannot help towards a stable nomenclature and will certainly upset the
BOOK REVIEWS 95
horticulturalists, weed scientists, etc. Some contributors might be seen to appear obsessed by the
number of times their name appears as the authority following a taxon.
It should be mentioned that an earlier version of the Pteridophyta alone was published in 1981,
in paperback format, and distributed free to members of OPTIMA. A substantial revision of this
group proved necessary and it has been republished as a second edition in this volume. It is to be
hoped a second edition of the rest will not be necessary, especially as the work costs 98 Swiss
Francs (about £32), and a further five volumes are planned. Many of us eagerly await these, as such
a vital checklist is long overdue. The work can be obtained from: The Med-Checklist Secretariat,
Botanischer Garten & Botanisches Museum, Berlin-Dahlem, Germany BDR.
Sal. Jury
Plant chemosystematics. J. B. Harborne & B. L. Turner. Pp. x + 562. Academic Press, London &
Orlando. 1984. Price £65.00 (ISBN 0-—12-—324640-7).
As the authors point out in their Preface, this book can be considered to be a revised version of
Alston & Turner’s Biochemical Systematics, published in 1963. Plant Chemosystematics covers the
same field and attempts to review its state and potential up to the end of 1982; it includes two of the
opening chapters from Biochemical Systematics, but most of the book is entirely new.
The first four chapters of Plant Chemosystematics are introductory. Chapters 1 and 4 are brief
discussions of the uses, advantages and disadvantages of chemical and biochemical characters in
plant systematics. Chapters 2 and 3, dealing with taxonomic and evolutionary principles and
perspectives, are reprinted almost without change from Biochemical Systematics; they were
excellent in the context of 1963, but, as elsewhere in the book, modern views of population
structure and the role of selection are not reflected. Chapter 5 opens the major chemosystematic
section of the book with a useful but sometimes rather dated account of plant scents and odours, a
rather diverse group of volatile compounds with a variety of functions. As in other chapters, most
of the material is based on earlier reviews and is in part outdated as a consequence; the section on
terpenoid scents and odours is particularly unfortunate, with references to routine techniques of
the late 1960s and a ‘recent’ review published in 1975. Chapters 6 and 7, valuable surveys of
alkaloids, some other plant toxins, and plant pigments, are more satisfactory in this respect.
Chapter 8, rather misleadingly entitled ‘Hidden Metabolites’, deals with fatty acids and some other
lipids, wax alkanes, polyacetylenes, sugars, and some related compounds. As elsewhere, the
impossibility of covering the whole field to everyone’s satisfaction is clear; there are, for example
no references to the biochemical functions of sugar alcohols and cyclitols.
Chapters 9 to 13 return to a major theme of Biochemical Systematics, the use of micromolecular
(chemical) characters in studies of plant populations and intraspecific variation, interspecific
hybridization, and relationships between species, genera and families. Systematists will find these
chapters valuable, although there are, as elsewhere, occasional lapses (for example, nomenclatural
confusion in Baptisia on pp. 322-323, and the description on p. 309 of the Psilotaceae as ‘‘a non-
flowering taxon which first appears in the fossil record during the early Palaeozoic’). More
cross-references between chapters are needed. Chapter 14, dealing with the comparative
biochemistry of metabolic pathways, is interesting but sometimes rather dated (most recent
references for lysine pathways 1974, C3; and C, photosynthesis 1978, CAM 1978). Chapter 15,
dealing with phytoalexin variation, and Chapter 16, on the analysis and interpretation of chemical
data, are brief but useful.
Chapters 17 and 18 deal with the use of macromolecular data from proteins and nucleic acids.
The accounts of each topic are relatively brief but provide a good introduction, with references to
recent reviews, though not to the excellent symposium edited by Jensen and Fairbrothers (Proteins
and Nucleic Acids in Plant Systematics, Springer-Verlag, 1983), which appeared too late to be
included. There are a few lapses; for example, few biochemists would describe cytochrome c as an
enzyme (p. 395 sqq.).
This book cannot be unreservedly recommended as a guide to plant chemosystematics, but it
does provide an excellent introduction to many aspects of the subject, with useful evaluations or
interpretations of many points. The authors have relied on reviews for most of their general
96 BOOK REVIEWS
coverage, and as a result much recent work has been omitted. Nevertheless, this book will be a
valuable basic reference source to those with access to a copy; unfortunately, its extremely high price
will deter sales even to the best-financed libraries.
Q. O. N. Kay
The living Earth. C. Back. 4 parts: 1. Plants and simple animals; 2. Animals with many legs. Animal
homes; 3. Animals with backbones; 4. Our planet. Pp. (each) 4 x 64, with numerous colour and
black & white photographs and illustrations. A. & C. Black, Ltd., London. 1985. Price (per volume)
£4.95 (hardback) (ISBN 0—7136 (ail): 2263-6, 2265-2, 2267-9, 2269-5); £2.95 (paperback) (ISBN
0—7136 (all): 2264-4, 2266-0, 2268-7, 2270-9).
These children’s books, which have beautiful cover photographs, present a great deal of information
in simple form and contain many suggestions for experimentation and investigation by the reader.
However, it is most important that instructive literature for children, as well as being simple,
should be accurate. Not every parent, teacher or nanny is able to assess the worth of a scientific book,
and it does no service to a young mind to give it wrong or only partly correct information. In these
books there are many careless, inaccurate or oversimple statements which may seriously mislead.
Some examples are:—
““. . during the day and night, plants and animals take in oxygen and give out carbon dioxide.
This is called respiration.” — ‘““Tubers are swollen stems which grow on the roots. Potatoes are
tubers.”’ — “‘Every fertilised flower can develop into a fruit or cone” — ‘“‘Living things which
aren’t plants must be animals’’. — ‘“‘Molluscs — is the word used to describe animals with shells”’.
The drawings are large and clear but contain mistakes and inaccuracies, for example an ivy plant
attaching itself to a wall with roots half as long as the leaves. The fern called Dryopteris on p. 22 is
shown with peltate indusia, and on the same page the captions for Hart’s-tongue Fern and a drawing
which is intended to be Broad Buckler Fern are reversed. No idea of scale is given — an Amoebaisa
quarter of the length of an earthworm illustrated on the same page; similarly Planaria and a leech are
the same size. There are also a number of spelling mistakes, for example “dulce” for dulse and
“scull”’ for skull!
There is a commendable emphasis on conservation — most of the time. One 1s enjoined not to pick
a leaf or gather a moss, but “if you ever keep tadpoles in a bow! don’t forget to put in a stone’!
The final part of The living Earth, Our planet, which deals with climate, geology, physical
geography and so on and which ends with a sensible plea for conservation on both a local and a global
scale, seems better than the others.
I am unable to recommend these books, although they are very well produced with clear drawings
and good photographs and look most attractive.
A. LEE
Flora of Jersey. Frances le Sueur. Pp. xlii + 244, with 18 colour plates, 18 black & white illustrations,
about 600 distribution maps and 8 maps of physical features. Société Jersiaise, St Helier, Jersey.
1984. Price £17.50 (plus £1.50 p. & p.).
The spice of the floras of the various Channel Islands is their remarkable differences (in animal life
too), united though they are by an abundance of special plants — and the absence, or rarity, of others
common north of the Channel, another aspect of their differences.
These differences are evidenced in this welcome work by a line after each species summarizing its
frequency in the other main islands, as was done in The wild flowers of Guernsey, but here is
updated. Thus this Flora is a useful guide in the other islands as well.
But of course it is the native and naturalized plants of the largest of the islands that are so well dealt
with here. For 30 years or more Mrs le Sueur has been the indefatigable leader of the botanical
section, which she revitalized, of La Société Jersiaise. Moreover, she had the advantage of close
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BOOK REVIEWS 97
personal contact with her two most important predecessors, I. W. Attenborough, who had worked
on the flora since 1911, and the contentious, curious, Frere Louis-Arséne, with his troublesome
herbarium (see Watsonia 14: 167—176, 1982). She organized the island mapping scheme from 1960-
1970, which produced the dot maps of all the main species. This is a feature lacking from The wild
flowers of Guernsey (although such maps existed and were used in writing the text). It is better also in
its telling half-page photos, by her elder son, of various parts of the island; and, most strikingly, in its
decorative cojour plates. It is no mean feat to have produced this Flora with so many of these for so
low a price. The whole production looks good, and is good.
The whole-page maps at the end are not all of real use. Most have few names on them and some,
e.g. hedgerows and farmland, none at all. The awful gap in this book is of any map showing the
numerous places that are mentioned in the text, which renders some of it relatively unhelpful. It is
small excuse to suggest that readers should buy an Ordnance Survey map, which inevitably fails to
serve the purpose for the opposite reason: there are too many places marked.
Mrs le Sueur is a scholarly, all-round naturalist and has shown great shrewdness in assessing the
facts, which are right up to date, of the 1500 or so taxa treated. I can hardly imagine the job being
better done. The book is a thoroughly reliable source, a pleasure to browse into, and warmly
recommended.
D. McCLintock
A simple field key to common British wild flowers. S. M. Arnold. Pp. 125, with 12 plates and many
text figures. Lockington Publishing Co., Ltd., North Ferriby, East Yorkshire. 1983. Price £5.00
(ISBN 0-905490-—23-1).
I approached this volume in the hope that it would be a very usable key but found it disappointing. Its
main problem to my eyes is that it is extremely eclectic without saying so. There is a glossary that is
fair, with useful illustrations. The Key, which forms the body of the book, is well laid out but suffers
badly from the fact that in only a few cases are you even given a hint that there may be plants other
than those mentioned. If one, for example, tried to run down Fringed Water-lily (Nymphoides
peltata) with this key, it comes out as Yellow Water-lily (Nuphar lutea), and there is no hint that this
identification may not be correct.
There is a good index of combined English and Latin names, with cross-referencing; but of course
it refers only to the plants included. There are twelve half-tone plates of variable quality, and here we
find a bad discrepancy. The plates have no legends; for identification of the plants depicted one must
turn to the plate index, where each half plate is indicated as simply ‘a’ or ‘b’. These letters, however
do not appear on the plates, and the plants indicated are inconsistent, e.g. Pl. 1a is the top plant
whereas PI. 3a is the bottom one. There ts a real need for a simple illustrated key to common wild
flowers in Britain, but unfortunately this book cannot be recommended as such.
J. M. MULLIN
The Cambridge encyclopaedia of life sciences. Edited by A. Friday & D. S. Ingram. Pp. 432, with
numerous colour and black & white illustrations and line drawings. Cambridge University Press,
Cambridge. 1985. Price £25.00 (IBSN 0-—521-—25696-8).
As science has become more and more the province of the specialist, we have seen in recent years a
number of books that attempt to give an overview of current thought in the subject. The Cambridge
encyclopaedia of life sciences is the latest and one of the best of the genre. Compiled by a panel of
distinguished contributors, mainly but not entirely working in Cambridge, it covers, in the first part
of the book, the natural sciences in a progression from the cell and its work through to the behaviour
and ecology of living things in general. The second section is on environments and takes the topic
from biogeographical zones to the living organism as host for others. The third section is concerned
with evolutionary processes and their results as seen in the fossil record down to recent times, and the
final pages give a classification of living organisms.
98 BOOK REVIEWS
In such a synthesis, and particularly one that concentrates on processes rather than objects, the
treatment is bound to be uneven. There is far more detail, for example, on cells than on the
environments that they ultimately form, which often have to be dealt with very superficially. The
list of further reading at the end of each chapter in general provides suggestions for filling in the
inevitable gaps, but does not provide a single title that would enable a serious student to pursue
an interest in coral reefs, to note but one such omission.
The question as to whom such a book is directed is perhaps answered when one glances at the
text, where certain words, particularly in the early sections, stand out in heavy type. These are
difficult or key words that a student might underline in his notes; and it is probably such readers
who will make most use of this book, which for VIth formers and perhaps undergraduate
students gives an excellent statement of our current knowledge and opinions of the world about
us. Librarians in schools and colleges should find it in great demand. The more general reader
might hesitate before spending £25.00, but would find that the basic text, backed up by the many
beautiful coloured illustrations and excellent diagrams should give him pleasure as well as
information. The botanist pure and simple might feel hard done by, as plants certainly get a
smaller percentage of space than they should, perhaps, warrant. Their importance is mentioned
in many places but is rarely described in detail; and although one of the editors is a botanist, the
zoologists seem to have called the shots. Perhaps this is inevitable, but it would be nice to see a
book of this kind in which the plants got a fair crack of the whip.
< J. Pope
Guide des fougéres et plantes alliées. R. Prelli, with the collaboration and drawings of A. Prelli.
Pp. 199, with 79 figures and 3 tables. Editions Lechevalier, Paris. 1985. Price not given.
For too long France has had no up-to-date account of its pteridophyte flora. Those works that
were published in the past have long been virtually unobtainable and of course way out of touch
with the taxonomic advances of the past few years. All this makes the appearance of this neatly
packaged volume most welcome.
This book is really a slim field guide. The 199 pages make up a book only 14 mm thick
including the hard cloth binding. It is not a ‘coffee table’ book, but a practical fern flora of
France, adequately supplied with workman-like drawings and photographs to demonstrate
diagnostic characters. In style therefore it is not unlike Welsh Ferns (Hyde, Wade & Harrison
1978), although, unlike that book, the balance is towards more illustrations and less text.
Introductory chapters cover the biology of ferns and their classification, reproduction and
evolution. The order of species roughly follows that given in Flora Europaea but differs in detail.
More significantly the Filicales are here treated as one group and not subdivided into the several
well known family names, e.g. Hymenophyllaceae, Aspidiaceae, etc. This approach is taken,
perhaps justifiably, while botanists sort out and agree on one classification. However, I found the
lack of structure to such a large section of the book confusing. Furthermore the arrangement of
entries is inconsistent — sometimes a heading and generic characters are given before a section,
e.g. Asplenium and Dryopteris, but more often not, e.g. Cheilanthes, Hymenophyllum and
Gymnocarpium. In the chapter on ecology, most interesting tables of altitude and pH
preferences are given for selected species. It would have been fascinating to see all taxa included
in these tables.
Within the systematic section of the book, many features will be of particular interest to the
British botanist, as the emphasis is on recognition of each taxon — it is not a guide to localities.
Clear drawings are given to show differences between the stems of Diphasiastrum alpinum and
D. issleri, among others. Within the genus Asplenium, differences between A. obovatum
(recorded from the U.K. in error) and A. billotii and between A. adiantum-nigrum and A.
onopteris are well illustrated. Also, differences between Polystichum setiferum, P. aculeatum, P.
braunti and P. lonchitis are clearly defined by photographs and line drawings. I found the
photograph of Dryopteris remota particularly welcome — I now know my spore-raising under this
label was not successful! This is one of the few relatively widely distributed hybrids that are
illustrated; other rarer ones are simply listed at the end of each genus.
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BOOK REVIEWS 99
I was sorry to see that the familiar Cystopteris alpina has been placed under C. atrovirens. This is
probably a legitimate change, but there are already too many changes going on — especially among
the ferns. A good example of this is the recent and highly confusing revival of Polypodium
cambricum for P. australe. P. cambricum is used here as in recent British books — all on the basis of
a sterile variety named by Linnaeus which occurred probably only once in the wild. If this name
must be used for the Southern Polypody should it not be qualified by adding ‘var. australe’?
Another controversial taxa included here is Cystopteris dickieana. This is not a fern widely grown
in gardens all over the temperate world originating from the Kincardine coast, but the type with
spores somewhat similar to those of the Kincardine plant. In such a difficult genus is one
approximating character sufficient to allow two morphologically distinct forms to be joined into
one species? For me C. dickieana is still endemic to the British Isles.
While protecting C. dickieana as one of our very few endemic fern species, it is perhaps
appropriate to comment that the North-west European Hymenophyllum wilsonii is here recorded
as occurring in Africa, Australasia and America. This is not an unusual statement but it is not true.
South African plants are placed under H. peltatum and certainly differ from European material; no
doubt other southern hemisphere material is also distinct.
The bibliography at the end of the book is interesting but not very complete. Two fairly recent
works which come to mind are not included, viz. Badré F. et al. (1982), Le genre Cheilanthes en
France, Webbia 36(1): 1-38; and Poirion, L. et al. (1967), Pteridophytes de la Céte d’ Azure, etc.,
Webbia 22(1): 21-37. This latter work lists Osmunda regalis var. plumieri as occurring in southern
France. I would have liked to see this variety defined in the present work. Does it differ from the
Osmunda regalis we have in Britain?
Overall, these minor criticisms will not detract greatly from the value of the book. It is a very
welcome new field guide to the ferns and fern allies of France which is likely to be of great value to
many British pteridological tourists in France, and elsewhere in western Europe.
M. H. RICKARD
Communicating in science: writing and speaking. Vernon Booth. Pp. 68. Cambridge University
Press, Cambridge. 1985. Price £3.95 (0—521—27771-X).
If I, as an editor, had dictatorial powers over the authors whose work I edit, I should insist on their
reading and inwardly digesting this small paperback. So much good sense regarding the writing of
scientific papers (not to mention the advice to speakers) is to be found in these few pages, that all
writers on scientific subjects would benefit from reading them and carrying out the instructions and
advice that they contain.
Dr Booth’s book originated in 1970 as an essay entitled Writing a scientific paper and has been
gradually enlarged through several editions. It is dedicated to Th. M’Fline, an acronym for The
Man whose First Language Js Not English, and its watchword is clarity — how to say exactly what
you mean to say without indulging in (to quote a well-known radio programme) hesitation,
repetition or deviation. The first and longest chapter, ‘Writing a scientific paper’, deals with the
basics of grammar, punctuation and style that are so often ignored nowadays, sometimes resulting
in such gems as “After standing in boiling water for two hours, examine the flask”’ or ambiguities
such as “Lions eat more than antelopes”. Then follows good advice on ‘Preparation of the
typescript and figures’ and ‘Speaking at a public meeting’. How I wish that everyone who stands up
to give a scientific paper (or even merely a slide talk) would read and digest the latter chapter first!
In the next two chapters, ‘Addressed to those for whom English is a foreign language’ and ‘An
appeal to North Americans’, the author comes to grips with difficulties of English idiom and usage,
and points out that caring Americans are themselves concerned about the trend towards inelegant
writing and pomposity in works emanating from that continent. Complex adjectival phrases
(stacked modifiers) such as ‘“‘Barley root tip cell chromosome aberrations’’, for instance, tend to
stop the reader in his tracks. The work ends with advice on ‘Preparation of a doctoral dissertation
or thesis’ and an annotated bibliography.
I found this a very sound book that was amusing and refreshing to read and well worth the cost.
N. K. B. RoBson
100 BOOK REVIEWS
Documents floristiques. Tome III. Edited by F. Vignon et al. Pp. 203. Institut Floristique Franco-
Belge, Station d’Etudes en Baie de Somme, F 80230, Saint-Valery-sur-Somme, France. 1982. Price
FF 60 (ISSN 0182-0788). Obtainable from F. Vignon, Secretaire Laboratoire de Biologie Végétale,
U.E.R. des Sciences exactes et naturelles, 80039 Amiens Cédex, France.
This part of the ““‘Documents Flioristiques”’ contains, besides some botanical records of more local
interest, a series of detailed grid maps of the present distribution of 151 plants in Northern France
(from the Cherbourg peninsula eastward) and in Belgium and Luxembourg.
These ‘‘précartes” (preliminary maps) are part of a series covering, or planned to cover, the
greater number of the rarer of more localized vascular plants of the regions concerned. They
incorporate, as far as was possible, all previous records, as well as the results of the survey currently
in progress. They include therefore the data already published in the Atlas de la flora belge et
luxembourgeoise(E. Van Rompaey & L. Delvosalle, 1979), and for the extreme south part of the
Netherlands covered by the maps used, data from the Atlas of the Netherlands Flora (J. Mennema,
A. J. Quene-Boterendbrod & C. L. Plate, 1980).
In the absence of any equivalent of our British National Grid, a grid (of 4 x 4km squares) has been
employed, based upon the Belgian “carte d’état-major” at the 1:40,000 scale, by dividing up each
sheet of that map into 40 equal squares; this grid has been extended into the area of France
concerned. It has the great disadvantage that it is not printed on the local maps of the territories
concerned; we are very lucky to be able to use our clear National grid for mapping purposes so
painlessly! 2
The maps under review cover, in alphabetical order of generic and then species names, the less
common species present in the region from Galeopsis speciosa to Lythrum salicaria. Perhaps the
most striking general point that emerges from their perusal is that, contrary to what is often supposed
in Britain, there has been an even greater decline in N. France and in Belgium than in lowland
Britain of many species. This is particularly true of heath and bog plants, for example Gentiana
pneumonanthe, Hypericum elodes, and Lycopodiella inundata, all of which are now much rarer than
about 50 years ago in N. France and in Belgium, and almost confined to limited areas in the Cotetin in
W. Normandy, the Pays de Bray in E. Normandy, the Belgian-Dutch Campine region, and a few
scattered sites elsewhere. Hammarbya paludosa, indeed, now seems to have disappeared from the
whole region covered by the map used (though still persisting in Brittany and the Netherlands); by
contrast it is still frequent in our New Forest.
On the other hand, Halimione pedunculata, now extinct with us, is still holding its own on the coast
between Abbeville and Calais, while Himantoglossum hircinum is far commoner in N. France (some
200 localities shown) than in S. England, both on the chalk and on coastal dunes.
Liparis loeselii, though extremely local in N. France, and now almost confined to the dune slacks
and fens of the coast from Abbeville to Calais, has there what may well now be the largest and most
viable populations of the typical species (as opposed to our western var. ovata) left in all Europe
except possibly in extreme southern Germany.
Most of the species covered in this work, however, display the distribution patterns one would
expect on habitat and climatic grounds; thus Luzula forsteri, an Atlantic-Mediterranean species, has
a fairly sharp N.E. limit from the mouth of the Somme across to Laon and the upper Seine Valley.
The interesting outliers of the central European calicole flora are well exemplified by the maps of
Geranium sanguineum and Helianthemum apenninum; these species and others are almost confined
in the region concerned to the cliffs and chalk grasslands of the lower Seine valley, though the latter
also occurs on the limestone cliffs of the Meuse Valley on the Franco-Belgium border. As in Britain,
these species appear to be relics of a wider distribution from a time when open, calcareous habitats
were probably more widespread, before the forests closed in.
The map of Gentianella germanica clearly shows how this species entirely replaces G. amarella as
the common species of French and Belgian calcareous grasslands. G. amarella in N. France is a very
rare and declining species of coastal dune slacks; indeed some reputed sites for it produce now only
G. uliginosa.
Many non-British species are of course included among those mapped, and these cannot be
discussed in detail here, except that it is worth remarking that very few of these actually reach the
Channel coast, though often common inland, either on the chalk or in such areas as the acid
Ardennes forest.
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BOOK REVIEWS 101
There seems to be few errors, though the outlying Luxembourg localities for Hymenophyllum
tunbrigense, mentioned in the rubric, are not actually shown on its map, and the occurrence of
Globularia punctata on the downs of the Somme valley above Amiens is omitted.
The point is made however that these are provisional maps, to which, it is hoped, further field
work will add records.
This is indeed an interesting work, which all British botanists interested in the wider distribution
of British plants should obtain; the whole series will be most valuable when complete. It is good to
see that our French colleagues are now getting to grips with detailed mapping of the flora of at least
the north of their country.
F. ROSE
Plant facts and fancies. Sylvia Woods. Pp. 93, with numerous line drawings. Faber & Faber Ltd.,
London. 1985. Price £5.50 (ISBN 0-—571-—13436—-X).
This short book is essentially Victorian in its approach, both to its subject and its readers. It is
suggested that the work should be catalogued under ‘Plant — Juvenile literature’, and in her effort
to write for young readers the author stoops to oversimplification and a persistent moralizing tone.
On the whole the material presented is accurate, although as in many books on plant-lore there
is a tendency to place in the past superstitions and customs that are still very much alive. Thus,
although it is implied that belief in ‘lone thorns’ died out with the advent of mechanical hedge-
trimmers, it is still possible to meet people who not only hold the belief that to damage such thorns
invites misfortune, but also claim to have seen the fairies associated with the trees. At times it is
difficult to know if inaccuracies are caused by a desire to oversimplify or by inadequate knowledge
of the subject. Certainly the opening paragraphs of the chapter ‘Naming the flowers’ seem to
suggest the latter: ““The official botanical names of plants were finally decided in the eighteenth
century by a Swedish botanist called Carl von Linné.”’
All in all we have a conscientious offering which, though neither inspired nor inspiring, provides
half-an-hour’s entertainment.
A. R. VICKERY
Holding your ground. An action guide to local conservation. A. King & S. Clifford. Pp. ix + 326,
with 39 black & white photographs and figures. Maurice Temple Smith, London. 1985. Price £5.95
(ISBN 0-—85117—250-4).
Here is a book for the desk-top of any conservation worker or group concerned with protecting
local wildlife, landscape and old buildings. Angela King and Sue Clifford have produced an ‘action
guide’ for Common Ground (an agency concerned with promoting and exploring our common
cultural heritage), which contains both character and a thoughtful compilation of valuable
information.
After emphasizing the importance of local environment to the fabric of our common culture and
explaining the major ways conservation can be put into practice locally, the main text is given over
to doing just that. Each chapter deals with a local habitat, for example trees and woodland,
summarizing its value. This value is given a broad interpretation: ancient woodlands not only
contain a wide diversity of plants and animals but also have landscape and amenity value and
historical and cultural interest. This refreshing perspective runs throughout the book and will
encourage anyone who picks it up to read on to the next stage: advice and examples of how to turn
interest into action.
The chapter on trees and woodland deals with Tree Preservation Orders; felling licences; the
survey, purchase and management of conservation areas; tree nurseries and planting; and town
trees and grants. At the end of the chapter the statutory agencies are described, and references and
further reading are prescribed. This pattern is repeated for other habitats and also for landscapes,
monuments and buildings. (There is also a short chapter on the conservation of plants and
102 BOOK REVIEWS
animals). The character of the book lies in the way the authors have blended together essential
information on, say, how to purchase a woodland with an illustration of how this can be achieved, in
this case the purchase of Hardings Wood. These examples are written by the local groups responsible
for the ‘action’ and contain not only the essential details but also the local flavour of a project. Ideas
and possibilities take shape that to the reader are often little more than pipe dreams. The examples
come from a range of trusts, projects, societies and individuals and describe both the successes and
the failures.
The final four chapters deal with the organization of local action, collecting and using information,
and understanding of local government, and how to achieve financial and physical help.
So often today, in the face of modern decision-making processes, there is a feeling of helplessness
— that the control of our environment is out of our hands. This book offers some sound and carefully
presented information on how to at least hold your ground. But more importantly, through carefully
used examples of local people achieving local success, it kindles the confidence in the reader to joinin
this success.
P. M. WADE
Ecological Flora of the Shropshire Region. C. A. Sinker, J. R. Packham, I. C. Trueman, P. H.
Oswald, F. H. Perring, & W. V. Prestwood. Pp. xvi + 344, with cover paintings and 8 colour plates
by Anne Gilbert, chapter headings by Lindsay Brown, 4 black & white photographs and endpapers,
48 text figures, 18 tables, numerous distribution maps and 12 overlay maps. Shropshire Trust for
Nature Conservation, Shrewsbury. 1985. Price £23.00 (ISBN 0-—9508637—0-X).
To some people, Shropshire may not spring to mind as one of the botanical aristocrats; but it is a big
county, and in the hands of this distinguished panel of authors it emerges as a fascinating and varied
area, a pleasant hunting ground for the field botanist, and a challenge to the ecologist.
This is not the first county Flora to profess an ecological bias, but there has been nothing on this
scale previously. The book is divided into three parts. The first gives an account of the methods used
in the present survey, followed by an extensive treatment of the history of botanical recording in the
county. The chapter on ‘Environmental background’ is readable and informative, covering climate,
topography, geology, soils and land use. These aspects are treated with considerable academic
vigour; for example, much recent work on Shropshire soils is discussed in some detail, and its
relevance to the vegetational patterns is considered. Of less value is the table of ‘Environmental
Profiles’, in which certain features are listed for each 10 km square. This seems to have limited
relevance, since there is no corresponding vegetational information on a 10 km basis. Twelve
selected environmental features have in fact been plotted on a tetrad basis, and these are given as dot
maps, which are also duplicated as a transparent overlay, a pocket for which would be useful. The
section ends with a short chapter on ‘Biogeographical elements’, in which one of the authors
(F.H.P.) indicates how much our knowledge of plant distribution has changed since the publication
of J. R. Matthews’ Origin and distribution of the British Flora in 1955.
Part Two is entitled ‘Habitats and plant communities’. The authors recognize four divisions: open
water and wetland; rock, heath and grassland; woodland, hedge and scrub; and disturbed and
ruderal habitats; and if you have forgotten what ruderal means, you are advised to think of ruins,
rubble, rubbish, railways, roadsides and runways! There follows a chapter in which the author
(C.A.S.) summarizes our knowledge of the vegetation of the region in the past, based on pollen
analysis, illustrates some fascinating ecological relationships on maps, and argues a convincing case
for conservation priorities.
The third part is the traditional systematic account of the vascular plants and their distribution.
The area surveyed is a large one, comprising 1100 tetrads; and it was decided to designate common,
intermediate and rare species for mapping purposes. About 200 of the common plants are not
mapped, neither are the rare plants that occur in fewer than three tetrads. The dot maps are placed
with the relevant text, and ecological notes include habitat, associated species, water regime, soil
texture, nutrient and base status, pH, microclimatic conditions, biotic factors and reproductive
strategies. The book ends with a list of herbaria containing Shropshire specimens, and a gazeteer of
place names.
BOOK REVIEWS 103
Perhaps the most radical feature of this new Flora is the analysis of the plant records by
computer methods. Two ordination techniques have been used: reciprocal averaging and indicator
species analysis. In each of the four habitat types, 42 stands are analysed, and the full species/stand
matrix is printed. The completed ordinations are then used as the basis for a discursive ecological
treatment of each habitat.
There are eight pages of colour illustrations (each dealing with a particular habitat), some
attractive sketches introducing each chapter, and a dozen or so aerial photographs with extensive
captions.
The area covered by the Flora is ‘““The Shropshire Region’’, interpreted as Shropshire, plus bits
of neighbouring counties to make up a ‘tidy’ rectangle. Interestingly, the few bits of Shropshire
which fall outside this rectangle are not omitted, so we end up with a rectangle plus warts! There
have been many critics of this pattern, whom the authors are not afraid to quote. I have some
sympathy with the sentiments of one of my fellow Welshmen who remarked “‘ ‘Greater Shropshire’
indeed; I’d make it the East Riding of Powys!” In fairness however, I must compliment the authors
on spelling the Welsh place-names correctly, e.g. Pumlumon.
Who was the cynic who said that the Bible was the only good book written by a committee? This
Flora has six authors, but the whole work hangs together effectively. Charles Sinker’s compelling
prose in the Preface sets the tone for a volume that is both scholarly and readable. It sets a new
standard for county Floras.
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Watsonia, 16, 105—108 (1986) 105
Obituaries
OLEG VLADIMIROVICH POLUNIN
(1914—1985)
Oleg Polunin died in his sleep on 2nd July 1985. A few weeks earlier he had been diagnosed as
suffering from Motor-neurone Disease, and he was thus spared much suffering. Indeed, he
continued to work, despite considerable weariness, until the day before he died. He was a
schoolmaster by profession, but he was widely known even beyond the botanical world as the
author of an acclaimed series of Field Guides to the flora of Europe and elsewhere. His travelling
and writing commitments had not allowed him to work actively on the British or Irish floras for
some time, but he had been a B.S.B.I. member for 40 years and he was undoubtedly a major
influence on British botanists learning about the flora of Europe and the Mediterranean region.
Oleg was born on 28th November 1914 at Checkendon near Reading in Berkshire. His Russian
father and English mother were both artists, who had collaborated with Diaghilev on stage sets for
his ballets, and Oleg himself was in many ways less a scientist than an artist with a passionate love
of plants and botany. He was educated at St Paul’s School, London, from where he went up to
Magdalen College, Oxford. He read Botany at Oxford at a time when the chair was held by Sir
Arthur Tansley, whose example in field studies and ecology was so great an influence on many
younger botanists. Oleg joined the staff of Charterhouse School, Godalming, in 1938 as a Biology
master, but was absent from the school during World War II when he served in the Intelligence
Corps, travelling widely in the Far East and elsewhere. In 1943 he married Lorna Venning, who
was then an actress (and who was to make a substantial contribution to production and costumes of
stage presentations at Charterhouse). Their long and happy marriage was a firm basis for Oleg’s
work and subsequent travels.
Following his return to Charterhouse in 1946, Oleg threw himself enthusiastically into teaching.
Many Carthusian naturalists and biologists owe a great deal to his encouragement, enthusiasm and
example, for his teaching went far beyond the bounds of the classroom. An important feature of
the post-war years at Charterhouse was a summer camp for Carthusian biologists run by Oleg and
Lorna, accompanied by their young family, [van and Natasha, at Lough Ine in Co. Cork. As well
as introducing the boys to the Irish countryside and its natural history, these visits resulted in a
_ valuable floristic inventory of Sherkin Island and other islands off the coast of south-western Co.
~ Cork (v.c. H3, West Cork), published in Watsonia 1: 359-363 (1950).
| Later Charterhouse biologists’ expeditions visited the Isles of Scilly, where Oleg was
accompanied by another great enthusiast, the late Percy Chapman, who taught the fabulous
mysteries of rockpools and their varied inhabitants. The Biology Department at Charterhouse
_ flourished during the era of this benign and scholarly partnership. Oleg made further floristic
_ contributions in Scilly and during the 1950s he was B.S.B.I. recorder for West Cornwall (v.c. 1).
_ He also wrote an account of the vegetation of the islands, Some plant communities of the Scilly Isles
_ (1953), in the form of a mimeographed handout to his pupils. This useful document is now very
_ rare and I know of only three copies! Oleg was an active contributor to the B.S.B.I. Atlas mapping
scheme, and ‘his’ 10 km square, based on Godalming, had a higher tally of species than any other
in the British Isles.
A consequence of this active field work was that Oleg collaborated with many professional
_ botanists and, during a visit to the British Museum in 1948, to check specimens from Sherkin
Island, he was invited by Sir George Taylor, then Deputy Keeper of Botany, to accompany Major
_H. W. Tilman’s 1949 expedition to Nepal as the group’s botanist. He thus became one of the few
botanists to visit that country since Sir J. D. Hooker one hundred years before, and the first
westerner to make extensive collections there since I. H. Burkill in 1907 and Col. F. M. Bailey in
_ the 1930s. In 1952 he took part in a wholly botanical expedition to Nepal, with W. R. Sykes and L.
\
i
106 OBITUARIES
H. J. Williams, under the joint auspices of the British Museum (Natural History) and the Royal
Horticultural Society. From then on Oleg travelled extensively, to Turkey in 1954 and 1956 (with
Peter Davis), Kashmir in 1956, Irag in 1958, Lebanon in 1959 and the Karakorum in 1960 (with the
Anglo-American Karakorum Expedition), collecting herbarium material and living plants and
seeds. These expeditions were written up in various articles in Journal of the Royal Horticultural
Society, Gardeners’ Chronicle and elsewhere. An eager traveller, he was happiest when on the
road, especially, from the 1960s onwards, in the company of Lorna in their camping van. This
vehicle was equipped with an outside rack for drying plant-presses, a small library, cooking
facilities, good food and drink as available locally, and various tables, chairs and awnings to make
an outdoor study. Many friends joined this long-running roadshow, sleeping in tents or even, as
Oleg preferred, in the open under the stars; such was my own introduction to the Greek flora in
1970. Oleg and Lorna made an excellent team, he writing, collecting and photographing, she
shopping in several languages and feeding and watering the absent-minded scientist. Both
absorbed with great vigour the countries through which they passed, making many new friends.
Further opportunities for travel were furnished from 1960 by a number of ‘special interest’ tour
operators, who engaged Oleg as guest lecturer to instruct their clients on holiday. This enabled him
to see large areas relatively cheaply. Oleg’s tours concentrated on cruises in the eastern
Mediterranean, notably in the Greek Islands, and pony-treks in Kashmir and Nepal in the
Himalaya. More recently he accompanied tours to the Caucasus, a successful reunion with Mother
Russia; sadly he never had the time to develop that area of interest, but he was very thrilled by the
region. .
He took early retirement in 1972 to devote himself to the writing of his famous series of Field
Guides. Flowers of the Mediterranean (1965, with Anthony. Huxley) and Flowers of Europe (1969)
set the pattern of these remarkable volumes, that combine scholarship with an appreciation of the
aesthetic side of botany, and an understanding of the needs of the amateur or novice. Flowers of
South-west Europe (1973, with Bill Smythies) was the first of the series to give descriptions of
individual areas where rare or interesting plants may be found, and it is these regional accounts,
above all, that reflect the author’s love and rapt enthusiasm for his material. Flowers of Greece and
the Balkans (1980) and Flowers of the Himalaya (1984, with Adam Stainton) were perhaps the
volumes that gave Oleg the greatest emotional satisfaction, and his fondness for the flora, scenery,
peoples and culture of both regions radiates vividly from the written page. Most of these Guides
have been translated into several languages and they comprise a worthy and lasting memorial to
their author. An important aspect of the Guides is their emphasis on conservation, a cause that
Oleg embraced long before it became as intellectually respectable as it is today.
The long sections of plates in these books, of colour photographs taken mostly by the author,
have set a new standard for publications of this type, and the line drawings, notably those
illustrating groups of species characteristic of an area or a habitat, were chosen with care and
executed by skilled artists. From the very first, Oleg was a champion of Flora Europaea, itself
another outstanding achievement of floristic botany, and the Field Guide series to a large extent
derived from, and forms a valuable accompaniment to, its volumes, to which it provides an
unofficial companion series of illustrations and background information. The Polunin books have
certainly served to introduce Flora Europaea’s overall view of the continent’s flora to a wider and
less specialized public. Oleg made a direct contribution to the Flora Europaea project by sending
his herbarium collections to Leicester University herbarium (LTR), where members of the Flora
team identified them ‘‘after tea’, and these collections remain an important element in that
herbarium. Oleg always worked closely with professional colleagues in Britain and abroad and
checked any queries or difficulties with them. Some professional botanists have occasionally
carped at the Field Guides for not giving fuller coverage of particular taxonomic groups, but these
books have been widely and profitably used by both amateurs and professionals. Flowers of the
Mediterranean and Flowers of Greece and the Balkans, for example, are the only accessible,
authoritative texts on the general Greek flora that are currently available. Oleg set out to educate,
not to have the final word on the subject.
His considerable achievements were honoured by the award of the Royal Horticultural Society’s
Veitch Memorial Medal in 1962, for introducing garden plants of quality, and their Grenfell Medal
in 1969 for the photographic plates in Flowers of Europe; and by the award of the Linnean Society
of London’s H. H. Bloomer award in 1983, for an outstanding contribution by an amateur
OBITUARIES 107
biologist. He is commemorated in the specific epithets of an Aristolochia, a Pedicularis, a Primula, a
Ranunculus and a Saxifraga, also those of an aphid and a frog!
When he was not travelling, Oleg had a secure home base in Godalming, for a long time at
“Dormers” (formerly the home of Julian and Aldous Huxley, whose father had taught at Charter-
house), and latterly since his retirement in a roomy flat at the top of Frith Hill (near to where Alfred
Russell Wallace had lived for a while). Here he worked, surrounded by books and cabinets of his
colour-slides, in a study that looked out on to the Surrey hills where he loved to walk in the
afternoons as a break from writing. In recent years he had less time to devote to our own flora, but it
was always a source of considerable pleasure to him. A walk around Charterhouse Copse or the
marshes of the River Wey in early spring would fill him with delight, and he would exclaim happily at
the first show of Sweet Violets or Marsh Marigolds. The artist in him was never far below the surface.
He lived amongst his collection of paintings, many by family or friends, and amongst rugs,
ornaments and sculptures picked up on his travels in the East. He was a gifted potter and enjoyed
presenting his friends with his wares — frequently robust cooking or eating utensils of indispensable
practical value as well as artistic elegance. In order to unwind from the discipline of writing, he had a
fondness for loud, wild dance music from the Balkans to India or beyond. It was splendid to enter
Oleg’s study after he had finished a day’s work and to see the scholar dance an impromptu jig to
bagpipes, zurna or bouzouki! He worked hard and played hard.
Above all, Oleg had a genius for friendship. The Polunins’ home has always been an open house to
family, neighbours and visitors, representing a wide range of interests. Many of his large circle of
friends were present at Charterhouse just before the end of 1984 for his 70th birthday celebrations.
One is left with a lasting image of a big, handsome, genial and kindly man, pleased to talk to
anybody, and deeply in love with everything he did. Those who knew him will miss him badly, but we
are all richer for his teaching, his friendship and his unique and special contribution to floristic
botany.
J. R. AKEROYD
ESTHER DORIS PUGH
(1910—1985)
Doris Pugh, a B.S.B.I. member since 1967 and Recorder for Monts., v.c. 47, since 1977, died on 16th
February 1985, having suffered a stroke the previous September.
Doris lived practically all her life in the village of Pant on the Shropshire—-Montgomeryshire
border. Daughter of the village postman, she taught at the Llanymynech primary school from the
early 1930s until her retirement, and she was also a pillar of her local Methodist Chapel.
Everyone in the botanical world seemed to know Doris. All over the country I have met people
who remember with pleasure the superlative flora of Llanymynech Hill or the Breidden or of almost
any other part of this unique and precious corner of Britain. They also remember with almost equal
pleasure taking tea and ‘grannie’s crunch’ on Doris’s lawn and the rarities which seemed to spring up
by magic in her garden. Doris seemed to know every plant in field and hedgerow as a personal friend,
and yet, at the end of a day in the field, her little posy of ‘bits’ nearly always included a new record or
the key to an old mystery. Always modest, and even shy, with a warm and wry sense of humour, she
taught us all, from expert to tyro, and generally made us think we were teaching her.
In 1977 the first ever comprehensive plant list for v.c. 47 was published. Edited by M. Hignett and
W. S. Lacey, it was largely compiled by Doris from the records of Janet Macnair, a previous
Recorder for v.c. 47. By this time she had become a prime but self-effacing mover in the Shropshire
Flora project, for which she was a tireless worker throughout the 1970s and 1980s. She did not quite
live to see the Flora published, but the editors never missed an opportunity to show her each new
_ chapter, each new illustration, as they were produced. The pages are liberally sprinkled with her
_ observations, and many of the recorders for the Flora learned their trade in Doris’s kindly hands.
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When this task was finished Doris plunged without hesitation into the preparation of a new Flora
of Montgomery, and those of us who have helped a little have been amazed at the skill and efficiency
_ with which she has conjured up a viable and enthusiastic team of recorders in this sparsely populated
area. A Flora for v.c. 47 will be yet another lasting monument to this extraordinary lady.
108 OBITUARIES
We will all remember Doris. I will remember her at the end of many a day’s recording, still fresh
after scrambling through thickets, climbing over fences, and protecting us from farmers (she
seemed to know them all). She would be miles behind down the path, she would have seen
everything we had seen and more, a dumpy little grey-haired lady of inestimable value. How we
will miss her!
I. C. TRUEMAN
Watsonia, 16, 109-120 (1986) 109
Reports
ANNUAL GENERAL MEETING, 11TH MAY 1985
The Annual General Meeting was held in the Department of Chemistry, University of Reading, at
12 noon, with 93 members present. Mr J. F. M. Cannon F.L.S., retiring President, took the Chair
and opened the meeting by thanking Professor V. H. Heywood and B.S.B.I. members on the staff
of the Department of Botany, Plant Science Laboratories for the use of the Department’s facilities
and for the excellent organization of the programme.
Apologies for absence were read, and the adoption of the Minutes of the last Annual General
Meeting, as published in Watsonia, 15: 295-296 (1985), was proposed by Dr H. J. M. Bowen,
unanimously approved by the meeting, and signed by the President.
REPORT OF COUNCIL
Following the adoption of the Report for the calendar year 1984, proposed by Mrs J. G. Dony,
seconded by Mr M. G. Young and carried unanimously by the meeting, the President invited Dr P.
M. Wade, Chairman of the B.S.B.I. Conservation Committee to report on the progress of plans to
appoint a botanical conservation officer. Dr Wade outlined the current position on the formation
of a Conservation Association of Botanical Societies, and answered questions from members on
this new development. The meeting gave general approval to the plans, indicating by show of
hands a clear majority in favour of an appointment as reported. The retiring President thanked Dr
Wade and the sub-committee for the great deal of work involved in the preparation of the final
proposals which had incorporated a diversity of ideas through the development of the planning.
TREASURER’S REPORT AND ACCOUNTS
The Treasurer proposed the adoption of his Report, which was seconded by Mrs M. Burnip and
carried unanimously by the meeting.
MEMBERSHIP SUBSCRIPTIONS
Due notice having been given of a recommended increased subscription rates as follows —
Ordinary £12.50, Junior £5.00, Family £1.00 and Subscriber £12.50, from Ist January 1986, the
Treasurer proposed adoption. This was seconded by Lady Anne Brewis and carried by the
meeting.
AMENDMENT TO RULES
The Honorary General Secretary introduced the recommended amendments to Rules 3, 4, 6 and 7,
to rationalize representation on Council and to reduce the size of Council:
— Rule 3, to read: The management of the affairs and property of the Society shall be in the
hands of a Council consisting of: (a) The Honorary Officers, nominated by Council and elected (or
re-elected) by members at the Annual General Meeting (length of service as in Rule 9). (b) The
Honorary Receiving Editor Watsonia, the Honorary Editor B.S.B.J. News and the Honorary Field
Secretary, appointed annually by Council. (c) The Honorary Secretaries of the five Permanent
Working Committees (Rule 4), elected annually by members of the appropriate Committee
(Schedule II, 2b). (d) Twelve members elected at the Annual General Meeting, according to Rule
10, and Regional Representatives according to Rule 11 (no change).
— Rule 4, Permanent Working Committees: delete (c) Junior Activities (but add to Meetings
Committee terms of reference, Schedule II. A representative to consider how the Society can
make known its activities to, and cater for, young botanists and beginners, shall be nominated by
the Committee as an ex officio member).
— Rule 6, to read: The Officers of the Society shall be the President, not more than four Vice-
Presidents, an Honorary General Secretary and Honorary Treasurer.
110 REPORTS
— Rule 7, Officers’ duties: delete (c) Honorary Meetings Secretary, (d) Honorary Field Secretary
and (f) Honorary Membership Secretary. (e) Editors, to read: The Honorary Receiving Editor of
Watsonia and the Honorary Editor of B.S.B.I. News shall be ex officio members of Council. All
Editors of the Society’s Journals shall be ex officio members of the Publications Committee.
The Treasurer explained the addition to Rule 24 for possible future implementation as follows —
Rule 24, add: Council to have the power to reduce the annual subscription for any member paying
by Direct Debit.
The adoption of the amendments to these Rules was proposed by Mr A. L. Grenfell, seconded
by Mrs M. E. S. Farrand, and carried by the meeting.
ELECTION OF OFFICERS
The retiring President, Mr J. F. M. Cannon, proposed the election of Mr D. E. Allen M.A., F.L.S.
as President. This was carried unanimously with applause and Mr Allen, taking the Chair,
commented that his election added a small facet to the history of the Society as he was the first ex-
Junior Member to be President. From the Chair Mr Allen proposed Dr N. K. B. Robson F.L.S.
for election as Vice-President; Mrs M. Briggs M.B.E., F.P.S., F.L.S. for re-election as Honorary
General Secretary and Mr M. Walpole for re-election as Honorary Treasurer, referring to these as
the ‘king-pin posts’ for the Society. These were carried unanimously with applause.
ELECTION OF COUNCIL MEMBERS
In accordance with Rule 10, nominations had been received for Dr J. R. aces F.L:S.; Dre. a
Cadbury and Mr H. J. Noltie. Their election was proposed from the Chair and carried
unanimously.
The new President expressed thanks on behalf of the Society to all the Honorary Editors and
Secretaries (now, following the amendment to the Rules above, ex officio members of Council) for
their very considerable work for the society:
Honorary Editors: Dr J. R. Akeroyd F.L.S., Dr B. S. Rushton, Dr R. J. Gornall, Mr C. D.
Preston, and Dr N. K. B. Robson F.L.S. (Editors Watsonia), and Dr S. M. Eden (retiring Editor
Watsonia), Mr D. H. Kent (Editor BSBI Abstracts) and Mr E. D. Wiggins (Editor B.S.B.I. News).
Honorary Secretaries: Mr J. F. M. Cannon F.L.S. (Co-ordinating Committee), Mrs J. Robertson
(Meetings Committee), Mr R. Smith (Field Meetings), Mr A. O. Chater (Publications Commitee),
Miss L. Farrell (Conservation Committee) and Mr D. J. McCosh (Records Committee).
Mr Cannon was thanked for his good offices during his Presidency, in particular for his guidance
during the planning and discussion on two major projects which had the potential for far-reaching
influence on the Society — the appointment of a Botanical Conservation Officer, and a new
B.S.B.I. Survey project.
RE-ELECTION OF HONORARY AUDITORS
The Honorary Treasurer, expressing our gratitude to Messrs Thornton Baker for auditing the
Society’s accounts, proposed their re-election. This was carried unanimously.
ANY OTHER BUSINESS
Mr R. M. Burton asked when a new list of members would be available, and Mr J. Ounsted
enquired if a larger typeface could be used. The Honorary General Secretary hoped that a list
could be printed before the end of the year, and the Honorary Treasurer would look into this
possibility. The print-size request was noted, but would be dependent on costs.
Dr P. Macpherson, Secretary to the Committee for Scotland, asked for news of the field
recording cards for Scotland in preparation by the Biological Records Centre and now three years
behind schedule. The Honorary General Secretary would again enquire for these.
The Honorary Treasurer reported that there would be a delay in publication of A Map Flora of
Mainland Inverness-shire. There being no further business the meeting closed at 12.59.
M. BricGs
REPORTS 111
PAPERS READ AT THE ANNUAL GENERAL MEETING
THE PARK AT WHITEKNIGHTS
For those with a sense of the whimsical, Whiteknights has a delightful history. This is particularly
true of the period 1798-1818 when the park was owned by the Marquis of Blandford, who
ornamented the estate in a most elaborate way with fountains, pavilions, bowers, vineyards, statues
and all the latest exotic plant introductions. By 1818 the Marquis was bankrupt, his debts to one
nurseryman alone amounting to £15,000. Much of this detail we know from a lavishly illustrated
book by a Mrs Hofland.
Today, the same site has 34 academic buildings and five Halls of Residence. The area totals 300
acres including woodlands, lakes and meadows as well as the more formal gardens around the many
buildings. In many ways, the University is a small town, with all the associated problems of access,
parking, services, etc., and many modern buildings requiring bold landscaping. The park also caters
for the recreation needs of both students and staff including formal sports as well as jogging, fishing
and walking. Many appreciate the conservation aspects of the park and in particular the many birds it
attracts. Although little of the original Blandford garden remains, some areas such as the Wilderness
with its grotto can still be seen as a historic landscape. Recent sponsorship by the Manpower Services
Commission for a two year community project has enabled us to partially restore the Wilderness and
carry out long-neglected conservation work.
In general the central areas have been treated formally, with mown lawns and neat shrub borders
of exotic plantings, all maintained using modern methods including much mechanization and
herbicides. The many mature and rare trees need constant care to keep them safe and healthy. New
plantings comprise shrubs, trees and groundcover for ease of maintenance, but the extensive range
of plants we use continues the tradition of imaginative planting which has always been encouraged by
the University because of its various plant-associated courses. By contrast, the conservation area is
relatively undisturbed, with the meadow areas being cut once a year for hay, brambles contained and
trees trimmed only when safety demands. New plantings are of predominantly native species and the
wild flora is encouraged to develop.
It can therefore be seen that any management plan has to be both complex and flexible to allow for
the many uses and needs of a community which is both genuinely critical and constantly changing.
I. K. S. COOKE
CHANGES IN THE BERKSHIRE FLORA
The plants of Berkshire, v.c. 22, were the subject of Floras by G. C. Druce in 1897 and myself in
1965. They have obviously been affected by many changes in land-use over the last century, for while
the area of woodland has remained more or less constant, at 10% of the total, much arable land
reverted to grassland during the period 1870-1939. Much grassland was ploughed in 1940 and has
remained arable ever since.
Woodland species have shown some changes which are not easily explained. Thus Sorbus
torminalis is much commoner than it was in Druce’s time, and Rhododendron ponticum (unrecorded
by Druce) has colonized half the grid squares in the county. However Colchicum autumnale has gone
from about half Druce’s sites, and Orchis militaris became extinct around 1907. Heath vegetation,
usually maintained by fire, has suffered from encroachment by farmers and builders. Few species
have been completely lost, apart from Orobanche rapum-genistae; the alien Odontites jaubertiana
colonized two heathy sites about 1945, but is nearly gone now. Bog vegetation is all declining as the
habitat is lost, and Gentiana pneumonanthe was lost over 100 years ago, though it survives nearby in
north Hampshire.
The better chalk grassland sites are partly protected in reserves, where Pulsatilla vulgaris and
Orchis ustulata survive, but O. simia is extinct and Spiranthes spiralis remarkably rare. Other
grassland species have declined due to ploughing (e.g. Genista tinctoria and Orchis morio), grazing
or lack of fallow (e.g. Echium vulgare and other biennials), but Lupinus arboreus is an introduced
species which is spreading.
Most plants of wetlands, including drying mud, are in decline, though Parentucellia viscosa
appeared for the first time in 1969 and Cyperus fuscus was refound in 1982. Ranunculus lingua and
Fritillaria meleagris are examples of declining species that it would be sad to lose. Many arable weeds
112 REPORTS
are vanishing due to improvements in agriculture. Adonis annua probably survives, but
Agrostemma githago, Arnoseris minima and Bupleurum rotundifolium have not been seen
recently. The total number of extinctions in the county lies between 40 and 55 species.On the other
hand, since 1965 eight new native species have been added to the flora and eleven ‘presumed
extinctions’ have been refound.
H. J. M. BOWEN
THE PLANT SCIENCE BOTANIC GARDEN
The Botanic Garden and the adjacent Experimental Grounds were founded in 1972 and
reorganized, following the acquisition of additional land, in 1975. Today the Garden occupies
some eleven acres and the Experimental Grounds two and a half acres. The Garden is set in what
was the home paddock of a Victorian house called “The Wilderness” (now demolished), which had
itself been part of the garden of the Marquis of Blandford. The main aims of the Botanic Garden
are to provide a scientifically organized collection of plants for teaching and research, to maintain
conservation collections of selected groups of plants, to play a general educational role for the
public, and to provide an amenity for the University and local residents.
The policy of the Garden is to build up its collections from wild material of known origin and it
specializes in plants of Europe and the Mediterranean region, together with North America,
temperate South America and Asia. The Garden is a member of I.U.C.N.’s Botanic Gardens
Conservation Co-ordinating Body, an organization set up to encourage exchange of information
between Botanic Gardens and the I.U.C.N. data-base on threatened plants at Kew. The Garden
holds, for example, a number of threatened, narrow endemic plants such as Cretan Date-palm
(Phoenix theophrasti), various arborescent species from the Canary Islands, and Brighamia citrina,
a curious, campanulaceous plant with a woody, bottle-shaped stem, from Hawaii. It is a member of
the National Council for the Conservation of Piants and Gardens National Collections Scheme,
and holds a collection of /ris.
Features of the Garden include a herb bed, planted with culinary herbs that are commonly used
in this country; a variegated plants bed, with a demonstration of variously coloured foliage and
‘true’ variegated plants; a rose border, underplanted with herbaceous and ground-cover plants; a
woodland walk, managed to encourage the native ground flora, including massed bluebells, wood
anemones and celandines, in spring; a fern glade, with a planting of early-flowering woodland
plants such as Helleborus, Galanthus and Polygonatum species, with species of Athyrium,
Cystopteris, Dryopteris, Osmunda, Polystichum and other ferns; and a meadow, down to grass for
many years and cut in late summer. There are many specimen trees, as elsewhere in the University
Park, including a magnificent Cypress Oak (Quercus robur forma fastigiata).
The Plant Science Botanic Garden holds a number of Open Days during spring and summer, but
is also open to interested visitors by arrangement.
V. H. HEywoop
EXCURSION TO STRATFIELD SAYE AND MOOR COPSE, BERKSHIRE, HELD IN CONJUNCTION WITH THE ANNUAL
GENERAL MEETING, 12TH MAY 1985
61 B.S.B.I. members and their guests made up the party for this meeting, which left the Plant
Science Laboratories, University of Reading, by coach at 11.30 hrs. Our morning stop was at
Stratfield Saye, where an area of alkaline water-meadow by the River Loddon supports a fine
population of Fritillaries (Fritillaria meleagris L.), a floristic ‘speciality’ of the Thames Valley. The
meadow clearly shows the pattern of former dykes and drains by which it would have at one time
been flooded during the early part of the year, in order to keep frost off the young grass and to
encourage growth (‘early bite’) on which grazing animals might feed in March and April (whilst
avoiding the toxic and unpalatable F. meleagris). The animals were then removed and a crop of hay
was harvested in July to August, by which time F. meleagris had shed its seeds. On our visit, the
meadow was dotted with the purple, and white, flowers of the fritillaries, an astonishing spectacle
for those who had not seen such a meadow before. Flowering had been held back by cold weather,
giving us a better display than is usual in mid-May. The continuing unseasonal weather gave
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REPORTS 113
overcast conditions which unfortunately acted against the photographers’ interests. Some fine
patches of Ophioglossum vulgatum were admired: this is another characteristic species of these
meadows. The party then moved on to Pangbourne where we sought lunch or ate our sandwiches
near the river. The organizers accept no responsibility for the Sunday closure of the public
conveniences in Pangbourne!
After lunch we drove 2 km to Moor Copse, a woodland reserve of the Berkshire, Buckingham-
shire and Oxfordshire Naturalists’ Trust (BBONT). This reserve is an area of some 27 hectares of
rather wet woodland of standard oaks and ashes, and coppiced alder, ash and hazel, together with a
section of the River Pang, an unpolluted chalk-stream. BBONT, who were fortuitously holding an
Open Day (with associated refreshments, exhibits and parking areas) on our visit, have maintained a
programme of regular coppicing since the reserve was established in 1975. The spring flora was a
magnificent sight, with sheets of Glechoma hederacea, Hyacinthoides non-scriptus, Orchis mascula,
Polygonatum multiflorum and Viola riviniana. For a while the sun shone, but at tea-time a heavy
shower drove us under cover. However, this did not spoil the day, which had been a most pleasant
mixture of the aesthetic and the scientific aspects of our subject. Following our return to Reading,
several enthusiasts carried on their botanical studies in the herbarium and the meeting did not finally
break up until 18.00 hrs.
We should like to express our thanks to Dr H. J. M. Bowen, who smoothed our path in the
Chemistry Department on the Saturday; to Mr D. Farmer, who handled the A.G.M. paperwork on
our Sirius micro-computer and who served refreshments on the Saturday; to Dr D. M. Keith-Lucas
and Mr D. J. N. Hind, BBONT as well as B.S.B.I. members, who acted as our liaison with the Trust
and who handled queries about Moor Copse from members; and, especially, to Mr M. F. Watson,
who helped with refreshments and carried out many of the seemingly trivial but essential jobs that
need be done behind the scenes at an A.G.M.
J. R. AKErRoypD & S. L. Jury
FIELD MEETINGS, 1984
ENGLAND
VALE OF AYLESBURY, BUCKINGHAMSHIRE. 26TH—27TH MAY
Due to the tragic death of Jeremy Milton, Field Secretary, less than two weeks before the meeting,
the arrangements were not finalized and in consequence there was a very poor attendance of only
four members during the weekend. The weather too was miserable, as it started to rain on the Friday
evening and never stopped until we left Waddesdon on Sunday evening. But while getting wetter and
wetter the four recorders were almost overwhelmed by the number of Black Poplars, mostly
pollards, in every tetrad examined; but this small team was able to cover only a fraction of the area
which needs to be surveyed.
In spite of the adverse circumstances the meeting was useful as it confirmed my feelings that the Vale
of Aylesbury is rich in Populus nigra, and I now have the task of trying to discover the reason why.
E. MILNE-REDHEAD
NORTH LINCOLNSHIRE. 17TH JUNE
Two areas were visited on the North Lincolnshire meeting: the National Nature Reserve at
Saltfleetby Theddlethorpe — with a great diversity of coastal sandy and wetland habitats, and the
local County Trust Reserve at Messingham — disused sand quarries with a fine series of inland sandy
and wetland habitats in early stages of colonization. In both reserves the writer, as leader, was aided
by the expertise of two friends, the wardens: Tim Clifford of The Nature Conservancy Council and
Vi Wilkin, Honorary Warden for Messingham. 14 members and friends were present.
Our field secretary, Jeremy Milton, was intending to make this his first Lincolnshire expedition,
and the party were saddened to hear of his tragic death.
The spectacular show of marsh orchids was a striking introduction to the Saltfleetby Reserve, with
Dactylorhiza incarnata, D. praetermissa and D. fuchsii and hybrids. Tim pointed out the new colony
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114 REPORTS
of Epipactis palustris (second site for both vice-counties) and commented on the objectives and
management of the reserve as a whole, including the problems of Natterjack Toads!
The extensive fresh-water marsh at Saltfleetby can be split into quite distinctive areas with
different associations and dominants. In the marsh orchid area other notable plants included Juncus
subnodulosus, Hydrocotyle vulgaris, Ranunculus flammula and Equisetum palustre, and on the drier
parts Ononis repens (not common on the Lincolnshire coast). In the parts where conditions are
brackish, large areas of Juncus maritimus with Juncus gerardi, Samolus valerandi, etc. occur and, in
the deeper, rich, fen-type areas, very extensive stands of Carex riparia and Rumex hydrolapathum,
with Jris pseudacorus, Scutellaria galericulata and Lycopus europaeus are found. (Other notable
plants included Triglochin palustris, Eleocharis palustris, E. uniglumis and, quite out of context, the
newly recorded Eriophorum angustifolium).
The party then crossed the large dune ridge, dodging through dense Hippophaé rhamnoides scrub
to the landward edge of the salt marsh where a new brackish slack is developing. Near the path,
Eleocharis quinqueflora was found and the party spotted some new sites for Dactylorhiza praeter-
missa. Returning on the seaward side, saltmarsh plants included Juncus maritimus, Armeria
maritima, Plantago maritima, Glaux maritima, Triglochin maritima, Limonium vulgare, Scirpus
maritimus and Parapholis strigosa.
After lunch, the party explored the northern edge of the reserve and found Carex extensa (not
common in the east), Blysmus rufus (southern limit) and Thalictrum minus subsp. arenarium. The
flora on the base-rich sand dunes included Rubus caesius, Carlina vulgaris, Anacamptis pyramidalis
and notably Cerastium arvense. The Bee Orchid monitoring site was also examined.
Then, taking leave of Tim and wishing him well on his new appointment, several members drove
across Lincolnshire for a brief visit to the British Industrial Sand Quarries at Messingham, a recently
acquired Lincolnshire Trust Reserve. This reserve supports one of the largest populations of
Pilularia globulifera in Great Britain. This species is colonizing new areas at an amazing rate. At its
original site the species had a very luxuriant growth and a large list of associated plants was made.
Altogether the plant list for the reserve was most extensive and with Vi’s enthusiasm one really
needed another day. Amsinckia lycopsoides, present in quantity last year, was not found but
Anagallis tenella, Isolepis setacea and Samolus valerandi were notable. So also were the large
specimens of Dactylorhiza praetermissa in newly colonized areas.
Perhaps it was ambitious to try to do justice to both these areas in a single meeting but it served as a
comparison of, and as an introduction to, two very rich sandy areas in Lincolnshire. The comments
of the wardens relating to management policies and to the problems of maintaining populations of
individual plant species were invaluable and very worthwhile, and the meeting was enjoyed by all.
I. WESTON
DALBY, YORKSHIRE. 23RD JUNE
24 members took part in the meeting, the object of which was to examine in detail the vegetation of
three fens on the southern edge of the North Yorkshire Moors National Park. Characteristic species
of these fens include Pinguicula vulgaris, Pedicularis palustris, Triglochin palustris , Juncus subnodu-
losus, Schoenus nigricans, Carex hostiana and C. lepidocarpa.
The first site visited at Seive Dale Fen is noted for its extensive population of Epipactis palustris.
Abundant leaves of this plant were noted, but it was too early for flowers. Orchids noted in flower
were Gymnadenia conopsea, Dactylorhiza fuchsii, D. maculata and D. incarnata subsp. pulchella.
At the second site in Sand Dale, Eleocharis quinqueflora and Eriophorum latifolium were noted
and Blysmus compressus was recorded for the first time. D. fuchsii and D. maculata were seen again,
but the main feature of the site was the extensive population of D. traunsteineri. A large population
of Cirsium dissectum was present in Molinia grassland adjacent to the fen, while dry ridges between
the springs supported an unusual community which included Calluna vulgaris, Carlina vulgaris, Salix
repens and Antennaria dioica.
In the afternoon, the third site, Haggs Wood Marsh, adjacent to Newtondale was visited. Here,
both Eriophorum latifolium and E. angustifolium were noted while Myrica gale, Menyanthes
trifoliata, Serratula tinctoria were all added to the list of species seen during the day. The main feature
of this fen is however the spectacular show of Trollius europaeus which was in full bloom at the time
of our visit.
REPORTS 115
Our thanks are due to the Forestry Commission, Mr R. Boyes and the Yorkshire Wildlife Trust
for permitting us to visit their land.
P. J. HorTOoN
NEW FOREST, HAMPSHIRE. 4TH AUGUST
This was a duplicate of the meeting held on 3rd September 1983 and which had been massively
oversubscribed. 21 members took part. Although the meeting was held a month earlier and in a
drought year, all the same plants were again recorded except for Spiranthes spiralis. Hammarbya
paludosa was found in the same bog but about 50 m away from the 1983 plants. (See Watsonia, 15:
297—298(1985)).
J. OUNSTED
WALES
TINTERN, MONMOUTH. IST—3RD JUNE
A small group of nine members met for this residential weekend at the Nurtons Field Centre in the
lovely wooded setting of the Lower Wye Valley. The object of the meeting was not to look for rare
or unusual plants. Quite the reverse in fact, as it was to concentrate on our common ‘weeds’ and
consider their relationship to modern vegetables, and in the evenings to sample their much
neglected edibility. The site chosen for this was the 3 ha of organic market garden of the 12 ha
holding. An area that has had no artificial sprays used on it in the last 20 years, this garden enjoys a
diverse community of both common and less common wild plants, as well as a large range of
cultivated vegetables. The regime of gardening allows all species a place to grow with and
alongside the crops, and members were impressed with the obvious health and abundance of the
crop plants. How many of our threatened species would grow freely if more farmland were like
this? The herb garden also provides a home for some of our disappearing favourites such as Woad
and Elecampane, as well as plants like Tansy, Sweet Cicely and Fennel.
At mealtimes we tasted the delights of classic dishes like Nettle soup garnished with Ramson
flowers, and of almost forgotten country greens — Fat Hen, Hop shoots, Hogweed sprouts, Ground
Elder and creamed Chickweed. A wild salad included White Dead Nettle, Yarrow, Salad Burnet,
garlic mustard and Dandelion leaves, and a dessert of Strawberries was scattered with Elder flowers.
Several members had come from the east of England, so we included in the weekend outings to
show something of this lovely area. Llywyn-y-celyn, a very compact calcareous marsh, was visited
on Saturday. This site is a Gwent nature reserve, with a rich sedge flora including Carex hostiana.
Menyanthes trifoliata and abundant Valeriana dioica were in flower, and fine Aconitum napellus
was found alongside the boundary stream. Locally common along the Welsh Marches, this plant
contains powerful alkaloids still used in equine and human medicine. On Sunday, the venue was
the area of magnificent limestone crags and woodland at Black Cliff near Chepstow. These broad-
leaved woodlands, still with some Ulmus glabra among Beech, Ash and Hazel, contain a high
percentage of ancient woodland indicator species including Paris quadrifolia in great quantity, and
Cardamine impatiens on ground disturbed by forestry work. It is possible to see Convallaria
majalis, Carex digitata and Melica nutans, together with the elusive Carex strigosa nearby, and
hybrids between Polystichum aculeatum and P. setiferum; and of course Euphorbia serrulata, the
Tintern Spurge, was visited. The outing ended with a short visit to Poors Allotment on the edge of
the Forest of Dean, a heathy area which shows a particularly clear geological division. A sudden
transition from sandstone to limestone makes it possible to stand with one foot by Carex binervis
and one by C. montana, and similarly with Nardus stricta and Helianthemum nummularium.
These outings were enjoyable, and of course did pay tribute to the botanical excellence of this
part of the world, and to its spectacular scenery, but both leaders wished that the weekend could
have been twice as long, so that the original subject of the relationship of common plants to man
could have been worked on in more depth, and more practical experience of their medicinal,
cosmetic and edible properties achieved. It would be very pleasing to both of us if we could take
these subjects further with B.S.B.I. members in the future.
R. FITZGERALD & E. Woop
116 REPORTS
LLANARMON-YN-IAL, DENBIGHSHIRE/FLINTSHIRE. 16TH JUNE
We met at Coed Talon to look at a sheltered valley in which coal is close to the surface. There were
no very rare plants, but different habitats were studied. The deciduous wood contained
Lamiastrum galeobdolon, Galium odoratum and Prunus padus. Recently felled areas were
colonized by Luzula pilosa, Veronica officinalis and Trifolium medium. In the marshy floor of the
valley were Carex acutiformis, Typha latifolia and Riccia fluitans (an aquatic liverwort). The dry
slopes yielded Blackstonia perfoliata, Phleum pratense subsp. bertolonii and Carex spp. (C. ovalis,
C. flacca. C. panicea, C. sylvatica, C. hirta, and C. demissa). At lunchtime, some of the party
visited the site at which Senecio cambrensis was first collected and saw plants in flower.
The afternoon was spent on meadow slopes with limestone outcrops near Eryrys. Numerous
plants of Coeloglossum viride were found. Three Alchemilla spp. (A. glabra, A. xanthochlora, and
A. filicaulis subsp. vestita) were growing together with Poa subcaerulea and Geum rivale. On the
cliffs were Vigla hirta, Cotoneaster microphyllus, Rosa pimpinellifolia, Cystopteris fragilis,
Saxifraga tridactylites and Scabiosa columbaria.
We searched an upland meadow unsuccessfully for Antennaria dioica which had been seen some
years before. However, we found many spikes of Botrychium lunaria and Gymnadenia conopsea,
and Polygala vulgaris was abundant in pale and dark blue, pink and white forms.
17 people attended, including members from South Wales and Wolverhampton.
J. A. GREEN & G. WYNNE
“
SCOTLAND
CREAG MEAGHAIDH, INVERNESSHIRE. 14TH-15TH JULY
The annual joint meeting of the Alpine Section of the Botanical Society of Edinburgh and the
Botanical Society of the British Isles attracted 23 people.
On the Saturday, we visited Coire Ardair, approaching by the path from Aberarder Farm. This
good path passed through some nice birch wood and open undulating moorland. A fine stand of
the three orchids Pseudorchis albida, Gymnadenia conopsea and Dactylorhiza maculata subsp.
ericetorum was found close to the path.
The large east-facing corrie still held a considerable amount of snow in the gullies and with mist
and occasional rain the impression of a wild and desolate place was obtained. A number of rare
plants was found including Gnaphalium norvegicum, which was almost in flower although most
plants were in a vegetative state. Flushes near the corrie contained Juncus castaneus, J. triglumis
and Carex saxatilis. The Holly Fern, Polystichum lonchitis could be found with the Parsely Fern,
Cryptogramma crispa as intimate ledge and crevice companions; this seemed unusual as they
normally prefer quite different soils and rock types. Athyrium distentifolium was probably the most
abundant fern. Other plants of note included Carex vaginata, Cerastium alpinum, Coeloglossum
viride, Galium boreale, Poa glauca, Saussurea alpina, Saxifraga aizoides, S. hypnoides, S.
oppositifolia, Sibbaldia procumbens, Silene acaulis, S. dioica, Trollius europaeus and Veronica
alpina.
Moy Lodge was the meeting place on the Sunday and the party soon split in two, one group
staying low, trying to locate an old record of Paris quadrifolia, but this they failed to find. They did
however find Hammarbya paludosa, Eriophorum latifolium, Drosera anglica and a wide range of
sedges in the bogs. Amongst birch scrub, rock outcrops and boulders they found Avenula pratensis
and Melica nutans.
The second party walked up to Moy Coire finding Cornus suecica, Parnassia palustris, Rubus
chamaemorus and some very free-flowering Vaccinium uliginosum on the moor. The Moy Coire is
a big horseshoe of rocks and cliffs encircling quite a large lochan. The chief finds here included
Alopecurus alpinus, Carex atrata, Cerastium cerastoides, Epilobium alsinifolium, Juncus castaneus,
J. triglumis, Phleum alpinum, Poa alpina, Polystichum lonchitis, Pyrola minor, Salix lapponum
and Tofieldia pusilla.
Alpine hawkweeds were much in evidence in both of the corries but no names are available.
R. J. D. MCBEATH
tis
ee et
REPORTS 7
IRELAND
ST JOHN’S WOOD, ROSCOMMON. 19TH MAY
St John’s Wood, on the western shore of Lough Ree, is one of the last native woods of any size in
the Irish midlands today. It is listed as a site of International Importance by An Foras Forbatha
(Areas of Scientific Interest in Ireland, 1981) — the only such site in Co. Roscommon. Ownership is
divided between the Forest and Wildlife Service and a number of private owners. The wood is
extensive (150 ha), and yet, apart from a few conifers and Sycamores in the western part, it is
composed entirely of native species.
The day threatened rain; it fell intermittently, but not enough to deter us. The party was about
15, with an unusually wide range of ages; it was led by J. Early (county recorder) and D. L. Kelly.
The junior members of the Northridge family proved adept at spotting Lathraea squamaria;
several colonies were located. Neottia nidus-avis, Ranunculus auricomus, Geum rivale X urbanum
and Dryopteris carthusiana were added to a list compiled from previous visits.
The major part of the wood is on shallow, well-drained soil over limestone. The woodland
canopy is unusually diverse, with plentiful Quercus robur, Fraxinus excelsior, Ulmus glabra,
Corylus avellana and Ilex aquifolium. A few Taxus baccata and Prunus avium were found, and
some fine trees of Sorbus hibernica. Sadly, there was much evidence of recent felling, especially of
oak, so that parts of the wood have been reduced to scrub. An area of several hectares had recently
been clear-felled. We made our way down to the rocky lakeshore, where we peered inconclusively
at Salix. Trying to get back to our cars, we found ourselves wandering in a real-life ‘Babes in the
Wood’. I draw two morals: a) this is a wood to be prized — one of the very few native woods left in
Ireland that is large enough to get lost in; b) be wary of getting separated from your lunch!
After our hard-earned repast, we proceeded to the adjacent wetland alongside Blackbrink Bay.
The fen flora included such characteristic midlands species as Cicuta virosa, Berula erecta,
Ranunculus lingua and Thalictrum flavum. Pin-eyed and thrum-eyed Menyanthes trifoliata were
admired (a trick not confined to Primula). The area was a complex of cut-away peats, with a series
of different mire types. Higher peaty ground supported Osmunda regalis, Prunus padus and
Frangula alnus. Crossing back into the adjacent part of St John’s Wood, we found a remarkable
swamp woodland community with F. alnus abundant in the understorey, accompanied by
Rhamunus catharticus and Prunus padus. F. alnus is rare in Ireland, but Tansley noted that the two
buckthorns may dominate young fen carr in East Anglia.
We left convinced that this is a very remarkable wood, deserving the highest priority for active
conservation; letters to this effect have been sent to the Chairman of Roscommon County Council
and to the Minister of Fisheries and Forestry.
Dil? Kerry
FRANCE
NORMANDY. 3RD—10TH JULY
Some 20 members attended this meeting in the part of France which most resembles southern
England in both flora and scenery, in beautiful sunny weather throughout the week. On 3rd July
the party crossed the Straits of Dover from Folkestone to Boulogne, and drove south to Rouen,
where we were to stay the first three nights. The opportunity was taken to spend some hours en
route in examining two of the magnificent and extensive calcareous fens near the coast of the Pas-
de-Calais at Cucq (Marais de Villiers) and at Merlimont (Marais de Balancon). These fens, among
the best left in western Europe, produced good colonies of Liparis loeselii (the large East Anglian
variant) and also Eriophorum gracile, Carex diandra, C. lasiocarpa, C. lepidocarpa, C. elata,
Schoenus nigricans, Cladium mariscus, Ranunculus lingua, Pedicularis palustris, Utricularia
australis, Orchis laxiflora subsp. palustris (unknown in the British Isles), Dactylorhiza trauns-
teineri, D. praetermissa, and D. incarnata subsp. ochroleuca and subsp. pulchella, besides the type;
this list however is merely a selection of the most interesting species seen in these fens.
On 4th July, we travelled south from Rouen to study the famous chalk slopes and cliffs of the
gorge of the River Seine near Orival. Among the enormous number of chalk grassland species seen
118 REPORTS
were many not known in the British Isles as natives, such as Gymnadenia odoratissima, Stachys recta,
Teucrium montanum, Globularia punctata, Digitalis lutea, Vincetoxicum hirundinaria, Bupleurum
falcatum, Linum tenuifolium, Euphorbia seguierana, Amelanchier ovalis, Prunus mahaleb, and
Koeleria pyramidata. These species were seen repeatedly in other similar sites. They have got as far
as Normandy, but have, for reasons not always clear, been unable to establish themselves on our side
of the Channel; presumably these steep south-facing chalk slopes are just a little warmer than any
sites we possess. In addition, many rare British calcicole species occurred, mostly in abundance, such
as Pulsatilla vulgaris, Phyteuma tenerum, Helianthemum apenninum, Carex humilis, Seseli libanotis,
Geranium sanguineum, Teucrium chamaedrys, Orobanche caryophyllacea, Himantoglossum
hircinum, Ophrys fuciflora, and Epipactis atrorubens, among dominant Sesleria albicans.
Orobanche species such as O. gracilis and O. teucrii also excited admiration. Other nearby chalk
cliffs and screes, at the Roche St Adrien, added Viola hispida, a beautiful Norman endemic, and
Helianthemum canum. The chalk downs of the Eure valley further south had a similar flora, with the
addition of Fumana procumbens, Astragalus monspessulanus (at the extreme northern limit for this
largely Mediterranean species), Coronilla minima, Anthericum ramosum, Ononis natrix, and O.
pusilla, besides the more familiar Rubia peregrina, and Orchis purpurea (in fruit). The Forét de Bray
on the Lower Greensand yielded a bog rich in Vaccinium oxycoccos, and the Forét d’Eu, again on
chalk, had Epipactis leptochila and Hordelymus europaeus in beech forest, besides Herminium
monorchis and much else that occurs on our southern chalk downs.
On 6th July we drove west to Carteret in the Cotentin peninsula by Bayeux (where the famous
Tapestry was inspected) to a very different type of country on ancient, acidic rocks, more like
Devonshire. The sand dunes here produced Bupleurum baldense, and Limonium auriculae-
ursifolium and Frankenia laevis on the saltmarsh edge; the more extensive dunes at Vauville to the
north were bright in places with Veronica spicata among Thesium humifusum, while the slacks had
Mentha pulegium, Teucrium scordium, Parentucellia viscosa, Carex punctata and Cyperus longus,
but some species were very scorched by the drought.
The coastline east of Cherbourg produced Otanthus maritimus, Lagurus ovatus, Euphorbia
peplis, Orobanche purpurea, Polycarpon tetraphyllum and Natterjack toads, besides much
Eryngium maritimum and E. campestre with Orobanche amethystea as their parasite. West of
Cherbourg, Centaurium scilloides was locally plentiful on the cliffs. Inland, a rocky oakwood at
Mesnil-au-Val had much Hymenophyllum tunbrigense, Dryopteris aemula, Sibthorpia europaea,
and Wahlenbergia hederacea on rocks and in flushes.
Perhaps the highlight of the week for many people, however, was the wet heathland soith of the
fine Norman Abbey of Lessay, where Spiranthes aestivalis still occurs as about 600-—700 individuals
in perhaps its finest remaining European site, together with Rhynchospora fusca, Lobelia urens,
Carum verticillatum, Galium debile, and Deschampsia setacea. Nearby, Gentiana pneumonanthe
(not in flower) and bogs with all three Drosera species and Narthecium ossifragum were seen in a
Nature Reserve at La Tourbiére de Mathon.
On our last day we visited the great cliffs of Cap de Flamanville, where Inula crithmoides,
Asplenium billotii and Limonium binervosum were seen, but Lotus subbiflorus and Juncus
capitatus, known here previously, seemed to have failed this summer due to drought.
Those who attended this excursion were left with the realization that though our own flora is
very interesting, it is only a pale shadow of the richness of that on the European continent, even as
near as Normandy.
F. ROSE
SWITZERLAND
LAUTERBRUNNEN. 25TH JUNE—2ND JULY
13 members assembled at Gatwick Airport for the flight to Ziirich, whence transfer to
Lauterbrunnen was made by rail via Bern and Interlaken. The party was based in self-catering
chalet accommodation just outside Lauterbrunnen, which lies about 12 km due south of Interlaken
in the heart of the Bernese Oberland. During the week, six highly successful field excursions were
made, each followed by lively evening identification and discussion sessions.
|
REPORTS 119
On the first morning the party travelled by rail through the awe-inspiring scenery of the
Lauterbrunnental up to Wengen and on to Kleine Scheidegg (2061m) for a walk through Alpiglen
to Grindelwald. Above us towered the lofty peaks of the Jungfrau (4158m), M6énch (4099m) and
the Eiger (3970m). After poor spring weather in the Alps, the season was fully three to four weeks
retarded and much more snow than is usual in late June lingered on in the vicinity of Kleine
Scheidegg, providing a fine opportunity to see Soldanella alpina and Crocus vernus subsp.
albiflorus, which flower as soon as the snow recedes (and often beneath it). Ranunculus
aconitifolius formed large white areas on the mountainsides and Caltha palustris vividly marked the
water courses. Within a few hundred metres of the village many members of our own impoverished
alpine flora were encountered, often in huge quantity, and on a well-worn pathside several plants
of the tiny Gagea fistulosa were located. On acid formations Gentiana acaulis, Homogyne alpina,
Potentilla aurea, Bartsia alpina, yellow-flowered Pulsatilla alpina subsp. apiifolia, Trifolium
alpinum, Luzula lutea and Loiseleuria procumbens were noted, with Gentiana verna, white-
flowered Pulsatilla alpina subsp. alpina, Silene acaulis, Androsace chamaejasme, Viola calcarata,
Primula farinosa and Plantago atrata on the outcropping limestone. The party progressed slowly
down through Alpiglen with numerous stops being made as members familiarized themselves with
the rich flora. Ranunculus alpestris, Globularia nudicaulis, Hutchinsia alpina and Tofieldia
calyculata adorned a stream-side; nearby Oxytropis jacquinii, Kernera saxatilis, Sedum atratum and
Biscutella laevigata were growing on bare stony ground with much Primula auricula on limestone.
Fine specimens of Cystopteris montana, Polystichum lonchitis and Gymnocarpium robertianum
abounded on the rock-strewn slopes. Richard Thomas, the party’s sedge expert, pointed out Carex
rupestris growing with Arctostaphylos alpinus; C. firma, C. ferruginea and C. sempervirens were
noted here (and throughout the week) but C. davalliana was found only on this day. A small patch
of the very local Moehringia ciliata beside the track was accompanied by tiny Veronica aphylla. The
wooded slopes of lower altitudes were rich in Moneses uniflora, Actaea spicata, Maianthemum
bifolium, Streptopus amplexifolius and sedges Carex digitata and C. ornithopoda.
The second morning took us to the summit of Schilthorn (2970m), widely known as the location
for the James Bond film ‘““On Her Mayjesty’s Secret Service’. Despite the lateness of the season,
Ranunculus glacialis was flowering well on crumbling rock surfaces, as were Saxifraga oppositifolia
and dense clumps of Androsace alpina and A. helvetica. Unfortunately, Geum reptans and
Artemisia genipi were yet to open. Few plant species are able to withstand the harsh conditions at
these altitudes and, not surprisingly, plant cover is probably as low as 5%. The considerably
reduced atmospheric pressure soon took its toll — the party becoming notably sluggish. Extensive
late snow fields made the planned descent through the Blumental to Mirren impossible, return to
Mirren by cable car followed by a leisurely walk down to Lauterbrunnen being substituted. Acinos
alpinus (in Mirren itself) and Veronica fruticans provided some compensation, however, and the
railway line to Grutschalp proved a fine hunting ground with Ervinus alpinus a notable feature.
Rhododendron ferrugineum was admired and photographed before the very steep descent to base
was undertaken. Cicerbita alpina, Stachys alpina and Lilium martagon were disappointingly only in
bud but a wealth of Polygonatum verticillatum, Stellaria nemorum and Saxifraga rotundifolia was in
flower.
A link-up with Mary Briggs’ Cox and King party on the next day preceded a coach drive to
Kandersteg for a leisurely perambulation of the unforgettable Gasterental. Led by Max Seiler, of
the Basle Natural History Society, the excursion was of exceptional botanical interest and,
moreover, after just half an hour’s steep climb, traversed almost level ground throughout the day!
Although only about 30 km west of the Lauterbrunnental, this beautiful valley supports a
somewhat different flora with Clematis alpina (scrambling over juniper), Amelanchier ovalis,
Aquilegia alpina and Cypripedium calceolus forming the highlights. Limestone scree yielded the
umbellifer Athamanta cretensis, Dryas octopetala, abundant Moehringia muscosa, a wealth of ferns
and local Senecio doronicum; in wet meadows, Pedicularis verticillata and Pseudorchis albida
abounded together with occasional plants of striking Pedicularis foliosa.
The lure of glaciers drew us to Obergletscher and its environs on the Thursday, reached by train
to Grindelwald and a 45-minute walk. The air temperature close to the glacier was considerably
depressed and, as a result, the vegetation was much retarded. Most noticeable here were
Epilobium fleischeri, its extensive roots binding moraine, and Gypsophila repens flowering on bare
glacial debris. Dense coniferous woodland supported fine colonies of Moneses uniflora,
120 REPORTS
Corallorhiza trifida and Orthilia secunda together with abundant Bellis sylvestris. Away from the
glacier, river shingle was, as always, a botanical paradise with a great many plants brought down
from higher altitudes; with Arabis alpina, Linaria alpina, Aconitum napellus, Epilobium collinum,
and grasses including Phleum alpinum and Poa alpina were Salix daphnoides, S. helvetica, Ribes
petraeum and much more besides. Carex flava and C. capillaris were noted amongst many
interesting species, including Arabis glabra, growing on the banks of a small stream, and adjoining
coniferous woodland had Listera cordata and several small populations of Cypripedium calceolus.
With the latter was Luzula luzulina, at first mistaken for L. forsteri which it much resembles. This
area has a very rich flora and is highly recommended, especially to those unable to climb. On the
return from Grindelwald, some members could not resist a look at a railway siding at
Zweilitschinen during a wait for a connection: here, on ballast, was a large colony of Astragalus
alpinus in flower and fruit, with Bromus inermis and Calamagrostis varia.
The fifth day, spent entirely in the Lauterbrunnental, proved to be the best of the week
weatherwise. The post-bus was taken to Stechelberg and the party walked leisurely back to
Lauterbrunnen. During a brief stop at the cable-car station, a tuft of Saxifraga cespitosa was
spotted on the river gravel ballast covering the long-term car park. Within a short while, upwards
of 20 species of alpines, largely in flower, had been located and identified; these included
Campanula cochlearifolia, Dianthus sylvestris, Erigeron glabratus, Artemisia genipi, A. umbellifor-
mis, Agrostis alpina, Globularia cordifolia, Verbascum lychnitis (the yellow-flowered form) and
Leontopodium alpinum, the last flowering some two months earlier than usual. A remarkable
assemblage of introductions indeed and at the low altitude of 868m! The walk continued along the
river and through adjacent broad-leaved woodland where Cardamine pentaphvllos in fruit, Senecio
nemorensis, Aconitum vulparia, Peucedanum ostruthium, Aruncus dioicus and Impatiens noli-
tangere were recorded. A picnic lunch was followed by a visit to the Triimmelbach Falls, the most
spectacular underground waterfalls in Europe. On the rocks outside were Luzula nivea,
Laserpitium siler and Valeriana montana. Nowhere can the richness of sub-alpine meadows be
more striking than in this valley — nor provide a starker contrast with our own ‘monocultures’.
After a short break, we continued beyond Lauterbrunnen to explore a Cephalanthera rubra site
near the falls on the Isenfluh road; here also were Epipactis atrorubens, Digitalis lutea, D.
grandiflora, Teucrium montanum and Saxifraga paniculata.
The next day saw an early departure for the Jungfraujoch. The party split at Eigergletscher
station (2320m), some continuing to the observatory above and others alighting to botanize.
Reunited at noon, we explored the ridges above Eigergletscher where Lloydia serotina, Draba
aizoides, Saxifraga moschata, Trifolium thalii, Hedysarum hedysaroides, Bupleurum stellatum and
diminutive Antennaria carpatica were in flower on alkaline soils, with Primula hirsuta and
Omalotheca supina on acid substrates. On moraine Doronicum grandiflorum, Achillea atrata,
Cirsium spinosissimum, Galium helveticum and Linaria alpina, in a variety of colour forms, were
noted. Towards Kleine Scheidegg the early flowers of the meadows were at their best with
Pulsatilla vernalis, Anemone narcissiflora, Minuartia verna, Carex firma, Gentiana verna, G.
brachyphylla, G. clusii (including a white-flowered form), abundant Primula auricula and P.
farinosa. Several scarlet and other variously coloured primulas probably represented hybrids
between P. auricula and P. hirsuta. Ibex (Capra hirca) grazing these slopes were not in the least
shy and were much photographed. The walk ended at Wengernalp after a concerted, and
eventually successful, search for Gentiana bavarica, the least common of the three local members
of the G. verna group.
On the last day, prior to the homeward journey, members went their separate ways. Two further
finds of note were Tolpis staticifolia and Anthericum ramosum, both on limestone scree near the
village.
This account would not be complete without thanks to Messrs Milton’s Travel of Bristol and the
proprietors of ‘Camping Jungfrau’, the family von Allmen-Jossi, for their assistance with travel
arrangements; and to Messrs A. O. Chater, R. W. David and E. J. Clement for taxonomic
assistance.
A. L. GRENFELL
INSTRUCTIONS TO CONTRIBUTORS
Papers and Short Notes concerning the systematics and distribution of British and European
vascular plants as well as topics of a more general character are invited.
Manuscripts must be submitted in duplicate, typewritten on one side of the paper only, with wide
margins and double-spaced throughout. They should follow recent issues of Watsonia in all matters
of format, including abstracts, headings, tables, keys, figures, references and appendices. Note
particularly use of capitals and italics. Only underline where italics are required.
Tables, appendices and captions to figures should be typed on separate sheets and attached at the
end of the manuscript. Names of periodicals in the references should be abbreviated as in the World
list of scientific periodicals, and herbaria as in Kent & Allen’s British and Irish herbaria. Line
drawings should be in Indian ink on good quality white card, blue-lined graph paper or tracing paper.
They should be drawn at least twice the final size and they will normally occupy the full width of the
page. Lettering should be done in Lettraset or by high-quality stencilling, though graph axes and
other more extensive labelling are best done in pencil and left to the printer. Photographs can be
accepted only in exceptional cases.
Contributors are strongly advised to consult the editors before submission in any cases of doubt. 25
offprints are given free to authors of papers and Short 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 articles.
Papers and Short Notes should be sent to Dr R. J. Gornall, Botany Department, The University,
Leicester, LE] 7RH. Books for review should be sent to Dr N. K. B. Robson, Botany Department,
British Museum (Natural History), Cromwell Road, London, SW7 5BD. Plant records should be sent
to the appropriate vice-county recorders. Reports of field meetings should be sent to Dr B. S. Rushton,
Biology Department, The University of Ulster, Coleraine, Co. Londonderry, Ireland.
Insects and Flowers
The Biology of a Partnership
Friedrich G. Barth
Translated by M.A. Biederman-Thorson
Includes forty-seven photographs in full colour, 16 full page
photographs made with an electron microscope and around
one hundred line drawings.
A superb new volume for botanists and zoologists alike.
Hardback £24.95
Allen & Unwin
Watsonia
January 1986 Volume sixteen Part one
Contents
Simpson, D. A. Taxonomy of Elodea Michx in the British Isles ... ts Ci
Crawrorp, T. J. & Jones, D. A. Variation in the colour of the keel petals in
Lotus corniculatus L., 2. Clines in Yorkshire and adjacent counties 15-19 . :
KADEREIT, Tow. & SELL, P. D. Variation in Senecio oe Li
(Asteraceae) in the British Isles... so ae i
WEBSTER, S. D. Two natural hybrids in ay. C subgenus aaching
(DC) A, Gray oye es oe Re
SARGENT, C., MOUNTFORD, O. & Goeae D. The dichibetion of Poa
angustifolia L. in Britain . bgp & a - a 31-36-/
Trist, P.J.O. A co iuadaeean of the taxonomic status of Poa balou ‘
.o Be
Parnell (Gramineae) a a a 37-42
Trist, P. J. O. The distribution, ecology, history a status Lai basnidiins
ventricosum (Gouan) Schinz & Thell. in the British Isles _... beg 43-54
BLAcKSsTocK, T. H. & Roberts, R. H. Observations on the morphology and
fertility of Juncus X ew Druce ex Stace & Lambinon in north- Ro
western Wales __... es fie 55-63
Dickson, J. H. James eho oad ne eee of Bue, 1768 oh 65-74 |
SHorT NOTES
Atkinson, M. D. & Codling, A. N. A reliable method for eee
between Betula pendula and B. pubescens _... 75-76
Cannon, J. F. M. More about sea- and lakeballs_... ie 76-77
Crackles, F. E. Dactylorhiza majalis (Reichb.) P. F. [inate Satnnievhages
subsp. cambrensis (R. H. Roberts) R. H. Roberts in $.B. Yorkshire 78-80
Leach, S. J. The rediscovery of Carex maritima Gunn. on the oy at St
Andrews links, Fife Bt 80-81
Matcham, H. W. & oe M. Flag pyramidata Le, seteitad in West
Sussex a ae y ee: 81-82
Parnell, J. A. N. Cacmeeene rignbers of Irish ae Li ae poe 82-83
Preston, C. D. An additional criterion for assessing native status ik 83
Rich, T. C. G. & Baecker, M. The distribution of Sorbus lancastriensis
E. F. Warburg — ... i 4 FA Ps 83-85
Tennant, D. J. Further discoveries of. Hieracium hackhonsel a 1, rails 86-87
Book REVIEWS ... Dy, Ma v5 Nt + i 7m ... 89-103
OBITUARIES i “ HS ey ie ne a He ... 105-108
REPORTS
Annual General Meeting, 11th May 1985... a Hiss f; ... 109-113
Field Meetings, 1984... Nd we i i ay io jah ede ZQ
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4 ft i gee
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Watsonia, 16, 121-129 (1986) a \ JAN 1 () 1990 121
The genus Cicerbita Wallr. in the British Isles
P. DaSELL
Botany School, Downing Street, Cambridge, CB2 3EA
ABSTRACT
Four species of Cicerbita Wallr., the native C. alpina (L.) Wallr. and the introduced C. macrophylla (Willd.)
Wallr. subsp. uralensis (Rouy) P. D. Sell, C. plumieri (L.) Kirschleger and C. bourgaei (Boiss.) Beauverd, are
recorded in the British Isles. A key to the species, citation of nomenclature, descriptions and an account of
their distributions are given.
INTRODUCTION
The genus Cicerbita Wallr. belongs to the family Asteraceae (Compositae) subfamily Lactucoideae
tribe Lactuceae. It consists of about 18 species native to the mountains of Europe, Asia, North
Africa and North America. They are perennial, rhizomatous or tap-rooted herbs with erect stems,
alternate, deeply divided leaves with a semiamplexicaul or more or less auriculate base and an
inflorescence of bluish, rarely yellow, 5—30-flowered capitula. The involucral bracts are in three or
more series and the achenes are linear or elliptic with a pappu' _—_1 two rows of hairs, the outer ring
consisting of very short, smooth hairs and the inner ring of « ng, scabrous hairs.
In the character of the pappus, Cicerbita is closely allied to Mycelis Cass., Cephalorrhynchus
Boiss. and Steptorhamphus Bunge. Mycelis is distinguished by its fewer involucral bracts, which
are in two series, the outer very inconspicuous, and by its yellow flowers. Cephalorrhynchus and .
Steptorhamphus are distinguished by the long beak to the achene. In Cicerbita the beak is absent or
not more than 0-5 mm. The large genus Lactuca L., in which all these small genera including
Cicerbita were once included, is distinguished by its pappus hairs all being of one kind and more or
less equal in length.
Only one species, C. alpina (L.) Wallr., is native in the British Isles, but three others, C.
macrophylla (Willd.) Wallr. subsp. uralensis (Rouy) P. D. Sell, C. plumieri (L.) Kirschleger and C.
bourgaei (Boiss.) Beauverd, have been recorded as naturalized.
KEY TO SPECIES
ees ARO ADEOUS fo esse eee vias sas asa ageeee etches Peenaas ison tontndecetedsmaubcatenes 3. C. plumieri
1. Peduncles and/or upper part of stem with glandular or simple hairs ........ pes
2. Stems and sometimes peduncles with rather rigid simple hairs, without
SlamubwlarirarS Ayo 12) LTA META A Ae aR lee Se SE CASS 4. C. bourgaei
2. At least the peduncles and usually also the upper part of the stem and
involuete, clandularhainy 2505 4 WS Se FR, 3
3. Lower leaves glabrous, with a triangular terminal lobe and few pairs of
small lateral lobes; capitula in an elongated panicle; achenes linear ..... 1. C. alpina
3. Lower leaves hairy on veins beneath, with a cordate terminal lobe and
usually only a single pair of lateral lobes; panicle wider, more or less
corymbose; achenes narrowly elliptical’ ...2). 2650).4o. 0. es 2. C. macrophylla
122 PD; SEEL
ACCOUNT OF SPECIES
1. C. ALPINA (L.) Wallr., Sched. Crit. 434 (1822).
Sonchus alpinus L., Sp. Pl. 794 (1753).
Mulgedium alpinum (L.) Less., Syn. Gen. Comp. 142 (1832).
Sonchus caeruleus Sm., Fl. Brit., 2: 815 (1800) nom. illegit. superfl. pro Soncho alpino L.
Lactuca alpina (L.) A. Gray, Syn. Fl. N. Amer., 1(2): 444 (1884).
Vernacular Name: Alpine Blue Sow-thistle
Illustrations: Ross-Craig, Drawings Brit. Pl., 18: pl. 35 (1963). Streeter, Wild FI. Brit. Is. 186, pl.
88 (1983).
Stem 50-250 cm, simple or branched, usually with dense, reddish glandular hairs on the upper
part. Leaves 8-25 x 2-12 cm, glabrous, glaucous beneath; lowest lyrate or runcinate-pinnatifid
with a large, broadly triangular, acuminate terminal lobe and a few pairs of much smaller
triangular lateral ones, with the base narrowed into a winged petiole; upper smaller and less
divided, with a winged petiole widened into a cordate-amplexicaul base. Capitula in an elongated
panicle; peduncles with dense reddish glandular hairs. Involucre 10-15 x 7-10 mm; bracts linear,
usually with numerous reddish glandular hairs. Ligules pale blue. Achenes 4.5—5 mm, linear.
C. alpina is a very rare plant in the British Isles, being confined to remote gullies and cliff ledges in
the mountains of Angus (v.c. 90) and Aberdeen (v.c. 92). Its past and present distribution, habitat
and general ecology and conservation are discussed in detail in the following paper in this journal
by Marren et al. (1986). In continental Europe it is locally abundant in Fennoscandia and the
mountains southwards to the Pyrenees, northern Apennini and Bulgaria.
2. C. MACROPHYLLA (Willd.) Wallr., Sched. Crit. 434 (1822).
Sonchus canadensis Froelich in Ann. Bot. (Usteri), 1: 29 (1791), non L., Sp. Pl. 793 (1753).
Sonchus macrophylla Willd. in L., Sp. Pl., 4th ed., 3(3): 1519 (1803).
Mulgedium macrophyllum (Willd.) DC., Prodr., 7(1): 248 (1838).
Lactuca macrophylla (Willd.) A. Gray, Syn. Fl. N. Amer., 1(2): 444 (1844).
subsp. URALENSIS (Rouy) P. D. Sell in Bot. J. Linn. Soc., 71: 249 (1976).
Sonchus hispidus Ledeb., Fl. Altaica, 4: 140 (1833), non Gilib., Fl. Lithuan., 1: 241 (1781) non rite
publ., non C. hispida (Bieb.) Beauverd in Bull. Soc. Bot. Genév. sér. 2, 2: 141 (1910).
Mulgedium hispidum (Ledeb.) DC., Prodr., 7(1): 250 (1838).
Mulgedium macrophyllum var. hispidum (Ledeb.) Korsh in Mem. Acad. Sci. Petersb. (Sci. Phys.
Math.) ser. 8, 1: 265 (1898).
Mulgedium uralensis Rouy, Ill. Pl. Eur. Rar., 16: 128, t. ccexc (1901).
C. uralensis (Rouy) Beauverd in Bull. Soc. Bot. Genév. ser. 2, 2: 123 (1910).
C. gmelinii Beauverd in Bull. Soc. Bot. Geneév. ser. 2, 2: 123 (1910) nom. nov. pro Soncho hispido
Ledeb., non C. hispida (Bieb.) Beauverd in Bull. Soc. Bot. Geneév. sér. 2, 2: 141 (1910).
Vernacular Name: Common Blue Sow-thistle
Illustration: Fig. 1.
Perennial with pale brown fibrous roots and long whitish rhizomes which send up flowering stems at
regular intervals. Stems 60-200 cm, pale green, rather slender, slightly angular, glabrous below with
an occasional very long, slender glandular hair in upper part. Leaves up to twelve, all cauline with up
to three sometimes close together near the base, becoming gradually smaller up the stem, rather
dull yellowish-green, glabrous or nearly so on upper surface, slightly glaucous with prominent
venation on lower surface and with numerous simple eglandular hairs on the veins; lowest 6-20 (—
40) x 5-17 (-21) cm, ovate, rounded-obtuse to subacute at apex, sinuate-glandular-denticulate,
more or less cordate at base, the petiole 10-30 cm with a denticulate, irregular wing for most or all
of its length, sometimes tinted purplish at base and with pale simple eglandular hairs; medium and
CICERBITA IN THE BRITISH ISLES 123
\ \\ SWZ
AY G
y
Me & = Cicerbita macrophylla
© del ‘Fred Rumsey 1182
FicureE 1. Cicerbita macrophylla (Willd.) Wallr. subsp. uralensis (Rouy) P. D. Sell; based on a specimen from
Dyfed, 10th July 1982, Jury & Rumsey 1382 (RNG).
124 Pr DISEEL
lower 16-30 x 8-12 cm, lyrate, the terminal lobe 12-16 x 8-12 cm, ovate, shortly acute or
acuminate at apex, sinuate-glandular-denticulate, often cordate at base, the midrib between lateral
and terminal lobes broadly winged with occasional small teeth, the basal lobes forming a subrotund
outline with a cordate-amplexicaul base; upper 6-14 x 1-5-6 cm, lanceolate to narrowly ovate,
long-acuminate at apex, sinuate-glandular-denticulate, rounded amplexicaul at base. Inflorescence
with a compact group of capitula near the apex and several long branches up to 16 cm from the
upper leaf axils. Capitula 40-50 mm in diameter; peduncles 3-25 mm, slender, with numerous,
yellowish glandular hairs. Involucre 8-15 x 4-9 mm, narrowly cylindrical; bracts 4-5—14 x 1-5-2
mm, pale yellowish-green with a very narrow pale margin and slightly purplish at apex, linear or
linear-lanceolate, more or less obtuse at apex, with numerous yellowish or slightly reddish
glandular hairs. Flowers 27-33; ligules 15-23 x 34 mm, pale lilac with slightly darker veins,
shallowly and irregularly dentate at apex. Stigmas pale lilac-blue. Receptacle shallowly pitted.
Achenes c. 5 mm, elliptical, flattened, narrowly winged, with 3 ribs on each face, glabrous.
As the only description of this species of any length is in Russian and as I have living material as
well as herbarium sheets available to me, I have taken the opportunity of giving a more detailed
account than for other species.
When preparing the account of Cicerbita for Flora Europaea (Sell 1976), I borrowed the
numerous specimens of C. macrophylla from the Leningrad (LE) herbarium and found they were
divisible into two subspecies. Plants from the Caucasus, subsp. macrophylla, had the main
branches of the panicle 2-5—3-5 (—5) mm in diameter and the ligules dark violet-while those from
the Urals, subsp. uralensis, had the main branches of the panicle 1-5—2-5 (—3) mm in diameter and
the ligules pale lilac. All the British naturalized material I have seen is referable to subsp. uralensis.
There is no record of when this plant was first introduced into British gardens. D. E. Allen informs
me there is a letter in the G. C. Druce correspondence at Oxford, from C. Bucknall dated 15th
May 1918, in which he states he found Mulgedium macrophyllum naturalized by the roadside at
Glenridding, Ullswater in August 1915. It was recorded as a naturalized escape in a field next to a
house, Kingsland, parish of Meole, Shropshire, v.c. 40, in July 1921 by Agatha Wilkinson. The
specimen in herb. Oxford (OXF) contains a note by Mr Oliver (the owner of the house) to say he
had seen it there for 30 years. These records were followed by one from Patterdale, v.c. 69, in 1922
by W. W. Mason and another by the same author from Melmerby, v.c. 70, in 1923. It has since
been recorded from most areas of Great Britain and has even reached the west of Ireland (Fig. 2)
(Akeroyd et al. 1983). The same subspecies seems to be widely naturalized in continental Europe.
In 1972 U. K. Duncan sent me plants from the grounds of Colliston Castle, near Arbroath,
Angus, v.c. 90, which I grew in my garden. Since then they have spread profusely over several
square metres but have not travelled any distance. Most naturalized colonies seem to be of this
size, although some spread over larger areas. It is likely that the species reaches a new area by
rhizomes thrown out from gardens, or rhizomes transported in piles of earth, and if the terrain is at
all suitable soon forms a clonal patch by rhizomatous spread. I have not seen viable seed produced
in this country and it is possible that each colony is a clone and the species is self-incompatible.
Moreover we do not know how many different introductions have been made to the country as a
whole. The flowers, which are open throughout the day from late June to August, are visited by
Bombus spp. and various Diptera.
3. C. PLUMIERI (L.) Kirschleger, Fl. Alsace, 1: 401 (1852).
Sonchus plumieri L., Syst. Nat., 10th ed., 2: 1192 (1759).
Mulgedium plumieri (L.) DC., Prodr., 7(1): 248 (1838).
Lactuca plumieri (L.) Gren. & Godron, Fl. Fr., 2: 322 (1851).
Vernacular Name: Hairless Blue Sow-thistle
Illustration: Fig. 3.
Plant glabrous. Stems 60-130 cm. Leaves 5-60 x 2-17 cm, lyrate-pinnatifid with a large triangular
terminal lobe and several pairs of more or less ovate lateral ones which are shorter than, but at
least as wide as, the terminal; lobes more or less undulate, with mammiform teeth; midrib between
lobes and petiole with a broad wing. Capitula in a wide, more or less corymbose panicle. Involucre
CICERBITA IN THE BRITISH ISLES 125
Ficure 2. The distribution of Cicerbita macrophylla subsp. uralensis in the British Isles.
10-17 x 9-12 mm; bracts lanceolate or linear-lanceolate. Ligules blue. Achenes 5-5—6-5 mm, flat,
linear to narrowly elliptical, conspicuously narrowed at apex.
Clapham (1962) stated that C. plumieri is often cultivated and sometimes escapes. I have been able
to trace the following records. The only specimen I have seen is the McCallum Webster one from
Tighnabruich.
v.c. 2, E. Cornwall. Saltash, c. 1975, E. Griffiths. Margetts & David, Rev. Cornish Fl. 370
(1981).
v.c. 4, N. Devon. Hedgerow near gardens, off Orchard Hill, Bideford, 11th July 1956, det. W.
Keble Martin.
v.c. 14, E. Sussex. Warren Wood, Borde Hill, near Haywards Heath, 13th Sept. 1982, K. E. Bull,
det. E. J. Clements.
126 PDI SELL
pes: em; Weak
Ly)
yh Ry.
FiGuRrE 3. Cicerbita plumieri (L.) Kirchleger; based on a specimen from Spain, Ist August 1982, Goyder & Jury
213 (RNG).
CICERBITA IN THE BRITISH ISLES 127
v.c. 32, Northants. Fletton brickyards, 1945, Herb. Oundle School Natural History Society. J. S.
Rees, A Flora of Oundle 131 (1969) (as Lactuca plumieri).
v.c. 59, S. Lancs. Preston neighbourhood, F. W. Stansfield. Gard. Chron. (1917). Sand-dunes,
Freshfield, 1960, Mrs F. W. Holder. J. P. Savidge, Travis’s Fl. S. Lanc. 346 (1963).
v.c. 76, Renfrews. Near Paisley. Proc. bot. Soc. Br. Isl., 1: 173 (1954).
v.c. 77, Lanarks. Tollcross. Proc. bot. Soc. Br: Isl., 1: 173 (1954).
v.c. 96, Easterness. Garden escape at Tighnabruich near Invermoriston, 1950 and 24th June 1975
(E). M. McCallum Webster, Fl. Moray, Nairn & E. Inverness 372 (1978).
v.c. 99, Dunbarton. Milngavie. Glasgow Nat., 17: 76 (1953); Proc. bot. Soc. Br. Isl., 1: 173 (1954).
v.c. H39, Co. Antrim. By railway arch at Lambeg, and near Drum Bridge, escapes from garden of
public-house at Drumbeg, where it was introduced about 1905, E. N. Carrothers. Stewart &
Corry, Fl. N.E. Ireland, 2nd ed., 311 (1938) (as Lactuca plumieri).
C. plumieri is a native of the Pyrenees, the mountains of France and western-central Europe and
south-western Bulgaria. There seem to be no records of when it was introduced to British gardens.
4. C. BOURGAE I (Boiss.) Beauverd in Bull. Soc. Bot. Genév. sér. 2, 2: 118 (1910).
Mulgedium bourgaei Boiss., Fl. Or., 3: 801 (1875).
Lactuca bourgaei (Boiss.) Irish & Taylor in L. H. Bailey, Standard Cycl. Hort. 1766 (1916).
Vernacular Name: Pontic Blue Sow-thistle.
Illustration: Fig. 4.
Robust perennial up to 300 cm with stout roots and many stems from one base. Stems erect, pale
green, slightly ridged, solid, with scattered pale simple eglandular hairs. Leaves 40-70, 8-26 x 2-7
cm, all cauline, lower soon dying off, gradually decreasing in size up the stem, yellowish-green,
thick, obovate or oblanceolate in outline, acute at apex, lower with a large terminal lobe and 1-2
pairs of small lateral ones, upper undivided, all shallowly and sharply dentate, narrowed to a
semiamplexicaul base, glabrous above, with numerous stiff white eglandular hairs beneath, particu-
larly on the prominent veins. Inflorescence long and narrow with up to 100 capitula, leafy in the
lower part; peduncles glabrous or with short simple pubescence. Involucre narrow and cylindrical;
bracts 1-12 x 1-7-2 mm, ovate-lanceolate to linear, obtuse at apex, pale green suffused purple,
glabrous. Ligules pale lilac with darker veins and a nearly white base. Stigmas pale lilac. Achenes 4—
4-5 mm, pale brown, many-ribbed.
I have traced the following localities for C. bourgaei as a naturalized British plant. There seems to be
no record of when it was first introduced to British gardens. The first record of it naturalized is a
specimen in CGE from Maldon in Essex in July 1950. It seems to be on the increase.
v.c. 7, N. Wilts. Kington Langley, near Chippenham, 1974, G. M. Kay and Mrs J. Swanborough.
Little Sodbury, Aug. 1978, Mrs E. Norman.
v.c. 17, Surrey. Warlingham, top of Bug Hill, a long established escape, 1958, S. Fletcher (BM).
1963, D. P. Young. Lousley, Fl. Surrey 327 (1976). Burton, Fl. London Area 161 (1983).
v.c. 18, S. Essex. Near Woodham Mortimer Place, roadside hedge, 1953, Miss G. Sparrow 1956-7.
Jermyn, Fl. Essex 162 (1974). Roadside, Maldon, 18th July 1950, O. Folkard (CGE).
v.c. 22, Berks. Boars Hill, 1963 and Whiteknights Park, 1965. Bowen, Fl. Berks. 264 (1968), as
Lactuca bourgaei. Poughley, Aug. 1968, H. J. M. Bowen (RNG).
v.c. 24, Bucks. Lower Winchcombe Priory, near Thame, 10th July 1959, J. F. May (RNG).
Naturalized by lane, Nether Winchendon, 12th Aug. 1968, J. G. & C. M. Dony.
v.c. 28, W. Norfolk. Roadside near a ford, Great Walsingham, Ist Aug. 1969, J. R. Palmer. Swann,
Suppl. Fl. Norfolk 73 (1975).
v.c. 34, W. Gloucs. Little Sodbury, Avon, naturalized on roadside, 1977, A. L. Grenfell & Mrs M.
M. C. Reiss. Westbury-on-Trym, Bristol, Avon, 1974, C. H. Cummins.
v.c. 64, Mid-W. Yorks. Pool, Otley, 4th Aug. 1947, R. P. Libbey (RNG).
v.c. 71, Man. Dumped rubble, Lezayre Curraghs, G.R. 24/4.9, 1984, Miss M. Devereau via D. E.
Allen.
v.c. 80, Roxburghs. Roadside verge, above Sprouston Burn, Newtown St Boswells, 2nd Aug. 1983,
Miss J. Blance (herb. R. W. M. Corner).
128 PD. SELL
Wert Ine
y] ° iy f
q Vi ff
VA
Y
SA \ =
y I) "y
Ficure 4. Cicerbita bourgaei (Boiss.) Beauverd; based on specimens from (A) Bucks.,
(RNG); and (B) (basal leaf) Berks., August 1968, Bowen 622 (RNG).
10th July 1959, May
CICERBITA IN THE BRITISH ISLES 129
v.c. 95, Moray. Formerly abundant outside the walled garden at Brodie Castle, now exterminated
with weed killer, 1967. M. McCallum Webster, Fl. Moray, Nairn & E. Inverness 372
(1978).
ACKNOWLEDGMENTS
I am grateful to John Akeroyd and Stephen Jury for making available to me all the notes they had
accumulated since their paper in B.S.B.I. News, and to Eric Clement for lending me the relevant
part of his card index of records. Others who have kindly sent records of Cicerbita are too
numerous to mention. Chris Preston and the Biological Records Centre have provided the map of
C. macrophylla subsp. uralensis. Fred Rumsey kindly allowed me to use his drawings to illustrate
this paper.
REFERENCES
Akeroyp, A. J. [J. R.], Jury, S. L. & Rumsey, F. J. (1983). Cicerbita macrophylla (Willd.) Wallr. B.S. B.I.
News, 33: 14-15.
CLaPHAM, A. R. (1962). Cicerbita Wallr. in CLAPHAM, A. R., Tutin, T. G. & Warsure, E. F. Flora of the
British Isles, 2nd ed., pp. 895—6. Cambridge.
MarreEN, P. R., PAYNE, A. G. & RANDALL, R. E. (1986). The past and present status of Cicerbita alpina (L.)
Wallr. in Britain. Watsonia, 16: 131-142.
SELL, P. D. (1976). Cicerbita L. in TuTin, T. G. et al., eds. Flora Europaea, 4: 331. Cambridge.
(Accepted May 1985)
sii eln’ >, it?
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Watsonia, 16, 131-142 (1986) 131
The past and present status of Cicerbita alpina (L.) Wallr. in
Britain
P. R. MARREN
Nature Conservancy Council, Northminster House, Peterborough, PEI 1UA
A. G. PAYNE
8 Esslemont Avenue, Aberdeen
and
R. E. RANDALL
Girton College, Cambridge
ABSTRACT
Cicerbita alpina (L.) Wallr. (Compositae) is a very rare plant in Britain, confined to remote mountain cliff
ledges and gullies in the counties of Angus (v.c. 90) and Aberdeen (S. Aberdeen, v.c. 92). Its past and present
distribution in Scotland is discussed in relation to the habitat of the species. Recent monitoring of populations
at Lochnagar and Caenlochan suggests that flowering performance varies from year to year in response to
climatic conditions in late summer and to the incidence of rockfalls. The grazing of sheep and red deer,
combined with the decline of natural submontane woodlands, has increasingly confined the plant to montane
cliffs, an atypical habitat to which it appears to be poorly adapted.
INTRODUCTION
Cicerbita alpina (L.) Wallr. (Compositae), the Alpine Blue Sow-thistle, is a very rare mountain
plant in Britain and is one of 62 species protected under Schedule 8 of the Wildlife and Countryside
Act of 1981. By contrast, it is a widespread and locally abundant species of submontane birch and
pine forests in Scandinavia and in European mountain ranges southwards to the Pyrenees,
northern Apennini and the mountains of the Balkan peninsula. C. alpina is a robust, perennial
herb growing to a height of about 130 cm in Britain. Flowering shoots rise annually from a rhizome
and the flower buds usually open in late July. The plant typically grows in large clumps and the
broad, sagittate leaves cast a heavy shade. When the flowers first appear, they are an attractive
shade of blue, but they soon wither to a dull bluish-violet, and the plants are often battered by
winds and rainstorms by late summer. The seeds are surmounted by a pappus of long, stiff hairs but
ripe seed is not always produced in Scotland. The habitat and autecology of C. alpina in Scotland
have been summarized by Roger (1943) and Randall (1977).
THE DISCOVERY OF THE SCOTTISH SITES OF CICERBITA ALPINA
Cicerbita alpina was discovered in Britain by the great botanical traveller George Don in September
1801 on one of the precipitous ledges in the north-eastern corrie of Lochnagar. Don sent
specimens gathered here to his correspondent Sir James Edward Smith, who published a
description of the plant as Sonchus caeruleus Sm. (Smith 1811). Some of Don’s specimens, which
132 P. R. MARREN, A. G. PAYNE AND R. E. RANDALL
are seldom dated, are housed in the Smith herbarium (LINN) while others were given to private
collectors, one of whose sheets is now in the herbarium of the Royal Botanic Gardens, Edinburgh
(E). Don left no account of his discovery, but in a paper published the year before his death he
mentions that he had also found “‘the Sonchus . . . on the rocks among the Clova mountains” (Don
1813; Druce 1904). He was thus also the first botanist to find the plant in one or more of its Angus
localities. Abundant herbarium material from Glen Doll and Caenlochan dates from the 1820s
onwards, and William Gardiner’s Flora of Forfarshire (1848) lists no less than six localities in the
former glen and “‘several”’ in the latter.
A further colony was subsequently found in Coire Kander, Glen Callater, although the precise
date of its discovery is uncertain. Members of the Scottish Alpine Botanical Club believed that
they had discovered this new site during a field excursion in 1878 (Boyd 1879). The 1866 edition of
Sowerby’s English Botany, however, lists C. alpina from “‘Glen Callater” (Ratcliffe 1958). George
Dickie (1860) was unaware of any Aberdeenshire site other than Lochnagar when he compiled his
Botanist’s Guide to the north-eastern Scottish counties. It is equally uncertain when the little-
known Angus station in Canness Glen first came to light. The earliest herbarium specimen that the
authors have seen dates from 1866 (ABD).
No other British station for C. alpina has yet been found. From the time of Don onwards, there
have been persistent reports of a blue-flowered sow-thistle from the Lake District and other upland
areas of Britain but whenever it is possible to check these reports, the plant in question is
invariably the widespread garden escape, Cicerbita macrophylla (Willd.) Wallr.* The British range
of C. alpina is therefore confined to the block of mountainous terrain bounded by Deeside, Glen
Muick, Glen Clunie and the glens of Angus, commonly known as the ‘Clova mountains’, although
more properly termed “The Mounth’. The reason for this unusually localized distribution is
unknown. Few other montane vascular plants are confined to the eastern highlands of Scotland
and, with the exception of Homogyne alpina, none are as local as C. alpina.
HABITAT REQUIREMENTS
Over 70 years ago, Smith (1911) classified Cicerbita alpina as a shade species that required a high
degree of humidity and pockets of deep, permanently moist soil. The present-day localities of C.
alpina lie either on precipitous north- and north-east-facing cliffs or in deep ravines. At Glen
Canness and Glen Doll the sites are (or were) the stony beds of gullies and stream gorges, at 530 —
670 m. The Caenlochan, Lochnagar and Coire Kander sites are broad, flat cliff ledges with
overhanging rocks, or sloping ‘ramps’, gullies and chimneys, at altitudes of 840-1090 m. Some,
possibly all, of these sites regularly carry snow into the spring and even early summer. Although
the famous inaccessability of the sow-thistle sites has been somewhat exaggerated by some authors,
most of its stations do indeed ‘“‘demand both a certain agility and a fair head for heights” (Raven &
Walters 1956).
The parent rock of the Scottish localities is either igneous (Lochnagar and Caenlochan) or
metamorphic (other sites). The geology of the C. alpina sites is complex; the parent rocks are all
acidic but they commonly contain base-rich intrusions. The parent material at the Coire Kander
locality is syenite, an acid pyritic rock (Ratcliffe 1958). The Caenlochan ledge lies in an area of
hard, fine-grained granite with porphyritic crystals of quartz and felspar. Ferreira (1958) found that
the flora of the quartz porphyry in this area closely resembled that of the syenite on Ben Loyal in
Sutherland. The Lochnagar sites occur within, or close to, gullies containing crushed rock
associated with fault zones where the typical grey or pink Lochnagar granite is replaced by cream
or greenish, heavily weathered rock in which the base-rich minerals epidote and calcite may be
frequent (Ferreira 1958). There is a strong likelihood that nutrients are transported to the C. alpina
sites from base-rich rocks, both by percolation and by rock-falls.
The soils of C. alpina sites were described by Roger (1943) as granitic gravels mixed with peaty
*The ‘C. alpina’ of Glenridding in the Lake District, for example, known to local botanists as ‘““Bob Nixon’s
Helvellyn Plant’, was correctly identified by Professor W. H. Pearsall as C. macrophylla (B.E.C. Report 1936,
p. 265). Don (1813) reported that “S. coeruleus is now lost at Howden Pans, in England” but he is unlikely to
have known either the plant or the site at first hand.
CICERBITA ALPINA IN BRITAIN 133
humus and possessing a good depth and moisture-retaining capacity. Ratcliffe (1958) characterized
the soil of the Coire Kander ledge as a “mull humus, poorer than the calcareous schists but
somewhat enriched by the drip of water from material containing some bases.” Although they
have not been exhaustively analysed, these soils are probably analogous to those of other tall herb
associations on mountain ledges, which are typically periodically irrigated, skeletal brown loams
with mull humus and slight podsolization (McVean & Ratcliffe 1962). The soil reaction of the C.
alpina sites varies from pH 4-8 to 5-9, that at Caenlochan showing a relatively high pH which is
attributed to the constant drip of base-enriched water falling onto the ledge from above. The
ledges on which C. alpina grows usually carry a luxuriant vegetation among which ptarmigan nests
are sometimes found. Bird droppings and nest remains are a further probable source of soil
nutrients locally.
C. alpina invariably grows in places where grazing is minimal, since it is extremely palatable to
large herbivores, notably sheep and deer. It typically occurs in association with equally palatable
plants, notably Hogweed (Heracleum sphondylium), and such luxuriant vegetation would offer an
open invitation to grazing animals were it not for its inaccessibility. The largest colony on
Lochnagar belies its altitude of c. 1000 m by possessing a lush plant community reminiscent of
lowland ditch-sides and open woodland in Norway. Athyrium distentifolium, a species of late
snow-beds, is a frequent associate of C. alpina in its higher sites, and other species which are
usually present in abundance are Alchemilla glabra, Deschampsia cespitosa, Dryopteris dilatata,
Luzula sylvatica, Oxalis acetosella, Rumex acetosa and Solidago virgaurea. A fuller list of close
associates of C. alpina is provided in Table 1. Some sites have their own individual features; the
Caenlochan ledge is characterized by the tall, robust stems of Hogweed, while in one of the
Lochnagar colonies, the sow-thistle leaves mask exceptionally tall specimens of Frog Orchid
(Coeloglossum viride). At a recently discovered station in Glen Doll there is a pronounced
calcicolous element in the flora, which includes Saxifraga aizoides, S. oppositifolia and
Polystichum lonchitis. We would agree with Ratcliffe (1958) that C. alpina appears to be tolerant
rather than genuinely calcifuge.
The phytosociological affinities of the fragmentary C. alpina sites in Scotland are uncertain.
McVean & Ratcliffe (1962) list C. alpina among the species exclusive to their Tall Herb nodum,
which is “‘probably best regarded as an unorganized or fragmentary Lactucion’’, an alliance
characterized by this species in Scandinavia. Huntley (1976) suggests that the Caenlochan ledge, at
least, is close to the Luzula sylvatica nodum which is, in turn, related to a treeless facies of the
woodland association, Betuletum Oxaleto-Vaccinietum. The Scottish sites for C. alpina, therefore,
have close affinities with woodland vegetation and, in the Angus glens at least, the plant may once
have been a constituent of submontane woods as in climatically comparable areas in Scandinavia.
The Glen Doll and Canness localities are still partially wooded, and even the high, remote shelf on
Lochnagar bears a flourishing rowan sapling.
The ecological amplitude of C. alpina in Scotland differs markedly from that in Europe, where it
is a mainly submontane species, more typical of damp, open, lightly grazed woodland and scrub
than the steep mountain crags of the Scottish sites. It is a characteristic species of Scandinavian tall
herb meadows and one of its synonyms, Lactuca alpina (L.) A. Gray, lends its name to the alliance
Lactucion alpina (McVean & Ratcliffe 1962). Such meadows are normally very wet and, in
Scandinavia, C. alpina is commonly a ditchside plant.
THE SCOTTISH SITES OF CICERBITA ALPINA
LOCHNAGAR
Byron’s “‘steep, frowning glories of dark Lochnagar” contain the largest remaining populations of
Cicerbita alpina in Britain. All the known sites are within the north-eastern corrie. The best known
station is a broad, sloping ledge at about 1000 m, mistakenly stated by Dickie (1860), following
Watson’s Cybele Britannica, to lie at 640-820 m. This was probably the ledge on which Don
discovered the species in 1801. The plant was gathered there at intervals by the hardier botanists of
the 19th century, including R. Mackay in 1845, J. Hutton Balfour (Balfour 1848, 1875), James
Backhouse the younger in 1852, F. J. Hanbury in 1886, F. Buchanan White in 1886, and P. Ewing
(Ewing 1887).
134 P. R. MARREN, A. G. PAYNE AND R. E. RANDALL
TABLE 1. SPECIES ASSOCIATED WITH CICERBITA ALPINA
Field data for vascular plants obtained by P.R.M. (1977-80, Lochnagar and Caenlochan), A.G.P. (1978-80, all
sites), and R.E.R. (1976-77, Lochnagar, Caenlochan and Coire Kander). Literature references to all plants
taken from McVean & Ratcliffe (1962) and Roger (1943).
Species
Vascular plants (co-dominants)
Athyrium distentifolium
Dryopteris dilatata
Alchemilla glabra
Oxalis acetosella
Heracleum sphondylium
Rumex acetosa
Vaccinium myrtillus
Solidago virgaurea
Luzula sylvatica
Deschampsia cespitosa
Festuca ovina/vivipara
Vascular plants (close associates)
Gymnocarpium dryopteris
Phegopteris connectilis
Blechnum spicant
Polystichum lonchitis
Anemone nemorosa
Ranunculus acris
Thalictrum alpinum
Silene dioica
Geranium sylvaticum
Geum rivale
Alchemilla alpina
Saxifraga stellaris
S. aizoides
S. oppositifolia
Rhodiola rosea
Epilobium angustifolium
Angelica sylvestris
Empetrum hermaphroditum
Oxyria digyna
Rumex acetosella
Euphrasia officinalis
Thymus praecox
Galium saxatile
Succisa pratensis
Gnaphalium norvegicum
Tussilago farfara
Saussurea alpina
Hieracium sp.
H. calenduliflorum
Coeloglossum viride
Carex bigelowii
Agrostis capillaris
Anthoxanthum odoratum
Deschampsia flexuosa
Total no. vascular plants
recorded
Bryophytes”
Sphagnum quinquefarium
Plagiothecium undulatum
Rhacomitrium lanuginosum
Dicranum majus
++
+++++ 44
++
Caenlochan Glen Doll
+++
++++
16
Canness
ye
Lochnagar
++4++4+
+++4+4++4
+++4++
21
Coire Kander Unspecified
+++ teeter+e+t
++
+++
CICERBITA ALPINA IN BRITAIN 135
Table 1 continued.
Species Locality
Caenlochan Glen Doll Canness Lochnagar Coire Kander Unspecified
D. scoparium ot
Mnium punctatum +
Rhytidiadelphus loreus ue
R. squarrosus
Thuidium tamariscinum
Hylocomium splendens
Polytrichum commune +
P. piliferum sp
Diplophyllum albicans 3°
Lichens
Cladonia rangiferina =f
C. cornuta +
+++
Other species recorded from the Lochnagar sites but less closely associated with C. alpina are Antennaria
dioica, Calluna vulgaris, Campanula rotundifolia, Cirsium heterophyllum, Cornus suecicus, Cryptogramma
crispa, Huperzia selago, Juncus trifidus, Luzula spicata, Nardus stricta, Oreopteris limbosperma, Polygonum
viviparum, Potentilla erecta, Salix lapponum, Sorbus aucuparia, Trichophorum cespitosum, Trollius europaeus,
Vaccinium vitis-idaea, V. uliginosum and Viola palustris.
* The small number of species recorded from Canness is almost certainly incomplete and does not necessarily
mean this site was unusually species-poor.
> Not recorded at Glen Doll, Canness or Lochnagar.
C. alpina seems to have been present in good quantity throughout these years: Balfour found it
“in great quantities”, Buchanan White was pleased to find it “‘still abundant” and Young (1908)
reported that it was “growing vigorously’. The only quantitative statement is that of Ewing (1887)
who counted about 150 plants. Although it is difficult to know what Ewing regarded as a single
‘plant’, this figure compares well with a count of 163 inflorescences in the most accessible station in
1983.
There are at least three main areas in which C. alpina grows on Lochnagar. These sites are all
damp, shady gullies and rock ‘chimneys’ or broad ledges, which are kept moist by more or less
constant percolation, and often carry snow until late in the summer. Some of the sites are on
climbing routes, and local climbers’ publications occasionally mention the plant in passing. One
climbing party, for example, referred to a ‘“‘veritable garden of ferns and flowers [which] included a
large plant not unlike a dahlia” (Ewan 1932). The site became known by the irreverent name of
“the potato patch’.
The flowering performance of these populations of C. alpina fluctuates from year to year. Table
2 records the number of flowering stems in the three known colonies between 1972 and 1983. ‘Site
A’ is a narrow, shady gully with a cave, at c. 1050 m, which contained a flourishing colony in the
1920s and 1930s. By the early 1970s, the population had declined, for reasons unknown, to four
small and widely spaced clumps. ‘Site B’ is a broad north-east-facing ramp sloping at about 45°.
This is one of the best known localities for C. alpina, for the flowers can easily be viewed from a
distance although access to the ramp itself is another matter. Between 1967-1974, the population
had fallen from just over 100 ‘plants’ to 42 (H. J. B. Birks in Randall 1977). The decline was due to
a natural rock-fall, a phenomenon of frequent occurrence in mountain gullies. Although such falls
cause temporary disturbance, from which C. alpina normally has the capacity to recover, they also
provide new nutrient supplies to the soil and may be beneficial in the long term. At any rate, C.
alpina soon increased from about 30 stems in 1973 to a maximum of 311 in 1980, before abruptly
decreasing again in the early 1980s. There was no fresh rock-fall to account for this but C. alpina
appears to flower more prolifically in cool, wet seasons than in the hot, dry summers of 1982-84,
and hence the reason for the second decline may be climatic. ‘Site C’, a narrow rock chimney
beneath the ramp and even more difficult of access, shows parallel increases and declines of
136 P. R. MARREN, A. G. PAYNE AND R. E. RANDALL
TABLE 2. FLOWERING STEMS OF CICERBITA ALPINA ON LOCHNAGAR, 1972-1983
Data of A.G.P. and Claire Geddes
Site A
Clumps
Date 1 2 3 4 Site B Site C Additional notes
24.8.72 0 1 few few many
30.8.73 30+
8.74 42
9.8.75 0 0 6 0 many
27.8.76 many 2+
14.9.77 = 0 0 3 0 120+ 22+
23.8.78 0 0 6 veg. 0 160 a1
shoots
4.8.79 222 75+ Most in bud
1.9.80 y 3 1 2 veg. 311 73.
shoots
17.8.81 33 50+
3.8.82 5 veg. 2 veg. 46 es
shoots shoots
28.7.83 1 2 veg. 163 85+ » All in bud
shoots
flowering shoots. The overall impression at Lochnagar is one of short term change but long term
stability. The plants appear particularly healthy and robust and, here at least, there seems to be
no immediate prospect of serious decline.
COIRE KANDER
The cliffs of Coire Kander in Glen Callater contain the second largest British population of
Cicerbita alpina. The main colony grows as a dense mass on a broad, north-facing ledge at about
840 m with a smaller colony in a perpendicular gully nearby. Since the site is difficult to reach,
there are few specimens from Coire Kander in herbaria. As a member of the Scottish Alpine
Botanical Club put it, ‘no member was sufficiently foolhardy to break his neck in an attempt to
scale the exact rock” (Boyd 1879). Ratcliffe (1958) counted 200 flowering stems but the popula-
tion had declined to only 40 flowering stems by August 1972. Since there was no obvious physical
cause, this colony was assumed to have “‘lost vitality” (Perring & Farrell 1983). Fortunately, as at
Lochnagar, a recovery has since taken place. The population was monitored annually between
1977-1979 and the results are reported in Table 3. The increase of C. alpina at Coire Kander
matches that at Lochnagar. A similar correlation between the drought years of 1975-76, which
may have been the real cause of the ‘lost’ vitality, and the subsequent increase in the cool, wet
summers of 1977-79 is suggested.
CAENLOCHAN GLEN
Caenlochan is probably the best known site for Cicerbita alpina in Britain. So far as the authors
are aware, only one colony is known today and successive botanical pilgrims have worn a track
along the base of the steep, north-facing cliff on which it grows. There were other stations in the
glen in the 19th century. According to Gardiner (1848) it grew “‘in several places by the sides of
streams at the head of Caenlochan glen’’. Ewing (1887) believed that there were no less than five
patches, three of which he knew personally, although he did “not believe one collector in a
hundred would undertake to gather a single specimen from any of the stations where I have seen
"| ae
These statements imply that C. alpina was once less rare in Caenlochan Glen. Gardiner implies
that it once occurred on the lower ground by the myriad burns which feed into Glen Isla, as well
as on cliff ledges. Unfortunately the fate of these colonies cannot be traced, for herbarium labels
seldom enable the exact location to be identified, and it is possible that C. alpina still occurs in
CICERBITA ALPINA IN BRITAIN 137
TABLE 3. CICERBITA ALPINA IN COIRE KANDER
Additional
Date No. flowering stems No. stems No. plants Recorder notes
30.7.77 67 160 30 R.E.R. In bud
9.8.78 ©2155 not recorded AGG: Mostly in bud
7.8.19 c.165 not recorded A.G.P.: Mostly in bud
some of the inaccessible places referred to by Ewing. Nevertheless, the present solitary ledge
seems always to have been the best known and consequently C. alpina was heavily collected there
for at least 160 years. This population occupies a broad, flat ledge at c. 840 m with steep,
overhanging rock-walls on the southern and western sides. The ledge has been visited annually for
many years, although population data have been available only since 1972. The plants suffer from
late summer wind and are often very battered by the time the flowers are in bloom. This
population can never have been very large, because of the physical limitations of the site, and only
10—40 flowering shoots are normally produced. The data in Table 4 reflect the changing fortunes of
this colony since 1975. After a dismal showing in the late 1970s, the Caenlochan plants are now
showing encouraging signs of recovery. In 1977 a ramp of exceptionally late snow enabled grazing
TABLE 4. CICERBITA ALPINA IN CAENLOCHAN 1975-1983
Date of No. flowering
Visit shoots Total no. stems Recorder Condition
1975 12 ‘60 plants”’ F. H. Perring Damaged by wind
T.OA9TS. 0 AvGiP: All flowering stems
clean cut at 30 cm
from ground.
Much late snow
forming ramp below
ledge
27.17.1976 3 in bud 30-40 C. Geddes
2 AGT] 12 in bud 80 J. Bevan &
J. M. Mullin
8.7.1977 12 in bud (about 24 plants) P.R.M. Poor condition with
browning leaves
301977 19 40 ROE.R; Ten grazed
stems at front of
ledge
£9:9.1977 30 A.G.P. Plants mainly
prostrate
10.8.1978 20 C. Geddes Poor condition, only
one well-developed
flowering stem;
others depauperate
with browning leaves
6.7:1979 No sign of buds 50+ A.G.P
12.8.1979 6 in bud A.G.P Many leaves browned
14.8.1979 8 N;G.€ Three plants
trampled at front
of ledge
7.10.1980 4] NCC Poor, but no signs of
interference
30.8.1981 14 C. Geddes Good
27.7.1983 45 in bud C. Geddes Good
138 P. R. MARREN, A. G. PAYNE AND R. E. RANDALL
animals to reach the ledge. Recovery appears to have been slow and perhaps hindered by violent
wind and human trampling. As with the Lochnagar populations, 1980 and 1983 were good years.
CANNESS GLEN
The Canness station of Cicerbita alpina is the least known and least visited — probably few botanists
are inclined to forego the splendours of Caenlochan for the comparatively dull flora of the
neighbouring Canness glen. Only one colony, a single large clump, was known and even this now
appears to have declined to extinction.
The population was presumably in a reasonably healthy state in 1885, when a large number of
flowering stems were removed (BM, OXF). The site was photographed by R. M. Adams in 1937,
and his picture, together with its surprising background of larch trees, is illustrated in Holden
(1952). The site.was on a pocket of deep soil by the rocky bank of the Canness burn in a shady
gorge which was normally isolated from the attentions of grazing sheep and red deer by a precipice
and a deep pool. The larches, growing here at an altitude of c. 535 m, are the remains of a 19th
century plantation.
In the late 1940s, possibly the dry summer of 1947, the Canness plants were grazed close by deer
(U. K. Duncan pers. comm.). A tiny population, which produced 2-4 flowering shoots annually
(Roger 1967; E. Jensen pers. comm), survived until 1976, when four flowering stems were
recorded (C. Geddes pers. comm). In the following year, however, only a single shrivelled shoot
remained and the colony now appears to be extinct. The hot, dry summer of 1976 seems to have
marked the final demise of the population. Perhaps the low water level of the burn again allowed
deer or sheep to reach the plants; or perhaps the drying and compaction of the soil resulting from
the drought was responsible for the loss of the last few plants. More recent searches have been
fruitless and any further sightings of C. alpina from Glen Canness would therefore be most
welcome.
GLEN DOLL
Glen Doll has more well-authenticated stations for Cicerbita alpina than any other Scottish site. A
number of streams, rising on the plateau, tumble down the steep, north-eastern faces of Craig
Maud, The Dounalt and Craig Rennet in gloomy, boulder-strewn ravines to join the White Water
in the glen below. At least six colonies of C. alpina once grew in gullies and on shelving rocks by
the old stations in streams (Gardiner 1848) at altitudes of 700-800 m. One colony at least was in the
bed of a burn and others grew on rocks kept permanently moist by waterfalls. In the 19th century,
Glen Doll was both the best known and the most accessible of the C. alpina sites, and there is more
herbarium material from this site than from any other.
By contrast, 20th century herbarium material is scarce and for many years it was feared that C.
alpina had disappeared from all its former stations in the glen. Mrs T. J. Foggitt gathered it from a
site on the Dounalt in 1921 and 1925 (BM), and U.K. Duncan (pers. comm) found a small
population in a different place in 1930. This locality was rendered more or less inaccessible by a
rock-fall in the early 1930s and, although the population seems to have survived, the plants had
produced no flowers when the site was revisited shortly afterwards. The Dounalt population is also
said to have survived a rock-fall but no subsequent observations are known to the authors. J.
Raven (Raven & Walters 1956) was informed that C. ges had disappeared in a flood from the
best known locality in Glen Doll.
In September 1979, one of us (A.G.P.) discovered a healthy population of C. alpina in a narrow,
north-facing gully blocked by a large ‘chockstone’. 40 flowering stems were counted on the
inaccessible floor of the gully above the chockstone and another four below. The majority of the
plants were visible only from above and were virtually inaccessible except to rock climbers. Rock-
falls and spates are evidently more frequent at Glen Doll than at the other localities and sudden,
violent change was probably always a feature of the C. alpina sites, although the diminished
numbers of the plant now render its survival more precarious. The blocking of gullies by rock-falls
may well have hidden from view other populations recorded by Gardiner, or possibly botanists
today are less adventurous climbers than were their 19th century counterparts. If so, the decline of
C. alpina in Glen Doll may be more apparent than real, although it should be added that several
parties have recently searched old localities in the glen without success.
CICERBITA ALPINA IN BRITAIN 139
FACTORS CONTRIBUTING TO THE PRESENT DAY RARITY OF CICERBITA ALPINA
LONG-TERM VEGETATION CHANGE
There are no fossil palynological records of Cicerbita alpina in Britain and, since the identification
of pollen of the Compositae at species level is rarely possible, it is unlikely that any such evidence
will be forthcoming. It is possible that, for unknown phytogeographical reasons, the species
reached only the restricted block of mountains south-east of the Cairngorms during the early post-
glacial. Equally, it may have been more widespread at first but became progressively restricted to
its present day localities. The total absence of the species from cliff-ledge vegetation elsewhere in
Britain points tentatively towards the former possibility.
C. alpina appears to have a narrower ecological amplitude in Britain than in more central parts
of its range, a circumstance it shares with certain other arctic-alpine species such as Cystopteris
montana. Whether the limited tolerance of this species in Britain is due to our Atlantic climate,
some process of genetic attenuation, or some other reason, is unclear. It is apparent, however, that
C. alpina did occur at lower altitudes in Scotland in the first few decades of the 19th century, and it
may once have occurred in sub-montane woodland in situations closer to its typical continental
habitats than the present day sites. The upland woods of eastern Scotland began to be exploited on
a large scale only in the late 17th century, and many now almost treeless glens such as upper Glen
Muick and Glen Isla are known to have possessed substantial woods of birch, juniper and pine in
the past. The introduction of sheep farming in the late 18th century resulted in the clearance or
neglect of many open, upland woods which, because of grazing and browsing, failed to regenerate.
The situation was further exacerbated when the stocking of the highland glens with red deer
became profitable in the 1850s and 1860s. Glens Muick, Callater, Isla (including Caenlochan and
Canness) and Doll all became deer forests during the middle to late 19th century. Under these
circumstances it is easy to imagine how a tall, palatable plant like C. alpina could be progressively
restricted to cliff sites inaccessible to deer and sheep. Botanical lore has it that red deer regard C.
alpina as a great delicacy and, having developed a taste for the plant, they will readily return to its
localities in subsequent years for another ‘bite of lettuce’. Although C. alpina has long been a
decidedly infrequent treat for Scottish deer, we have direct evidence from the ill-fated colony in
Canness Glen to underline the extreme vulnerability of this species to grazing.
C. alpina could never have been a common montane cliff plant in Scotland, for suitable cliff sites
are relatively few and highly localized. Since its discovery in 1801, it is doubtful whether the total
British population has ever exceeded a few hundred plants and, apart from a few apomictic species
of hawkweed, it is probably the rarest mountain vascular plant in Britain. The fundamental cause
of the present day rarity of C. alpina can be attributed to long term change in the vegetation of the
upland glens whose land-use militates against the plant’s survival except on inaccessible crags.
Nevertheless the plant has continued to decline even in its remote refuges, for which other factors
must be responsible. Some of the possible causes are examined below.
OVER-COLLECTING AND OTHER HUMAN DISTURBANCE
Cicerbita alpina, like many mountain plants, was heavily collected in the past. Gardiner (1848)
noted that it was “eagerly sought for on account of its being one of the rarest and stateliest of our
native alpine plants’’. Barton (1858) regarded it as “‘a great prize’. The records suggest, however,
that many who went in search of the plant failed in their objective or were deterred from collecting
by the inaccessibility of its haunts. The herbaria of Oxford University Botany Department (OXF),
the British Museum (Natural History) (BM), the Royal Botanic Gardens, Edinburgh (E) and the
Dept of Botany, Aberdeen University (ABD) have been examined for dated gatherings and,
although these collections may represent only a fraction of the total, some general conclusions can
be drawn (Table 5).
The data presented in Table 5 suggest that collecting was at its most intensive in the middle
decades of the 19th century and that the Glen Doll plants were by far the most intensively
collected. Heavy collecting is not necessarily evidence for the cause of a plant’s decline, although it
is tempting to see some correlation between the very precise locality details given for Glen Doll by
Gardiner (1848) and the rarity of the plant there today. Very few herbarium specimens, however,
include portions of root or rhizome, and this species is unlikely to have been popular with alpine
gardeners, for its ungainly habit and aggressive growth make it unsuitable for the rockery. Most
140 P. R. MARREN, A. G. PAYNE AND R. E. RANDALL
TABLE 5. ANALYSIS OF DATED GATHERINGS OF CICERBITA ALPINA FROM SCOTLAND AT
ABD, BM, E AND OXF
pre- post-
(a) Decade 1820 1820s 1830s 1840s 1850s 1860s 1870s 1880s 1890s 1900s 1910s 1920s 1930s 1940
Number of
gatherings 1 3 19 11 5) jin 11 11 4 i} 3 3 3 2
(b) Locality ‘Glen Clova’ Glen Doll Caenlochan Coire Kander Lochnagar Canness
Number of
Gatherings 12 4] 21 + 11 4
collectors seem to have been satisfied with a flowering stem and a radical leaf. Since such stems are
produced annually from a perennating organ and the plant probably does not rely on regular seed
production to maintain its numbers, it is unlikely that collecting ever made much difference in the
long term.
Disturbance by trampling or photographic ‘gardening’ takes place on some sites today.
Bootprints on and below the Caenlochan ledge are not infrequent, although those who use this
ledge may not always be botanists (see next section). Two of the Lochnagar sites lie on a classic but
now little-used climbing route, but we have no evidence that climbers damage the plants.
GRAZING AND CUTTING
The basal leaves of Cicerbita alpina are almost succulent and the plant’s extreme sensitivity to
grazing is well-attested. Its restriction to cliff ledges and other inaccessible sites is attributed to the
attentions of red deer and sheep, but even remote ledges are not always free from grazing pressure.
Boyd (1879) saw several sheep apparently trapped on precipitous ledges in Corrie Kander. Adam
(1930) counted 60 goats on the same ledges and Lochnagar, whose name means ‘Loch of the
Goats’, also held a substantial goat population in the past. Mountain hares and voles seem to find
little difficulty reaching cliff ledges, probably aided by ramps of late snow, and hare damage was
suspected at Caenlochan in 1969 (F. H. Perring, pers. comm.). The authors have seen voles and
hares in a Lochnagar colony. It is believed that deer occasionally manage to reach and graze the
Caenlochan ledge in exceptionally late seasons when snow forms a ramp below the C. alpina
colony at the time when it is coming into leaf. This appears to have happened in 1977 when the
plants on the near side of the ledge showed clear signs of nibbling. The plants on this ledge were
also cut in 1975 but probably not by deer. On 7th August 1975, the flowering stems of the sow-
thistle and those of neighbouring hogweeds were neatly severed about 30 cm from the ground.
Lying amid the ruins was a whisky bottle whose former contents had clearly been consumed very
recently! Such occasional events clearly influence the plants’ ability to flower and set seed, but
whether or not they are contributing factors to the apparent long-term decline is less certain.
FAILURE TO SET RIPE SEED
Cicerbita alpina is pollinated by nectar-seeking insects such as bees and butterflies, which possess
sufficiently long tongues to reach the hidden, deep-seated nectaries (Willis & Burkhill 1902). One
of us (A.G.P.) has seen Small Tortoiseshell butterflies (Ag/ais urtice) fluttering about and settling
on the flowers of a Lochnagar colony. The flowering period is comparatively late and, in some
years, perhaps most years, the flowers fail to set ripened seed. In 1972 and 1974, Dr G. Miller
collected C. alpina seed from the Caenlochan colony, under licence from the Nature Conservancy
Council, but it failed to germinate and appeared to be shrunken and not fully formed. Equally
malformed, unripened seed was found at Lochnagar in 1979 and 1980. No conclusions can be
drawn from so few collections and C. alpina evidently does succeed in ripening seed in Scotland on
occasion, since both young plants and seedlings have been reported recently from Lochnagar and
Coire Kander (Hobbs 1980) and at least one herbarium specimen (BM) was grown from wild seed.
CICERBITA ALPINA IN BRITAIN 141
The extreme isolation of the Scottish colonies of C. alpina must mean that cross-fertilization
between populations is unlikely to occur, but whether this circumstance might have led to some
self-infertility through reduction of the gene-pool is not presently known. From the clumped habit
of the plants at Caenlochan, it is possible that this colony, at least, is a clone (G. Miller, pers.
comm.).
NATURAL DISASTERS
The present-day localities of Cicerbita alpina are vulnerable to the extreme conditions associated
with mountain gullies and ledges. Strong winds and storms batter the tall inflorescences in late
summer, while some localities are particularly prone to flash floods, rock-falls and drought. A
rock-fall reduced the size of the largest population on Lochnagar for a time, and the Glen Doll
localities have suffered a number of severe falls, some of which may have extirpated a colony. The
vigorous, rhizomatous growth of C. alpina normally enables the plant to recover well after such
events but in small or weakened populations this capacity for recovery may be diminished, and the
isolation of most sites means that natural recolonization is unlikely once a colony has become
extinct.
CONCLUSIONS
Cicerbita alpina is usually a plant of sub-montane woods, and does not appear to be well adapted to
Scottish mountain cliff conditions. It is dependent on pockets of deep soil in sheltered, humid
conditions which approximate to that of woodland. Here, erratic flower and seed production,
natural spates, droughts, strong winds and rockfalls, and occasional grazing and human
disturbance continue to affect the few remaining populations. Of the 15 or so populations recorded
since 1801, only seven are known to survive. The losses have all taken place in the Angus glens,
mainly on sites of relatively low altitude where the populations were perhaps more vulnerable to
occasional grazing. Loss of natural woodland, which characterizes the land-use history of the
highland glens from the 17th century onwards, may have confined this species to remote,
inaccessible sites and possibly reduced its usual insect pollinators. The Glen Doll populations may
have been over-collected in the 19th century but the main reason for their decline appears to have
been chance events such as spates and rockfalls. On the other hand, there is no reliable evidence to
suggest that the total populations in Coire Kander and the north-eastern corrie of Lochnagar have
greatly declined since their discovery. We are unlikely to witness the extinction of C. alpina in
Scotland in the immediate future, but the long-term prospects of the species are not favourable. C.
alpina was given legal protection from collecting or wilful destruction in the Conservation of Wild
Creatures and Wild Plants Act of 1975 and the Wildlife and Countryside Act of 1981. The Scottish
localities all lie within Sites of Special Scientific Interest; two are also within the Caenlochan
National Nature Reserve, whilst Lochnagar is a nature reserve of the Scottish Wildlife Trust.
However, in order to conserve the plant we require a better understanding of its biology. A useful
avenue of research would be to establish the breeding mechanism of the species in Britain in order
to test whether or not the evident infertility of the seed has a genetic cause. If it has, artificial
cross-pollination between the isolated populations might produce some improvement in vigour
although the remoteness and inaccessibility of the sites of this species provide practical problems.
Introductions or re-introductions from wild propagules could be attempted in suitable areas which
have been fenced against grazing animals, and an interesting experiment would be to introduce the
species into an ungrazed, sub-montane wood. A possible site is the National Nature Reserve of
Morrone Birkwood, near Braemar, in which a number of arctic-alpine species such as Juncus
alpinus, J. triglumis and Potentilla crantzii still grow in open birch woodland, in conditions recalling
early post-glacial times.
ACKNOWLEDGMENTS
The authors are grateful to the following for helpful advice and information or for allowing us to
use unpublished material: D. Bayne, A. O. Chater, Ursula Duncan, Lynne Farrell, J. A. Forster,
Claire Geddes, E. Jensen, G. Miller, D. A. Ratcliffe, J. Grant Roger and Rosalind Smith.
142 P. R. MARREN, A. G. PAYNE AND R. E. RANDALL
REFERENCES
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Edinb., 30: 246-249.
Ba.rour, J. H. (1848). Notes of a botanical excursion with pupils, to the mountains of Braemar, Glenisla and
Clova, and to Ben Lawers. Edinb. Phil. J., 45: 122-128.
Ba.rour, J. H. (1875). Scottish Alpine Botanical Club field meeting, 1874. Trans. Proc. bot. Soc. Edinb., 12:
204-208.
Barton, T. (1858). Glens Canlochan and Doll. Phytologist, N.S., 2: 581-590.
Boyp, W. B. (1879). Scottish Alpine Botanical Club field meeting, 1878. Trans. Proc. bot. Soc. Edinb., 13:
103-106.
DickiE, G. (1860). The botanist’s guide to the counties of Aberdeen, Banff and Kincardine. Aberdeen.
Don, G. (1813). Account of the native plants in the country of Forfar, and the animals to be found there, in J.
Headrick. General View of the Agriculture of the County of Angus or Forfarshire, reprinted in Notes R.
Bot. Gdn. Edinb. , 3: 208-255 (1905).
Druce, G. C. (1904). The life and work of George Don. Notes R. Bot. Gdn. Edinb., 3: 53-88.
Ewan, W. A. (1933). Cairngorm Club Journal, 13: 90-93.
Ewinc, P. (1887). Notes on some alpine plants from Forfarshire and Aberdeenshire. Proc. Nat. Hist. Soc.
Glasgow, 1: 286-289.
FERREIRA, R. E. C. (1958). A comparative ecological and floristic study of the vegetation of Ben Hope, Ben
Loyal, Ben Lui and Glas Maol in relation to geology. Ph.D. thesis, University of Aberdeen.
GARDINER, W. (1848). Flora of Forfarshire. Edinburgh and London.
Ho.peEN, A. E. (1952). Plant life in the Scottish Highlands. Edinburgh. ms
Hosss, R. (1980). The flora of the N.E. coire of Lochnagar. A study of certain species undertaken for the
Scottish Wildlife Trust. Brathay Exploration Group Field Studies Report 37.
Hun t_ey, B. (1976). The past and present vegetation of Morrone Birkwoods and Caenlochan National Nature
Reserves. Ph.D. thesis, University of Cambridge.
McVEan, D.N. & RatcuirFe, D. A. (1962). Plant communities of the Scottish Highlands. Monographs of the
Nature Conservancy 1. London.
PERRING, F. H. & FARRELL, L. (1983). British red data books: 1. Vascular plants, 2nd ed. Lincoln.
RANDALL, R. E. (1977). Cicerbita alpina (L.) Wallr. in Scotland 1976-7. Unpublished World Wildlife Fund
report.
RATCLIFFE, D. A. (1958). Cicerbita alpina (L.) Wallr. in Glen Callater. Trans. Proc. bot. Soc. Edinb. , 37: 287-
289.
RAVEN, J. E. & Watters, S. M. (1956). Mountain Flowers. London.
Rocer, J. G. (1943). Ecological notes on Lactuca alpina Benth. (Mulgedium alpinum Less.) in Scotland.
Trans. Proc. bot. Soc. Edinb., 33: 404-406.
Rocer, J. G. (1967). Day excursion to Canness Glen. Scottish Alpine Botanical Club report. Trans. Proc. bot.
Soc. Edinb., 40: 353.
SmiTH, J. E. (1811). An account of several plants, recently discovered in Scotland by Mr George Don A.L.S.
Trans. Linn. Soc. Lond., 10: 333-346.
SmiTH, W. G. (1911). Arctic-alpine vegetation, in TANSLEY, A. G., ed. Types of British Vegetation, pp. 324—
325. Cambridge.
WuiteE, F. B. (1886). Report of an excursion to Glen Shee, Lochnagar and Glen Callater. Scot. Nat., 8: 339.
Wixus, J. C. & Burkitt, I. H. (1902). The phanerogamic flora of the Clova mountains in special relation to
flower biology. Trans. Proc. bot. Soc. Edinb., 22: 109-125.
YounG, W. (1908). The alpine flora and rarer plants of the Glenshee District. Trans. Proc. bot. Soc. Edinb.,
23: 83-91.
(Accepted May 1985)
Watsonia, 16, 143-151 (1986) 143
Natural hybrids between Festuca and species of Vulpia section
Vulpia
M. M. AINSCOUGH, C. M. BARKER and C. A. STACE
Department of Botany, The University, Leicester, LE] 7RH
ABSTRACT
The occurrences and characteristics of the intergeneric hybrids between Festuca rubra L. agg. and Vulpia
bromoides (L.) S. F. Gray and V. myuros (L.) C. C. Gmelin (Poaceae) are detailed. The hybrid of V.
bromoides has been found on five occasions in three localities in England, involving both F. rubra and F.
nigrescens as the other parent. The hybrid of V. myuros has been found on five occasions in four localities in
England and Wales, and once in Holland, also involving both F. rubra and F. nigrescens as the other parent.
Studies of meiosis in the hybrids show that the chromosomes of F. rubra can exchange genetic material with
those of both V. bromoides and V. myuros, although both hybrids are very highly sterile. The significance of
these facts to the evolution of F. rubra agg. is discussed.
INTRODUCTION
THE GENUS FESTUCA
Festuca contains several hundred species, which were placed in six sections by Hackel (1882). One
of these sections, Ovinae (= Festuca), consists largely of the F. rubra and F. ovina aggregates, and
included eleven species in Hackel’s classification. Resulting from the extensive taxonomic splitting
which has taken place in the last 100 years within the above two aggregates, Markgraf-Dannenberg
(1980) recognized 129 species (out of a total of 170 for the genus) that are referable to section
Festuca. 91 of these fall into F. ovina sensu Hackel and 21 into F. rubra sensu Hackel.
14 of the latter 21 form our concept of the F. rubra aggregate. These are species 65 to 78 in Flora
Europaea (Markgraf-Dannenberg 1980), which correspond to Hackel’s F. rubra subspp. eu-rubra,
pyrenaica, dumetorum and nevadensis. Five of Markgraf-Dannenberg’s 14 species occur in Britain:
F. rubra (with five subspecies: rubra, litoralis (G. F. W. Meyer) Auquier, arenaria (Osbeck) Syme,
pruinosa (Hackel) Piper and juncea (Hackel) Sod), F. richardsonii Hooker, F. juncifolia St
Amans, F. diffusa Dumort. and F. nigrescens Lam. All members of the F. rubra aggregate are
perennials with extra-vaginal non-flowering shoots (often as rhizomes), and have leaf-sheaths
which are closed to the mouth (hence lacking infolded thin margins). Chromosome number reports
vary from diploid to decaploid (2n = 14, 28, 42, 56, 70); the hexaploids are by far the commonest
and octoploids are the only other frequently occurring plants. Some taxa do exist at more than one
ploidy level, but a number of reports of this phenomenon are based on misidentifications. All the
taxa are chasmogamous and behave as outbreeding diploids; figures above 10% self-fertility are
very exceptional (Auquier 1977; Barker & Stace 1982).
Taxa of the F. rubra aggregate are the only members of Festuca which have been found to
hybridize with the genus Vulpia.
THE GENUS VULPIA
Vulpia consists of about 20 species, which are currently placed in five sections (Cotton & Stace
1977; Stace 1978, 1981). Species of section Vulpia differ markedly from all species of Festuca in
their annual habit, cleistogamous florets with usually only one or two stamens, and usually
markedly unequal glumes. In addition they have overlapping leaf-sheaths and only intravaginal
innovations, and are fully self-fertile. Each species is characterized by a single chromosome
number at the diploid (e.g. V. bromoides (L.) S. F. Gray), tetraploid (e.g. V. ciliata Dumort.) or
hexaploid (e.g. V. myuros (L.) C. C. Gmelin) levels (2n = 14, 28, 42).
144 M. M. AINSCOUGH, C. M. BARKER AND C. A. STACE
However, species of the other four sections approach Festuca more closely, but to varying
degrees. Species of the western Mediterranean section Loretia (Duval-Jouve) Boiss. are highly
self-incompatible diploids having chasmogamous florets with three large anthers, and one species
(V. sicula (C. Presl) Link) is a perennial. In other respects (e.g. the markedly unequal glumes) V.
sicula is easily separable from Festuca, but without such a combination of characters it is impossible
to separate the two genera. However, their taxonomic distinctness or otherwise is not a matter for
discussion here.
Species of the remaining three sections of Vulpia are intermediate between those of sections
Vulpia and Loretia in several ways. Sections Spirachne (Hackel) Boiss. and Apalochloa (Dumortt.)
Stace each contain one diploid self-fertile annual species, exhibiting semi- chasmogamous and
chasmogamous flowering respectively. Section Monachne Dumott. contains four semi-chasmoga-
mous self-fertile annual species, all but V. fasciculata (Forsskal) Fritsch, which is tetraploid, being
diploids.
INTERGENERIC HYBRIDS (X FESTULPIA MELDERIS EX STACE & COTTON)
An extensive programme of crossing carried out by C. M. Barker (Barker & Stace 1982, 1984, in
press) obtained the following intergeneric combinations (female parents first): V. sicula (diploid)
x hexaploid F. rubra agg.; hexaploid F. rubra agg. Xx V. sicula (diploid); V. sicula (diploid) x
octoploid F. rubra agg.; V. fasciculata (tetraploid) x hexaploid F. rubra agg.; and V. myuros
(hexaploid) X hexaploid F. rubra agg. Hence the Vulpia parents included species from three
different sections (all of those used), both annuals and perennials, and all three ploidy levels. The
Festuca parents included both ploidy levels used and six of the eight segregate taxa used: F. rubra
subspp. rubra, arenaria and pruinosa, F. nigrescens, F. diffusa and F. juncifolia. Earlier, R. Cotton
had synthesized the hybrid V. fasciculata x F. rubra subsp. rubra with considerable success (Stace
& Cotton 1974).
In Barker’s experiments two of the 20 plants obtained of hexaploid F. rubra agg. x V. sicula
(diploid) (both involving the same plant of F. nigrescens as female parent) were unexpectedly
heptaploid, presumably resulting from unreduced female gametes. These two plants exhibited
about 60% pollen stainability, though no seed-set has been detected. All the other hybrids
possessed the expected intermediate chromosome numbers and were highly sterile, with very
rarely as high as 1% pollen stainability.
Morphologically the artificial hybrids were more or less intermediate between their parents in
appearance and, in the cases involving Vulpia myuros and V. fasciculata, they closely resembled
wild hybrids.
Wild hybrids are known between the Festuca rubra aggregate and three species of Vulpia: V.
fasciculata, V. bromoides and V. myuros.
Many details of the natural intergeneric hybrids involving V. fasciculata (as V. membranacea
TABLE 1. DIAGNOSTIC MEASUREMENTS OF FOUR xX FESTULPIA COMBINATIONS AND THE
1961 COLLECTION FROM LITTLEHAMPTON
The ranges given for F. rubra X V. myuros and F. rubra x V. bromoides are derived from the wild collections
detailed in Tables 4 and 2 respectively. Figures are ranges of means per inflorescence. Glume lengths include
awns; lemma lengths exclude awns and refer to only the first and second lemmas of each spikelet; awn lengths
refer to the longest in each spikelet.
F. rubra F. rubra Littlehampton F. rubra F. juncifolia
x x hybrid x x
V. myuros V. bromoides 1961 V. fasciculata V. fasciculata
Lower glume length (mm) 1-5-3-3 2:0-3-4 3-0-3:-7 2:4—4-4 5:2-8:-0
Upper glume length (mm) 3-2-5-0 3-4-5-9 6:2-7:°4 3-5-7:2 8-0-11-5
Glume ratio 0-48-0-7 0-5-0-75 0-45-0-58 0-55—0-69 0-6—0-69
Lemma length (mm) 4-5-6:2 4:5-7-0 7:5-8:4 6:0-9-5 9-5-10-5
Awn length (mm) 3-0-6-0 3:2-6:0 5:7-7:2 2:0-5-5 3-5-5-0
1-6-1-7 1-5-2:0 1-5-2-0
Anther length (mm) 0-6-1-5 0-8—1-7
NATURAL FESTUCA xX VULPIA HYBRIDS 145
(L.) Dumort.) were given by Stace & Cotton (1974) and Willis (1975), and further data have
been provided by Barker & Stace (1984, in press), and by Stace & Ainscough (1984) on the
progeny of one of these hybrids. A summary of diagnostic measurements is provided in Table 1.
The number of taxa of the F. rubra aggregate which have produced natural hybrids with V.
fasciculata is uncertain, but both hexaploids and octoploids are involved, producing pentaploid
and hexaploid hybrids respectively. On the basis of morphological characters Stace & Cotton
(1974) deduced that the British hexaploid and octoploid Festuca taxa were F. rubra and F.
juncifolia respectively, and they named the two hybrids x Festulpia hubbardii Stace & Cotton
and X F. melderisii Stace & Cotton respectively. However, it is possible that the octoploid parent
of X F. melderisii was actually F. rubra subsp. arenaria, which exists as both hexaploids and
octoploids, and in fact the precise distinction between F. rubra subsp. arenaria and F. juncifolia is
unclear. Which hexaploid taxa of F. rubra agg. are involved in the parentage of x F. hubbardii in
Britain is also uncertain, but the restricted habitat (sand-dunes) involved suggests only F. rubra
subspp. rubra and arenaria.
Pentaploid xX Festulpia hubbardii occurs in many places on the coasts of southern Britain, from
E. Kent to S. Lancashire, while hexaploid x F. melderisii has been confirmed from only two
localities in south-eastern England (Fig. 1).
This paper presents data on natural hybrids between the Festuca rubra aggregate and the other
two species of Vulpia: V. bromoides and V. myuros. Their known occurrences in the British Isles
are shown in Fig. 1.
FESTUCA RUBRA AGG. X VULPIA BROMOIDES
OCCURRENCE
1. Coastal sand-dunes at Littlehampton, W. Sussex, 1961, A. Melderis (BM) (Melderis 1965),
close to plants of both parents as well as of V. fasciculata, x F. hubbardii and X F. melderisii.
Festuca rubra subspp. rubra and arenaria and F. juncifolia occur in the immediate vicinity. We
have not traced a voucher specimen of the original collection, but a garden-grown specimen
(coll. 1964) of the original plant is in BM. We have no doubt that it is F. rubra x V. fasciculata.
Its measurements fall within the range of the latter hybrid and outside those of F. rubra x V.
bromoides (Table 1). Most of the upper glumes have a distinct awn and the pedicels are
distinctly dilated distally (both characters of the V. fasciculata hybrid but not of the V.
bromoides hybrid). Searches for F. rubra X V. bromoides in the locality have not been
successful.
2. Ten plants on fixed shingle at Shingle Street, E. Suffolk, 1969, P.J.O. Trist (K, det. C. E.
Hubbard). The only other species of Vulpia in the area is V. myuros, whose hybrids with F.
rubra are of different appearance. The hybrid has been sought there in several subsequent years
by Trist, but was refound only in 1973 (two plants) and 1976 (one plant). A specimen collected
on the last occasion is in cultivation at Leicester. All material of F. rubra near the hybrid at
Shingle Street is subsp. rubra, though subsp. litoralis occurs within a few hundred metres.
3. On ballast by railway sidings, East Ella, Hull, S.E. Yorkshire, 1980, J.E.L. Spencer (LTR). It
was growing with Vulpia bromoides, V. myuros and Festuca nigrescens, and was determined by
C. A. Stace as probably F. nigrescens x V. bromoides. It has been searched for in subsequent
years by Spencer, but without success.
4. One plant on cinders of a disused railway line at Leire, Leicestershire, 1983, C.A. Stace (LTR).
This is being cultivated at Leicester. Festuca rubra subsp. rubra and F. nigrescens occur in close
proximity, but V. bromoides is the only species of Vulpia in the area. The Festuca parent is
probably F. rubra subsp. rubra as the hybrid has a limited rhizome development.
CHARACTERISTICS
Festuca rubra X Vulpia bromoides closely resembles X F. hubbardii in most characters.
Vegetatively the characters of the Festuca parent are dominant: the plants are densely caespitose
perennials with many non-flowering shoots, some of which are extravaginal in origin and in some
plants form short rhizomes. The leaf-sheaths are closed almost to the mouth (not overlapping as in
Vulpia), and are pubescent in those hybrids involving a pubescent Festuca parent (always glabrous
M. M. AINSCOUGH, C. M. BARKER AND C. A. STACE
146
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NATURAL FESTUCA x VULPIA HYBRIDS 147
in V. bromoides). The leaf-blades are intermediate in anatomy and morphology, being more
slender and with less sclerenchyma than in F. rubra. The floral characters are intermediate. The
inflorescence is more one-sided than in F. rubra, and the awns are longer. The glume ratio is the
same as in V. bromoides, i.e. lower than that in F. rubra. There are three stamens, as in F. rubra,
but they are indehiscent and shorter than those of F. rubra, though longer than the usually single
stamen of V. bromoides.
It is best distinguished from x F. hubbardii by its thin, not distally thickened, pedicels; by the
spikelets usually having only one ovary-less floret at the apex, the lemmas of the other florets
varying little in length; and by the awn-less, rather than distinctly awned, upper glumes. In x F.
hubbardii there are usually two or three apical ovary-less florets, and the lemmas diminish
markedly in length acropetally. In fact there is no Vulpia fasciculata (the parent of x F. hubbardii)
in localities 2—4 above.
The characters of F. rubra x V. bromoides vary not only according to the characteristics of the
Festuca parent, but also with environmental conditions. Living plants grown at Leicester have
shown significant differences in some characteristics from year to year. An indication of the range
of variation shown by this hybrid is given in Table 2.
Chromosome counts have been made of the E. Suffolk (1976) and Leics. (1983) plants; in both
cases the surprising count of 2n = 42 was obtained, whereas 2n = 28 would have been expected.
Stace & Ainscough (1984) have attempted to explain this anomaly on the basis that the two plants
in question are actually backcrosses to Festuca rubra, not F, hybrids. Pollen stainability is less than
1% (0-1-0-7%) in all cases, and no seed-set has been observed.
Attempts to synthesize hybrids between V. bromoides and a range of taxa of F. rubra agg. have
been unsuccessful.
CHROMOSOME BEHAVIOUR
Meiotic analyses were undertaken on the plants from E. Suffolk (collected 1976) and Leics. (Table
3). The former plant gave significantly different results in 1978 and 1984, the later ones agreeing
very closely with results obtained from the Leics. plant in 1984. The 1984 results produced means
of 16-0 and 14-38 bivalents per cell, of which 11-2 and 11-75 respectively were ring bivalents,
amounting to a rather high degree of pairing. Multivalents were very rare or absent. The 1978
results showed a much lower pairing affinity, with a mean of only 7-8 bivalents and many more
univalents, although there were more trivalents, quadrivalents and quinquevalents. More technical
problems were encountered in 1978, and the results are less accurate than those of 1984, but the
differences are so marked as to be undoubtedly real (note, for example, mean numbers of
univalents of 20-3 and 9-7 respectively). Chiasma frequency (26—28 per cell in 1984) is high for a 2n
= 42 hybrid. Despite this, pollen grain stainability is very low and no seed-set has been observed.
TABLE 2. DIAGNOSTIC MEASUREMENTS OF FIVE WILD COLLECTIONS OF FESTUCA RUBRA
AGG. xX VULPIA BROMOIDES
Figures are ranges of means per inflorescence. Glume lengths include awns; lemma lengths exclude awns and
refer to only the first and second lemmas of each spikelet; awn lengths refer to the longest in each spikelet.
Dates indicate year of collection.
Shingle Street
1969 LOTS 1976 Hull Leire
Lower glume length (mm) 2:0-3-1 2-1-3-0 2:5-3-4 2-1-3-1 2-0-2:5
Upper glume length (mm) 3-9-5-8 4-0-4-6 4-5-5-9 3-4—5-2 3-5-4-4
Glume ratio 0-5—0-62 0-5-0-65 0-52—0-62 0:51-0:75 0-53-0-71
Lemma length (mm) 5-7-6:5 4-5-6:2 5:9-7-0 5-1-6-4 5-0-5:-7
Awn length (mm) 5-0-6-0 3-6-5-4 3-2-5:0 3-4-5-1 3-7-5:3
Anther length (mm) 1-1-1-6 1-5-1-7 1-1-1-3 0-8—1-2 0-9-1-5
148 M. M. AINSCOUGH, C. M. BARKER AND C. A. STACE
TABLE 3. MEIOTIC ANALYSES OF FIVE x FESTULPIA HYBRIDS
Figures represent ranges and (in brackets) means. The term ‘chromosome pairs’ represents the number of
bivalents plus a minimum number to take account of the multivalents. Dates represent year of observation.
F. rubra X V. bromoides
F. rubra V. myuros
Shingle St Shingle St Leire x x
Chromosome configurations 1978 1984 1984 V. myuros _ V. diffusa
Vv 0-1 ( 0-1) 0 0 0 0
IV 0-4 ( 0-4) 0 0 0-2 (0-3) 0-1 ( 0-2)
Ill 0-3 ( 1:2) 0-1 ( 0-1) 0 0-1 (0-1) 0-1 ( 0-2)
II (total) 5- a 7-8) 13-19(16-0) 11-18(14-4) 9-18(13-6) 15-20(17-8)
II (ring) 8-18(11-2) 9-15(11-8) 4-16( 9-2) 9-20(14-4)
II (chain) — 1-8 ( 4-8) 1-5 ( 2-6) 0-9 ( 4-4 0-8 ( 3-4)
I 12-27(20-3) 4-16( 9-7) 6-20(13-3) 6-24(13-3) 2-12( 5-4)
Chromosome pairs 7-14(10-0) 13-19(16-1) =11-18(14-4) 9-18(14-3) 15-20(18:-2)
Chiasmata per cell — 22-37(27:9) 20-32(26:1) 16-32(24-3) 26-40(33-2)
FESTUCA RUBRA AGG. X VULPIA MYUROS
OCCURRENCE
1. On rubble by road just east of Mawddach Crescent, Arthog, Merioneth, 1957, P.M. Benoit
(LTR, NMW); and on disused railway track 690 m south-east of this locality, 1970, P.M. Benoit
(LTR). Searched for unsuccessfully in the area in other years. In both cases one plant appeared
with V. myuros and F. rubra subsp. rubra, which are presumed to be the parents even though
both hybrids show no rhizome development. Both are still (1984) grown in P. M. Benoit’s
garden in Barmouth, Merioneth.
2. On cinders in railway sidings immediately north of Stockport Station, S. Lancashire, 1974, R.
Cotton & C.A. Stace (LTR). The only species of Vulpia present was V. myuros, and the
commonest Festuca immediately adjacent was F. nigrescens; these are presumed to have been
the parents.
3. On sandy shingle bank near sea at Snettisham, W. Norfolk, 1974, R.P. Libbey (LTR). A
sample of this plant is in cultivation at Leicester. The commonest related plants in the
immediate vicinity are Festuca rubra subsp. litoralis and Vulpia myuros, which are presumed to
be the parents.
4. Between cobble-stones at old railway station, Woensdrecht, Noord Brabant, Netherlands,
1978, E.J. Weeda (L, LTR). This is the only confirmed example of x Festulpia from outside
Britain. It was determined originally as F. rubra agg. x V. bromoides by C. A. Stace, but it is
probable that V. myuros rather than V. bromoides was involved. It has not been refound. The
only other relevant taxa found at the same site were V. myuros and F. nigrescens, which were
the likely parents.
5. Two plants on disused railway line near Oswestry, Salop, 1983, P.M. Benoit (in litt.). Festuca
rubra subsp. rubra and Vulpia myuros were the only possible parents in the area.
CHARACTERISTICS
This hybrid is extremely similar to the last in all characters, and may not be always distinguishable
from it. However, all specimens seen lack any rhizome development. Unfortunately, whereas
glume ratio is a very important character for separating V. myuros and V. bromoides (as well as V.
fasciculata), the dominance of the subequal glumes of F. rubra produces a similar glume ratio in
hybrids between F. rubra and all three species of Vulpia. The best character for distinguishing the
V. bromoides and V. myuros hybrids is the inflorescence shape, which is very long and narrow in
the latter, but shorter and often more spreading in the former. The lemmas are usually narrower
and more gradually tapering in the V. myuros hybrid. Some characteristics of this hybrid are given
in Table 4. Again, variation is probably an effect both of the different Festuca parent involved and
NATURAL FESTUCA xX VULPIA HYBRIDS 149
TABLE 4. DIAGNOSTIC MEASUREMENTS OF FIVE WILD COLLECTIONS AND ONE ARTIFICIAL
HYBRID OF FESTUCA RUBRA AGG. X VULPIA MYUROS
Rubric as in Table 2.
Arthog V. myuros
*
1957 1970 Stockport Snettisham Holland F. diffusa
Lower glume length (mm) 2-4-3:-2 2-0-2:4 1-5-—3-0 2:0-3:-3 2-3-3:-0 1-3-2-2
Upper glume length (mm) 3:7—4-7 4-0-4-6 3°2-4:5 4-0-5-0 3-9-4-4 3-2-4:3
Glume ratio 0-6-0-7 0-5-0:55 0-48-0-67 0-5-0-67 0:57-0:68 0-45-0-6
Lemma length (mm) 5:3-5:6 4-6-5:5 4-5-5-0 5-5-6:2 5:5-6:1 4:7-6-4
Awn length (mm) 3-6-4:3 3-2—4:2 3-5-4:5 3-0-6-0 3-6—4:-4 4-0-4-9
Anther length (mm) 0-8-1-0 1-0-1-1 0-6—-0-9 1-0-1-3 1-3-1-5 1-0-1-5
of environmental conditions. For example, the longer lemma length of the W. Norfolk hybrid
compared with the S. Lancs. hybrid indicates that in the former case F. rubra subsp. litoralis
(lemmas mostly 6-0—7-5 mm) was the likely parent whereas in the latter case F. nigrescens (lemmas
mostly 5-0—6-5 mm) was involved. On the other hand the Dutch hybrid, probably involving F.
nigrescens, has lemmas as long as those of the W. Norfolk hybrid. Of the two plants collected at
Merioneth (1957 and 1970) one has glabrous and one pubescent leaf-sheaths.
The chromosome number of the W. Norfolk hybrid is, as expected, 2n = 42. The pollen
stainability is c. 0-4¢% and no seed-set has been observed.
Two plants of Vulpia myuros xX Festuca diffusa were synthesized by Barker (1980), and
measurements of the diagnostic features were presented by Barker & Stace (1984) (see also Table
4). In all cases the characteristics were very close to those of the wild hybrids (despite the very
different Festuca parents involved). Pollen was 0% stainable.
CHROMOSOME BEHAVIOUR
Meiotic analysis of the W. Norfolk hybrid (Table 3) showed a slightly lower degree of pairing than
the wild hybrids involving Vulpia bromoides showed in 1984. A mean of 13-6 bivalents, of which
9-2 were ring bivalents, and a mean chiasma frequency of 24-3 per cell, were obtained. In view of
the markedly different results obtained in 1978 and 1984 for the E. Suffolk plant of F. rubra x V.
bromoides, the differences between the V. bromoides and V. myuros hybrids are probably not
significant.
For comparison the results of meiotic analysis of the artificial hybrid Vulpia myuros (female) X
Festuca diffusa are given in Table 3. Pairing affinity in this hybrid is slightly higher than in any of
the wild hybrids mentioned above, emphasizing the point that there are probably no significant
differences between any of the data.
DISCUSSION
Wild hybrids occur in Britain and the Channel Isles in the following combinations: Festuca rubra x
Vulpia fasciculata, F. juncifolia x V. fasciculata, F. rubra X V. bromoides, F. nigrescens X V.
bromoides, F. rubra X V. myuros and F. nigrescens X V. myuros. Probably three subspecies
(rubra, arenaria and litoralis) of F. rubra are involved. The three species of Vulpia represent three
ploidy levels (diploid, tetraploid, hexaploid) and two sections of the genus (Vulpia, Monachne).
Two ploidy levels of Festuca (hexaploid, octoploid) are involved. Because of the cleistogamous or
semi-cleistogamous nature of the Vulpia parents, and the small amount of pollen that they
produce, all the wild hybrids are probably formed by crosses of male Festuca with female Vulpia.
Artificial hybrids (Barker & Stace 1982) include a number of the above as well as crosses
involving a third (self-incompatible, chasmogamous) section of Vulpia (Loretia). In the latter case
reciprocal hybrids were raised, but in the crosses between Festuca and Vulpia sections Vulpia and
Monachne hybrids were raised only when the Vulpia parent was used as female. Whether this is
150 M. M. AINSCOUGH, C. M. BARKER AND C. A. STACE
due to the small amount of Vulpia pollen available for crosses, or to unilateral interspecific
incompatibility, is unknown.
Elsewhere the only recorded occurrences of X Festulpia are F. nigrescens X V. myuros in
Holland, detailed above, and the report by Patzke (1970) of F. rubra subvar. microphylla x V.
membranacea found in 1966 by Peter at Suances, Santander, Spain. According to the
nomenclature of Flora Europaea the latter hybrid is presumably F. nigrescens subsp. microphylla
(St Yves) Markgr.-Dannenb. x V. fasciculata. We have not seen the specimen.
The relative rarity of Festuca <x Vulpia crosses in the wild is probably due largely to the
cleistogamous or semi-cleistogamous and self-compatible floral biology of the Vulpia species in the
British Isles, so that stigmas are rarely available to foreign pollen, together with the very small
amounts of pollen (often none of it air-borne) produced by these species. The lack of records of
hybrids between F. rubra and chasmogamous, self-incompatible species of Vulpia section Loretia
in the Mediterranean region is less easily explained. However, both F. rubra and suitably
experienced field-botanists are much less common there than in north-western Europe.
Festuca rubra X Vulpia fasciculata occurs in most sand-dune areas of south-western Britain and
the Channel Isles where V. fasciculata is common. We have never failed to find it in such places,
although the same is not true of similar localities in East Anglia or Jersey. Hybrids involving V.
bromoides and V. myuros are much less certain in occurrence. The much greater regularity of
complete cleistogamy (compared with the normal semi-cleistogamy of V. fasciculata) in these two
species might explain this. The S. Lancs. hybrid of F. nigrescens x V. myuros was, in fact, found in
a population of V. myuros exhibiting a high (and very unusual) degree of chasmogamy, with
dehiscing anthers and receptive stigmas partially exposed.
In cultivation the hybrids appear not to be very long-lived (a few years only), and not all of them
flower every year. These facts might contribute to the rarity of records in the wild. The E. Suffolk
locality of F. rubra x V. bromoides has yielded hybrids on three occasions (1969, 1973, 1976), but
it is as likely that, in years when unsuccessful searches were made, the hybrids occurred there in the
vegetative state as the locality is especially conducive to hybridization. Moreover the number of
hybrids found dropped from ten to one over the eight-year period, and the hybrid found in 1976
(having 2n = 42) was not an F, plant (Stace & Ainscough 1984).
Unfortunately it might not be possible to distinguish the hybrids involving V. bromoides from
those involving V. myuros on all occasions. The best character is the inflorescence shape. Since the
only two plants of F. rubra x V. bromoides with a known chromosome number are hexaploids, it
remains a possibility that they are F. rubra X V. myuros hybrids. However, in both cases the
inflorescence is very short, quite unlike the elongated panicles of known natural and artificial
hybrids of F. rubra X V. myuros, and in one case (Leics.) there is no V. myuros known in the
vicinity.
Similarly, it is very difficult to distinguish hybrids involving F. rubra from those involving F.
nigrescens (the same is also true of those involving F. juncifolia). If the hybrid is rhizomatous,
obviously F. rubra (not F. nigrescens) is involved, but the known hybrids of F. rubra X V. myuros
from Merioneth and W. Norfolk are not rhizomatous.
Despite the relative rarity of hybridization, and the high degree of sterility of the hybrids, there
remains a considerable degree of genomic homology between species of Vulpia and the Festuca
rubra aggregate. In the hexaploid hybrids involving V. bromoides and V. myuros (Table 3) more
than 14 bivalents were rather regularly observed, and there were often more than 14 ring-bivalents.
Therefore both homogenetic (F-F and V-V) and heterogenetic (F-V) pairing, often with chiasmata
in both chromosome arms, occurs. Similarly, in the hexaploid derivative of F. rubra x V.
fasciculata reported by Stace & Ainscough (1984), both homogenetic and heterogenetic pairing
takes place. The same is also true of artificial hybrids between the diploid V. sicula and hexaploid
F. rubra agg. (Barker & Stace in press). Hence there is genomic homology between F. rubra agg.
and species of all three sections (Loretia, Monachne, Vulpia) of Vulpia investigated.
We interpret the main evolutionary trend in Vulpia as being away from a perennial,
chasmogamous outbreeder (section Loretia) to an annual, cleistogamous inbreeder (section
Vulpia), the former being derived from a Festuca-like ancestor similar in many respects to F. rubra
agg. Whereas F. rubra agg. has developed into a declining polyploid complex, with very few (if
any) diploid representatives still extant, Vulpia has retained a substantial number of diploids as
well as developing tetraploidy and hexaploidy. The apparently ‘closed’ genetic system of F. rubra
NATURAL FESTUCA x VULPIA HYBRIDS 151
agg. is, however, alleviated by the retention of its ability not only to hybridize with Vulpia but also
for its chromosomes to exchange genetic material with those of Vulpia. Stace & Ainscough (1984)
have demonstrated that this level of genomic homology can result in backcrossing and the
introgression of Vulpia genes into F. rubra, which thus retains a partially ‘open’ genetic system
normally out of the reach of a declining polyploid complex. This is a good illustration of the
evolutionary importance of relatively rare hybridization events between superficially dissimilar
taxa, even where the primary hybrids are highly sterile.
ACKNOWLEDGMENTS
We are grateful to P. M. Benoit, R. P. Libbey, G. Renaud-Nooy v.d. Kolff, J. E. L. Spencer and
P. J. O. Trist for kindly sending us material and for supplying valuable information on sites, to
Professor A. J. Willis and the Biological Records Centre for help with the records of V. fasciculata
hybrids, and to D. Halsall for preparing Fig. 1.
REFERENCES
AvuagulgR, P. (1977). Biologie de la reproduction dans le genre Festuca L. (Poaceae), 1. Systémes de
pollinisation. Bull. Soc. r. bot. Belg., 110: 129-150.
Barker, C. M. (1980). Investigation into the relationships and ancestry of Vulpia and Festuca. Ph.D. thesis,
University of Leicester.
Barker, C. M. & Stace, C. A. (1982). Hybridization in the genera Vulpia and Festuca: the production of
artificial F, plants. Nord. J. Bot., 2: 435-444. |
Barker, C. M. & Stace, C. A. (1984). Hybridization in the genera Vulpia and Festuca (Poaceae): the
characteristics of artificial hybrids. Nord. J. Bot., 4: 289-302.
Barker, C. M. & Stace, C. A. (in press). Hybridization in the genera Vulpia and Festuca: meiotic behaviour of
artificial hybrids. Nord. J. Bot.
Cotton, R. & Stace, C. A. (1977). Morphological and anatomical variation of Vulpia (Gramineae). Bot.
Notiser, 130: 173-187.
HackEL, E. (1882). Monographia Festucarum europaearum. Kassel.
MARKGRAF-DANNENBERG, I. (1980). Festuca, in Tutin, T. G. et al., eds. Flora Europaea, 5: 125-154.
Cambridge.
MELpkeRIS, A. (1965). Festuca rubra X Vulpia bromoides, a new hybrid in Britain. Proc. bot. Soc. Br. Isl., 6:
172-173.
PaTzkE, E. (1970). Untersuchungen tiber Wurzel fluoreszenz von Schwingelarten zur Gliederung der
Verwandtschaftsgruppe Festuca Linné. Senkenberg. biol., 51: 255-276.
StAcE, C. A. (1978). Changing concepts in the genus Nardurus Reichenb. (Gramineae). Bot. J. Linn. Soc. , 76:
344-350.
STAcE, C. A. (1981). Generic and infrageneric nomenclature of annual Poaceae: Poeae related to Vulpia and
Desmazeria. Nord. J. Bot., 1: 17-26.
STACE, C. A. & AinscouGu, M. M. (1984). Continuing addition to the gene-pool of the Festuca rubra aggregate
(Poaceae: Poeae). PI. Syst. Evol., 147: 227-236.
STACE, C. A. & Cotton, R. (1974). Hybrids between Festuca rubra L. sensu lato and Vulpia membranacea (L.)
Dumort. Watsonia, 10: 119-138.
Wis, A. J. (1975). Festuca L. X Vulpia C. C. Gmel. = X Festulpia Melderis ex Stace & Cotton, in Stace, C.
A., ed. Hybridization and the flora of the British Isles, pp. 552-554. London.
(Accepted March 1985)
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Watsonia, 16, 153-162 (1986) 153
Hybridization in the genus Atriplex
section Teutliopsis (Chenopodiaceae)
P. M. TASCHEREAU*
Department of Botany, University of Manchester
ABSTRACT
A study based on experimental hybridization, culture and field work reports the occurrence of Atriplex hybrids
in the flora of the British Isles, and discusses the taxonomic implications of these findings. The following
hybrids were synthesized experimentally: Atriplex littoralis x longipes, A. littoralis x praecox, A. littoralis x
patula. Segregation of wild A. littoralis x prostrata hybrids collected from two localities in W. Norfolk, v.c. 28,
was observed in the botanic garden. Some of the segregants were similar in leaf morphology to A. patula, and
others were identical to the formerly recognized taxon A. littoralis var. serrata. Evidence is presented that A.
calotheca (Raf.) Fries, a species endemic to Scandinavia and the Baltic coasts of adjacent countries, did not
originate from A. littoralis x A. prostrata as claimed by G. Turesson. Literature reports of the putative hybrids
A. glabriuscula X littoralis and A. patula X prostrata in the British Isles are probably wrong.
INTRODUCTION
Section Teutliopsis Dum. includes all the native British Atriplex species except A. laciniata L. The
members of this section are morphologically similar, genetically highly variable and phenotypically
plastic. Because of this, the recognition of hybrids can be extremely difficult. Without evidence
from carefully controlled experimental hybrids, assumptions about hybridization in Atriplex can be
no more than speculation.
Turesson (1925) produced the first artificial hybrids in Atriplex. He used two techniques: one,
which he called ‘‘free crossing’’, consisted in surrounding a plant of one species with several plants
of another species; the other was to isolate the inflorescences of the parent species together in the
same pergamin bag. He used these uncontrolled techniques because, as he stated, “. . . castrations
unfortunately cannot be made in the genus Atriplex because of technical difficulties...”
(Turesson 1925). Hulme (1957, 1958), however, succeeded in producing controlled experimental
hybrids.
Gustafsson (1972, 1973a, 1973b, 1974) made a large series of carefully controlled experimental
hybrids between all the members of the A. prostrata group in Scandinavia. He examined the
cytology and fertility of natural and artificial hybrids and variation in hybrid offspring. He later
(Gustafsson 1976) provided morphological descriptions of the species in this group with notes on
the morphology, frequency and distribution of the hybrids in Scandinavia. The Scandinavian
representatives of the group include all those found in the British Isles and an additional species,
A. calotheca (Raf.) Fries, indigenous to the Baltic region. Gustafsson’s studies form the basis for
understanding this complex as it exists in Britain.
Partially fertile artificial hybrids have been made between species of sections Teutliopsis and
Sclerocalymma, but none have been found in nature (Bjérkman et al. 1969, 1971). Nobs (1976)
summarized the results of a number of attempted intersectional crosses (including A. prostrata X
A. laciniata and A. glabriuscula x A. laciniata) which produced strong F, progenies, but these
were less than 10% fertile. The other results were highly variable with some crosses yielding a
strong F, but others producing no seed or only a sub-lethal F,;. None of the crosses, however, was
sufficiently fertile to produce a second generation.
*Present address: Institute for Resource and Environmental Studies, Dalhousie University, Halifax, Nova
Scotia, Canada, B3H 3E2
154 P. M. TASCHEREAU
The present study is based on experimental hybridization and culture work in the botanic garden
and on field studies in Britain between 1974 and 1978. It examines the crossing relationships
between A. littoralis and other members of the section Teutliopsis occurring in Britain and
elsewhere. It reports on segregation in hybrid specimens of the A. prostrata group collected in
northern and western Scotland and it discusses the taxonomic implications of the results of these
studies.
MATERIALS AND METHODS
EXPERIMENTAL HYBRIDS
The technical difficulties of making artificial crosses in Atriplex involve: a) the small flower size (c.
1 mm just before-anthesis); b) the close proximity of the staminate to the pistillate flowers (both
types occur together in tightly compressed glomerules); and c) the ease with which self-pollination
can occur in this self-fertile, primarily autogamous group.
Two characteristics of Atriplex floral development make experimental crossing possible: the
occasional production of isolated axillary pistillate flowers, and the occurrence in some species of a
degree of protogyny.
The plants used as female parents in these experiments were all at least slightly protogynous.
Some also produced exclusively pistillate flowers in the upper leaf axils. The branches of these
plants, all of which would have produced terminal inflorescences, were clipped before the flowers
opened. Only two or three branches were left for controlled crossing. The staminate buds, which in
normal glomerules occurred immediately above the pistillate ones, were removed with fine
forceps. Pollen, freshly collected from mature newly-opened flowers, was applied to the receptive
stigmas of the female parent with a no. 000 fine sable hair brush. Each morning the inflorescences
were examined before 08:00 hours with a x14 lens, and successively forming staminate buds
removed. After each examination the emerging new stigmas were repeatedly brushed with fresh
pollen. The process was continued for about five weeks by which time the first seeds were
beginning to ripen and flower formation had ceased. Plants used as female parents were isolated in
separate, screened greenhouse cubicles. Emasculated plants, not pollinated, did not develop seed.
By these methods, 35 crosses within section Teutliopsis were made, 14 of which produced an F,
generation. The following species were used: A. littoralis L. from England, Scotland, Finland,
Norway, Romania and U.S.S.R.; A. praecox Hilphers from Scotland and Norway; A. prostrata
Boucher ex DC. from England; A. longipes Drejer from England; A. glabriuscula Edmondston
from England; A. calotheca (Raf.) Fries from Norway and Denmark; A. patula L. from England,
Hungary and Argentina. The crossing combinations are given in Table 1. In addition, one
intersectional cross between A. littoralis (Romania) of section Teutliopsis and A. rosea L.
(Romania) of section Sclerocalymma was attempted, but no seed was produced.
NATURAL HYBRID SEGREGATION
A. littoralis x A. prostrata
Three plants of this putative hybrid were collected from two localities in Norfolk. Seed was taken
from each of these plants and sown separately in sterilized compost. A total of 332 F, plants was
scored.
A. prostrata Group Hybrid Derivatives
Seeds of putative hybrid plants believed to involve A. prostrata, A. longipes, A. glabriuscula and
A. praecox from three localities on the northern and western coasts of Scotland were sown in the
botanic garden. About 40 plants from each hybrid were raised.
POLLEN AND SEED FERTILITY
Pollen fertility was estimated as the percentage of well-developed pollen grains deeply stainable
with trypan blue in lactophenol. From 500 to 1,500 grains were counted for each individual. Seed
fertility was estimated as percentage of seeds germinating. In addition to pollen and seed fertility,
seed production was examined in the hybrids and hybrid progeny.
Voucher specimens of experimental hybrids, putative hybrids and hybrid segregants were
deposited in MANCH.
HYBRIDIZATION IN ATRIPLEX 155
TABLE 1. ATTEMPTED CROSSES WITH A. LITTORALIS
Origin of seed is given in parentheses after the species.
No. of No. of crosses Total
attempted producing no. of F,;
Female parent Male parent crosses an F, plants
Diploid 2n=18 Diploid 2n=18
A. littoralis (England) x _ A. prostrata (England) 1 0 0
A. littoralis (U.S.S.R.) X A. prostrata (England) 1 0 0
A. littoralis (Finland) x _ A. glabriuscula (England) 1 0 0
A. littoralis (England) x A. longipes (England) 1 0 0
A. longipes (England) x_ A. littoralis (England) 2 0 0
A. littoralis (Finland) x A. longipes (England) 4 4 45
A. littoralis (Finland) x _ A. praecox (Scotland) 2 2 9
A. praecox (Scotland) x_ A. littoralis (Norway) 4 0 0
A. littoralis (Romania) < A. calotheca (Norway) 1 0 0
A. littoralis (U.S.S.R.) X A. calotheca (Norway) T 0 0
A. littoralis (U.S.S.R.) X A. calotheca (Denmark) 2 0 0
A. praecox (Norway) xX _ A. littoralis (England) 1 0 0
Diploid 2n=18 Tetraploid 2n=36
A. littoralis (Romania) X A. patula (England) 1 1 13
A. littoralis (U.S.S.R.) X A. patula (Argentina) 1 1 32
A. littoralis (U.S.S.R.) X A. patula (England) 5 > 91
A. littoralis (U.S.S.R.) X A. patula (Hungary) 1 1 40
RESULTS AND DISCUSSION
EXPERIMENTAL HYBRIDS
A. littoralis x A. prostrata Group
The results of attempts to cross diploid A. /ittoralis with members of the diploid A. prostrata group
are summarized in Table 1. The pollen fertility of the species and hybrids is compared in Table 2.
Although A. littoralis hybridizes with A. prostrata in nature, as discussed below, attempts to
obtain artificial hybrids were unsuccessful. In all attempted crosses, however, the A. prostrata
parents were inland ruderal biotypes. The natural hybrids involved only the coastal halophytic
biotypes.
TABLE 2. POLLEN FERTILITY (% STAINABLE GRAINS) IN SPECIES,
NATURAL HYBRID PROGENY, AND EXPERIMENTAL HYBRIDS
N=number of plants sampled
0-10 11-20 21-30 31-40 41-50 51-60 61-70 71-80 81-90 91-100 N
Species
A. littoralis - - - = = ts - cs 1 3 4
A. patula ~ _ = = ss = = = ve
A. praecox - - - - a = = = = 1 1
_—_
—
Wild hybrid segregants
A. littoralis x A. prostrataF, —- ~ - 1 1 3 1 3 3 4 16
Experimental hybrids
A. littoralis x A. longipes F, ~ _ 5
A. littoralis X A. praecox F,
A. littoralis x A. patula F,
A. littoralis x A. patula F,
A. littoralis X A. patula F,
lL mep |
— |
ke Te]
Inte o
Ie | BN
|
|
|
|
156 P. M. TASCHEREAU
TABLE 3. A. LITTORALIS x A. PATULA: SEQUENCE OF PLANTS CULTIVATED
1975-1978 FROM A SINGLE CROSS
One cross yielding 40 hybrid plants Number %
Fy
Plants cultivated to maturity 36 90
Plants surviving to produce some seed 12 33
Total seed production of all seed-bearing plants 35
Seeds planted 35 100
F,
Seeds germinating 17 49
Plants surviving to produce some seed 17 100
Total seed production of all seed-bearing plants c.900
Seeds planted 20 2
F,
Seeds germinating 13 65
Attempts to synthesize the hybrid A. littoralis x glabriuscula were also unsuccessful.
A. littoralis was successfully crossed with A. longipes only when Finnish A. /ittaralis was used as
the female parent and English A. longipes as the male parent. Reciprocal crosses using English
plants only were unsuccessful. The F; hybrids were morphologically intermediate between the
parent species and distinctly different from A. littoralis x prostrata hybrids.
A. littoralis (female) was crossed with A. praecox (male) and produced morphologically
intermediate hybrids. Four attempts at the reciprocal cross failed. Crosses using two strains of A.
praecox (one from Norway and one from Scotland) as the female parent produced many well-
formed seeds but none of them germinated.
Unsuccessful attempts were made to cross A. littoralis with A. calotheca, a member of the A.
prostrata group almost entirely restricted to the coasts of Scandinavia. This cross was of particular
interest because of Turesson’s (1925) belief, based on experimental findings, that A. calotheca
itself originated from the hybrid A. littoralis x A. prostrata.
Except for A. littoralis X prostrata, discussed below, no hybrids between A. littoralis and
members of the A. prostrata group have been found in nature.
A. littoralis X A. patula Crosses
All crosses between diploid A. littoralis (female) and tetraploid A. patula (male) were successful
and 176 triploid F,; plants were obtained (Table 1). In cultivation, the F,; hybrids were large (up to
TABLE 4. A. LITTORALIS X A. PATULA: SUMMARY OF CULTIVATED PLANTS AND
FERTILITY CHANGES IN THREE GENERATIONS
Total no. of crosses 8
Total no. F, plants produced 176
Total no. F, plants grown to maturity 140
Total no. F; plants producing seed 18
% seed-bearing F, plants 13
Range of seeds per F, plant (0-) 1-8 (-32)
Total no. of F, plants grown to maturity 42
Total no. of F, plants producing seed 33
% seed-bearing F, plants 79
Range of seeds per F, plant (O—) 15-175 (-—c.300)
Total no. of F; plants grown to maturity 11
Total no. of F; plants producing seed 9
% seed-bearing F, plants 82
Range of seeds per F, plant (0—) c.300-c.1000 (—c.2000)
HYBRIDIZATION IN ATRIPLEX 157
TABLE 5. HYBRID DERIVATIVES IN THE A. PROSTRATA GROUP.
SUMMARY OF CULTIVATION RESULTS
Hybrid Locality Progeny
A. glabriuscula X praecox Tongue, v.c. 108 One of the progeny had bracteoles with stalks
suggesting A. longipes may be involved
A. X kattegatensis Ullapool, v.c. 105 No segregation. Plants very similar to each other
(A. longipes X prostrata)
A. glabriuscula x longipes Oban, v.c. 98 Parental characters apparent in several of the
progeny, i.e., long-stalked, very thick bracteoles
1.5 m high) and robust, but mostly sterile. Pollen fertility varied from 5% to 9% stainable grains
and the seed set varied from one to eight seeds per plant (to 32 in one plant). Only 13% of the
plants produced any seed. Hulme (1957) examined the cytology of this hybrid and reported
irregular meiosis: bivalents and univalents occurred and trivalents were frequent; tetrads contained
from three to five units that varied considerably in size. In later generations (Tables 3 and 4),
pollen fertility, seed production and fertility increased but this was not matched by an increase in
vegetative vigour. The F, and F; plants were weak and morphologically distorted and probably
would not have survived outside of greenhouse cultivation.
The experimental F, hybrid is readily made and vigorous. Therefore, one might expect the wild
hybrid to be frequent in nature. This, however, is not the case. Despite careful searches in several
coastal localities where A. patula and A. littoralis were observed growing together in disturbed
ground, only one putative hybrid population was found.
The wild hybrids were much smaller than the experimental hybrids, with shorter leaves and
much more condensed inflorescences. A characteristic feature of this hybrid was the presence on
most inflorescences of one or two expanded, well-formed, fertile bracteoles that stood out amongst
a mass of compressed, mis-shapen, sterile bracteoles.
NATURAL HYBRIDS
A. prostrata Group
Segregation in Putative Hybrids. Segregation in the putative hybrid offspring was not clear. The
wild hybrids seemed to be hybrid derivatives rather than first generation hybrids and it was not
obvious which of the four species in this group were involved. Their progeny often showed the
same characters as the parents, only in different combinations. Characters derived from the
original parent species were not seen together in any single offspring but could sometimes be
observed in several plants derived from one hybrid. The development of stalked bracteoles on
several plants indicated, for example, that A. Jongipes was involved. One progeny from Ullapool
plants showed little variation and compared well with a relatively stable variant of A. longipes X A.
prostrata that is common on the western coasts of Sweden. The results of cultivation are
summarized in Table 5.
Effectiveness of Cultivation as a Technique. Previous authors have commented on the problems of
using cultivation to determine the parentage of putative hybrids ir the A. prostrata group.
Turesson (1925), who raised up to 200 individuals from putative hybrids in the group, observed
that segregation often took place only as regards subtle, small characters that allowed no
conclusion to be drawn as to the parental species. Gustafsson (1973a) noted that most of the genes
governing the taxonomic characters in this group, even those conditioning the different characters
of the bracteoles, are inherited unlinked.
The following practical reasons limited the use of cultivation to investigate the parentage of wild
hybrids in this group. Firstly, the progeny of hybrids were often extremely luxuriant, producing
tangled masses of branches up to 1 m long, and requiring large areas for cultivation. Secondly, in
this group it was essential to grow all the plants to maturity to study the characters of the
bracteoles. These, however, were very easily lost from the plants. Indoors, the branches of
adjacent plants became tangled and most of the bracteoles fell off in separating them; outdoors,
birds removed most of the bracteoles.
158 P. M. TASCHEREAU
TABLE 6. ATRIPLEX LITTORALIS X A. PROSTRATA NATURAL HYBRIDS,
F, SEGREGATION AND SEED GERMINATION
No. of No. of No. of No. of No. of
% seeds plants prostrata intermediate littoralis sickly
Locality germinating examined types types types plants
Wolferton, Norfolk
76-24A 86.5 82 10 61 5 6
76-24B 86.1 46 4 36 6 0
Burnham Deepdale, Norfolk
76-20 50.7 204 56 85 “) 56
«
~
Occurrence in the British Isles. A. longipes X A. prostrata is one of the most frequent hybrids on
the coasts of Britain. A true-breeding variant is known from northern and north-western Scotland
and Shetland where it occupies a habitat uncolonized by either of the parent species. Gustafsson
(1973b) has made artificial hybrids between this variant and the following species: A. longipes, A.
praecox and A. prostrata, but these hybrids have not been reported from nature. Field studies
indicate that other hybrids and hybrid derivatives involving A. longipes are very frequent in Britain
and of key importance in understanding variation in the A. prostrata group here.
A. littoralis X A. prostrata
Segregation in Putative Hybrids. A. littoralis x A. prostrata putative F, hybrids collected in nature
were relatively fertile. About 70-80% of the bracteoles contained well-developed seed. Both
large-brown and small-black seed morphs were frequent in the same plant and germination of both
morphs was high (80 to 90%). Pollen fertility of F, plants is given in Table 2, and segregation is
summarized in Table 6.
From 3% to 13% of the plants segregated toward A. littoralis, 9% to 27% toward A. prostrata
and from 42% to 78% of the segregants were morphologically intermediate. The percentage of
sickly and distorted plants ranged from 0% to 27%. In all three hybrids the number of progeny that
segregated back toward the parental types left no doubt as to the parentage of the hybrid. Pollen
fertility varied from less than 40% to more than 90%, but in 56% of the plants the fertility values
were less than 80%.
In contrast to hybrid segregation in the A. prostrata group (Gustafsson 1973b), morphological
segregation towards the parent species appears to be combined with the restoration of male
fertility. With few exceptions, plants segregating toward parental types, including the serrata leaf
form of A. littoralis, had pollen with well-formed, equal-sized grains and fertility values of over
90%. Morphologically intermediate plants showed considerable variation both in equality of grain
size and stainability. Most of the plants in this category had pollen grains of unequal size with the
amount of stained pollen ranging from 40% to 74%.
Segregants showing a wide range of character combinations occurred in cultivation and were
observed growing with F, plants in the field. Many of the segregants were largely sterile and often
weak-stemmed, but some were vegetatively very vigorous. The following three variants appeared
in cultivation and were later found to be common where the hybrids occurred in nature:
1. Patula-leaf form (Fig. 1B). Plants with leaves like A. patula that possess the falcate basal lobes
so characteristic of this species.
2. Serrata-leaf form (Fig. 1A). Plants with markedly sinuate-dentate leaves like extremes of plants
formerly called A. /ittoralis var. serrata.
3. Gigantic form. Plants up to 1 m (2 m in cultivation) high with thick stems and gigantic leaves
with ovate-lanceolate, irregularly-lobed laminae up to 15 cm long and 4 cm wide.
Occurrence in the British Isles. A. littoralis and A. prostrata commonly occur together. A. littoralis
is frequently a dominant, forming dense stands, whilst A. prostrata is an associated subdominant.
The two species are reproductively isolated from each other by differences in flowering time. A.
littoralis flowers earlier. Cropping the terminal inflorescence in this species induces re-flowering in
the new branches that arise. The later-flowering branches overlap in flowering time with A.
prostrata. In cultivation, when the terminal inflorescences are removed, the axillary flowers that
HYBRIDIZATION IN ATRIPLEX
weybyd
oaeee
408-3
10. JULY.77
396-6
10. JULY.77
TCT SRL RSET Re SE) FEA DENT REIS SER a a RR)
Ficure 1. Leaf morphology of a variety of F, segregants cultivated from the wild hybrid Atriplex littoralis x A.
prostrata. A, A. littoralis types (boxed) and leaves similar to those of the formerly recognized taxon A. littoralis
var. serrata. B, Leaves morphologically + identical to those of A. patula. C, Various intermediate leaf forms.
D, A. prostrata types.
HERBARIUM
MANCHESTER MUSEUM
Abhiglis Lda XL -prartoale la
_ Kaktaratecl 197-77 ad...
sate, Brame. £). Ay lriale SAY
eckbsclialnt lebfentarn, Nerpeth an...
Coll LB: SFOS F LG yg erovvnvicee Datezuhy= Augi977
Der Tontdeneain. a Kibhey J6-2¥@
CULT. MOS. YOR-S, 394-16, 408-3, 396-6, 395-4
160 P. M. TASCHEREAU
form are largely pistillate. In nature, this phenomenon coupled with protogyny would further
favour out-crossing in A. /ittoralis. The hybrid A. littoralis x A. prostrata is known from the eastern
and western coasts of England in v.cc. 28, 58, 60, and 66. It can be expected in disturbed habitats
where the parent species are present in abundance: banks of estuaries recently dredged, earthen
sea walls less than three years old, and salt marshes disturbed by rabbit cropping.
Resemblance of Hybrid Segregants to A. patula. In cultivation, numerous F, segregants of A.
littoralis X A. prostrata were identical in leaf morphology to A. patula. Plants with this leaf
morphology (Fig. 1B) were later found to occur in populations of wild hybrids in Britain. The
plants bear a close resemblance to A. patula and in the vegetative state are very difficult to
distinguish from that species. I have seen more than one sheet of the hybrid in British herbaria
labelled “A. patula’’.
The mature hybrid plants may be readily distinguished from A. patula by the bracteoles. In the
hybrid these are spongy-thick while in A. patula they are herbaceous and thin. Unlike the hybrid,
A. patula does not grow in the same habitat as A. littoralis. The hybrid tends to be more succulent
than A. patula and the lower leaves and branches on the hybrid are commonly alternate while in A.
patula they tend to be opposite to subopposite.
ORIGIN OF SERRATE-LEAVED VARIANTS OF A. LITTORALIS
Most populations of A. littoralis have some plants with leaves that exhibit a degree of toothing on
their margins. The development of marginal teeth varies from leaves that have only a few short
irregular teeth in their distal portions to leaves that are coarsely and irregularly sinuate-dentate
throughout and may have pronounced basal lobes. In their extreme form, the serrate-leaved
variants of A. littoralis are highly distinctive plants. Hudson (1762), Linnaeus (1771) and pre-
Linnean authors such as Petiver (1713) recognized such variants as distinct species.
For excellent examples, see the following sheets in the British herbarium BM: v.c. 15, Sandwich
Bay, R. Meinertzhagen, 10. VIII.32; Pegwell Bay, A. J. Wilmott, 20/9/1912; v.c. 18, Bank of Thames
at Tilbury Fort. There are also specimens from v.cc. 13, 16, 18. In LD there are specimens of this
variant collected from Borgholm, Oland, Sweden by different collectors in successive years: 1874
by C. F. Elmgqvist; 1912 by B. J. Holmgren; June 1932 by A. Vilke and July 1932 by H. Hylander.
The repeated appearance of this extreme variant as well as less extreme plants amongst the F,
segregants of A. littoralis X A. prostrata suggests that the serrate-leaved plants of A. littoralis
originated through hybridization with A. prostrata. However, cultivation experiments have
demonstrated that the expression of this leaf character is dependent on relatively high nitrogen or
optimum salt concentration in the soil (Ahmad in Taschereau 1979). For example, 50 mM NaCl in
the nutrient led to the development of leaves with marked serrations but increasing the salt
concentration to 400 mM NaCl produced plants with entire leaves. Control plants, grown in diluted
standard nutrient without salt, produced leaves with a few short teeth such as are commonly found
in A. littoralis populations.
Moss & Wilmott (1914) and various other authors have treated plants with more or less serrate
leaves as a taxonomic variety: A. littoralis L. var. serrata (Huds.) S. F. Gray. Such plants occur in
most populations of A. littoralis and the degree of toothing varies greatly within and between
individual plants. Owing to this and the environmental component of the variation and the fact that
there is no consistent correlation with bracteole morphology, it is better to recognize this variation
in a description rather than by means of a formal epithet.
ORIGIN OF A. CALOTHECA
Turesson (1925) stated that Atriplex calotheca probably originated from the cross A. littoralis x A.
prostrata. He based this on presumed artificial hybrids morphologically similar to A. calotheca. The
pollen fertility of these plants was low and a great many of the bracteoles empty. Turesson
suggested that the laciniate-leaved individuals he obtained originated from crosses of A. prostrata
with the serrate-leaved variant of A. littoralis.
I was unable to find specimens in LD or S of the plants Turesson assumed represented the hybrid
A. littoralis * A. prostrata. The photograph, however, in Fig. 8 and the drawings in Fig. 9 of
Turesson’s (1925) paper as well as the description are of specimens that are clearly identical to A.
calotheca. They bear no resemblance to the plants I have described as A. littoralis x A. prostrata.
The plants that Turesson described as this hybrid may have arisen by segregation from A. prostrata
HYBRIDIZATION IN ATRIPLEX 161
plants that were themselves of hybrid origin involving A. calotheca. The following facts support
this explanation. Firstly, hybrid plants in the A. prostrata group can resemble one or other of the
parents (Gustafsson 1973a). When the seeds from such plants are cultivated, the effects of
hybridization usually become evident in some of the progeny. Secondly, Turesson did not succeed
in emasculating the parent plants, and in his attempt to obtain hybrids between A. prostrata and A.
littoralis by ‘free crossing’, he harvested only the A. prostrata plants. Thirdly, introgressive
hybridization between A. prostrata and A. calotheca is a common phenomenon in western
Scandinavia and extensive hybrid swarms are frequent throughout the entire range of A. calotheca
(Gustafsson 1976). The exact geographical origin of the plants Turesson used as parents is
unknown, but hybrids between A. prostrata and A. calotheca are common at the sites Turesson
investigated (M. Gustafsson pers. comm. 1975).
ERRONEOUS AND UNCONFIRMED REPORTS OF HYBRIDS
The identification of hybrid material in Atriplex, without evidence from experimental studies, is
unsatisfactory. The following reports, all unconfirmed, are probably wrong.
A. glabriuscula X A. littoralis
Jones (1975) reported this hybrid based on plants she observed at Gibraltar Point, v.c. 54. My
attempts to synthesize it were unsuccessful. Jones’s report is probably incorrect because A.
glabriuscula, one of the putative parents, is absent from Gibraltar Point. I searched carefully for it
there in 1975 and the species later proved to be entirely absent from the coasts of Yorkshire,
Lincolnshire and Norfolk, v.cc. 61, 54, 28, 27 (Taschereau 1979). The description of Jones’s
putative hybrid agrees with that of A. littoralis x A. prostrata and both of the parent species of this
hybrid are present at Gibraltar Point. Jones (pers. comm. 1975) was herself not at all certain of her
identification and was not aware of the existence of A. littoralis X A. prostrata.
A. patula X A. prostrata
Jones (1975) reported this hybrid as “‘very doubtfully recorded from v.cc. 3, 10 and 14 and from
Germany on the basis of apparently intermediate specimens’’. Atriplex patula and A. prostrata very
frequently grow together, often with their branches intertwined, in disturbed ground on vacant lots
and demolished building sites throughout the British Isles. The hybrid was synthesized by Hulme
(1957, 1958) who reported that the plants did not resemble any wild plants she had seen. Although
the wild hybrid may yet be found in Britain, Jones’s report is probably wrong. Despite numerous
searches for the hybrid in situations where it might be expected, I have not found it. All specimens
of apparently intermediate morphology, on cultivation, proved to be one of the species, usually A.
prostrata. The basis for at least some reports of plants intermediate between A. patula and A.
prostrata is probably the hybrid A. longipes X prostrata, only recently reported to occur inland in
waste places (Taschereau 1985). Derivatives of this hybrid frequently have leaves with obtuse to
cuneate bases that make them appear intermediate between A. patula and A. prostrata, previously
the only taxa known from this habitat.
A. glabriuscula X A. praecox
Putative hybrids between A. glabriuscula and A. praecox, reported from Tongue, v.c. 108 and
Ullapool, v.c. 105 (Taschereau 1977), remain unconfirmed. Plants grown from seed taken from
these specimens yielded a range of variants (Table 5), none of which showed a clear segregation
toward either of the putative parents. Other reports of this hybrid (Taschereau 1985) are based on
plants clearly intermediate with small leaves morphologically identical to those of A. praecox and
with bracteoles and seeds morphologically like those of A. glabriuscula.
A. glabriuscula X A. prostrata
This hybrid is not as common in the British Isles as Moss & Wilmott (1914) believed and as
suggested by Jones (1975) who stated that intermediates between A. glabriuscula and A. prostrata
“appear to be common in the British Isles where the two species are in contact. This occurs when
weedy habitats are introduced into maritime areas by landslides or the building of sea-defences
etc.” Specimens I examined in BM identified by A. J. Wilmott as this hybrid were variants of A.
glabriuscula.
162 P. M. TASCHEREAU
ACKNOWLEDGMENTS
Barbara Hulme’s important pioneer experimental hybridization work, except for a brief note,
remains unpublished. From her thesis, I obtained much valuable information. I am grateful to her
for allowing me to copy it and for her hospitality and help during my visit. I gratefully acknowledge
the assistance and hospitality of Mats Gustafsson during my visits to Sweden and I am grateful to
Richard P. Libbey, with whom I found the first wild hybrids in Britain.
REFERENCES
BJORKMAN, O., GAUHL, E. & Noss, M. A. (1969). Comparative studies of Atriplex species with and without B-
carboxylation-photosynthesis and their first-generation hybrid. Carnegie Inst. Washington Yb., 68: 620-
633.
ByORKMAN, O., Noss, M. A. & Berry, J. A. (1971). Further studies on hybrids between C3 and C, species of
Atriplex. Carnegie Inst. Washington Yb., 70: 507-511.
Gustarsson, M. (1972). Distribution and effects of paracentric inversions in populations of Atriplex longipes.
Hereditas, 71: 173-194.
Gustarsson, M. (1973a). Evolutionary trends in the Atriplex triangularis group of Scandinavia, 1. Hybrid
sterility and chromosomal differentiation. Bot. Notiser, 126: 345-392.
Gustarsson, M. (1973b). Evolutionary trends in the Aériplex triangularis group of. Scandinavia, 2.
Spontaneous hybridization in relation to reproductive isolation. Bot. Notiser, 126: 398-416.
Gustarsson, M. (1974). Evolutionary trends in the Atriplex triangularis group of Scandinavia, 3. The effects of
population size and introgression on chromosomal differentiation. Bot. Notiser, 127: 125-148.
Gustarsson, M. (1976). Evolutionary trends in the Atriplex prostrata group of Scandinavia, 4. Taxonomy and
morphological variation. Op. bot. Soc. bot. Lund., 39: 1-63.
Hupson, G. (1762). Flora Anglica, p. 377. London.
Hume, B. A. (1957). Studies on some British species of Atriplex. Ph.D. Thesis, University of Edinburgh.
Hume, B. A. (1958). Artificial hybrids in the genus Atriplex. Proc. bot. Soc. Br. Isl., 3: 94.
Jones, E. M. (1975). Atriplex, in Stace, C. A., ed. Hybridization and the flora of the British Isles, pp. 185-186.
London.
LINNAEUS, C. (1771). Mantissa Plantarum, p. 300. Stockholm. Facsimile reprint Weinheim, (1961).
Moss, C. E. & Wimott, A. J. (1914). Atriplex, in Moss, C. E. The Cambridge British flora, 2: 168-182, plates
172-188. Cambridge.
Noss, M. A. (1976). Hybridizations in Atriplex. Carnegie Inst. Washington Yb., 75: 421-423.
Petiver, J. (1713). Herbarij Britannici. London.
TASCHEREAU, P. M. (1977). Atriplex praecox Hilphers: a species new to the British Isles. Watsonia, 11: 195-
198.
TASCHEREAU, P. M. (1979). Taxonomy of the genus Atriplex in Great Britain. Ph.D. thesis, University of
Manchester.
TASCHEREAU, P. M. (1985). Taxonomy of Atriplex species indigenous to the British Isles. Watsonia 15: 183-209.
Turesson, G. (1925). Studies in the genus Atriplex. Lunds Univ. Arsskr., N.F. Adv. 2, 21 (4): 1-15.
(Accepted April 1984)
Watsonia, 16, 163-172 (1986) 163
Further notes on the flora of Bedfordshire
J. G. DONY and C. M. DONY
9 Stanton Road, Luton, LU4 OBH
ABSTRACT
Vascular plant records from Bedfordshire are updated post-1969, and further notes are added on some species.
Notes are given on the status, native or otherwise, of many species, including the status in Britain of
Petrorhagia prolifera (L.) P. W. Ball & Heywood and Melampyrum arvense L.
INTRODUCTION
The Flora of Bedfordshire (Dony 1953) was the result of 18 years of field study. It has since been
supplemented by a list of additional records, with notes on changes in the flora (Dony 1969), and
by the publication of the Bedfordshire plant atlas (Dony 1976) which showed the distribution of 800
species on a tetrad basis. One of us (J.G.D.) has recently relinquished the B.S.B.I. recordership of
vascular plants in the county (v.c. 30) after 40 years. His successor, C. R. Boon, has been an active
co-worker in recent years. It would now seem to be an appropriate time to make a further appraisal
of the natural vegetation and flora of the county. Since 1969 there have been no important changes
in the boundary of the administrative county. Events of historic interest have been the presentation
to Luton Museum (LTN) of the previously missing sixth volume of Charles Abbot’s herbarium and
his Flora Selecta, a smaller herbarium in four volumes which, unlike his main herbarium, has some
of the specimens localized. The Museum has also received ten volumes of excellent drawings of
plants made by Caroline Gaye (1804-1883). The drawings are all localized and dated, so providing
the first evidence of the occurrence of some plant species in the county.
Since 1969 there have inevitably been many changes in habitat. The most unfortunate has been
the loss of the rich water-meadows at Eaton Socon, Habitat Study 14a (Dony 1953), which have
been covered with imported topsoil and converted into an amenity area. Improvements to the
River Ouse, to make it navigable once more as far as Bedford, have had an adverse effect on its
vegetation. Flitwick Moor, still a very important site when Habitat Studies 38-40 (Dony 1953)
were made 36 years ago, continues to dry out notwithstanding the efforts of the Bedfordshire and
Huntingdonshire Wildlife Trust, which now owns or manages 21 Bedfordshire reserves, to delay
this. Two rich ancient woodlands were clear-felled and turned over to arable cultivation in 1973 —
“plant a tree year” — but fortunately all the other, too few, important woods remain little changed.
The same is true of the chalk downland, the county’s most distinctive natural feature. Barton Hill,
Habitat Study 60 (Dony 1953), has become the second Bedfordshire National Nature Reserve.
CHANGES IN THE NATIVE FLORA
Needless to say, some plant species may be presumed to have become extinct in the county during
the last 30 years, although there is always a hope that they might return, even if not to their
previously known site. Potentilla palustris (L.) Scop., Menyanthes trifoliata L., Stellaria palustris
Retz and Dactylorhiza maculata (L.) So6 have not been seen at their last remaining station,
Flitwick Moor, for over 20 years and it is as long since Sium latifolium L. was seen by the River
Ouse, where it once seemed to be abundant. Orobanche rapum-genistae Thuill. had been
apparently well established at Rowney Warren, at least since Caroline Gaye drew it from there in
1832, but Cytisus scoparius (L.) Link on which it was parasitic failed to survive the severe winter of
164 J. G. DONY AND C. M. DONY
1962-3. C. scoparius has now returned, but without the broomrape. Otherwise, the ten species
(Dony 1953, p. 139) that give a distinctiveness to the county’s natural vegetation are unchanged in
their abundance, with the exception of Carum carvi L., not seen since 1967, and Bunium
bulbocastanum L., a particularly notable Bedfordshire plant species. This has diminished mainly
on account of the continued growth of Luton and Dunstable rather than because of its absence now
as a weed of arable land. It is, however, still common within a limited area, giving no immediate
cause for concern for its future.
THE ALIEN FLORA
Most of the species added below are of alien origin. During the period 1953-1969 an additional 200
alien species weré accounted for (Dony 1969), but in the subsequent period of equal length the
number to be added is less than half of this. There are two main reasons:
(a) Much of the alien flora of Bedfordshire has in the past consisted of plants introduced with wool
shoddy, which was used extensively on sandy soils devoted to market gardening, now a diminishing
agricultural practice in the county. Few of the remaining market gardeners now use shoddy, which
in any case is not so readily available. It is also more costly than before as it is now delivered by
road instead of rail. Its diminishing use has, however, shown that wool aliens may continue to
occur in fields for as long as eight years after the application of shoddy. é
(b) Changes in local government in 1974 allowed the collection of refuse to remain the
responsibility of district councils but its disposal that of county councils. In Bedfordshire there are
now only three relatively large refuse disposal sites instead of eight. The disposal is by a new
method described as ‘earth filling’, which reduces plant growth on the site. Nevertheless, plants of
foreign origin, mainly bird seed aliens, may occur. |
At the same time, fewer visits have been made to the county since 1969 by botanists resident
elsewhere. Until his death in 1976, J. E. Lousley came annually to study wool aliens, which also
brought visits from H. J. M. Bowen, E. J. Clement, C. G. Hanson, J. L. Mason and T. B. Ryves.
Longer, valuable visits were made each year until 1974 by P. M. Benoit, and E. Milne-Redhead
returned to a county of his wartime residence to study its Black Poplars. Other records were made
by A. C. Leslie and C. A. Stace.
LIST OF SPECIES
Listed below are species additional to the flora of Bedfordshire since 1969, together with notes on
the extension of range of some rarer species. Notes are also added giving additional information
for some species to that given in the Flora of Bedfordshire (Dony 1953).
The nomenclature and sequence of families follow Flora Europaea. The genera and species are
listed in alphabetical order within families. Standard herbarium abbreviations are used. ‘!’ after a
station indicates that the taxon has been seen there by one or both of the authors and after a
person’s name indicates that it is a joint record with one or both of the authors. The stations given
for the records are in the part of v.c. 30 now in the administrative county of Bedfordshire, except
when given in the form, e.g. v.c. 24 [Beds.], which represents a station in v.c. 24 but now in the
administrative county of Bedfordshire. This form is used to indicate similar boundary changes.
Tetrad references (e.g. 02B) are those used in Bedfordshire plant atlas (Dony 1976).
Pteridaceae
Pteris cretica L.: garden wall, Luton, 02B, 1981 (LTN), H. B. Souster!
Aspidiaceae
Gymnocarpium dryopteris (L.) Newm.: railway wall, Radwell, 05D, 1972 (LTN). Second record.
This is one of a number of species found in a similar habitat at Leagrave in 1950 (Dony 1953,
p. 197), but only Cystopteris fragilis (L.) Bernh. survived when the wall was reconstructed.
Polystichum falcatum (L.f.) Diels: in basement of house, Dunstable, 02B, 1982, J. Schneider.
P. setiferum (Forssk.) Woynar: Holcot Wood, 04K, 1976 (LTN).
FLORA OF BEDFORDSHIRE 165
Azollaceae
Azolla filiculoides Lam. This has appeared spontaneously in some sites, e.g. at Hatch, 14N, 1981, but
in no case becoming established.
Salicaceae
Populus nigra L. The account of this species given in Dony (1953) is misleading but, thanks largely to
the fieldwork done by E. Milne-Redhead, it is now known in 21 tetrads — but shows no simple
distribution pattern.
Salix pentandra L.: meadow, Milton Bryan, 93Q, 1978 (LTN). Second record.
Polygonaceae
Reynoutria sachalinensis (F. Schmidt) Nakai: derelict railway station, Willington, 1SA, 1973 (LTN).
Rumex conglomeratus Murr. X R. crispus L. (R. xX schulzei Hausskn.): meadow, Langford,
14V, 1972 (LTN), P. M. Benoit!
R. conglomeratus Murr. X R. obtusifolius L. (R. < abortivus Ruhmer.): meadow, Langford,
14V, 1972 (LTN), P. M. Benoit!
Chenopodiaceae
Atriplex muelleri Benth.: Beeston, 14U, 1950 (LTN, RNG), J. E. Lousley!; Maulden, 03N, 1976, E.
J. Clement & T. B. Ryves! Wool alien.
Chenopodium capitatum (L.) Aschers.: spontaneously in garden, Stevington!, 9SY, 1980 (LTN),
Mrs Robinson. Only previous record, 1950.
C. glaucum L.: arable field subject to periodic flooding, Shefford, 13P, 1979 (LTN). This has
appeared regularly for six years. Last record, unconfirmed, c. 1906.
C. nitrariacum F. v. Muell.: Flitton, 03M, 1979 (LTN), det. E. J. Clement. Wool alien.
Kochia scoparia (L.) Schrader var. subvillosa Moq.: Maulden, 03N, 1967 (LTN). Wool alien.
Amaranthaceae
Amaranthus albus L.: Maulden, 031, 1976 (LTN), det. J. P. M. Brenan. Wool alien.
A. cruentus L.: landfill site, Sundon, 02J, 1985 (LTN), det. E. J. Clement. Bird seed alien.
A. deflexus L.: Maulden, 03N, 1969 (LTN). Wool alien.
Phytolaccaceae
Phytolacca acinosa Roxb.: waste ground, Aspley Guise, 93H, 1981 (LTN), J. Morris!
Portulacaceae
Monitia sibirica (L.) Howell: well established, Bedford Cemetery, 05K, 1985 (LTN), C. R. Boon!
The only previous record was as a garden weed, c.1950.
Caryophyllaceae
Petrorhagia prolifera (L.) P. W. Ball & Heywood: south-facing bank and track of dismantled
railway, Potton, 24E, 1974 (LTN), det. J. R. Akeroyd. This has been constant in its population
here for twelve years and it cannot be considered to be a casual. There are records of it, or a closely
allied species, from West Norfolk, v.c. 28, causing us to wonder if this species is, like P. nanteuilii
(Burnat) P. W. Ball & Heywood, a native species in southern England.
Ranunculaceae
Ranunculus lingua L.: Harlington!, 03F, 1983 (LTN), L. G. Adams; R. Ouse, Sharnbrook!, OSE,
1983, Bedfordshire Natural History Society Field Meeting. These are the first records for 100
years of a species doubtfully native in the county.
Papaveraceae
Fumaria capreolata L.: waste ground, Stanbridgeford!, 92G, 1984 (LTN), B. M. Inns, conf. P. M.
Benoit. A casual.
F. densiflora DC. X F. officinalis L. subsp. officinalis: Dunstable, 022A, 1972, P. M. Benoit. For a
description of this hybrid see Stace (1975, p. 135).
166 J. G. DONY AND C. M. DONY
F. muralis Koch subsp. boraei (Jord.) Pugsley: arable field, Pavenham, 95X, 1972 (LTN), P. M.
Benoit!. A casual.
Cruciferae
Eruca vesicaria (L.) Cav.: Bedford R.D.C. refuse tip, OSF, 1970 (LTN). A casual.
[satis tinctoria L.: spontaneously in garden, Oakley!, 05C, 1970 (LTN), Mrs Smart. The last certain
record was in 1695.
Neslia paniculata (L.) Desv.: Bedford R.D.C. refuse tip, OSF, 1971 (LTN). A bird seed alien,
previously recorded in 1876.
Crassulaceae
Sedum dasyphyllum L.: wall of mill, Eaton Socon, 15U, v.c. 30 [Cambs.], 1967, but no longer
there; wall, Flitwick Manor, 03H, 1975 (LTN).
Saxifragaceae
Chrysosplenium alternifolium L. This, perhaps the most surprising native species claimed for
Bedfordshire, is based on correctly named specimens in two herbaria compiled by William
Crouch (1818-1846) and labelled “Lidlington, May 1844, W. Crouch’’. Crouch was aware of the
closely allied C. oppositifolium L. represented by two further specimens correctly named and
labelled ‘“‘Eversholt, May 1843, W. Crouch”. The smaller of these herbaria has 33 specimens
from counties other than Bedfordshire, the names of the collectors being added. Ten were from
Caroline Gaye (1804-1883), who lived five km from Crouch when not away from home as a
governess, but the almost 800 vascular plants she drew included only C. oppositifolium.
Although some doubt must remain with regard to the validity of the C. alternifolium record, it is
based on apparently sound evidence.
Grossulariaceae
Ribes aureum Pursh: chalkpit, Houghton Regis, 02B, 1976 (LTN). Of garden origin, but well
established and increasing.
Rosaceae
Cotoneaster multiflorus Bunge: Bison Hill, Whipsnade, 91Z, 1981 (LTN), det. E. J. Clement.
Remote from houses.
Prunus cerasus L.: edge of wood, Stopsley, 12C, 1983 (LTN). The first certain record, see Dony
(1953, p. 260).
Rosa rubiginosa L.: Home Wood!, 14N, 1971 (LTN), N. Dawson.
Sorbus hupehensis Schneid.: Speedwell Farm, Woburn, 93K, 1981 (LTN). Remote, with no
evidence of being planted.
S. torminalis (L.) Crantz: Maulden Wood, 03V, 1969 (LTN); Home Wood, 14N, 1972; Palmers
Wood, 14H, 1973. The first records.
Leguminosae
Cicer arietinum L.: Bedford R.D.C. refuse tip, OSF, 1971 (LTN). Bird seed alien.
Coronilla scorpioides (L.) Koch: Flitwick, 03G, 1976 (LTN), E. J. Clement & T. B. Ryves. Wool
alien.
Cytisus striatus (Hill) Rothm.: bank of M1 Motorway!, 03C, 1977 (LTN), C. A. Stace. Originally
planted, but now well established.
Hedysarum glomeratum F. G. Dietrich: Flitwick, 03G, 1978 (LTN), det. E. J. Clement. Wool
alien.
Lathyrus grandiflorus Sibth. & Sm.: gravel pit, Cople!, 14E, 1981 (LTN), 7. C. E. Wells. Garden
origin.
Medicago scutellata (L.) Miller: Flitwick, 03G, 1985 (LTN), C. G. Hanson & B. S. Wurzell, det. E.
J. Clement. Wool alien.
Psoralea bituminosa L.: Flitton, 03T, 1981 (LTN), det. D. E. Coombe. Wool alien.
Trifolium lappaceum L.: Maulden, 03N, 1968 (LTN). Wool alien.
T. leucanthum Bieb.: Flitton, 03T, 1979 (LTN), det. E. J. Clement. Wool alien.
FLORA OF BEDFORDSHIRE 167
Ulex minor Roth: remnant of heath, Heath and Reach, 93P, 1985 (LTN), C. R. Boon!. The first
record of a species which is a feature of heaths a short distance away in north Hertfordshire.
Vicia tenuifolia Roth: railway bank, Westoning, 03G, 1971 (LTN). Well established.
Geraniaceae
Erodium gruinum (L.) L’Hér.: Flitton, 03T, 1970 (LTN), E. J. Clement & T. B. Ryves. Woo) alien.
E. stephanianum Willd.: Maulden, 03N, 1971 (LTN). Wool alien.
Malvaceae
Anoda cristata (L.) Schlecht.: Maulden, 03N, 1976 (LTN), det E. J. Clement. Wool alien.
Lavatera punctata All.: Maulden, 03N, 1980 (LTN), H. J. M. Bowen! Wool alien.
L. trimestris L.: roadside, Luton, 01Z, 1982 (LTN). Garden origin and second record.
Malope trifida Cav.: The Lodge, Sandy, 14Y, 1976 (LTN). Bird seed alien.
Malva verticillata L.: Flitwick, 03G, 1971 (LTN). Wool alien.
Modiola caroliniana (L.) G. Don: Maulden, 03N, 1980 (LTN), det. E. J. Clement. Wool alien.
Sida spinosa L.: Maulden, 03N, 1976 (LTN). Wool alien.
Urocarpidium shepardae (Johnst.) Krapov.: Maulden, 03N, 1969 (herb. E. J. Clement), E. J.
Clement, J. L. Mason & T. B. Ryves, det. C. C. Townsend. Wool alien.
Guttiferae
Hypericum maculatum Crantz subsp. maculatum: waste ground, Aspley Guise, 93N, 1972 (LTN),
H. B. Souster!, det. N. K. B. Robson.
Umbelliferae
Bupleurum falcatum L.: garden, Ickwell, 14M, 1980, N. Dawson. Spontaneous and increasing.
Heracleum mantegazzianum Somm. & Levier: roadside, Ridgmont, 93T, 1979. See note in Dony
(1953, p. 299).
Oenanthe crocata L.: canal bank, Linslade!, v.c. 24 [Beds.], 92C, 1981 (LTN), H. B. Souster. Not
recorded from v.c. 30 since 1881.
O. lachenalii C. C. Gmel.: the record of O. silaifolia Bieb. in Dony (1969) was in error for this, the
station being the only one now known for O. lachenalii in the vice-county.
Trachyspermum ammi (L.) Sprague: Bedford R.D.C. refuse tip, OSF, 1969 (LTN). Bird seed alien.
Convolvulaceae
Convolvulus erubescens Sims: Maulden, 03N, 1977 (LTN), det. E. J. Clement. Wool alien.
Boraginaceae
Anchusa azurea Mill.: waste ground, Luton, 12A, 1950 (LTN), det. E. J. Clement. Previous
records of A. officinalis L. should probably be referred to this species.
Symphytum grandiflorum DC.: garden, Segenhoe Manor, 93Y, 1978 (LTN). Naturalized.
Labiatae
Salvia viridis L.: landfill site, Sundon, 02J, 1980 (LTN). Garden origin.
Solanaceae
Capsicum annuum L.: landfill site, Potton, 24E, 1983 (LTN); landfill site, Sundon, 02J, 1985. Bird
seed alien.
Hyoscyamus albus L.: Bedford R.D.C. refuse tip, 05F, 1979 (LTN). Origin not known.
Physalis peruviana L.: landfill site, Sundon, 02J, 1983 (LTN), det. E. J. Clement. Bird seed alien.
Solanum laciniatum Aiton: landfill site, Sundon, 02J, 1985 (LTN), det. E. J. Clement. Origin not
known.
S. nigrum L. X S. sarrachoides Sendtn. (S. X procurrens Leslie): Maulden, 03T, 1977 (LTN),
H. J. M. Bowen! This hybrid was noted by Leslie (1978) from market-gardens at Potton and
Sutton.
168 J. G. DONY AND C. M. DONY
Scrophulariaceae
Melampyrum arvense L.: spoil heap, Lidlington!, 93U, 1975, A. J. Martin. This species has an
interesting history in the county. The population of the most long-standing site, which had been
maintained by periodic ditching that has now ceased, became reduced to two plants in 1983,
since when none has been seen. In a second site at Bidwell there were 1000+ plants when it was
first observed in 1946 but there followed a regular diminishing in the number of plants, the last
ones being seen in 1970. At the Lidlington site an estimate of 5000+ plants was made by one
observer, since when there has been a large annual reduction. Similar large populations in
Essex, Wiltshire and the Isle of Wight, the only other counties in which it has appeared in recent
years, have shown the same diminution, ending in all cases but one in the complete
disappearance of the species. There can be little doubt that it is an alien introduction into Britain
with impure grain seed, a view expressed long ago by Bromfield (1856) and many others since,
including Salisbury (1961) and Webb (1985). It is, however, of considerable ecological interest
as a hemiparasite.
Veronica hederifolia L. subsp. hederifolia: this subspecies appears to be common in the county,
occurring most frequently on arable land.
V. hederifolia L. subsp. lucorum Klett & Richter: churchyard, Woburn, 93L, 1977 (LTN), conf. P.
M. Benoit. This subspecies appears to be as common as the preceding and was observed in 24
additional tetrads, growing in shady places as well as in open areas.
Martyniaceae
Ibicella lutea (Lindl.) Van Eseltine: Flitton, 03T, 1976 (LTN). Wool alien.
Lentibulariaceae
Utricularia australis R.Br.: gravel pit, Felmersham, 95Z, 1951 (LTN), det. P. Taylor. Recorded in
error as U. vulgaris L. in Dony (1953). It was first observed here, in a vegetative condition, in
1949 and is now well established, flowering annually in some quantity.
Rubiaceae
Asperula arvensis L.: The Lodge, Sandy, 14Y, 1982 (LTN), J. K. Dawson. Bird seed alien.
Galium pumilum Murr.: spontaneously in garden, Luton!, 02W, 1982 (LTN), E. B. Rands. The
only previous record was from v.c. 20 [Beds.].
Caprifoliaceae
Lonicera caprifolium L.: hedgerow, Streatley, 02U, 1976 (LTN), P. Ford! Possibly bird-sown; only
previous record 1889.
Campanulaceae
Downingia elegans (Douglas) Torrey: bank of mineral workings, Wavendon Heath, v.c. 24
[Beds.], 93H, 1981 (LTN), conf. E. J. Clement. Introduced with imported grass seed.
Compositae
Amellus microglossus DC.: Maulden, 03N, 1969 (herb. E. J. Clement), E. J. Clement & T. B.
Ryves, det. C. C.Townsend. Wool alien.
A. strigosus Less.: Maulden, 03N, 1969 (herb. E. J. Clement), J. L. Mason. Wool alien.
Arctium tomentosum Mill.: landfill site, Sundon, 02J, 1970 (LTN). Origin not known.
Bidens connata Muhl.: Grand Union Canal, Linslade, v.c. 24 [Beds.], 92D, 1980 (LTN).
Chrysanthemum myconis L.: Flitton, 03T, 1968 (LTN), det. J. E. Lousley. Presumed to be a wool
alien.
Cicerbita macrophylla (Willd.) Wallr.: roadside, Whipsnade, 011, 1981. A remote site for a species
known previously only as a garden escape.
Cichorium endivia L.: Flitwick, 03G, 1957 (LTN), det. E. J. Clement. Origin not known.
Cosmos bipinnatus Cav.: Maulden, 03N, 1982 (LTN). Wool alien.
Crepis setosa Haller f.: spontaneously in garden, Barton!, 03V, 1970 (LTN), W. L. Stevens. The
only previous recent record was from v.c. 20 [Beds.]. The last record from v.c. 30 was in 1930.
Hieracium diaphanum Fr.: Rowney Warren, 14A, 1971 (LTN), det. P. D. Sell.
FLORA OF BEDFORDSHIRE 169
H. grandidens Dahlst.: railway bank, Souldrop, 96A, 1971 (LTN), det. P. D. Sell.
H. maculatum Sm.: railway bank, Harlington, 03F, 1971 (LTN).
H. vulgatum Fr.: Rowney Warren, 14A, 1971 (LTN), det. P. D. Sell.
Iva xanthifolia Nutt.: Maulden, 03N, 1976 (LTN), det. E. J. Clement. Wool alien.
Pilosella praealta (Vill.) C. H. & F. W. Schultz subsp. arvorum (Naegl. & Peter) P. D. Sell & C.
West: railway bank, Moor End, OSE, 1971 (LTN), det. P. D. Sell. Origin not known.
Rudbeckia hirta L.: landfill site, Sundon, 02J, 1983 (LTN). Garden origin.
Hydrocharitaceae
Elodea nuttallii (Planch.) St. John: lake, Russell Park, Bedford, 04U, 1978 (LTN). This has since
been recorded in other sites in the north of the county.
Potamogetonaceae
Potamogeton crispus L. xX P. friesii Rupr. (P. x lintonii Fryer): water-storage tank, Roxton,
15M, 1973 (BM), det. J. E. Dandy. P. crispus is a common species in Bedfordshire but P. friesii
rare: it would appear that P. pusillus L., which is common, could be a parent rather than P. friesii.
P. obtusifolius Mert. & Koch: Battlesden Lake, 92P, 1976 (LTN); Mermaids Pond!, 93H, 1981, J.
Morris; flooded sandpit, Heath and Reach, 92J, 1982 (LTN).
Liliaceae
Allium oleraceum L.: Shelton, 06E, 1973 (LTN). The first confirmed record; still there in 1985.
Juncaceae
Juncus acutiflorus Ehrh. X J. articulatus L. (J. X surrejanus Druce ex Stace & Lambinon): marsh,
Dropshort, 02D, 1972 (LTN), P. M. Benoit!
J. effusus L. X J. inflexus L. (J. X diffusus Hoppe): marsh, Dropshort, 02D, 1972 (LTN), P. M.
Benoit!; marsh, Tebworth, 92Z, 1973 (LTN), P. M. Benoit!
J. tenuis Willd.: pond, The Lodge, Sandy, 14Y, 1976 (LTN); Kingshoe Wood, 93X, 1982 (LTN);
Wavendon Heath, v.c. 24 [Beds.], 93H, 1984. This species is no longer at the station given in Dony
(1953) and, while appearing to extend its range in the county, is not permanent.
Gramineae
Aegilops speltoides Tausch: Flitwick, 03G, 1977, ex hort. 1979 (LTN), det. C. E. Hubbard. Wool
alien.
Agrostis castellana Boiss. & Reuter: bank of mineral workings, Wavendon Heath, v.c. 24 [Beds.],
93H, 1981 (LTN). Introduced with imported grass-seed.
A. exarata Trin.: bank of mineral workings, Wavendon Heath, v.c. 24 [Beds.], 93H, 1981 (LTN),
det. E. J. Clement. Introduced with imported grass-seed.
A. scabra Willd.: Flitwick, 03G, 1977 (LTN), det. C. E. Hubbard. Wool alien.
Alopecurus antarcticus Vahl: Maulden, 03N, 1968 (herb. E. J. Clement), E. J. Clement & T. B.
Ryves, det. C. E. Hubbard. Wool alien.
Anthoxanthum aristatum Boiss. (A. puelii Lecog & Lamotte): bank of mineral workings, Wavendon
Heath, v.c. 24 [Beds.], 93H, 1982 (LTN). Introduced with imported grass-seed. The last record
from v.c. 30 was in 1919.
Apera interrupta (L.) Beauv.: dismantled railway, Henlow, 13M, 1973 (LTN). Casual.
Avena barbata Pott ex Link: Maulden, 03N, 1964 (LTN), det. C. E. Hubbard. Wool alien.
Bromus arvensis L.: field border, Dunton, 24G, 1970 (LTN), conf. C. E. Hubbard. Established for
15 years, but the site has now been ploughed.
B. lanceolatus Roth var. lanuginosus (Poir.) Dinsm.: Maulden, 03N, 1965 (K), det. C. E. Hubbard.
Wool alien.
Cenchrus echinatus L.: Maulden, 03N, 1969 (K), det. C. E. Hubbard. Wool alien.
C. incertus Curtis: Maulden, 03N, 1967 (K), J. L. Mason, det. W. D. Clayton. Wool alien.
Danthonia racemosa R.Br.: Maulden, 03N, 1967 (LTN). Wool alien.
Deschampsia danthonioides (Trin.) Munro: Woburn Park, 93R, 1977 (LTN), det. C. E. Hubbard.
Introduced with imported grass-seed and found subsequently at other sites.
Digitaria ctenantha (F. Muell.) Hughes: Maulden, 03N, 1967 (K), det. C. E. Hubbard. Wool alien.
170 J. G. DONY AND C. M. DONY
Diplachne muelleri Benth.: Flitton, 03T, 1983 (LTN), det. E. J. Clement. It is possible that
previous records of D. fusca (L.) Beauv. may be referred to this. Wool alien.
Echinochloa turneriana (Domin) Black: Maulden, 03N, 1967 (herb. E. J. Clement), E. J.
Clement & T. B. Ryves, det. C. E. Hubbard. Wool alien.
Eleusine tristachya (Lam.) Lam.: Maulden, 03N, 1976 (LTN), det. C. E. Hubbard. Wool alien.
Eragrostis curvula (Schrad.) Nees: Maulden, 03N, 1967 (LTN), det. C. E. Hubbard. Wool alien.
E. molybdea Vicary: Maulden, 03N, 1976 (LTN), det. C. E. Hubbard. Wool alien.
E. procumbens Nees: Flitton, 03T, 1983 (LTN), det. E. J. Clement. Wool alien.
E. schweinfurthii Chiov.: Maulden, 03N, 1966 (K), det. C. E. Hubbard. Wool alien.
E. virescens J. & C. Presl: Flitton, 03T, 1982 (LTN). Wool alien.
Eremopyrum bonaepartis (Spreng.) Nevski: Flitwick, 03G, 1978 (LTN), det. E. J. Clement.
Wool alien. .
Eriochloa creba S. T. Blake: Maulden, 03N, 1967 (LTN), det. C. E. Hubbard. Wool alien.
Festuca heterophylla Lam.: shady drive, Podington, 96G, 1973 (LTN). Well established, but
possibly introduced.
Hordelymus europaeus (L.) Harz. The only current record given in Dony (1953) was from v.c. 20
[Beds.]. It has now been found at four sites in v.c. 30, from which it was last recorded in 1906.
Panicum effusum R.Br.: Maulden, 03N, 1967 (K), J. L. Mason, det. C. E. Hubbard. Wool alien.
Parapholis incurva (L.) C. E. Hubbard: Maulden, 03N, 1969 (K, herb. E. J. Clement), E. J.
Clement & T. B. Ryves, det. C. E. Hubbard. Presumed to be a wool alien here, but it is native
elsewhere in Britain in dry saltmarshes.
Pennisetum clandestinum Hochst. ex Chiov.: Maulden, 03N, 1968 (herb. T. B. Ryves), E. J.
Clement & T. B. Ryves. Wool alien.
Phalaris angusta Nees: Maulden, 03N, 1977 (LTN). Wool alien.
Poa schimperiana Hochst. ex A. Rich.: Maulden, 03N, 1966 (K), det. C. E. Hubbard. Wool
alien.
P. sterilis Bieb.: Maulden, 03N, 1968 (LTN), det. C. E. Hubbard. Wool alien.
Puccinellia distans (L.) Parl.: waste ground, Thurleigh, 0SP, 1973 (LTN). A coastal species that
has been recorded from 66 tetrads in the county since 1973, on the margins of main roads. For
an account of the increase of this species inland in recent years, see Scott and Davison (1982)
and Scott (1985). There was one previous record from v.c. 30 in 1798.
Schismus arabicus Nees: Flitwick, 03G, 1978 (LTN), det. E. J. Clement. Wool alien.
Sorghum bicolor (L.) Moench: landfill site, Sundon, 02J, 1971 (LTN). Bird seed alien.
Sporobolus fimbriatus Nees: Maulden, 03N, 1976 (K, LTN, herb. T. B. Ryves), E. J. Clement &
T. B. Ryves!, det. C. E. Hubbard. Wool alien.
Stipa brachychaetoides Speg.: Maulden, 03N, 1973 (herb. T. B. Ryves), E. J. Clement & T. B.
Ryves, det. C. E. Hubbard. Wool alien.
S. falcata Hughes: Maulden, 03N, 1966 (K, herb. T. B. Ryves), E. J. Clement & T. B. Ryves,
det. C. E. Hubbard. Wool alien.
S. formicarum Delile: Maulden, 03N, 1962 (LTN), det. C. E. Hubbard.
S. juergensii Hack.: Maulden, 03N, 1960 (K), M. McCallum Webster, det. C. E. Hubbard. Wool
alien.
S. philippii Steud.: Maulden, 03N, 1969 (herb. T. B. Ryves), E. J. Clement & T. B. Ryves, det.
C. E. Hubbard. Wool alien.
Lemnaceae
Lemna minuscula L.: moat, Hatch, 14N, 1985 (LTN), conf. A. C. Leslie.
Typhaceae
Typha angustifolia L. x T. latifolia L.: ditch, Lidlington!, 93Z, 1984 (LTN), A. C. Leslie.
T. latifolia was the only parent present.
Cyperaceae
Carex hostiana DC.: Cow Common, 92W, 1952 (K), E. Nelmes (1955). This is the only con-
firmed record from v.c. 30. The site was ploughed shortly after Nelmes’ visit.
C. hostiana DC. Xx C. lepidocarpa Tausch: as above. This is the only record from v.c. 30.
FLORA OF BEDFORDSHIRE 171
C. lepidocarpa Tausch: marsh, Toddington, 02E, 1979 (LTN); marsh, Dropshort, 02D, 1984;
marsh, Eggington, 92S, 1985. This is no longer present at the sites given in Dony (1953, 1969).
C. pulicaris L.: Knocking Hoe!, 13F, 1978, J. L. Mason, old chalk quarry, Barton Hill!, 03A,
1982, M. Massey; Sundon Hill, 02P, 1978 (LTN). These are interesting additional records, as
all three sites are on chalk downland, the species not having been recorded from the county in
its normal habitat of ‘“‘boggy places” since 1798. The only known record since then was made
by C. Crouch (1898) on a chalk downland site near Sundon Hill where he found it growing
“with Pinguicula vulgaris and Parnassia palustris’. One of us (J.G.D.) remembers Parnassia
growing there from 1917 until 1925. The three new records made independently over a short
period may be the result of a minor climatic change, it being of some significance that C.
panicea L. was observed in 1983 on another chalk downland (Habitat Study 12A), (Dony
1953). They also coincide with the records of Epipactis palustris (L.) Crantz from chalk
downland (Dony 1969) and more recently Dactylorhiza praetermissa (Druce) So6 nearby (see
below). These could be the result of one of the all too frequent orchid transplants, but
transplants seem less likely to be the case with sedges over so wide an area.
Eleocharis palustris (L.) Roemer & Schultes subsp. palustris: side of pond, Woburn, 93K, 1981
(LTN), det. S. M. Walters. The only site now known in v.c. 30.
Scirpus maritimus L.: sandpit, Tiddenfoot, v.c. 24 [Beds.], 92B, 1973 (LTN); side of lake,
Ampthill Park, 03J, 1974. For a previous record see Dony (1953). This species survives well in
inland sites, but the means of introduction is not known.
Orchidaceae
Dactylorhiza incarnata (L.) So6: marsh, Felmersham!, 9SY, 1978, R. K. Saxton. This is a wel-
come discovery of a healthy colony of a species feared to be lost in the vice-county through the
filling-in of the Eaton Socon water-meadows and its non-appearance at its only other recently
known site.
D. praetermissa (Druce) Sod. There have been four recent records for this species, which was
feared lost in the vice-county. One, in 1980, was by R. M. Bateman who reported it on chalk
downland (see below), near the site where Epipactis palustris (L.) Crantz was found in 1966
(Dony 1969). Non-flowering plants of E. palustris were seen in 1967 but none since.
D. praetermissa (Druce) So6 Xx D. fuchsii (Druce) So6: chalk downland, Dunstable!, 1980, R.
M. Bateman.
Gymnadenia conopsea (L.) R. Br.: pasture, Thurleigh!, OST, 1982, G. Dennis. This is the first
record from v.c. 30 since 1884, other than on chalk downland, where it is still common.
ACKNOWLEDGMENTS
The authors are indebted to an increasing number of local naturalists in v.c. 30 who have drawn
their attention both to new sites and to plants of apparent interest. They are grateful for the
assistance of many in plant identification, but most especially to E. J. Clement and the late C. E.
Hubbard.
REFERENCES
BROMFIELD, W. A. (1856). Flora Vectensis. London.
Croucn, C. (1898). Carex pulicaris on chalk. J. Bot., Lond., 36: 399.
Dony, J. G. (1947). William Crouch (1818-1846). J. Bedf. Nat. Hist. Soc., 1: 50-51.
Dony, J. G. (1953). The Flora of Bedfordshire. Luton.
Dony, J. G. (1957). The drawings of Caroline Gaye. Bedf. Nat., 10: 14-15.
Dony, J. G. (1969). Additional notes on the flora of Bedfordshire. Proc. bot. Soc. Br. Isl., 7: 523-535.
Dony, J. G. (1976). Bedfordshire plant atlas. Luton.
Lesuiz, A. C. (1978). The occurrence of Solanum nigrum L. x S. sarrachoides Sendtn. in Britain. Watsonia,
12: 29-32.
NELMES, E. (1955). Carex hostiana in Bedfordshire. Proc. bot. Soc. Br. Isl., 1: 314-315.
SALISBURY, E. J. (1961). Weeds and aliens. London.
172 J. G. DONY AND C. M. DONY
Scott, N. E. (1985). The updated distribution of maritime species on British roadsides. Watsonia, 15: 381-386.
Scott, N. E. & Davison, A. W. (1982). De-icing salt and the invasion of road verges by maritime plants.
Watsonia, 14: 41-52.
Stace, C. A., ed. (1975). Hybridization and the flora of the British Isles. London.
WeEsB, D. A. (1985). What are the criteria for presuming native status? Watsonia, 15: 231-236.
(Accepted January 1986)
Watsonia, 16, 173-182 (1986) 173
Short Notes
SARRACENIA IN THE NEW FOREST
Members of the genus Sarracenia (pitcher plants) are native to the eastern seaboard of North
America. One species, S. purpurea, is known to have been introduced to Europe: in
particular it has naturalized from plantings in western Switzerland and central Ireland (Webb
1964). It has also been recorded during the past 20 years from three localities in northern
England.
On 16th July 1984, the authors found a clump of a Sarracenia species in the New Forest, S.
Hants., v.c. 11. This plant had erect, slender, trumpet-shaped pitchers which were yellow-green
and broadened gradually from their bases to their orifices. It was not S. purpurea, which has
decumbent, inflated pitchers that vary from green, suffused with red or purple, to almost
completely red. Up the front of each pitcher was a distinct wing which was broadest at the base and
became progressively narrower as the trumpet itself became wider. No flowers were present, but
reference to Godfrey & Wooten (1979) led to the conclusion that the plant was S. flava L., a
species of seepage zones, cypress swamps and bogs in the coastal plain area of south-eastern North
America. With assistance from Miss C. Whitefoord, subsequent comparison with herbarium
material held at BM confirmed the identification. North American populations of this species are
variable: wings are usually narrow, but may be up to 3 cm wide towards the pitcher base and the
pitchers themselves vary from yellow-green to almost entirely red, with a range of venation
patterns (Schnell 1978). In the New Forest, the clump of pitchers (which appeared to originate
from a single rootstock and was about 15 cm in diameter) was growing between tussocks of Molinia
caerulea (L.) Moench on a seepage terrace at the side of a valley mire. Associated species were:
Myrica gale L., Calluna vulgaris (L.) Hull, Erica tetralix L., Juncus acutiflorus Ehrh. ex Hoffm.,
Carex echinata Murr., Eriophorum angustifolium Honck., Equisetum palustre L. and Sphagnum
palustre L. The area was well-frequented by ponies, as evidenced by the extent of trampling and
grazing and by the quantity of dung in the vicinity.
At the time of the first visit, the clump consisted of three grazed leaves, six mature but ungrazed
pitchers and six smaller leaves, either immature pitchers or phyllodia. Phyllodia are small, sword-
shaped leaves, not growing to more than about one third the length of mature pitchers, which do
not develop into pitchers. In S. flava they develop during the latter half of the growing season and
persist through the winter, though in other species they may grow in the spring before pitchers are
produced. On a second visit (24th August 1984), there were 19 grazed leaves, six mature pitchers
and five small leaves (phyllodia or young pitchers). Mature pitchers were 34-36 cm tall, compared
with maximum lengths of c. 90 cm found in material from North America.
There is a long history of cultivation of S. flava in Britain and different variants have been
recognized (Masters 1881). The New Forest clump undoubtedly originated from introduced,
cultivated stock, though it is not possible to determine how it arrived at this locality. The site is at
least 200 m from the nearest road or track, although it is close to a walkway used by ponies. Apart
from damage caused by these animals, there are no signs of recent disturbance; associated plants
are well-grown and the vegetation cover is complete. Thus it would seem that direct planting in the
recent past is unlikely. The plant appears to be selectively grazed and, on a third visit to the site in
October 1984, no mature pitchers remained ungrazed. On 30th July 1985 the clump contained five
mature pitchers, five developing pitchers or phyllodia, and 13 grazed leaves. Grazing pressure may
be responsible for keeping the size of the clump in check and it is possible that it has existed in this
area for a considerable time.
174 SHORT NOTES
REFERENCES
Goprrey, R. K. & Wooten, J. W. (1979). Aquatic and wetland plants of the south-eastern United States.
Athens, Georgia.
Masters, M. T. (1881). Sarracenias. Gard. Chron., 16: 11-12, 150-156.
SCHNELL, D. (1978). Sarracenia flava L.: infraspecific variation in eastern North Carolina. Castanea, 43: 1-19.
Wess, D. A. (1964). Sarracenia, in Tutin, T. G. et al., eds. Flora Europaea, 1: 349. Cambridge.
R. E. DANIELS & A. J. CRANE
Institute of Terrestrial Ecology, Furzebrook Research Station, Wareham, Dorset, BH20 5AS
CAREX MURICATA L. SUBSP. MURICATA NOT IN DEVON
Ivimey-Cook (1984) has reported that Carex muricata L. subsp. muricata ‘is thought to occur on
the chalk in the vicinity of Branscombe and has also been recorded for Bideford’’. This is a sedge of
northern and eastern Europe, largely replaced in the west by its close ally, the subspecies
lamprocarpa Celak. (= C. pairaei F. W. Schultz). Its presence in Britain was established by
Nelmes (1947) who found herbarium specimens from four widely separated stations. To these only
one more has since been added.
Although one of Nelmes’ stations was in Gloucestershire (where the plant, though refound in
1973, now appears to be at least temporarily extinct), an occurrence in Devonshire is extremely
improbable, for in Britain the habitat is closely restricted to very well drained limestone scree.
Opportunities for error are, moreover, great: determination is not so very difficult — though the
differences between the two subspecies are small they are quite definite (David 1979) — but the
nomenclature has been very much confused.
L. J. Margetts has kindly and thoroughly investigated these Devonshire records. For ‘Bideford’
a specimen exists (now in CGE) and this turns out to be not a ‘muricata’ at all but Carex otrubae
Podp. For ‘Branscombe’ no specimen has been preserved, but the finder was unaware that there
are two subspecies of C. muricata proper and followed Jermy & Tutin (1968) in using the name
merely to distinguish her plant from C. muricata subsp. leersii Aschers. & Graebn., now known
(Chater 1980) as C. divulsa subsp. leersii (Kneucker) W. Koch. It is Margetts’ opinion that the
plant found was most probably C. spicata Huds., yet another member of the complex.
There is no reason, then, to suppose that C. muricata L. subsp. muricata is, or could be, in
Devon. And I would beg all recorders not to publish records of this (in Britain) extremely rare
plant before making the most thorough check of their authenticity.
REFERENCES
Cuater, A. O. (1980). Carex, in Tutin, T. G., et al., eds. Flora Europaea, 5: 290-323. Cambridge.
Davin, R. W. (1979). Another British locality for Carex muricata L. sensu stricto. Watsonia, 12: 335.
IvimEY-CooK, R. B. (1984). Atlas of the Devon flora. Exeter.
Jermy, A. C. & Tutin, T. G. (1968). British Sedges. London.
NewMES, E. (1947). Two critical groups of British sedges. Rep. botl Soc. Exch. Club Br. Isl., 13: 95-105.
R. W. DAvip
50 Highsett, Cambridge, CB2 INZ
PETRORHAGIA NANTEULII (BURNAT) P. W. BALL & HEYWOOD IN
MID-GLAMORGAN
In August 1985, Petrorhagia nanteulii (Burnat) P. W. Ball & Heywood (Kohlrauschia prolifera
auct.) was discovered by the author in the Dare Valley Country Park, Aberdare, v.c. 41. It was
found to be growing in a large established colony along with other grassland plants on a well
drained embankment of a disused railway line.
SHORT NOTES 175
P. nanteulii is a rare species found growing in sandy and gravelly places in south-eastern England
and is also established as an alien. There are only two previous records for this species in Wales,
both from Cardiff Docks (Ellis 1983). The most recent record was 1980, the previous record was
1926. Thus this site is the second in Wales so far discovered. Its presence on what was a busy
railway line serving the coal mines and Gadlys Ironworks in the Dare Valley, ultimately connected
with Cardiff Docks, gives a clue as to its possible origin. Coal mining operations ceased by the
1950s and the line fell into disuse. As the colony is well established it is conceivable that it has
existed at this site for 30 years or more after being carried there, presumably on railway wagons
loaded with imported iron ore.
ACKNOWLEDGMENTS
I am grateful to Dr J. R. Akeroyd of the Botany Department, Reading University and Mr R. G.
Ellis of the National Museum of Wales for confirming the identification of this species.
REFERENCE
Exuis, R. G. (1983). Flowering plants of Wales. Cardiff.
H. J. DAwson
Department of Botany, Birkbeck College, Malet Street, London, WCIE 7HX
DACTYLORHIZA MACULATA (L.) SOO x D. TRAUNSTEINERI (SAUTER) SOO IN
N.E. YORKS.
In mid-June 1985, I examined what may possibly be a previously unrecorded colony of
Dactylorhiza traunsteineri (Sauter) So6 at a site east of Pickering, N.E. Yorks., v.c. 62, (GR 44/
8.8). The location is a very small, isolated calcareous flush (approx. 10 m diameter), where D.
traunsteineri and its putative hybrid with D. maculata (L.) So6 grow along with such normal
associates as Primula farinosa, Pinguicula vulgaris and Schoenus nigricans. Dactylorhiza maculata
grows close by and certainly within 30 m. The hybrid, D. maculata x D. traunsteineri, for which
this is a new vice-county record and second English record, is rare in the British Isles.
The typical hybrid shows many features intermediate between the parents and is a relatively
robust and distinctive plant generally 20—26 cm tall (at this site typical D. traunsteineri is 6-12 cm),
with broader leaves than D. traunsteineri, marked with well-spaced dark blotches. The stem is
stouter than in D. traunsteineri and the upper stem and bracts are not so deeply stained red-brown
with anthocyanin. The flower colour is a very distinctive pale bluish-red, intermediate between the
parents, and the lateral sepals are not held erect but arched forward into a semi-hood. The
labellum is large and is lightly flecked with dots and broken loops, with the lateral lobes reflexed
and the central lobe smaller than the laterals but of similar length, although perhaps closer in shape
to D. maculata. The spur is shorter and more conical than in D. traunsteineri.
Although the characteristics as outlined above are found to be in close accord with those given
by Roberts (1962), it is interesting that some of the plants had unspotted (rather than spotted)
leaves and, in some cases, the usual pronounced reflexing of the lateral lobes of the labella was
absent or reduced and, often, the flower spike was distinctly less lax than is usual.
A detailed description and colour photographs were sent to R. H. Roberts, who noted in these
plants a greater leaf width and wider spur opening than in examples of the hybrid which he had
observed, but agreed that the arched forward position of the lateral sepals, the shape and colour of
the labellum, and the reduced anthocyanin staining of the upper stem and bracts indicate the
hybrid. It is in fact quite possible that a hybrid swarm is present, with back-crossing and/or
segregation causing some degree of variability, but such swarms, whilst demonstrated to occur for
instance between D. fuchsii and D. majalis subsp. purpurella (Lord & Richards 1977), appear not
to have been recorded for D. maculata X D. traunsteineri.
176 SHORT NOTES
Herbarium specimens were not gathered, but colour photographs and detailed field notes are
retained by the author.
ACKNOWLEDGMENT
I wish to thank Mr R. H. Roberts for his help with these plants.
REFERENCES
Lorp, R. M. & Ricwarps, A. J. (1977). A hybrid swarm between the diploid Dactylorhiza fuchsii (Druce) S06
and the tetraploid D. purpurella (T. & T. A. Steph.) So6 in Durham. Watsonia, 11: 205-210.
Roserts, R. H. (1962). Dactylorchis maculata subsp. ericetorum Xx D. traunsteineri. Proc. bot. Soc. Br. Isl., 4:
418.
M. J. Y. FOLEY
87 Ribchester Road, Clayton-le-Dale, Blackburn, BB1 9HT
SEED MORPHOLOGY IN ARUM MACULATUM L.
A recent study of seed morphology in Arum maculatum L. has shown some interesting differences
between hedgerow and woodland populations of this species. Seven discrete populations of A.
maculatum were sampled from the Deeside area (GR 33/29.69) of north-eastern Flints., v.c. 51.
Four were from woodland and three from hedgerow habitats. In all, 77 fruiting spikes were
collected during August 1981. In each case, the ripe berries were removed from the spike and a
random subsample of twelve berries per population was selected. The enclosed seeds were
separated from the pulp of each berry and the largest seed was chosen for further examination.
Seed length and width were measured across the widest points respectively. The total sample was
therefore represented by 84 seeds, i.e. twelve seeds per population. To determine overall and
habitat mean seed weights, each sample of twelve seeds per population was weighed. This
provided a total of seven population weights on which to calculate means.
The results are shown in Table 1. The overall means for seed length and width are similar to
those obtained by Sowter (1949), i.e. c. 6 X c. 5 mm. Both seed length and width show continuous
variation and are strongly correlated (r=0-74, p=0-001); this indicates that seed shape is fairly
constant irrespective of absolute size.
TABLE 1. SEED MORPHOLOGY IN WOODLAND AND HEDGEROW POPULATIONS OF ARUM
MACULATUM L.
Population characters Woodland Hedgerow Total
No. of populations 4 3 7
No. of spikes 52 25 77
No. of seeds sampled 48 36 84
Mean seed length (mm) 6-25+0-72 5:25+0-68 5-82+0-86
and S.D. (n=48) (n=36) (n=84)
Mean seed width (mm) 5-02+0-66 4-36+0-54 4-74+0-69
and S.D. (n=48) (n=36) (n=84)
Mean L/W ratio 1-3+0-1 1-2+0°1 1:2+0-2
and S.D. (n=48) (n=36) (n=84)
Mean weight of 12 seeds (g) 0-44+0-07 0-33+0-02 0-39+0-08
and §.D. (n=4) (n=3) (n=7)
SHORT NOTES 177
There are, however, statistically significant differences between the mean values of woodland
and hedgerow populations (Table 1). The differences involve (a) mean seed length (t=6-5,
p<0-001); (b) mean seed width (t=5-0, p<0-001); (c) mean seed weight (t=10-3, p<0-001); and
(d) mean seed length/width ratio (t=4-5, p<0-001).
I therefore conclude that there is a relationship between seed size and habitat in A. maculatum.
Salisbury (1942) was able to demonstrate a broad correlation between the average seed weight of a
species and its preferred habitat. In particular, species from closed, woodland communities have
larger seeds than those of more open habitats such as hedgerows. This example of intraspecific
variation in seed size was not discussed by Prime (1960) nor by other students of Arum, e.g. Sowter
(1949).
REFERENCES
Prime, C. T. (1960). Lords and ladies. London.
SALISBURY, E. J. (1942). The reproductive capacity of plants. London.
SowTer, F. A. (1949). Arum maculatum L. in Biological Flora of the British Isles. J. Ecol., 37: 207-219.
P. HARMES
21 Newthorn Place, Buckley, Clwyd, CH7 2EY
OPHRYS APIFERA HUDSON SUBSP. JURANA RUPPERT FOUND IN BRITAIN
Every year since 1980, I have visited a site on the Wiltshire downs (v.c. 8) and, among a range of
plants typical of chalk downland, I have recorded five species of orchid, Listera ovata, Orchis
mascula, Ophrys insectifera, O apifera and Dactylorhiza fuchsi. O. apifera first appeared in June
1983, but the single specimen was not closely examined.
On 17th June 1984, a single spike of O. apifera was again found, in approximately the same
position as the 1983 plant, and close examination of the flowers revealed that the two upper, inner
perianth segments were of similar form and colour to the three outer perianth segments.
Photographs were taken on 21st June and again on 23rd June, by which time four flowers had
opened fully. These agreed in all details with the descriptions of Ophrys apifera Hudson subsp.
jurana Ruppert given by Lang (1980) and So6 (1980) and also the descriptions and illustrations of
this plant found in Duperrex (1961), Williams et a/. (1978), Kohlhaupt (1981), Baumann &
Kiinkele (1982) and Davies et al. (1983). Six flowers ultimately opened, and the spike reached a
height of 28 cm. Photographs were sent to D.C. Lang, who agreed with the identification, and
copies have been placed in LTR.
A visit to the site on 16th May 1985 revealed a developing rosette of leaves. Additional visits on
23rd June and 2nd July were made by D. C. Lang and myself, flowering being delayed to the latter
date due to the late season.
The site is on an east-facing bank above a sunken track on the north side of a chalk hill. The flora
is typical of chalk downland, and species recorded within 3 m of the plant include Lotus
corniculatus, Polygala vulgaris, P. calcarea, Helianthemum nummularium, Linum catharticum,
Poterium sanguisorba, Thymus praecox, Asperula cynanchica, Cirsium acaule, Leontodon
hispidus, Listera ovata and Dactylorhiza fuchsii.
The plant had three basal leaves and two stem leaves, and reached a height of 26-5 cm with four
mature flowers. Individual flowers varied slightly in size, the outer perianth segments averaged 13
mm long by 6 mm wide at the widest point, and the upper, inner perianth segments 10 mm long by
4mm wide. The labellum was, on average, 8 mm wide. The upper, inner perianth segments were
pink, with a green central rib, blunt and noticeably hairy on the margins. They tended to curl
forward at the tips. The labellum pattern differed from that of the normal variant of O. apifera.
The typical U-shaped pattern at the base was missing and, halfway down the labellum, an irregular
horizontal yellow bar stretched across the entire breadth.
Seed-set was not observed in 1984. In 1985, self-pollination was evident in two of the flowers
178 SHORT NOTES
and, on 30th August, three well-developed seed capsules were present. All known sites for O.
apifera within 6 km have been searched, but no specimens resembling subsp. jurana have been
found.
The distribution of subsp. jurana is described generally as western-central and southern Europe
(Williams et al. 1978; Sod 1980; Davies et al. 1983), but more specifically it has been found in parts
of south-eastern France (Duperrex 1961) and south-western Germany (Baumann & Kiinkele
1982). Although no previous record of this subspecies in Britain appears to exist, it has recently
been discovered in northern Germany and Holland by M. Ebbens (D. C. Lang pers. comm.).
Careful examination of O. apifera flowers might reveal that this subspecies is more widely spread
than appears at present.
ACKNOWLEDGMENT
I am indebted to D. C. Lang for help in identification, and for encouragement and suggestions in
writing this note.
REFERENCES
BAUMANN, H. & KUNKELE, S. (1982). Die wild-wachsenden Orchideen Europas, pp. 186-187. Stuttgart.
Davies, P., Davies, J. & Hux ey, A. (1983). Wild orchids of Britain and Europe, p. 178. London.
DuperrEx, A. (1961). Orchids of Europe, p. 68. London.
KouHLHaupt, P. (1981). Mittel- und stideuropdische Orchideen, pp. 32-33. Bozen.
LanG, D. C. (1980). Orchids of Britain, p. 14. Oxford.
Sod, R. (1980). Ophrys L., in Tutin, T. G. et al., eds. Flora Europaea, 5: 349. Cambridge.
WiLuiaMs, J. G., WittiaMs, A. E. & Artortt, N. (1978). A field guide to the orchids of Britain and Europe, pp.
44-45. London.
R. J. LAURENCE
The Gables, Murtry, Frome, Somerset, BAI1 3NP
THE STATUS OF ORCHIS FRANCIS-DRUCEI WILMOTT
Wilmott (1936) described a new species of orchid, Orchis francis-drucei, following its discovery
whilst he was on an excursion with Francis Druce in north-western Scotland. This new species was
described as being slender in habit, with few narrow leaves, and a short, lax-flowered
inflorescence, with a long, projecting, median lobe of the labellum. Wilmott recognized that these
characters were shared by a species then known as O. traunsteineri Sauter, which at that time had
not been recognized from the British Isles; he distinguished the Scottish plants as being smaller and
having a very long median labellum lobe, which also had intense reddish-purple markings on a
white background.
O. francis-drucei was originally described from a single colony found on a hill slope above Loch
Maree, W. Ross, v.c. 105. Since its discovery on 23rd June 1935, the plant has apparently remained
undetected until a colony of similar plants was found by M.R.L. in June 1983 near Loch Maree. A
study of this colony of some 40 plants was undertaken by the authors the following year, and
specimens sent to Mr R. H. Roberts were confirmed beyond any doubt as Dactylorhiza
traunsteineri (Sauter) Soo.
The detailed description, illustrations and plates published by Wilmott (1936) enable a
comparison of the 1935 and 1984 populations with each other and with a generalized description of
D. traunsteineri (Table 1). In spite of the differences, the 1984 plants are clearly conspecific both
with the 1935 plants described as O. francis-drucei and with D. traunsteineri. It is assumed that the
variation between the two collections is either because the 1984 population was a different one or
that the 1935 population is the same but consisted of variants which are no longer dominant in this
population.
It has been demonstrated by Roberts & Gilbert (1963) that small isolated populations of D.
Character
Height (cm)
Total no. leaves
No. non-sheathing leaves
Length of longest leaf (cm)
Max. width of longest leaf (cm)
Leaf spotting
Leaf colour
Bracts
Floral spike
Labellum length x width (mm)
Labellum shape and colour
Spur length x width (mm)
Length of peripheral bract cells
SHORT NOTES
TABLE 1. COMPARISON OF O. FRANCIS-DRUCEI WITH D. TRAUNSTEINERI
Figures in brackets show mean values
O. francis-drucei*
11-13
4
0
4
0-4-0-5
unspotted
pale green
deeply suffused
purple
sub-secund with 5-8
flowers
7X6
sub-deltoid obtrian-
gular with a long
projecting mid-lobe,
intense reddish pur-
ple markings on a
white ground
6 X 2:3
179
D. traunsteineri®
West Ross
10-18 (14-2)
3-5 (3-7)
0-1 (0-8)
4-10 (7-6)
0-8-1-3 (1-1)
mainly lightly spot-
ted with small dots
or bars
pale green
deeply suffused
purple
secund with 4-11
flowers
(7-2) X (8-5)
more or less deltoid
with a prominent
mid-lobe, intense
markings on a back-
ground of magenta
to pale pink
(7:7) X (2-4)
(c. 110)
D. traunsteineri®
Generalized
0-6-1-5
markings light or absent
pale green
usually deeply suffused
purple
frequently secund with
2-20 flowers
6:5-—8-5 x 0-7-1
more or less deltoid with
less distinct mid-lobe than
column (b), colour as col-
umn (b)
8—9 x 3-0—3-5
(c. 110)
(um)
* Wilmott (1936).
> Field measurements on 20 plants by the authors in 1984 near Loch Maree.
© Heslop-Harrison (1953), Roberts & Gilbert (1963) & R. H. Roberts (pers. comm.)
traunsteineri may be uniform but possess features at variance with other populations in the British
Isles. Such local variation, when considered with Heslop-Harrison’s (1953) concept of ‘anthocy-
anin high’ and ‘anthocyanin low’ modes for flower colour and leaf markings, offers an explanation
for the differences between Wilmott’s O. francis-drucei and D. traunsteineri in the British Isles.
The only character which distinguishes both the 1935 and 1984 W. Ross plants from typical D.
traunsteineri is the somewhat narrower spur (Table 1), which is not thought to be of high
significance. We therefore consider that O. francis-drucei Wilmott should be regarded as wholly
synonymous with D. traunsteineri and not be given even infra-specific status.
In 1983, D. traunsteineri was removed as a species known with certainty to occur in Scotland
(Tennant & Kenneth 1983). The 1984 Loch Maree record therefore re-establishes D. traunsteineri
as a Scottish plant beyond doubt. Moreover, a population has recently been recognized from
Westerness, v.c. 97, (Mr D. C. Lang and Miss L. M. Watson, pers. comm.) that should be referred
to D. traunsteineri, and a solitary plant found by the authors in 1984 in Knapdale, Kintyre, v.c. 101,
is also this species; there is little doubt that further fieldwork will reveal it elsewhere in Scotland.
Specimens and photographs of the 1984 Loch Maree dactylorchids have been placed in E.
The 1984 population occurred in a series of open flushes dominated by Molinia caerulea and
Schoenus nigricans with penetration of Erica tetralix, Salix aurita and Myrica gale from the
surrounding heath community. A full list of the species recorded showed the vegetation to be
referable to the Pinguiculo-Caricetum Jones described by Wheeler (1980). (This unit includes the
Cariceto-Saxifragetum and Carex panicea-Campylium stellatum syntaxa of McVean & Ratcliffe
(1962), both widespread in base-rich flushes in Scotland). The vegetation is also similar to the W.
Ross site for D. incarnata subsp. cruenta (Kenneth & Tennant 1984). D. incarnata subsp. incarnata
and subsp. pulchella were both present at the 1984 Loch Maree site, and D. maculata subsp.
ericetorum occurred in more acid parts nearby. Several rather robust dactylorchids were also noted
180 SHORT NOTES
near the periphery of this site and were initially suspected to be the hybrid D. incarnata x D.
traunsteineri, although this could not be confirmed; no other dactylorchids were found in the
immediate vicinity.
ACKNOWLEDGMENTS
We should like to thank Mr R. H. Roberts for his assistance with the identification and his
observations on peripheral bract cells, also to Dr B. D. Wheeler for his guidance with the
vegetation descriptions.
. REFERENCES
HEsLop-Harrison, J. (1953). Studies in Orchis L., II. Orchis traunsteineri Saut. in the British Isles. Watsonia,
2: 371-391.
KENNETH, A. G. & TENNANT, D. J. (1984). Dactylorhiza incarnata (L.) So6 subsp. cruenta (O. F. Mueller)
P. D. Sell in Scotland. Watsonia, 15: 11-14.
McVEAN, D.N. & Ratc.irFe, D. A. (1962). Plant communities of the Scottish Highlands. London.
Roserts, R. H. & GILBerT, O. L. (1963). The status of Orchis latifolia var. eborensis Godf. in Yorkshire.
Watsonia, 5: 287-293. <
TENNANT, D. J. & KENNETH, A. G. (1983). The Scottish records of Dactylorhiza traunsteineri. Watsonia, 14:
415-417.
WHEELER, B. D. (1980). Plant communities of rich fen systems in England and Wales, II. Communities of
calcareous mires. J. Ecol. , 68: 405—420.
WivtmotTrt, A. J. (1936). New British marsh orchids. Proc. Linn. Soc. Lond., 148: 126-130.
M. R. Lowe, D. J. TENNANT & A. G. KENNETH
School House, Brancepeth, Durham, DH7 8DG
CROSS-POLLINATION BY WASPS IN EPIPACTIS LEPTOCHILA (GODF.) GODF. S.L.
This note describes what I believe to be the first published observation of cross-pollination in those
species of Epipactis which are generally regarded as being autogamous, i.e. automatically self-
pollinated. In the flora of the British Isles, these comprise E. leptochila (Godf.) Godf., its very
close relative E. dunensis (T. & T. A. Stephens.) Godf., E. phyllanthes Sm. and E. youngiana
Richards & Porter.
In the other species of Epipactis in the British Isles, it is believed that within-flower pollination
rarely occurs and that pollen transfer is usually between flowers (allogamy), either on the same
(geitonogamy) or different (xenogamy) genets. It was Darwin (1862) who first showed that the
allogamous species (E. helleborine (L.) Crantz, E. purpurata Sm., E. atrorubens (Hoffm.) Schult.)
are usually pollinated by wasps (Vespa spp.); E. palustris (L.) Crantz is also allogamous but is
usually visited by bees. These species are, however, self-compatible, so geitonogamous selfing is
possible. Wasps visit the flowers to drink nectar, which is located in the proximal, hemispherical
and usually purplish segment of the labellum (hypochile). To reach the hypochile, the head or top
of the thorax of the wasp rubs against the projection on the outer end of the stigma (rostellum),
which is covered by a sticky, whitish membrane (viscidium). The two pollinia drop from the anther
into grooves (clinandria) shortly after the flower opens, in which position they adhere apically to
the viscidium. When a wasp leaves a flower, the viscidium together with one or both pollinia
adhere to the wasp. When a subsequent flower is visited, part or all of one or both pollinia may be
deposited in the wet secretion of the stigma surface (just inside the position of the rostellum),
thereby achieving allogamous pollination. However, Hagerup (1952) noted that when pollinia are
not removed, some self-pollination occurs within the flower (autogamy) as the viscidium ages and
flows onto the stigma surface, carrying the pollinia with it.
In contrast, in the autogamous species, the rostellum is small (except in E. youngiana) and the
viscidium is missing, or is present in bud but disappears before or shortly after the flower opens.
The flower is usually greenish, inclined, and less fully open than in the allogamous species. The
SHORT NOTES 181
pollinia rest uneasily in the shallow clinandria, and rapidly break up and fall onto the stigma
unimpeded because the viscidium is absent. It is usually considered that such plants are
invariably self-pollinated, and this must certainly be so for cleistogamous variants, in which the
flower never opens, and is probably so for the variants of E. phyllanthes in which self-pollination
occurs before the flower opens. However, Proctor & Yeo (1973) and Richards (1982) have
speculated that some casual allogamy in the supposedly autogamous E. leptochila, E. dunensis
and E. youngiana might be caused by insect visitors, to which parts of disintegrating pollinia
might adhere. The biological importance of the allogamous and autogamous breeding systems
has been illustrated by Richards (1982), who showed that four populations of three allogamous
species are more variable than five populations of four autogamous species for each of ten metric
characters.
On 10th August 1985 at 16.00 hrs, I visited a population of the inland Northumberland variant
of E. leptochila (Richards & Swan 1976), many individuals of which closely resemble E.
dunensis. This population grows on lead-mine spoil under birch trees beside a road leading into a
caravan site. Some 30 individuals were flowering, with 5—35 flowers per spike, approximately half
of which were open and not withered. Within an area of 1 m? were seven spikes next to the road
in filtered sunlight, where the shade temperature was approximately 20°C. For at least 30
minutes, these received the intermittent but urgent attention of at least five workers of the wasp
Vespa germanica Fab. The wasps were drinking nectar from the hypochile for periods of 5—60
seconds per flower. For twelve recorded spike visits, from three to five flowers were visited;
these were usually adjacent to each other and in the middle of the region of the open flowers on
a spike. Some feeding took place at every flower which the wasp visited, and the wasps alighted
on every flower that was approached; however, wasps usually visited adjacent flowers by
crawling.
Although viscidia were present within large buds (determined by dissection), almost all open
flowers lacked a viscidium, and the rostellum was poorly marked, the outer edge of the stigma
being rather straight with only a slight rostellar peak. For three spikes, only the topmost open
flower had a detectable viscidium. Pollinia remained in the anther or on the clinandria in 15 out
of a sample of 61 open flowers in which the stigma was greenish (and therefore presumed to be
receptive) rather than brownish. On six occasions, a wasp was observed to visit a flower with
pollinia in situ. On each occasion, all or most of both pollinia adhered to the head or to the front
of the top of the thorax of the wasp when it left the flower, despite the apparent absence of a
visible viscidium of the type found in allogamous species. When a wasp visit was simulated using
the sharpened end of a pencil, the pollinia adhered to the bare wood surface of the pencil. All
the five wasps that were closely observed were carrying pollinia on, or just behind, the head; a
minimum of three and a maximum of at least eight pollinia were observed on different wasps at
various times. In some cases, pollinia had adhered on the top of other pollinia, making an
accurate estimation difficult.
Wasps carrying pollinia were observed visiting about 30 flowers with apparently receptive
stigmas. In no cases were whole pollinia or parts of pollinia large enough to be detected by the
naked eye on stigmas, although in every case that was closely observed, the pollinia on the wasp
were clearly brought into contact with the stigma as the wasp drank. It seems very likely however
that some pollen was deposited on stigmas, either as individual tetrads or as very small
fragments. In between flower visits, wasps tried vigorously to remove pollinia from their heads
using their front legs, but in this they were apparently unsuccessful. However, it is likely that
such efforts would loosen tetrads and massulae from the pollintum, making them available for
pollination.
From the same sample of 61 flowers with apparently receptive stigmas, whole pollinia or parts
of pollinia large enough to be observed with the naked eye were seen in 19 flowers. It is assumed
that these resulted from within-flower pollination, because between-flower visits by wasps
apparently did not leave large pollinium fragments on stigmas. It follows that for 42 of 61 flowers
in this ‘autogamous’ population, within-flower pollination had not occurred; in most the pollinia
were missing.
Within the 1 m? that was intensively studied, the minimum distances between spikes were 5,
10, 15, 20 and 50 cm respectively. Nine out of 30 between-flower transfers by wasps were
between spikes and should therefore have resulted in at least some cross-pollination. As pollinia
182 SHORT NOTES
apparently adhere to wasps for some time, releasing pollen slowly, pollen carry-over should be
large, and the potential for cross-pollination should be considerable.
It is concluded that in apparently autogamous populations of Epipactis in which the viscidium is
absent in open flowers and large parts of pollinia fall onto the stigma of the same flower at an early
stage, substantial amounts of cross-pollination may occur. The high level of pollinator activity
(observed fortuitously and briefly on this occasion) might be ascribed to a pleasantly warm, sunny,
still interlude in a dismal summer. Out of many observations of populations of E. leptochila s.1. in
different years, I have seen wasps visiting the flowers only on rare occasions, and then casual
visiting by single individuals seemed to be occurring.
REFERENCES
~
Darwin, C. (1862). The various contrivances by which wild orchids are fertilised. London.
Hacervup, O. (1952). Bud autogamy in some northern orchids. Phytomorphology, 2: 51-60.
Proctor, M. C. F. & YEo, P. F. (1973). The pollination of flowers, p. 54. London.
RicHarps, A. J. (1982). The influence of minor structural changes in the flower on breeding systems and
speciation in Epipactis Zinn. (Orchidaceae), in ARMSTRONG, J. A., POWELL, J. M. & RicHarps, A. J., eds.
Pollination and evolution, pp. 47-53. Royal Botanic Gardens, Sydney.
RicHarps, A. J. & SWAN, G. A. (1976). Epipactis leptochila (Godf.) Godf. and E. east ca G.E.Sm.
occurring in South Northumberland on lead and zinc soils. Watsonia, 11: 1-5.
A. J. RICHARDS
Department of Plant Biology, The University, Newcastle upon Tyne, NEI 7RU
Watsonia, 16, 183-198 (1986) 183
Plant Records
Records for publication must be submitted to the appropriate vice-county Recorder (see Vice-county Recorders
(1985)), and not the Editors. The records must normally be of species, hybrids or subspecies of native or
naturalized alien plants belonging to one or more of the following categories: Ist or 2nd v.c. record, 1st post-
1930 v.c. record; only extant v.c. record, or 2nd such record; a record of an extension of range by more than
100 km. Such records will also be accepted for the major islands in v.cc. 102-104 and 110. Only Ist records can
be accepted for Rubus, Hieracium and hybrids. Records for subdivisions of vice-counties will not be treated
separately; they must therefore be records for the vice-county as a whole. Records of Taraxacum are now being
dealt with separately, by Dr A. J. Richards, and will be published at a later date.
Records are arranged in the order given in the List of British vascular plants by J. E. Dandy (1958) and his
subsequent revision (Watsonia, 7: 157-178 (1969)). All records are field records unless otherwise stated. With
the exception of collectors’ initials, herbarium abbreviations are those used in British and Irish herbaria by D.
H. Kent & D. E. Allen (1984).
The following signs are used:
* before the record: to indicate a new vice-county record.
+ before the species number: to indicate that the plant is not a native species of the British Isles.
+ before the record: to indicate a species which, though native in some parts of the British Isles, is not so in the
locality recorded.
[] enclosing a previously published record: to indicate that the record should be deleted.
1/4. LYCOPODIUM CLAVATUM L. 59, S. Lancs.: Little Lever, Bolton, GR 34/74.07. Waste
ground near moors. V. Gordon, 1985. 1st post-1930 record.
4/1. EQUISETUM HYEMALE L. 93, N. Aberdeen: Tarty, GR 38/97.26. Flushed scrub. D.
Welch, 1984. 2nd extant record.
4/9 X 5. EQUISETUM ARVENSE L. X E. FLUVIATILE L. *78, Peebless.: W. of N. Esk Reservoir,
West Linton, GR 36/15.58. H. McHaffie, 1983, det. C. N. Page.
4/10. EQUISETUM TELMATEIA Ehrh. 109, Caithness: Below Niandt, GR 39/20.33. Coastal cliff.
J. Mackintosh, 1983. 2nd record.
7/1. HYMENOPHYLLUM TUNBRIGENSE (L.) Sm. 47, Monts.: Near Aberllefenni, GR 23/7.1.
Shaded rock by stream. P. M. Benoit et al., 1985. 2nd record.
7/2. HYMENOPHYLLUM WILSONII Hook. 76, Renfrews.: Derol Glen, Port Glasgow, GR 26/
31.74. Boulders in stream. A. McG. Stirling, 1981. 2nd record.
15/5 tri. ASPLENIUM TRICHOMANES L. subsp. TRICHOMANES *73, Kirkcudbrights.: Near
Borgan, GR 25/36.74. Granite boulder. O. M. Stewart, 1985, E.
16/1. CETERACH OFFICINARUM DC. 25, E. Suffolk: Mendham, GR 62/26.82. Church wall. F.
W. Simpson, 1984. 2nd extant record.
21/8. DRYOPTERIS AEMULA (Ait.) Kuntze 47, Monts.: Afon Crewi, GR 22/8.9. Wooded
gorge. A. J. Morton, 1984, conf. P. M. Benoit. 2nd record.
25/int. X 1. POLYPODIUM INTERJECTUM Shivas X P. VULGARE L. *41, Glam.: Kenfig Burrows,
GR 21/7.8. Sand dunes. G. Hutchinson, 1985, det. R. H. Roberts.
+27/1. AZOLLA FILICULOIDES Lam. *52, Anglesey: Near Llyn Alaw, GR 23/40.86. Pond. T.
Blackstock, 1984.
28/1. BOTRYCHIUM LUNARIA (L.) Sw. *76, Renfrews.: Near Old Hall, Kilmacolm, GR 26/
36.71. Old pastures. K. M. Calver, 1976.
184 PLANT RECORDS
29/1a. OPHIOGLOSSUM VULGATUM L. subsp. VULGATUM 76, Renfrews.: Skiff Wood, GR 26/
40.60. Woodland edge. A. J. Silverside, 1978. Only extant record.
+43/3. ANEMONE APENNINA L. *49, Caerns.: Treborth, GR 23/5.7. E. T. Jones, 1936.
46/7. RANUNCULUS SARDOUS Crantz *32, Northants.: Finedon, GR 42/91.70. Infilled
ironstone workings. I. Cameron, 1981. Duston Mill Lane, Northampton, GR 42/72.60. Disturbed
ground. A. Robinson, 1982. 1st and 2nd records.
46/10. RANUNCULUS AURICOMUS L. 46, Cards.: Near Coedmore Mansion, GR 22/19.43.
Ash-beech wood. A. O. Chater, 1985, NMW. 2nd record.
46/cal. RANUNCULUS CALCAREUS Butcher *25, E. Suffolk: Tattingstone, GR 62/12.35.
Stream. E. M. Hyde, 1984, herb. E. M. & M. Hyde, det. N. T. H. Holmes.
57/2. CERATOPHYLLUM SUBMERSUM L. *38, Warks.: Near Warwick, GR 42/27.68. Pond. J. C.
Bowra, 1985, det. R. J. Pankhurst.
+64/for. DICENTRA FORMOSA Walpers *44, Carms.: Near Cwmann, GR 22/62.46. Hedgebank.
A. O. Chater, 1985. 99, Dunbarton: Clachan Woods, Rosneath, GR 26/22.82. Woodland
bank. A. McG. Stirling & A. Rutherford, 1982. 2nd record.
66/2. FUMARIA CAPREOLATA L. *93, N. Aberdeen: Eden, GR 38/69.59. Farmyard. D. Welch,
1985, ABD. i
66/8 wir. FUMARIA OFFICINALIS L. subsp. WIRTGENII (Koch) Arcangeli *46, Cards.: Between
Byrlip and Coybal, GR 22/36.58. Disturbed soil. J. R. Akeroyd & C.D. Preston, 1985, CGE, det.
PD Sell)
+72/1. DIPLOTAXIS MURALIS (L.) DC. +76, Renfrews.: Paisley College, GR 26/48.63. A. J.
Silverside, 1977, PSY.
79/6. LEPIDIUM LATIFOLIUM L. +*32, Northants.: Collyweston Great Wood, GR 53/00.01.
Disturbed soil by A47. G. M. Gent, 1985.
[+79/7. LEPIDIUM GRAMINIFOLIUM L. 59, S. Lancs.: Delete record in Watsonia 15: 392; plant is
L. sativum L., conf. E. J. Clement]
+80/2. CoRONOPUS DIDYMUS (L.) Sm. *99, Dunbarton: Near Craigendoran Pier, GR 26/
31.81. Waste ground. A. Rutherford, 1985, E.
+81/1. CARDARIA DRABA (L.) Desv. 76, Renfrews.: R. Cart, Renfrew, GR 26/49.68. River
bank. A. McG. Stirling, 1981. 2nd record. *98, Main Argyll: Newtown Bay, Inveraray, GR
27/05.05. Shingle. B. H. Thompson, 1984, herb. B.H.T.
+ AETHIONEMA GRANDIFLORUM Boiss. *29, Cambs.: Sidney Sussex College, Cambridge, GR
52/43.58. Wall. P. D. Sell, 1966, CGE. Still present in 1977.
+90/2. BUNIAS ORIENTALIS L. *12, N. Hants.: Woolmer Forest, GR 41/8.3. Dr Drake, 1917,
RDG.
102/5. RORIPPA AMPHIBIA (L.) Bess. *76, Renfrews.: Newlands, Glasgow, GR 26/57.60.
Roadside hedge. P. Macpherson, 1970, herb. P. M., det. R. D. Meikle.
+105/1. ERYSIMUM CHEIRANTHOIDES L. *76, Renfrews.: Paisley, GR 26/47.65. Disturbed
ground. A. J. Silverside, 1977.
112/2. RESEDA LUTEA L. *98, Main Argyll: Glencruitten Road, Oban, GR 17/8.2. Rubble.
V. Hyslop, 1984, det. M. McC. Webster.
113/6. VIOLA CANINA L. 79, Selkirks.: Ettrickbridge End, GR 36/39.24. Rocks by river.
1984. 80, Roxburghs.: Scaw Knowe, Ancrum, GR 36/62.25. Grassland. 1985. Both R. W. M.
Corner, herb. R.W.M. Corner. 2nd records.
113/6 & 4. VIOLA CANINA L. X V. RIVINIANA Reichb. *70, Cumberland: Greengill, GR 35/
53.36. Sandy soil. R. W. M. Corner, 1985, LANC.
PLANT RECORDS 185
121/1. FRANKENIA LAEVIS L. 11, S. Hants.: Tournerbury, Hayling Island, GR 40/7.9.
Saltmarsh. Lady A. Brewis, 1984. 2nd extant record.
122/1. ELATINE HEXANDRA (Lapierre) DC. *76, Renfrews.: Loch Libo, GR 26/43.55. Sandy
loch bay. A. J. Silverside, 1979.
124/2. LYCHNIS VISCARIA L. 85, Fife: G. H. Ballantyne & M. Benstead, 1985. Locality
confidential. 1st record since 1903.
130/nan. PETRORHAGIA NANTEUILLI (Burnat) P. W. Ball & Heywood +41, Glam.: Dare
Valley Country Park, GR 22/99.02. Grassy bank. H. J. Dawson, 1985, NMW. 2nd record.
131/10. CERASTIUM DIFFUSUM Pers. 43, Rads.: Bach Howey viaduct, GR 32/10.42. Roadside.
R. G. Woods, 1985. 2nd record.
132/1. MyosoTON AQuaATICUM (L.) Moench +*70, Cumberland: Salta, GR 35/08.45. Rubbish
dump. M. Milne, 1985, LANC.
133/3. STELLARIA PALLIDA (Dumort.) Piré 51, Flints.: Graig, Tremeirchion, GR 33/0.7.
Limestone grassland. J. A. Green, 1985. 2nd record. 73, Kirkcudbrights.: Cardoness, GR 25/
56.52. Sandy ground near shore. O. M. Stewart, 1985. 2nd record.
136/2. SAGINA APETALA Ard. subsp. APETALA +*77, Lanarks.: Between Glasgow and
Renfrew, GR 26/51.67. Bare ground. P. Macpherson & E. Teasdale, 1984, herb. P. M., conf. F.
N. Hepper.
137/4. MINUARTIA HYBRIDA ( Vill.) Schischk. +61, S. E. Yorks.: Kipling Cotes, GR 44/92.43.
Railway track. Dr Grant, 1985. 2nd record.
143/1. SPERGULARIA RUBRA (L.) J. & C. Presl *98, Main Argyll: Balinoe, Loch Feochan, GR
17/85.20. Glen Lonan, GR 17/95.25. Both on edge of forest road. B. H. Thompson, 1984. Ist and
2nd records.
149/1b. MONTIA FONTANA L. subsp. CHONDROSPERMA (Fenzl) Walters 47, Monts.: Pen-y-
graig, Darowen, GR 23/84.01. Seasonally damp soil over rock. P. M. Benoit et al., 1985. 2nd
record.
7149/2. MonrTIA PERFOLIATA (Donn ex Willd.) Howell *52, Anglesey: Beaumaris, GR 23/6.7.
A. J. Armitstead, 1926. 76, Renfrews.: Paisley College, GR 26/48.63. Garden weed. A. J.
Silverside, 1977. 1st post-1930 record.
+154/17. CHENOPODIUM CAPITATUM (L.) Aschers. *26, W. Suffolk: Lakenheath R.A.F. base,
GR 52/7.8. Sandy ground. A. Brown, 1983, det. C. J. King.
156/1. ATRIPLEX LITTORALIS L. 75, Ayrs.: Monkton, Ayr, GR 26/34.27. Muddy tidal flats. J.
Swarbrick, 1964, herb. G. W. Swarbrick. 2nd record.
156/3. ATRIPLEX PROSTATA Boucher ex DC. *77, Lanarks.: R. Clyde, Shieldhall, GR 26/
54.66. P. Macpherson, 1985, herb P.M.
156/lon. x 3. ATRIPLEX LONGIPES Drejer X A. PROSTRATA Boucher ex DC. *25,\.E.i Suffolk:
Landguard Common, GR 62/28.32. Sandy ground. A. Copping, 1984, herb. E.M. & M. Hyde, det.
P. M. Taschereau.
4163/3. MALVA NICAEENSIS All. *70, Cumberland: Silloth, GR 35/11.53. N. Botham & M.
Milne, 1985, LANC, det. G. Halliday.
TABUTILON THEOPHRASTI Medicus *77, Lanarks.: King George V dock, Glasgow, GR 26/
52.66. Waste ground. P. Macpherson & E. Teasdale, 1984, herb. P. M.
167/1. RADIOLA LINOIDES Roth *98, Main Argyll: Degnish, Loch Melfort, oe LFTSAO.
Sandy soil by track. B. H. Thompson, 1984, herb. B.H.T.
168/7. GERANIUM SANGUINEUM L. *76, Renfrews.: S. of Gourock, GR 26/2.7. Grassy bank
above shore. A. McG. Stirling, 1979.
186 PLANT RECORDS
168/9. GERANIUM PYRENAICUM Burm. f. *+*77, Lanarks.: Carmunnock, GR 26/60.57. Waste
ground. P. & B. C. M. Macpherson, 1985, herb. P.M. +*93, N. Aberdeen: Leslie, GR 38/
59.24. Waste ground. D. Welch, 1984, ABD.
168/10. GERANIUM COLUMBINUM L. 80, Roxburghs.: Smailholm Crags, GR 36/63.34.
S.-facing ledge. R. W. M. Corner, 1985, herb. R.W.M.C. 1st record since 1853.
+168/ibi. X pla. GERANIUM IBIRICUM Cav. X G. PLATYPETALUM Fischer & C. A. Meyer *76,
Renfrews.: Lochwinnoch, GR 26/35.61. Quarry. A. J. Silverside, 1976, PSY. S. of Newton
Mearns, GR 26/52.54. Waste ground. P. Macpherson, 1978, herb. P.M., det. E. J. Clement. 1st
and 2nd records.
169/3. ERODIUM CICUTARIUM L’Her. +76, Renfrews.: Near Erskine Hospital, GR 26/45.72.
A. McG. Stirling, 1983. Only extant record.
4170/4. OXALIS EUROPAEA Jord. 43, Rads.: Argoed Mill, Doldowlod, GR 22/99.62. Refuse
tip. R. G. Woods, 1985. 2nd record.
171/1. IMPATIENS NOLI-TANGERE L. +67, S. Northumb.: Bolam, GR 45/08.81. Wet wood. C.
Crawford, 1985, herb. G.A. Swan. 2nd record.
7183/2. LUPINUS ARBOREUS Sims *73, Kirkcudbrights.: Near Meikle Culloch, GR 25/84.63.
Roadside bank. O. M. Stewart, 1985. <
185/2. GENISTA ANGLICA L. 51, Flints.: Llyn Helyg, GR 33/1.7. Marsh. J. A. Green, 1985.
2nd extant record.
190/1. MEDICAGO FALCATA L. +38, Warks.: Stonydelph, GR 43/24.01. Waste ground. M. A.
Arnold & H. Morrall, 1985. 2nd record.
+191/2. MELILOTUS OFFICINALIS (L.) Pall. *78, Renfrews.: Lonend, Paisley, GR 26/48.63.
Disturbed ground. A. McG. Stirling, 1980.
+191/4. MELILoTUsS INDICA (L.) All. *77, Lanarks.: Kelvinhaugh, GR 26/55.66. Waste
ground. P. Macpherson & A. McG. Stirling, 1985, herb. P.M. Ist record of established population.
192/1. TRIFOLIUM ORNITHOPODIOIDES L. 85, Fife: Balmerino, GR 37/35.24. Rocky spur by
shore. M. Benstead, 1985. Ist post-1930 record.
192/10. TRIFOLIUM STRIATUM L. +*76, Renfrews.: Paisley, GR 26/48.64. Lawn weed. A. J.
Silverside, 1980.
195/2. Lotus TENUIS Waldst. & Kit. ex Willd. 67, S. Northumb.: Near Newlands, GR 45/
09.55. Roadside bank. G. A. Swan, 1985, herb. G.A.S. 1st post-1930 record.
200/1. ASTRAGALUS DANICUS Retz. *93, N. Aberdeen: Sands of Forvie, GR 48/00.235.
Rabbit-grazed bank. B. Davis, 1981, conf. D. Welch.
202/1. ORNITHOPUS PERPUSILLUS L. 99, Dunbarton: Duntocher, Glasgow, GR 26/48.72.
Sandy trackside. A. McG. Stirling & A. Rutherford, 1985, E. 2nd record, Ist since 1891.
+203/1. CORONILLA VARIA L. *67, S. Northumb.: Near Wylam, GR 45/12.64. Hedge by
disused railway. G. A. Swan, 1985, herb. G.A.S.
206/2. VICIA TETRASPERMA (L.) Schreb. 73, Kirkcudbrights.: Balmae Ha’en, GR 25/67.44.
Rocky ground by shore. O. M. Stewart, 1985, E. 1st record since 1843 record from
Balmae. +80, Roxburghs.: Hardies Hill, Hawick, GR 36/50.13. Railway ballast. A. J. Smith,
1985, herb. R.W.M. Corner. Ist record since 1917.
4206/6. VICIA VILLOSA Roth 70, Cumberland: Silloth, GR 35/11.53. M. Milne, 1985, LANC,
det. G. Halliday. 2nd record, Ist this century.
+206/6 vill. VicIA VILLOSA Roth subsp. vILLOSA *77, Lanarks.: Kelvinhaugh, GR 26/55.66.
Waste ground. A. McG. Stirling, 1985, GL.
206/16. VICIA LATHYROIDES L. +76, Renfrews.: Paisley College, GR 26/47.63. Garden weed.
A. J. Silverside, 1980. 2nd record.
PLANT RECORDS 187
206/seg. VICIA SEGETALIS Thuill. *4, N. Devon: Braunton Burrows, GR 21/45.33. Sand
dunes. L. J. Margetts, 1985.
207/2. LATHYRUS NISSOLIA L. 57, Derbys.: Lower Hartshay, GR 43/39.51. Grassy roadside
bank. Mr Wilson, 1985, det. R. Smith. Ist post-1930 record.
+209/2 x 1. SPIRAEA DOUGLAS! Hook. X S. SALICIFOLIA L. *69, Westmorland: N. end of
Grasmere, GR 35/33.06. Lakeside. G. Halliday, 1984, LANC, det. A. J. Silverside.
+ARUNCUS DIOICUS (Walter) . Fernald *77, Lanarks.: Shiels, Glasgow, GR 26/52.66.
Grassland. P. Macpherson & E. Teasdale, 1984, conf. E. J. Clement. Male plants. Ibrox, GR 26/
55.64. Scrubby woodland. P. Macpherson, 1985. Female plants. Both herb. P.M. Ist and 2nd
records.
211/1. RUBUS CHAMAEMORUS L. *76, Renfrews.: Misty Low Range N.W. of Queenside Loch,
GR 26/29.64. Calluna moor at 400 m altitude. H. Galbraith, 1971.
211/11/2. RusBus scissus W.C.R. Wats. *59,S. Lancs.: Withnell, GR 34/61.23. Moor. A.
Newton & M. Smith, 1984, det. A. N.
211/11/139. RuBus PRocerus P. J. Muell. *76, Renfrews.: Between Paisley and Linwood,
GR 26/46.65. Waste ground. A. McG. Stirling, 1981.
211/11/178. RuBus CRINIGER (E. F. Linton) Rogers *59, S. Lancs.: Heapey Road, Heapey,
GR 34/60.20. Roadside. A. Newton & M. Smith, 1984, det. A. N.
211/11/204. RuBUS RADULA Weihe ex Boenn. *77, Lanarks.: Overtown, GR 26/80.52. A.
McG. Stirling, 1981.
211/11/213. RuBUS ECHINATOIDES (Rogers) Sudre -597S) dbanes.: Birkacre, GR. 34/57 .15,
Waste ground. A. Newton & M. Smith, 1984, det. A. N.
211/11/356. RuBUS DASYPHYLLUS (Rogers) Rogers *76, Renfrews.: Braidbar, Giffnock, GR
26/56.59. Waste ground. A. McG. Stirling, 1980.
¥211/11/coc. RUBUS COCKBURNIANUS Hemsley *77, Lanarks.: Between Glasgow and
Renfrew, GR 26/51.67. Side of old railway. P. Macpherson & E. Teasdale, 1983, herb. P.M.
211/11/new. RUBUS NEWBOULDI Babington *59_S. Lancs.: Withnell, GR 34/61.23. Moor. A.
Newton & M. Smith, 1984, det. A. N.
211/11/rob. RuBus Rosi (W. C. R. Wats.) A. Newton *59,S. Lancs.: Coppull, GR 34/56.15.
Sidings. A. Newton & M. Smith, 1984, det. A. N.
211/11/vill. RUBUS VILLICAULIFORMIS A. Newton. *6, N. Somerset: Blackdown, Mendip, GR
31/48.57. Moorland. R. D. Randall, 1982, herb. R.D.R., det. A. Newton.
4212/8. POTENTILLA NORVEGICA L. 76, Renfrews.: Langbank, GR 26/38.73. T. Wise, 1905,
GL. 2nd record.
4212/9. POTENTILLA INTERMEDIA L. *77, Lanarks.: Cambuslang, GR 26/63.60. Rough
grassland. P. Macpherson & E. L. S. Lindsay, 1985, herb. P.M., conf. R. J. Pankhurst.
212/14 X 15. POTENTILLA ANGLICA Laichard. X P. REPTANS L. *80, Roxburghs.: Scaw Knowe,
Ancrum, GR 36/62.25. Bare turf. R. W. M. Corner, 1985, herb. R.W.M.C., det. B. Harold.
220/3/2 ALCHEMILLA VESTITA (Buser) Raunk. *93, N. Aberdeen: Pitcaple, GR 38/73.26.
Disused railway by wood. D. Welch, 1985, ABD.
4222/2. SANGUISORBA CANADENSIS L. 99, Dunbarton: Garscadden, GR 26/52.71. Waste
ground. J. H. Dickson, 1984, GL. 2nd record.
4223/2. POTERIUM POLYGAMUM Waldst. & Kit. *70, Cumberland: Denton Holme, GR 35/
39.50. Rough ground. R. E. Groom, 1985, LANC.
4227/2. COTONEASTER SIMONSII Bak. *77, Lanarks.: Linn, Glasgow, GR 26/58.58. River
bank. P. Macpherson, 1984, herb. P.M.
188 PLANT RECORDS
+227/3. COTONEASTER HORIZONTALIS Decne. *77, Lanarks.: Shieldhall, Glasgow, GR 26/
53.65. P. Macpherson, 1984, herb. P.M. Ist record of established population.
+227/bul. COTONEASTER BULLATUS Boiss. 44, Carms.: Laugharne, GR 22/30.10. Wooded
cliff. J. Rees & T. S. Crosby, 1985. 2nd record.
*+231/lam. AMELANCHIER LAMARCKII F. G. Schroeder *70, Cumberland: E. of Thurstonfield
Lough, GR 35/32.56. R. E. Groom, 1978, LANC. Beacon Hill, Penrith, GR 35/52.31. R. W. M.
Corner, 1985, LANC. 1st and 2nd records.
232/5/3. SORBUS EMINENS E. F. Warb. 6, N. Somerset: King’s Wood, Yatton, GR 31/45.64.
Wood. R. G. B. Roe, 1984, herb. P.J.M. Nethercott, det. P.J.M.N. 2nd record.
232/5/5. SORBUS PORRIGENTIFORMIS E. F. Warb. *5, S. Somerset: Yearnor Wood, Porlock,
GR 21/84.48. Wood. J. Bevan & Lady R. FitzGerald, 1984, CGE, det. P. D. Sell & P. J. M.
Nethercott.
+235/3. SEDUM SPURIUM Bieb. 38, Warks.: Near Arley, GR 42/28.89. Concrete debris. J.
Oliver, 1971, det. J. G. Hawkes. 2nd and earliest record.
+237/hel. CRASSULA HELMSII (T. Kirk) Cockayne *5, S. Somerset: Clapton Court, W.
Crewkerne, GR 31/41.06. Ornamental pond. J. Ounsted, 1983.
+240/1. TELLIMA GRANDIFLORA (Pursh) Dougl. ex Lindl. *77, Lanarks.: Linn, Glasgow, GR
26/58.58. River bank. P. Macpherson, 1984, herb. P.M. Cambuslang, GR 26/63.61. River bank. E.
L. S. Lindsay & P. Macpherson, 1985. Ist and 2nd records.
246/2. RIBES SPICATUM Robson *93, N. Aberdeen: Pitcaple, GR 38/73.26. Wood. D. Welch,
1985, ABD.
250/1. LYTHRUM PORTULA (L.) D. A. Webb 80, Roxburghs.: Smailholm Craigs, GR 36/63.34.
Pool in hollow. R. W. M. Corner, 1985, herb. R.W.M.C. Ist record since 1915.
251/1. DAPHNE MEZEREUM L. +*51, Flints.: Ddol Uchaf Nature Reserve, GR 33/1.7. Wood.
B. Ing, 1985.
254/3 xk 9. EPILOBIUM MONTANUM L. X E. OBSCURUM Schreb. *47, Monts.: Llwydiarth Hall,
Aberllefenni, GR 23/77.10. Vegetable garden. P. M. Benoit et al., 1985, NMW.
4254/6. EPILOBIUM CILIATUM Rafin. *93, N. Aberdeen: Gask, GR 38/72.47. Garden. D.
Welch, 1984, ABD.
254/+6 X 3. EPILOBIUM CILIATUM Rafin. X E. MONTANUM L. *4, N. Devon: Cove, Bampton,
GR 21/95.19. Roadside bank. L. J. Margetts, 1985, herb. L. J. M.
254/+6 4. EPILOBIUM CILIATUM Rafin. X E. LANCEOLATUM Seb. & Mauri *4, N. Devon:
Cove, Bampton, GR 21/95.19. Rocky slope by road. L. J. Margetts, 1985, herb. L.J.M.
259/1. MYRIOPHYLLUM VERTICILLATUM L. 67, S. Northumb.: Reigh Burn near Throckley, GR
45/14.65. Pond. G. A. & M. Swan, 1985, herb. G.A.S. Ist post-1930 record and most northerly
English locality.
+AUCUBA JAPONICA Thunb. *44, Carms.: Gellideg, Llandyfaelog, GR 22/42.10. Wood. T. S.
Crosby, 1985. Ist Welsh record.
+271/1. ASTRANTIA MAJOR L. *111, Orkney: North Walls, GR 39/27.89. Roadside. H. Smith,
1985.
272/2. ERYNGIUM CAMPESTRE L. +*5, S. Somerset: Hurstone Farm, Waterrow, GR 31/05.25.
Pasture. R. G. Corns, 1984.
274/1. ANTHRISCUS CAUCALIS Bieb. 67, S. Northumb.: Hadston Links, GR 46/27.00. G. A. &
M. Swan, 1985, herb. G.A.S. 1st localized post-1930 record.
+274/3. ANTHRISCUS CEREFOLIUM (L.) Hoffm. *43, Rads.: Boughrood, GR 32/1.3. Roadside
verge. T. G. Evans, 1985.
PLANT RECORDS 189
+280/1. SMYRNIUM OLUSATRUM L. 70, Cumberland: E. of Anthorn Chapel, GR 35/19.58.
Open scrub by stream. R. E. Groom, 1984, LANC. 2nd record.
285/1. APIUM GRAVEOLENS L. *77, Lanarks.: Yorkhill, Glasgow, GR 26/55.66. River bank.
P. Macpherson, 1985, herb. P.M.
285/4. APIUM INUNDATUM (L.) Reichb. f. 77, Lanarks.: Carmunnock, Glasgow, GR 26/
61.57. Reservoir. P. Macpherson, 1985. 2nd record.
286/2. PETROSELINUM SEGETUM (L.) Koch *49, Caerns.: Bardsey Island, GR 23/1.2. P. M.
Butler, 1933, det. A. J. Wilmott.
297/1. BERULA ERECTA (Huds.) Coville *98, Main Argyll: Near Creagantairbh, GR 17/80.00.
Slow-flowing stream. B. H. Thompson, 1984.
300/2. OENANTHE PIMPINELLOIDES L. *35, Mons.: Tynewydd N.W. of Bettws, GR 31/27.91.
Roadside verge. T. G. Evans, 1985, herb. T.G.E. 1st Welsh record.
300/3. OENANTHE SILAIFOLIA Bieb. *57, Derbys.: Priestcliffe Lees, GR 42/15.72. N.-facing
slope. G. & G. Wheeldon, 1985, DBY, det. A. Willmot.
311/42 X 1. HERACLEUM MANTEGAZZIANUM Somm. & Levier X H. SPHONDYLIUM L. wee
Lanarks.: East Kilbride, GR 26/66.55. River bank. P. Macpherson & E. L. S. Lindsay,
1983. *80, Roxburghs.: Ormiston Mill, GR 36/70.27. Riverside. F. Stewart & J. Grace, 1976.
*319/6. EUPHORBIA DULCIS L. *98, Main Argyll: Craignish Castle, GR 17/75.00. Shrubby
bank. B. H. Thompson, 1984.
319/12. EUPHORBIA PORTLANDICA L. 11, S. Hants.: Stanswood Bay, Calshot, GR 40/4.9.
Shingle beach. R. P. Bowman & J. Venner, 1984. 1st post-1930 record.
320/1/4. POLYGONUM ARENASTRUM Boreau. *98, Main Argyll: Glen Lochy, GR 27/20.25.
Roadside verge. B. H. Thompson, 1984.
320/13. POLYGONUM MITE Schrank 25, E. Suffolk: North Cove, GR 62/47.90. Wet ride. F. W.
Simpson, 1984. Only extant record. 68, Cheviot: Detchant Pond, GR 46/08.36. G. A. Swan,
1985, herb. G.A.S., det. J. R. Akeroyd. 2nd record.
320/14. POLYGONUM MINUS Huds. *76, Renfrews.: Pilmuir, Newton Mearns, GR 26/51.54.
Loch margin. R. Mackechnie & P. Macpherson, 1975. 1st post-1930 record.
4320/19. REYNOUTRIA JAPONICA Houtt. *111, Orkney: Ramsdale Burn, Orphir, GR 57/34.01.
Burnside, Kirkwall Bourgh, GR 57/45.11. Waste land. Both E. R. Bullard, 1983. 1st and 2nd
records.
+320/20. REYNOUTRIA SACHALINENSE (F. Schmidt) Nakai *76, Renfrews.: Pollok, GR 26/
54.62. Edge of wood. P. Macpherson, 1980, herb. P.M.
325/1/3. RUMEX TENUIFOLIUS (Wallr.) Love *75, Ayrs.: Bogside, Stevenston, GR 26/31.40.
Sandy waste ground. A. McG. Stirling, 1979, E.
4325/5. RUMEX ALPINUS L. *93, N. Aberdeen: Blairfowl, GR 38/81.38. Roadside. D. Welch,
1984, ABD.
325/8 RUMEX LONGIFOLIUS DC. +76, Renfrews.: Cowglen, Glasgow, GR 26/54.61. Central
reservation of road. P. Macpherson, 1983. 1st post-1930 record.
343/15. SALIX PHYLICIFOLIA L. *93, N. Aberdeen: Whinnyfold, GR 48/08.33. Coastal cliff. D.
Welch, 1985, ABD. .
343/16b. SALIX REPENS L. subsp. ARGENTEA (Sm.) G. & A. Camus *49, Caerns.: Near
Caernarfon, GR 23/4.6. G. C. Druce, 1919. Bardsey Island, GR 23/1.2. P. M. Butler, 1933, det.
A. J. Wilmott. 1st and 2nd records.
358/4. VACCINIUM OxyYcoccus L. *93, N. Aberdeen: Strathbogie, GR 38/47.23. Raised bog
R. E. C. Ferreira & J. G. Roger, 1983.
190 PLANT RECORDS
358/5. VACCINIUM MICROCARPUM (Rupr.) Hook. f. *93, N. Aberdeen: Cairnbrallan, GR 38/
33.24. Blanket peat. D. Welch, 1984, ABD.
+358/cor. VACCINIUM CORYMBOSUM L. *11, S. Hants.: Ashley Heath, GR 41/10.04. Heath. R.
P. Bowman, 1980, herb. R.P.B., det. E. J. Clement.
359/1. PYROLA MINOR L. 52, Anglesey: Newborough Forest, GR 23/4.6. Pine plantation. Mr
& Mrs J. Davidson, 1984, NMW, det. R. H. Roberts & N. Brown. Ist record since 1813.
370/3. LYSIMACHIA VULGARIS L. *109, Caithness: Reisgill Burn, GR 39/24.35. Wet grassland.
D. Wells, 1983.
371/1. TRIENTALIS EUROPAEA L. 69, Westmorland: Firbank Fell, GR 34/6.9. Under Calluna.
T. & K. Wilson, 1985. 1st record since 1909.
372/4. ANAGALLIS MINIMA (L.) E. H. L. Krause 43, Rads.: Road to Pentre Caeau, Llandeilo
Graban, GR 32/08.44. Damp soil by stream. R. G. Woods, 1985, NMW. 2nd record. *47,
Monts.: Manafon, GR 33/1.0. H. H. & S. Haines & R. D. Tweed, 1938.
387/1. NYMPHOIDES PELTATA (S. G. Gmel.) Kuntze +*4, N. Devon: Halberton, GR 31/00.13.
Canal. B. Benfield & L. J. Margetts, 1984.
+390/1. OMPHALODES VERNA Moench *76, Renfrews.: Rouken Glen Park, Glasgow, GR 26/
54.58. Bank above stream. A. McG. Stirling, 1981, E.
392/6. SYMPHYTUM TUBEROSUM L. +*52, Anglesey: Marianglas, GR 23/50.84. Waste ground.
R. H. Roberts, 1985.
+392/7. SYMPHYTUM GRANDIFLORUM DC. *67, S. Northumb.: Diptonfoot, GR 35/99.60.
Wood. G. A. Swan, 1985, herb. G.A.S.
+AMSINCKIA INTERMEDIA Fischer & C. A. Meyer *5,_S. Somerset: Taunton, GR 31/2Z3.23.
Golfcourse. T. T. Freeston, 1984. *14, E. Sussex: Malling Down, Lewes, GR 51/43.10.
Disturbed ground. A. O. Chater, 1984, BM, det. M. Hyde.
400/8 umb. MyosoTIs ARVENSIS (L.) Hill subsp. UMBRATA (Rouy) O. Schwarz *76, Renfrews.:
Pollok estate, GR 26/54.61. Wood. A. J. Silverside, 1979, herb. A. J. S.
402/1. MERTENSIA MARITIMA (L.) Gray 73, Kirkcudbrights.: Dundrennan Army Range, GR
25/70.43. Shingle beach. D. Hawker, 1985. 1 plant. 1st record since 1882 record near here.
406/1 ros. CALYSTEGIA SEPIUM (L.) R.Br. subsp. ROSEATA Brummitt +*80, Roxburghs.: Old
station yard, Newcastleton, GR 35/48.87. Embankment. R. W. M. Corner, 1985, herb. R.W.M.C.
Probably present since 1960 (cf. Proc. bot. Soc. Br. Isl. 6: 290).
+406/2. CALYSTEGIA PULCHRA Brummitt & Heywood *76, Renfrews.: Road to Kerse, GR 26/
33,55. Hedgerow. C: BPead’& E. Rv?) Conacher,-198!. *93, N. Aberdeen: Eden, GR 38/
69.59. Grassland. D. Welch, 1985, ABD.
411/1. HyoscyAMUS NIGER L. +99, Dunbarton: Duntocher, Glasgow, GR 26/49.71. Waste
ground. A. McG. Stirling, 1985, E. 1st post-1930 record.
+413/nit. SOLANUM NITIDIBACCATUM Bitter *25, E. Suffolk: Landguard Common, GR 62/
28.32. Sandy shingle. A. Copping, 1983. Chillesford, GR 62/38.51. Red Crag pit. F. W. Simpson,
1984, det. A. C. Leslie. 1st and 2nd records.
416/7 X +pyr. VERBASCUM NIGRUM L. X V. PYRAMIDATUM Bieb. *26, W. Suffolk: Kentford
Heath, Kennett, GR 52/70.68. Waste ground. G. M. S. Easy, 1971, herb. G.M.S.E.
+416/10. VERBASCUM VIRGATUM Stokes 70, Cumberland: Silloth, GR 35/11.53. N. Botham &
M. Milne, 1985.
+416/pyr. VERBASCUM PYRAMIDATUM Bieb. *26, W. Suffolk: Kentford Heath, Kennett, GR
52/70.68. Waste ground. G.M.S. Easy, 1971.
$420/2. LINARIA PURPUREA (L.) Mill. *72, Dumfriess.: Hightae, Lockerbie, GR 35/09.78.
Disturbed ground. L. Kungu & M. E. R. Martin, 1984.
PLANT RECORDS 19]
420/+2 x 3. LINARIA PURPUREA (L.) Mill. xX L. REPENS (L.) Mill. *48, Merioneth:
Penrhynedeudraeth, GR 23/61.38. By railway ballast. A. P. Conolly, 1983, herb. A.P.C., det. C.
A. Stace.
420/3. LINARIA REPENS (L.) Mill. +*75, Ayrs.: Ardrossan B. P. Terminal, GR 26/22.42. Rock
and gravel. B. Simpson, 1984.
420/3 x 4. LINARIA REPENS (L.) Mill. x L. VULGARIS Mill. *47, Monts.: Machynlleth railway
station, GR 23/74.01. B. Gale et al., 1985. *75, Ayrs.: Ardrossan B. P. Terminal, GR 26/22.42.
Rock and gravel. B. Simpson, 1984.
422/2. KICKXIA ELATINE (L.) Dumort. +43, Rads.: Below Yr Allt, Llandeilo Graban, GR 32/
08.44. Rubble. R. G. Woods, 1985, NMW. 2nd record.
+424/5. SCROPHULARIA VERNALIS L. *32, Northants.: Ferry Meadows, Peterborough, GR 52/
15.98. Wood. S. Wells, 1982, ABRN.
+425/1 x 2 x var. MIMULUsS GuUTTATUS DC. X M. LuTEUS L. X M. vARIEGATUS Loddiges "aT:
Lanarks.: Near Carmunnock, GR 26/60.56. Streamside. P. Macpherson, 1985, herb. P.M., det. A.
J. Silverside.
4430/14. VERONICA PEREGRINA L. *76, Renfrews.: Newlands, Glasgow, GR 26/57.60. Garden
weed. P. Macpherson, 1972, herb. P. M., det. J. E. Lousley. Finlaystone House, GR 26/36.73.
Nursery weed. A. J. Silverside, 1975.
430/20 hed. VERONICA HEDERIFOLIA L. subsp. HEDERIFOLIA *4, N. Devon: N. of Waytown
Tunnel, Greenham, GR 31/07.19. Hedgebank. L. J. Margetts, 1984. Near Fairlynch Quarry,
Braunton, GR 21/47.37. Hedgebank. W. H. Tucker, 1985. 1st and 2nd records.
430/20 luc. VERONICA HEDERIFOLIA L. subsp. LUCORUM (Klett & Richter) Hartl *4, N. Devon:
Between Tidcombe and Manley Bridge, GR 21/98.12. Hedgebank. L. J. Margetts, 1984. *76,
Renfrews.: Finlaystone estate, GR 26/36.73. Woodland path. A. J. Silverside, 1977.
435/1/13. EUPHRASIA NEMOROSA (Pers.) Wallr. *77, Lanarks.: Browncastle, GR 26/62.43.
Heath. P. Macpherson, 1981, herb. P. M., det. A. J. Silverside.
435/1/15 x 13. EUPHRASIA CONFUSA Pugsl. X E. NEMOROSA (Pers.) Wallr. *35, ‘Mens. :
Chepstow Park Wood, GR 31/50.98. Woodland path. T. G. & U. T. Evans, 1985, herb. T. G. E.,
det. A. J. Silverside.
437/1. PARENTUCELLIA VISCOSA (L.) Caruel +35, Mons.: Bank of R. Usk, Llanbadoc, GR 31/
37.99. S. J. Tyler, 1985, herb. T.G. Evans.
440/2. OROBANCHE PURPUREA Jacq. +*70, Cumberland: Near Senhouse Dock, Maryport, GR
35/0.3. Waste ground. M. Milne, 1985, LANC, conf. D. J. Hambler.
440/4. OROBANCHE ALBA Steph. ex Willd. *6, N. Somerset: Berrow, GR 31/29.52. Fixed
dunes. J. R. Comley, 1985.
440/10. OROBANCHE HEDERAE Duby *13, W. Sussex: N.W. of Chichester, GR 41/7.1.
Parkland. Rev. E. A. Pratt, 1985, conf. M. Briggs.
440/11. OROBANCHE MARITIMA Pugsl. 4, N. Devon: Croyde, GR 21/43.39. H. J. M. Bowen,
1984. 2nd record.
441/3. PINGUICULA VULGARIS L. 11, S. Hants.: Acres Down, Lyndhurst, GR 41/2.0. Basic
flush. A. Bolton, 1984. 1st post-1930 record.
441/4. PINGUICULA GRANDIFLORA Lam. +*48, Merioneth: Brithdir near Dolgellau, GR 23/7.1.
Basic boggy rill. P. M. Benoit, 1983, NMW. Ist Welsh record.
451/3. CALAMINTHA NEPETA (L.) Savi *49, Caerns.: South Shore, Llandudno, GR 23/7.8. F.
T. Jones, 1930.
459/3. STACHYS ARVENSIS (L.) L. 93, N. Aberdeen: Eden, GR 38/70.60. Turnip field. D.
Welch, 1985, ABD. Ist post-1930 record.
192 PLANT RECORDS
462/1. LAMIUM AMPLEXICAULE L. *42, Brecs.: Glandulas, Llanafan Fawr, GR 22/94.53.
Roadside verge. R. G. Woods, 1985, NMW.
462/2. LAMIUM MOLUCCELLIFOLIUM Fr. *52, Anglesey: Beaumaris, GR 23/60.76. Waste
ground. R. M. Burton, 1985, NMW, conf. R. G. Ellis.
462/3. LAMIUM HYBRIDUM Vill. 93, N. Aberdeen: Sandford Bay, GR 48/12.43. Grassland
track. D. Welch, 1985, ABD. ist post-1930 record.
+464/rus. PHLOMIS RUSSELLIANA (Sims) Bentham *85, Fife: Coultra, GR 37/35.32. Roadside.
M. Benstead, 1981.
465/4/2 X 1. GALEOPSIS BIFIDA Boenn. X G. TETRAHIT L. *47, Monts.: Pont ar Byllfa, GR 23/
89.12. Roadside bank. P. M. Benoit & M. Wainwright, 1985, NMW.
465/4/2. GALEOPSIS BIFIDA Boenn. *47, Monts.: Pont ar Byllfa, GR 23/89.12. Roadside
bank. Ola Luest, Cwm Nant-y-Meichiaid, GR 33/12.14. Wood. Both P. M. Benoit & M.
Wainwright, 1985. 1st and 2nd records. *77, Lanarks.: Shieldhall, GR 26/53.66. 1984. Sheils,
Glasgow, GR 26/52.67. 1985. Both rough grass. P. Macpherson, herb. P. M. 1st and 2nd records.
469/2. SCUTELLARIA MINOR Huds. 51, Flints.: Above Ffynnon Beuno, Tremeirchion, GR 33/
0.7. Marsh. J. A. Green & G.Wynne, 1985. 2nd extant record.
472/2. PLANTAGO MEDIA L. +*111, Orkney: Graemshall, Holm, GR 57/48.01.E. Meek, 1984.
475/2. CAMPANULA TRACHELIUM L. +*77, Lanarks.: Between Glasgow and Renfrew, GR 26/
51.67. By old railway. P. Macpherson & E. Teasdale, 1983, herb. P. M., det. E. J. Clement.
1475/3. CAMPANULA RAPUNCULOIDES L. *5, S. Somerset: Ashill, GR 31/33.18. Railway
cutting. W. L. Landsell, 1985, herb. R.G.B. Roe. *§2, Anglesey: Baron Hill, Beaumaris, GR
23/5.7. A. J. Armitstead, 1926.
485/14. GALIUM PARISIENSE L. *25, E. Suffolk: Westleton, GR 62/45.69. Sandy heath. G. W.
Maybury & E. Beaumont, 1985, herb. E. M. & M. Hyde.
+491/cae. LONICERA CAERULEA L. *57, Derbys.: Wye Dale, GR 43/09.72. Bank by R. Wye.
L. Storer, 1985.
494/1. VALERIANELLA LOCUSTA (L.) Betcke 43, Rads.: Knocklas Castle, GR 32/25.74. Shale
below castle. J. & C. Port, 1984. 2nd record.
494/2. VALERIANELLA CARINATA Lois. *44, Carms.: Ammanford, GR 22/63.12. Chapel
graveyard. A. M. Pell, 1985, NMW, det. R. G. Ellis.
495/1. VALERIANA OFFICINALIS L. subsp. SAMBUCIFOLIA (Mikan fil.) Celak. *49, Caerns.:
Dinas Duille, GR 23/4.5. Aberglaslyn, GR 23/5.4. Both N. Woodhead, 1929. Ist and 2nd records.
+CEPHALARIA GIGANTEA (Ledeb.) Bobrov 25, E. Suffolk: Fen Lane, East Bergholt, GR 62/
0.3. Laneside. I. Rose, 1984. 2nd record.
498/1. KNAUTIA ARVENSIS (L.) Coult. 98, Main Argyll: Connel Airstrip, GR 17/90.36.
Disused railway. P. Wormell, 1983.
502/1. BIDENS CERNUA L. 43, Rads.: Near Cloggie, Llangunllo, GR 32/22.71. Edge of small
pond. P. Richards, 1964. 2nd record.
+504/1. ARTEMISIA ARTEMISIIFOLIA L. 77, Lanarks.: Shieldhall, GR 26/53.66. Waste ground.
P. Macpherson, 1984, herb. P.M. Ist record since casual occurrence in 1919.
506/2 * 1. SENECIO AQUATICUS Hill * S. JACOBAEA L. *79_ Selkirks.: Ettrick Water above
Howden cauld, GR 36/44.27. Riverbank. *80, Roxburghs.: Ettrick Water above Overscloss,
GR 36/47.30. Riverbank. Both R. W. M. Corner, 1983, herb. R.W.M.C., det. D. R. McKean.
506/+4 X 7. SENECIO SQUALIDUS L. X S. viscosus L. *32, Northants.: Wellingborough, GR
42/90.67. Old railway sidings. A. Robinson, 1983, herb. A.R.
PLANT RECORDS 193
+506/18. SENECIO BICOLOR (Willd.) Tod. subsp. CINERARIA (DC.) A. O. Chater *46, Cards.:
S. of Aberaeron harbour, GR 22/45.62. Festuca rubra sward on shingle. A. O. Chater, 1985.
506/+18 X 1. SENECIO BICOLOR (Willd.) Tod. subsp. CINERARIA (DC.) A. O. Chater x S. JACOBAEA
| ee *47, Monts.: Felin-Gerrig, E. of Machynlleth, GR 23/76.00. Disturbed soil. C. A. Small,
1985, det. P. M. Benoit.
+506/ver. SENECIO VERNALIS Waldst. & Kit. *29, Cambs.: Fen Drayton gravel pit, GR 52/
34.69. Recently sown grass. G. M. S. Easy, 1985, herb. G.M.S.E.
+507/2. DORONICUM PLANTAGINEUM L. *32, Northants.: Between West Haddon and Long
Buckley, GR 42/62.71. Roadside. I. Cameron, 1981. *43, Rads.: W. of Stow Bridge near
Milebrook, GR 32/30.72. Wood. J. Roper, 1978. 70, Cumberland: W. of Penton Station, GR
35/43.76. Roadside. R. W. M. Corner, 1985, LANC. 2nd record.
+518/3. SOLIDAGO GIGANTEA Ait. 77, Lanarks.: Between Glasgow and Renfrew, GR 26/
51.67. Clearing in scrub. E. Teasdale & P. Macpherson, 1984, herb. P. M., det. A. J. Silverside.
2nd record.
519/1. ASTER TRIPOLIUM L. *77, Lanarks.: R. Clyde, Yoker, Glasgow, GR 26/53.66. A.
McG. Stirling et al., 1985. R. Clyde, Shieldhall, GR 26/53.66. P. Macpherson, 1985, herb. P. M.
1st and 2nd records.
526/3. ANTHEMIS ARVENSIS L. +99, Dunbarton: Clydebank, GR 26/49.70. Landscaped area.
A. McG. Stirling, 1985, E. 1st record since 1891.
+533/3. CHRYSANTHEMUM MAXIMUM Ramond *77, Lanarks.: Between Glasgow and Renfrew,
GR 26/51.67. Waste ground. P. Macpherson & E. Teasdale, 1983.
539/1. CARDUUS TENUIFLORUS Curt. *75, Ayrs.: Horse Island, GR 26/21.42. Sandy ridge. B.
Simpson, 1984.
540/4 X 3. CIRSIUM ARVENSE (L.) Scop. X C. PALUSTRE (L.) Scop. *99, Dunbarton: Gartlea
Farm, Gartocharn, GR 26/45.83. I. Christie, 1976, det. A. McG. Stirling.
550/3. LEONTODON TARAXACOIDES (Vill.) Mérat *93, N. Aberdeen: Fraserburgh, GR 48/
00.65. Moist dunes. D. Welch, 1985, ABD.
555/1. MYCELIS MURALIS (L.) Dumort. 79, Selkirks.: Corbielinn, Selkirk, GR 36/45.28.
Wall-top. R. W. M. Corner, 1985. 2nd record.
558/1/45. HIERACIUM LASIOPHYLLUM Koch *44, Carms.: Craig Clyngwyn, GR 22/77.47.
Base-rich cliffs. N.C.C. Field Unit, 1982.
558/1/56. HIERACIUM SUBRUDE (Arv.-Touv.) Arv.-Touv. *78, Peebless.: Windy Gowl,
Stonypath, W. Linton, GR 36/14.54. Dry rocks. D. J. McCosh, 1979, herb. D.J.McC., det. P. D.
Sell.
558/1/83. HIERACIUM PSEUDOSARCOPHYLLUM Pugs]. *77, Lanarks.: Lang Gill, Culter Fell, GR
36/04.27. Rocks by pool. D. J. McCosh, 1984, herb. D. J. McC., det. P. D. Sell.
558/1/103. HiERACIUM STENSTROEMII (Dahlst.) Johans. *44, Carms.: Caeo, GR 22/67.39.
Bridge parapet. I. M. Vaughan, 1965, NMW, det. J. Bevan.
558/1/125. HTERACIUM AURATIFLORUM Pugsl. *83, Midlothian: Arthur’s Seat, Edinburgh, GR
36/2.7. G. Taylor, 1934, BM, det. P. D. Sell & C. West.
558/1/133. HIERACIUM STENOPHYES W. R. Linton *78, Peebless.: Williamslee Burn, Leithen,
GR 36/32.46. D. J. McCosh, 1981, herb. D. J. McC., det. P. D. Sell.
558/1/144. HIERACIUM ANGUSTISQUAMUM (Pugsl.) Pugsl. *46, Cards.: Nant Rhuddnant gorge,
GR 22/80.78. Rocks by stream. A. O. Chater, 1984, NMW, det. P. D. Sell. 1st Welsh
record. *57, Derbys.: Rowsley, GR 43/26.64. Bridge, disused railway sidings. R. Smith, 1985,
herb. R. S., det. D. J. McCosh & P. D. Sell.
558/1/206. HIERACIUM LATOBRIGORUM (Zahn) Roffey *78, Peebless.: Neidpath Castle,
194 PLANT RECORDS
Peebles, GR 36/23.40. Basalt rocks near R. Tweed. D. J. McCosh, 1981, herb. D.J.McC., det. P.
D. Sell:
558/1/222. HIERACIUM SALTICOLA (Sudre) Sell & West *44, Carms.: Cynghordy, GR 22/
80.40. Railway embankment. I. M. Vaughan, 1965, NMW, det. J. Bevan. *77, Lanarks.:
Between Glasgow and Renfrew, GR 26/51.67. Waste ground. P. Macpherson & E. Teasdale,
1983, herb. P. M., det. A. McG. Stirling.
558/1/223. HIERACIUM VAGUM Jord. *76, Renfrews.: Elderslie, GR 26/45.63. Waste ground.
A.McG. Stirling, 1981, E.
+558/2/5. HIERACIUM FLAGELLARE Willd. *77, Lanarks.: Kenmuirhill, GR 26/66.60. Old pit
bing. J. H. Dickson, 1985, GL.
+559/2. CREPIS VESICARIA L. subsp. TARAXACIFOLIA (Thuill.) Thell. 77, Lanarks.: Linthouse,
Glasgow, GR 26/54.66. Rough grassland. P. Macpherson, 1985, herb. P.M. 2nd record.
559/8. CREPIS PALUDOSA (L.) Moench 44, Carms.: Valley of Afon Merchon, GR 22/73.41.
Rock outcrops. P. Day, 1980. 2nd record, Ist since c. 1840.
+570/2. ELODEA CALLITRICHOIDES (Rich.) Casp. *13, W. Sussex: Chichester, GR 41/86.05.
Canal basin. M. Fowler, 1979. Hunston, GR 41/86.02. Disused canal. M. Briggs, 1980. Both
LANC. Ist and 2nd records, det. D. A. Simpson.
+571/1. LAGAROSIPHON MAJOR (Ridl.) Moss *69, Westmorland: Bankhead Pond, Helsington,
GR 34/49.89. K. Raistrick, 1984, LANC, det. D. A. Simpson.
576/1. ZOSTERA MARINA L. *29, Cambs.: River at Wisbech, GR 53/4.1. R. S. Adamson, 1908,
BM, det. T. G. Tutin.
577/3. POTAMOGETON COLORATUS Hornem. *11, S. Hants.: The Moors, Bishop’s Waltham,
GR 41/56.16. Stream. F. Rose, 1984, herb. A. Brewis. 68, Cheviot: Dunstanburgh, GR 46/
25.22. Pond on golf links. G. A. & M. Swan, 1984, herb. G.A.S., conf. N. T. H. Holmes. 1st
record since 1847 record from Dunstanburgh.
577/16. POTAMOGETON TRICHOIDES Cham. & Schlecht. *11, S. Hants.: Mans Bridge,
Southampton, GR 41/44.15. Old reservoir. R. P. Bowman, 1975, BM, det. J. E. Dandy.
577/20. POTAMOGETON FILIFORMIS Pers. 93, N. Aberdeen: Cotehill Loch, GR 48/02.29. D.
Welch, 1985, conf. N. T. H. Holmes. 1st post-1930 record.
580/1. ZANNICHELLIA PALUSTRIS L. 43, Rads.: Llynheilyn Pool, Llanfihangel-nant-Melan,
GR 32/16.58. I. Soane & A. C. Powell, 1985, NMW. 2nd record.
+589/3 X 2. POLYGONATUM MULTIFLORUM (L.) All. X P. ODORATUM (Mill.) Druce KE
Lanarks.: Between Glasgow and Renfrew, GR 26/51.67. Scrubby woodland. P. Macpherson & E.
Teasdale, 1984, herb. P.M. *99, Dunbarton: West Helensburgh, GR 26/29.83. Railway
embankment. R. R. Mill, 1967.
600/+2 * 1. HYACINTHOIDES HISPANICA (Mill.) Rothm. x H. Non-scripTA (L.) Chouard ex
Rothm. *77, Lanarks.: Meikle Dripps near Glasgow, GR _ 26/57.55. River bank. P.
Macpherson, 1984, herb. P.M., conf. C. A. Stace.
+605/2. JUNCUS TENUIS Willd. *80, Roxburghs.: Old station yard, Newcastleton, GR 35/
48.87. Damp track bed. R. W. M. Corner, 1985, herb. R.W.M.C.
605/amb. JUNCUS AMBIGUUS Guss. *61, S.E. Yorks.: Spurn, GR 54/42.15. Near Kilnsea
Beacon Lane Pond, GR 54/41.18. Both F. E. Crackles & M. Nicholls, 1985, herb. F.E.C. 1st and
2nd records.
605/fol. JuNcuUS FoLIOSUS Desf. *44, Carms.: Cilyblaidd, Pencarreg, GR 22/54.45. Wet
pasture. BSBI meeting, 1985, NMW, det. R. G. Ellis.
605/15. JUNCUS ACUTUS L. 11,S. Hants.: Sandy Point, Hayling Island, GR 40/7.9. Saltmarsh.
P. H. Colebourn, 1983. Only extant record.
PLANT RECORDS 195
605/18 x 19. JUNCUS ACUTIFLORUS Ehrh. ex Hoffm. * J. ARTICULATUS L. *14, E. Sussex:
Chailey Common, GR 51/38.21. Boggy hollow. London N.H.S., 1984, det. B. R. Radcliffe.
606/2 x 1. LUZULA ForSTERI (Sm.) DC. x L. piLosa (L.) Willd. *4, N. Devon: Withleigh,
Tiverton, GR 21/91.11. Woodland bank. L. J. Margetts, 1985, herb. L.J.M.
606/3. LUZULA SYLVATICA (Huds.) Gaudin *25, E. Suffolk: Reydon, GR 62/47.78. Ancient
woodland. P. G. Lawson, 1984, herb. E.M. & M. Hyde, conf. F. W. Simpson.
+606/4. LUZULA LUZULOIDES (Lam.) Dandy & Wilmott *47, Monts.: Powis Castle, GR 33/
21.06. Grassy bank in wood. M. Wainwright, 1984, NMW.
607/1. ALLIUM AMPELOPRASUM L. *52, Anglesey: Near South Stack, GR 23/2.8. Field border.
D. B. Hornby, 1975, conf. P. Day & R. H. Roberts.
607/3. ALLIUM SCORODOPRASUM L. *78, Peebless.: Near Kirklawhill, GR 36/08.37. Roadside.
D. J. McCosh, 1985, E, conf. D. R. McKean.
607/6. ALLIUM OLERACEUM L. 5. S..\Somerset:,:Cleeve Hill,’ GR. .31/05.42. Limestone
grassland. R. G. B. Roe, 1985. Ist record since 1924.
+607/nig. ALLIUM NIGRUM L. *29, Cambs.: Milton gravel pits, GR 52/47.61. Waste ground.
G. M. S. Easy, 1985, herb. G.M.S.E.
611/1. LEUCOJUM VERNUM L. +*29, Cambs.: Sheep’s Green, Cambridge, GR 52/44.57. Damp
ground near brook. P. F. Yeo, 1985.
+614/4. NARCISSUS X INCOMPARABILIS Mill. *77, Lanarks.: Between Glasgow and Renfrew,
GR 26/51.67. Bank. E. Teasdale & P. Macpherson, 1984, herb. P.M., det. D. McClintock.
+614/ < bar. NARCISSUS X BARRII Baker *77, Lanarks.: Between Glasgow and Renfrew, GR
26/51.67. Bank. E. Teasdale & P. Macpherson, 1984, herb. P.M., det. D. McClintock.
+616/2. IRIS VERSICOLOR L. *73, Kirkcudbrights.: New Galloway, GR 25/63.77. Stream under
willows. D. McClintock, 1943. Rascarrel Bay, GR 25/80.45. Bog by shore. O. M. Stewart & M.
McC. Webster, c. 1970. Ist and 2nd records.
616/3. IRIS FOETIDISSIMA L. +*99, Dunbarton: Cove, Loch Long, GR 26/22.82. Shore bank.
A. McG. Stirling & A. Rutherford, 1984.
+618/spe. Crocus sSPECcIOsUS Bieb. *17, Surrey: Chiddingfold, GR 41/96.35. Grassy
churchyard. A. C. & J. F. Leslie & E. J. Clement, 1985, herb. A.C.L. Abundantly naturalized.
633/1. CORALLORHIZA TRIFIDA Chatel. *76, Renfrews.: Kilmaculm Moss, GR 26/36.69. Fen
under trees. R. Kennedy, 1985, herb. E.R.T. Conacher.
638/2. PLATANTHERA BIFOLIA (L.) Rich. *111, Orkney: Sandwick, GR 57/21.14. Short cliff-
top turf. C. & J. Booth, 1985, conf. J. J. Wood.
642/7 x 5. ORCHIS MASCULA (L.) L. X O. Moro L. *69, Westmorland: Hazelslack, Arnside,
GR 34/4.7. Limestone pasture. M.T.Y. Foley, 1985, det. J. J. Wood.
643/1 x 5. DaAcryLorHizaA FUCHsII (Druce) So6 X D. PURPURELLA (T. & T. A. Stephenson)
So6 *77, Lanarks.: Shiels, Glasgow, GR 26/52.65. Damp scrub. E. Teasdale & P.
Macpherson, 1985, herb. P.M.
643/2b x 5. DACTYLORHIZA MACULATA (L.) So6 subsp. ERICETORUM (E. F. Linton) P. F. Hunt &
Summerhayes X D. PURPURELLA (T. & T. A. Stephenson) So6 *77, Lanarks.: Near Coulter,
GR 36/0.3. D. J..McCosh; 1967.
643/3. DACTYLORHIZA INCARNATA (L.) Sood *93, N. Aberdeen: Corsemaul, GR 38/39.40.
Flush. D. Welch, 1984, ABD.
643/6. DACTYLORHIZA MAJALIS (Reichenb.) P. F. Hunt & Summerhayes subsp. OCCIDENTALIS
(Pugsl.) Sell *109, Caithness: Near Thurso Castle, GR 39/12.68. Neutral grassland. J. K.
Butler, 1983, E, det. R. H. Roberts.
196 PLANT RECORDS
+646/1. AcoRUS CALAMUS L. *46, Cards.: S.W. of Hendy, Highmead, GR 22/49.42.
Overgrown pond in copse. “A. O. Chater, 1985.
+649/2 ita. ARUM ITALICUM Mill. subsp. ITALICUM 73, Kirkcudbrights.: Munches, GR 25/
83.58. O. M. Stewart, 1985. 2nd record. *99, Dunbarton: Dunglass Castle, Bowling, GR 26/
43.73. Rocky slope. A. McG. Stirling, 1972. Near Knockderry, Cove, GR 26/21.83. Reynoutria
thicket. A. McG. Stirling & A. Rutherford, 1983, E. 1st and 2nd records.
665/9. SCHOENOPLECTUS TABERNAEMONTANI (C. C. Gmel.) Palla 98, Main Argyll: Bagh Ban,
Craignish, GR 17/75.00. Seaward edge of swamp. B. H. Thompson, 1984, herb. B.H.T.
656/2. ELEOCHARIS ACICULARIS (L.) Roem. & Schult. 76, Renfrews.: Loch Libo, GR 26/
43.55. Sandy loch bay. A. J. Silverside, 1979. First post-1930 record.
656/6. ELEOCHARIS UNIGLUMIS (Link) Schult. *5, S. Somerset: West Sedge Moor, GR 31/
35.26. By rhine. R. S. Cropper, 1984, herb. R.G.B. Roe.
658/1. CYPERUS LONGUS L. 49, Caerns.: Criccieth, GR 23/50.37. Wet unstable cliff. L. J.
Larson, 1985, NMW. 2nd record.
663/2. CAREX DISTANS L. *99, Dunbarton: Mambeg, Gare Loch, GR 26/23.89. Rocks on
shore. A. McG. Stirling, 1985, E.
663/8 * 4. CAREX DEMISSA Hornem. X C. HOSTIANA DC. *47, Monts.: E. of Pont Crugnant,
GR 23/89.95. Boggy rill. P. M. Benoit et al., 1985.
663/10. CAREX SEROTINA Mérat 25, E. Suffolk: Lound, GR 63/50.00. Edge of ponds. F. W.
Simpson, 1984. 2nd extant record.
663/11. CAREX EXTENSA Gooden. *76, Renfrews.: Cardwell, GR 26/20.74. Saltmarsh turf. A.
McG. Stirling, 1979.
663/12. CAREX SYLVATICA Huds. 93, N. Aberdeen: Meet Hill, Peterhead, GR 48/11.44.
Wood. D. Welch, 1984, ABD. 2nd extant record.
663/20. CAREX RIPARIA Curt. 69, Westmorland: N.W. of Kirkby Stephen, GR 35/74.13.
Rough pasture. K. Raistrick, 1985, LANC, conf. A. O. Chater. Ist post-1930 record.
663/28. CAREX LIMOSA L. 93, N. Aberdeen: Strathbogie, GR 38/47.23. Flushed mire. R. E.
C. Ferreira & J. G. Roger, 1983. 1st post-1930 record.
663/35. CAREX ERICETORUM Poll. *32, Northants.: Barnack Hills and Holes, GR 53/07.04.
Limestone grassland. P. S. Lusby, 1978.
663/47. CAREX ACUTAL. ~° 50, Denbs.: R. Clwyd near Denbigh, GR 33/08.65. Edge of ditch. J.
A. Green, 1985, NMW, det. A. O. Chater. 2nd record.
663/67. CAREX SPICATA Huds. *46, Cards.: Cors Caron, E. of Cruglas, GR 22/70.65. Verge of
disused railway. A. O. Chater, 1985, NMW, det. A. O. C. & R. W. David.
663/72 X 73. CAREX CURTA Gooden. X C. LACHENALII Schkuhr 92, S. Aberdeen: Cairn Toul,
GR 27/95.97. D. J. Tennant, 1975, BM, conf. A. O. Chater & R. W. David. 2nd v.c. and British
record.
663/73. CAREX LACHENALII Schkuhr 92, S. Aberdeen: Record published in Watsonia 15: 404
is not only extant record.
+ARUNDINARIA JANSAURENSIS Gamble *46, Cards.: S.W. of Highmead mansion, GR 22/
49.43. Damp woodland by pond. A. O. Chater, 1985, NMW, det. D. McClintock. Ist Welsh
record.
670/1 X +671/2. FESTUCA PRATENSIS Huds. X LOLIUM MULTIFLORUM Lam. *17, Surrey:
Between Old Woking Sewage Works and Newark, GR 51/03.57. By path in meadow. R. & N.
Sherlock, 1985, herb. A. C. Leslie, conf. A.C.L. *61, S.E. Yorks.: N.E. of Firby, GR 44/
75.66. E. Chicken, 1985, det. T. A. Cope.
PLANT RECORDS 197
670/3 X 671/1. FESTUCA GIGANTEA (L.) Vill. x LOLIUM PERENNE L. *29, Cambs.: Balsham,
GR 52/59.51. Meadow. P. J. O. Trist, 1974, herb. P.J.O.T.
670/5. FESTUCA HETEROPHYLLA Lam. *69, Westmorland: Appleby Castle, GR 35/68.19.
Woodland above R. Eden. R. E. Groom, 1985, LANC.
670/6. FESTUCA RUBRA L. subsp. MEGASTACHYS Gaudin *76, Renfrews.: Williamwood, GR
26/56.58. P. Macpherson, 1978, herb. P. M., det. C. E. Hubbard.
672/3. VULPIA MyuROS (L.) C. C. Gmel. *77, Lanarks.: King George V dock, Glasgow, GR
26/53.66. P. Macpherson & E. L. S. Lindsay, 1985, herb. P. M. Bunhouse, Glasgow, GR 26/56.66.
Old bridge. J. H. Dickson, 1985, GL. Ist and 2nd records.
673/1 xX 5. PUCCINELLIA MARITIMA (Huds.) Parl. xX P. RUpPESTRIS (With.) Fernald &
Weatherby *11, S. Hants.: R. Test, Nursling, GR 41/36.14. Brackish pasture. R. P. Bowman,
1977, herb. R.P.B., det. A. Melderis.
673/2. PUCCINELLIA DISTANS (L.) Parl. *32, Northants.: Wansford, GR 52/07.99. Roadside
verge. N. E- Scott, 1980.
676/11. POA ANGUSTIFOLIA L. *76, Renfrews.: Dunrod near Inverkip, GR 26/22.72. Near
Quarriers Homes, GR 26/37.66. Both Railway. I.T.E. Railway Survey, 1980. Ist and 2nd records.
676/12. POA SUBCAERULEA Sm. 76, Renfrews.: Barrangary near Bishopton, GR 26/44.69.
Railway. I.T.E. Railway Survey, 1980. 2nd record.
+676/15. Poa cHAIxI Vill. *93, N. Aberdeen: House of Daviot, GR 38/74.27. Policy
woodland. D. Welch, 1984, ABD.
683/13. BRoMuUS LEPIDUS Holmberg 77, Lanarks.: Cadder, Glasgow, GR 26/61.72. Riverside
path. B.S.B.I./Glasgow N.H.S. Field Meeting, 1982, herb. P. Macpherson, conf. P.J.O. Trist. 2nd
record.
683/10 x 13. BROMUS HORDEACEUS L. X B. LEPIDUS Holmberg *77, Lanarks.: East Kilbride,
GR 26/66.55. Edge of meadow. P. Macpherson & E. L. S. Lindsay, 1983, herb. P. M., det. P. J. O.
Trist.
+683/jap. BROMUS JAPONICUS Thunb. *70, Cumberland: Silloth, GR 35/1.5. E. J. Glaister,
1877, CLE, det. P. M. Smith.
685/3 xX 5. ELYMUS FARCTUS (Viv.) Runemark ex Melderis X E. REPENS (L.) Gould *69,
Westmorland: Newbiggin, GR 34/26.69. Sandy shore. *70, Cumberland: Beckfoot, Silloth,
GR 35/09.49. Sandy gravel. Both G. Halliday, 1978, LANC, det. A. Melderis.
685/5. ELYMUS PYCNANTHUS (Godron) Melderis *67, S. Northumb.: N. Blyth, GR 45/31.82.
Sand dune. G. A. Swan, 1978, herb. G.A.S., conf. T. G. Tutin.
+687/jub. HORDEUM JUBATUM L. *77, Lanarks.: Provanmill, Glasgow, GR 26/62.66.
Roadside. J. H. Dickson, 1982, GL.
4692/2. AVENA LUDOVICIANA Durieu *76, Renfrews.: Near Inchinnan, GR 26/4.6. J. H.
Penson, 1970, herb. J.H.P.
700/1. CALAMAGROSTIS EPIGEJOS (L.) Roth 46, Cards.: W.S.W. of Traeth y Mwnt, GR 22/
19.51. Grassy sea cliff. A. O. Chater, 1985, NMW. 2nd record. *98, Main Argyll: S. of Bagh
Ban, Craignish, GR 17/75.00. Bushy cliff. B. H. Thompson, 1984.
1701/7. AGROSTIS SCABRA Willd. *76, Renfrews.: Govan Docks, GR 26/57.64. Old railway
yards. A. J. Silverside & C. Tavendale, 1977, E. Well established.
707/1. PHLEUM BERTOLONID DC. *99,| Dunbarton: Near Cardross Station, GR 26/34.77.
Sandy ground near shore. A. McG. Stirling & A. Rutherford, 1985, E.
708/4. ALOPECURUS AEQUALIS Sobol. *5, S. Somerset: Scott’s Nurseries, Merriott, GR 31/
44.12. Damp ground. R. G. B. Roe, 1985, herb. R.G.B.R.
198 PLANT RECORDS
708/5 x 3. ALOPECURUS BULBOSUS Gouan X A. GENICULATUS L. *11, S. Hants.: R. Test,
Nursling, GR 41/3.1. Brackish pasture. R. P. Bowman, 1980, K, det. T. A. Cope. 1st British
record.
709/1. MILIUM EFFUSUM L. 93, N. Aberdeen: Blairfowl, GR 38/80.38. Wet woodland. D.
Welch, 1984, ABD. 2nd record.
+713/4. PHALARIS PARADOXA L. var. PRAEMORSA Coss. & Dur. *26, W. Suffolk: Hawkedon,
GR 52/79.51. Wheat field. Boxted, GR 52/80.50. Barley field. Both M. A. Hyde, 1985, herb. E. M.
& M. Hyde. Ist and 2nd records, former conf. E. J. Clement.
714/2. PARAPHOLIS INCURVA (L.) C. E. Hubbard *49, Caerns.: Porth Ysgaden, GR 23/21.37.
Eroding drift above beach. J. R. Akeroyd, 1985, RNG, conf. C. A. Stace.
Watsonia, 16, 199-209 (1986) 199
Book Reviews
Bee Orchids (3). Stephen Blackmore. Pp. 24, with 12 colour & 8 black & white illustrations. 1985.
Price £1-25 (ISBN 0-85263-745-4). Buttercups (6). Stephen Blackmore. Pp. 24, with 13 colour and
7 black & white text illustrations. 1985. Price £1-25 (ISBN 0-85263-763-2). Willows of the British
Isles (8). Theresa Brendell. Pp. 24, with 16 colour and 14 black & white text illustrations. 1985.
Price £1-25 (ISBN 0-85263-765-9). Shire Natural History Series. Shire Publications Ltd,
Aylesbury.
These AS-sized, glossy paperbacks are, it is hoped, the first in a long line of botanical ‘mini-
monographs’ of groups of plants found in the British Isles. The publishers define the aim of this
new series, which also covers zoological topics, as “‘to fill the gap between brief guides of general
interest and full-length books for the specialist . . . for the interested layman or student”. They
combine informative accounts of the biology of each group with an identification guide.
Bee Orchids covers the four Ophrys species found in this country. Few ordinary folk have ever
seen a wild orchid in this country, but they will all have heard of the Bee Orchid. This little guide
with its intimate details of life history, including the latest data on the nature of the association with
mycorrhizal fungi and the bizarre pollination mechanism of the Bee Orchids — which actually emit
pheromones to attract their male insect pollinators, as well as visually resembling their mates — is
sure to enthral both layman and dedicated botanist alike. The side-by-side sketches and colour
photographs should ensure accurate identification of the various Ophrys species and their varieties.
Why no colour plate of the Late Spider Orchid (O. fuciflora), which some people confuse (perhaps
fortunately) with the Bee?
Buttercups deals with our members of the genus Ranunculus. The yellow-flowered species are
dealt with in some detail, the white-flowered ones more cursorily. The chatty comparative
narrative of these guides with accurate side-by-side line drawings and character tables would have
been the ideal mode for sorting out the difficult water crowfoots. To claim that infertility in R.
ficaria subsp. bulbifera is due to tetraploidy per se is misleading, and obscures a fascinating if as yet
speculative story. These are topics, perhaps, for an expanded second edition, and their omission is
no reason to decry a splendid account of our buttercups — a must for all school biology libraries.
Willows compliments nicely the B.S.B.I.’s own Willows & Poplars and gently introduces both
layman and perplexed botanist alike to our willows. It has an excellent character-matrix key for
identification and colour photographs of whole-tree and branch closeups. Side-by-side leaf outlines
of all the species and colour shots of more catkins would be a welcome inclusion for a later edition,
as the willows are perhaps more likely to appeal to the discerning layman.
These and future botanical booklets in the series should be in all 6th form libraries. They form
the ideal medium for introducing the recreational pleasure that field botany can bring to young
receptive minds.
K. J. ADAMS
Cambridge and Clare. Sir Harry Godwin. Pp. xxi + 230, with 43 black & white illustrations.
Cambridge University Press, Cambridge. 1985. Price £19-50 (ISBN 0—-521-30765-1).
In the last few years there has been a sudden rush of books on Cambridge botany and botanists,
greatly enriching our knowledge of the more recent periods in particular. First came the collective
tribute to Humphrey Gilbert-Carter by some of his past pupils and friends, then the similar volume
in memory of the late John Raven, then the history of the Botanic Garden by its then Director,
Max Walters. Now the late Sir Harry Godwin has given us his memoirs of his sixty-odd years in
residence — to the best of my knowledge the first former occupant of a British botanical chair to
have published an autobiography, at any rate of book length, since F. O. Bower in 1938.
200 BOOK REVIEWS
As the title indicates, the book is addressed to two overlapping readerships: to past and present
members of Clare College (of which Sir Harry was successively an undergraduate, research student
and Fellow) and to those interested in the University’s history in general and in its botanical teaching
in particular. This dual aim was perhaps over-ambitious, for satisfying the one lot of readers must
inevitably have meant disappointing in some degree the other, in view of the fact that (doubtless by
decree of the publisher) the book is a comparatively short one. As it is, much of the matter can hardly
help but be prohibitively parochial to anyone not of the College, who would surely have preferred
much more on the author’s botanical career and to have had the benefit of his reflections on how the
discipline has developed.
Nevertheless what we are given on botany is full of interest, even though it touches on the concerns
of this Society only tangentially. For while Sir Harry was always a field man, it was ecology and
Quaternary studies, never taxonomy and only indirectly floristics, that held his allegiance. Even so,
B.S.B.I. members predictably crop up here and there in his pages. Our first President, John
Gilmour, was, for example, his first pupil when he embarked on College ‘supervisions’ (Cambridge
for tutorials), and the Demonstrator for his early first-year practical classes turns out to have been
none other than Dr Cyril West. Another with whom the author literally rubbed shoulders — for he
was one of the outsider M.A.s with dining rights in Clare — was the author of that pre-war Flora of
Cambridgeshire and specialist in Arctium, A. H. Evans. Evans, we learn, was crippled by locomotor
ataxia and in his later years depended on undergraduates to do the collecting that nourished his final
passion, Rubus. 2
As one of the more towering of his early colleagues and co-nurse of the infant British Ecological
Society, Sir Arthur Tansley expectedly features prominently — indeed has a whole chapter to himself.
Tribute is paid to him, inter alia, as a superlative editor and as the possessor of “‘remarkable powers
of timely innovation.”” The two were additionally brought together by a shared interest in
psychoanalysis. Tansley’s fascination with that newly-emergent field of knowledge went so far
indeed that in 1923, taking advantage of his private means, he resigned his University post and went
to live with his family in Vienna in order to study under Freud. Tansley, the founder of the New
Phytologist, was subsequently the author of The New Psychology, a minor bestseller in its day.
Fortunately for botany, however, that proved only a passing phase and a mere four years later he was
back in academic harness, summoned by Oxford to the Sherardian Chair. On arriving at Oxford he
was soon quarrelling bitterly with G. C. Druce, for reasons that still remain obscure (Sir Harry,
regrettably, is silent on the matter), and as a result was permanently alienated from ‘Druce’s society’,
the B.S-B.I.’s ancestor, the B.E.C.
Much the most intriguing revelation in the book is the extraordinary paranoia with which one of
Sir Harry’s predecessors in the Cambridge Chair, F. T. Brooks, was afflicted. So violent was the
animosity Brooks felt towards his Professor, Sir Arthur Seward, that he once went so far as to
denounce him to the police as a German spy. For years it was quietly accepted among their
colleagues that on no account could the two be left alone in a room together. Yet despite his
notorious instability Brooks was appointed Seward’s successor and went on to head the department
for the next twelve years. It is just such quirks of personality, such chance conjunctures of
individuals, that account for so many of the otherwise inexplicable twists and turns of history, and it
is to autobiographers above all that we look to reveal them.
Sir Harry’s life was clearly an enviably happy and productive one. From comparatively humble
origins he emerged, under the wing of the customary influential schoolteacher, to pursue the academic
career as an ecologist that he had resolved upon even ahead of his arrival at university. It would surely
have been a successful career whatever direction it had taken. All students of floristics, however, must
ever be grateful that, eight years into teaching and research, he was guided by Tansley (or rather,
initially, his wife was) into the then-virgin field in Britain of pollen analysis. The rest is history—or, to be
specific, The History of the British Flora. He needs no other monument than that.
D. E. ALLEN
The Victorians and their flowers. Nicolette Scourse. Pp. 195, with 17 colour plates, 102 black &
white text illustrations and | table. Croom Helm, London and Canberra. 1983. Price £12-95 (ISBN
0-7099— 2377-5).
BOOK REVIEWS 201
Anyone who has had the mischance to borrow, buy or be given Lynn Barber’s The Heyday of
Natural History will find this book a refreshing antidote. Both authors have similarly fallen under
the spell of those popular illustrated works that survive in such numbers from the Victorian era and
that seem so peculiarly redolent of the ideas and attitudes of the age; but where the one felt obliged
to scoff and to jeer the other approaches her subject with evident admiration and sympathy. The
product of wide reading and with numerous well-chosen illustrations, this attractive new
contribution to what might be termed ‘Saturday Book social history’ will deservedly be a much-
favoured choice for presents.
Like so many members of that genre, though, it is essentially a confection, made out of material
that has been forced into an unreal shape. For it is founded on a fallacy: that there exists a single
entity, the admiration-cum-study of flowers, of which field botany and horticulture merely form
scarcely distinguishable facets. In reality, of course, the two are largely discrete and, even though
they spring from the same tap-root, have long developed independently of one another. The
garden wall is a very solid dividing-line and it is idle to pretend otherwise. By disregarding its
existence, the author has produced a book which, inasmuch as it keeps almost entirely to the
horticultural side of the wall, is strictly speaking of very limited relevance to the field botanist
reader. The main exception is Chapter Five, in which extensive recourse is had to the journals and
superb herbarium (sample sheets from which are reproduced as illustrations) of Robert Dick, the
poverty-stricken Thurso baker who first detected Hierochloé borealis in Britain.
But perhaps it was wise for the author to penetrate into our territory no further than this, for
once at the limits of her reading her touch noticeably falters. On page 139, for example, we are told
that ‘‘museums only came into being after the Great Exhibition in 1851 — the Natural History
Museum in London only being opened in 1881 — and natural history was not taught in schools until
the 1880s.” There are three misleading statements there in that single sentence alone. What a pity
the text was not submitted to the scrutiny of a specialist. What a pity, too, that a book that is
otherwise so pleasing visually should have had the captions to two of the figures (5.6 and 5.16)
transposed so obtrusively.
D. E. ALLEN
The names of plants. D. Gledhill. Pp. viii + 159. Cambridge University Press, Cambridge. 1985.
Prices: paperback £6-95 (ISBN 0—521—31562-X); hard covers £20-00 (ISBN 0-521-30549-7).
This small book, not unreasonably priced as a paperback, is in two parts. The first 48 pages are a
history of the naming of plants, concentrating on the Latin names and the development of the rules
_ of nomenclature. The account of nomenclature up to Linnaeus is an interesting summary, but the
exposition of the current International Code of Botanical Nomenclature is very confusing and
unlikely to enlighten readers. Although, for example, Wardian cases and chromosomes are
described in some detail, illegitimate names are mentioned only in a misleading aside. Errors
abound, from Linnaeus’s Systema Naturalis (vice Naturae) to a statement that the name
Papilionaceae refers to the flowers being adapted to pollination by butterflies. Most of the
references in the text, e.g. Grew (1672) and Sutton (1902), are not in the Bibliography, and one of
the few that seems to be, Farr (1980), is given there as Farr (1979).
The rest of the book is a sensibly constructed glossary, mostly of generic and specific names, and
of elements used to make up such names; thus a large number of names can be interpreted by
looking up their constituent parts. Its potential usefulness to B.S.B.I. members is indicated by the
fact that 90% of the first 300 names of angiosperms in A. R. Clapham, T. G. Tutin & E. F.
Warburg, Excursion Flora of the British Isles, 3rd ed. (1981) are covered by it, more than in any
Other currently available glossary. This is in spite of the statement by the author that he has in
general omitted commemorative and geographical epithets and anagrams. But he also says “I
make no claim that the meanings which I have listed are always the only meanings which have been
put upon the various entries’’, and this leads to one of the chief drawbacks of the book. He is so
partial in some of the meanings that the results are very misleading, e.g. ‘‘fluitans floating on
water’, “natans floating under water’; “‘aquaticus growing in water’, “‘aquatilis growing under
water”. These antitheses are entirely spurious. In other cases, e.g. ‘“‘runcinatus saw-toothed,
202 BOOK REVIEWS
sharply cut’’, the essential definition, in this instance that the teeth or lobes point towards the base of
the organ, is missing. Some words are quite wrongly translated, quadratus for example does not
mean “‘into four, in fours” but square or rectangular. It is startling to see under the Cambridge
imprint “‘saepius-a-um of hedges”’; saepius of course means more often, and only saepium or sepium
is the genitive plural of saepes or sepes, a hedge.
With thorough editing this book could have been useful, but as it is it cannot be recommended. For
an exposition of nomenclature, C. Jeffrey, Biological nomenclature (1973), is a model of clarity and
helpfulness, but for a glossary there seems to be nothing entirely satisfactory currently in print.
A. O. CHATER
The English landscape past, presentand future. Edited by S. R. J. Woodell. Pp. x + 240, with 43 black
& white photos and 56 text figures. Oxford University Press, Oxford. 1985. Price £15-00 (ISBN
0-19-211621-S5).
This book is the text of the 1983 Wolfson College Lectures and consists essentially of brilliantly
written review chapters on many of the factors that shape the English landscape. It expounds recent
ideas that may in many cases be unfamiliar to the general botanical reader and forms probably the
best up-to-date introduction to the subject. The chapter by O. Rackham on ancient woodland and
hedges, lucidly summarizing his own and others’ work on these features, will be of particular value to
botanists. He begins by deploring that it is still necessary to refute the myth that our present hedged
and walled landscape is the result of eighteenth and nineteenth century agriculture changes, and
ends by indicating that this landscape is vastly older than most of us realize, the Romans and their
predecessors having reduced well over half of England to farmland and moorland, so that the
Anglo-Saxons probably took over an at least partly hedged landscape. B. W. Cunliffe, in the chapter
on man and landscape 6000 BC-AD 400, even refers to the British landscape at the end of the
Roman period as “‘over-used”’. The chapter on the geomorphological background by D. K. C. Jones
contains a Landsat image of the area from Bournemouth to Builth Wells under snow, showing the
underlying structural features with astonishing clarity, and this alone is almost worth the price of the
book. (Some of the other illustrations, notably photographs of Bamford Edge and of a South Downs
dry valley, are of quite deplorable quality.) Joan Thirsk, in an account of fads and fashions in the
agricultural landscape, tells us that rapeseed, which many consider today a garish and unwelcome
novelty, was warmly welcomed and widely cultivated here in the sixteenth century and continued
into the early nineteenth century. Among other crops whose history she recounts are various dye
plants. A madder (Rubia tinctoria) plantation at Deptford in 1660 was said to employ 1000 people,
and several hundred acres of safflower (Carthamus tintorius) were growing in Oxfordshire in 1673.
The history of our climate and its effect on the landscape 1s described by H. H. Lamb witha mass of
illustrative detail, and when we learn, for example, that in the late seventeenth century the growing
season at Oxford was probably 3-5 weeks shorter than in the warmest decades of the present
century, we can understand why he warns us to understand well what we attempt to conserve
“because conservation is often not nature’s way’. A tentatively optimistic final chapter by M. E. D.
Poore on the present state of agriculture, forestry and conservation, with some emphasis on the
effects of E.E.C. policy, is somewhat countered by the epilogue. In this the editor suggests that
although earlier chapters have shown that our landscape has undergone more changes, and reflects
much more of the past, than we previously realized, today’s changes are so drastic that the future,
unlike the past, probably really does have in store loss and impoverishment rather than evolution
and enrichment.
A. O. CHATER
Riches of the rain forest. An introduction to the trees and fruits of the Indonesian and Malaysian rain
forest. W. Veevers-Carter. Pp. 103, with c. 22 line drawings and 12 colour plates. Oxford University
Press, Singapore. 1985. Price £13-50 (ISBN 0—-19-582576-4).
BOOK REVIEWS 203
The author, dedicating this slim book to the memory of the Dutch conservationist, Marius Jacobs,
has attempted to illustrate the richness of the Malesian rain forests not in terms of diversity alone
but also in relation to the economic products obtained from the wild. She has used the examples to
develop a passionate plea for conservation. There is a foreword by Md. Khir Johari, President of
W.W.F. Malaysia stressing the timely appearance of the book and its value as non-academic
publicity for the cause of conservation. Fifteen short chapters or essays deal with particular plant
groups, their natural history and their economic importance: dipterocarps, figs, rattans, ironwood
(Eusideroxylon spp), Eugenia spp, nutmegs, rambutans and relatives, durians, Artocarpus spp,
Agathis, Rafflesia, climbers, orchids and Pinus. There is an epilogue commenting on forest
dwellers, and a glossary. There are twelve slightly ‘primitive’ colour plates and several line
drawings prepared by the Indonesian artist, Md. Anwar.
There is much to praise in this book. It is clearly written and focusses attention directly on some
of the most important problems in the exploitation of Malesian rain forest. The choice of examples
is good but there are some unfortunate generalizations and inaccuracies, the most glaring of which
is the statement that curare comes from a species of Menispermaceae rather than Strychnos in
Loganiaceae. The account of the Dipterocarps is spoiled by the oversimplified statement
concerning stocks of these trees. Anyone in Peninsular Malaysia can still see many dipterocarps
within 30 km of the centre of Kuala Lumpur, thus undermining the credibility of the author’s
message concerning over-exploitation.
For whom is the book intended? The author states it is written for the layman, but whether in
Europe or South East Asia it is not clear. The price is much too high if it was intended for an
Indonesian market. Beautiful though the production is, I feel the cause of conservation would have
been more compellingly served with a more dynamic book design, the use of photographs and, if
possible, a lower price.
J. DRANSFIELD
The wildlife of the Thames Counties, Berkshire, Buckinghamshire and Oxfordshire. Edited by
Richard Fitter. Robert Dugdale in association with Berkshire, Buckinghamshire and Oxfordshire
Naturalists’ Trust, Oxford. 1985. Price £4-95.
Eleven contributors under the skilful editorship of naturalist and wildlife author (and former
B.B.O.N.T. president) Richard Fitter makes this an authoritative book with something for
everybody interested in this part of the Thames Valley. Richard Fitter opens with a chapter
entitled ‘The Physical Background and the Influence of Man’, John M. Steane contributes ‘Land
Use History’ and S. R. J. Woodell ‘Vegetation’. H. J. M. Bowen’s chapter, ‘The Flora’, follows
with notes on interesting species and their localities. I found this chapter particularly gripping
reading, wanting to set off and visit the interesting plants and sites immediately! It includes
bryophytes, lichens, fungi and even algae. Pages 70-161 are then devoted to animals. After this
come smaller chapters on ‘Conservation Achievements and Problems’ by Richard Fitter (very
much a history of conservation bodies and B.B.O.N.T.), ‘A Note on Environmental Education’ by
Ursula Bowen, and finally ‘The Warburg Reserve, Bix Bottom: A Case History’ by Vera Paul.
This interesting guide mentions the highly successful public appeal for £23,000 to buy the reserve,
but of course, with the author’s modesty, omits any mention of the spectacular work done to
achieve this, work for which she was awarded the O.B.E. The book ends with an appendix on
museums with natural history collections, a gazetteer of principal wildlife sites and a bibliography
(divided for each chapter).
I would have liked lists of species found in the area covered, but this would perhaps have made
the book too long. H. J. M. Bowen’s Flora is out of print and not really readily available, as
mentioned, and I fear the Floras of Buckinghamshire and Oxfordshire are some years off
publication. Checklists would surely help the recording of the flora.
The book is very well illustrated, very readable and well-produced, and at £4-95 very good value
for over 200 pages!
Spb: Jury
204 BOOK REVIEWS
Essential oils and aromatic plants. A. Baerheim Svendsen & J. J. Scheffer. Pp. 246 with numerous
illustrations and diagrams. Martin Nijhoff, Dortrecht. 1985. Price £24-95 (ISBN 90-247-3195-X).
This book represents the Proceedings of the 15th International Symposium on Essential Oils, held
in July 1984 in the Netherlands. Like a previous volume reviewed by me (Watsonia, 14: 440 (1983))
its contributions reflect the growing importance of essential oils and aromatic plant ingredients for
science, industry and, above all, medicine. Again the potential buyer must be warned that this is
not ‘a book’ but an assembly of loosely connected lecture-papers of a highly specialized nature (in
some cases perhaps the sort of paper written by people hastily in order to justify their attendance at
a conference!). Maybe only the last contribution of this compilation is of interest to a general
reader, namely an essay on medicinal plants mirrored in paintings from the Netherlands (in
German, nota bene).
E. LAUNERT
The correspondence of Charles Darwin, Vol. 1, 1821-1836. Edited by F. Burkhardt & S. Smith.
Pp. 702. Cambridge University Press, Cambridge. 1985. Price £30-00 (ISBN 0—521—25587-2).
These letters, written by and to Charles Darwin, encompass 15 years of his life, from the time when
he was a twelve-year-old schoolboy at Shrewsbury to the period immediately aftér his return from
the famous five-year voyage aboard Beagle. The relative amounts of correspondence in the various
phases of this period are reflected in the number of pages each phase occupies — Shrewsbury School
(18 pp.), medical student at Edinburgh University (22 pp.), undergraduate at Cambridge (136
pp.), naturalist aboard Beagle (318 pp.), traveller returned to England (31 pp.). Five appendices,
occupying 34 pages, provide a ‘Chronology’ from his Journal for the years 1809-1836, notes on his
Beagle records, lists of the ‘Persons on board the Beagle, 1831-6’, books available aboard Beagle (a
mouth-watering treasurehouse to the bibliophile!) and early notes on coral reefs. Sixteen pages
detail alterations to the manuscript letters, the bibliography occupies a further 18 pages, a
biographical register and index to correspondents is covered in 50 pages, while a comprehensive
index (39 pp.) completes the volume.
The bald outline of this book, just given, provides little indication of the wealth of its contents.
Nor can any review. Whilst reading it, I identified with the young schoolboy collecting rocks, plants
and animals, surely the experience of most readers of Watsonia. I shared the excitement of the
undergraduate trying to understand the diversity on this planet and the wonder of a young scientist
exposed to the manifold experience and observations provided during a long voyage to foreign
parts. Looking back over the years we see Darwin as a Great Man, but these letters reveal him as a
very human person as well. To be sure, there are numerous indications of his powers of
observation and spirit of scientific inquiry, mirrored by the sound advice and information offered
him by many of his correspondents. There is also, however, much in his letters of these 15 years
which reflects the understandable concerns of a young man away from his home and, later,
country. The Victorian formality of the letters cannot disguise Darwin’s easy relations with his
family and friends, and their support and affection for him.
This is a strikingly well edited book. The full and helpful footnotes, the clear documentation and
the useful appendices all underpin a most scholarly work. The printing is of high quality and largely
free from errors. The correspondence is a treasurehouse of the scientific views of Darwin and his
friends, of insights into the social scene in which he and they were involved, and of observations on
new lands seen through the fresh eyes of youth. Whilst enjoying it one is tempted to reflect on what
is being lost by current use of the telephone rather than the letter.
D. M. Moore
The Oxford illustrated encyclopedia. 2. The natural world. Edited by M. Coe. Pp. vi + 375, with
numerous colour and black & white photographs and diagrams. Oxford University Press, Oxford.
1985. Price £15-95 (ISBN 0-19-869134-3).
BOOK REVIEWS 205
The natural world forms the second part of an eight-volume encyclopaedia, spanning the arts, the
social, biological and pure sciences and claiming overall to give a map of contemporary knowledge.
It is clearly intended for the layman for it is arranged alphabetically under vernacular names, with
Latin or jargon terms kept to a minimum. The subjects are dealt with briefly, few meriting more than
three hundred words, but there is an extensive cross referencing. Thus although rats and mice both
have their own entry, both of these refer back to more general information under the heading of
rodents. Most pages are illustrated with a good proportion of attractive colour photographs.
Overall, however, the book gives the feel of having been assembled by people less interested in the
natural world for its own sake than as a background for man. Few of the entries on plants fail to
include details of their food, medicinal or pest value, sometimes to the exclusion of other more
purely botanical information, while the pest aspect of many animals is stressed to the detriment of
other possible detail. One might perhaps have expected this when looking at the plan for the eight
volumes, five of which are directly concerned with human evolution, present day societies and
achievements. In line with this we find a substantial number of entries are potted biographies of
scientists or medical men. Useful, perhaps, but more questionable in a book entitled The natural
world is to give space to ailments described from a purely human standpoint. Even if other animals
do not suffer from club foot, cleft palate or piles, these entries might have been justified with a note
to that effect. Acupuncture and homeopathy find places, veterinary medicine and vivisection do not.
Perhaps related to this anthropocentric approach there seems to be some carelessness in dealing
with other organisms. The front endpaper refers to reptiles as the first animals to breed on land, in
spite of previous remarks about the emergence of invertebrates from the seas. The back endpaper
shows an abbreviated scheme of plant, animal and protist classification that includes bacteria, but
not blue-green algae, with cryptogams. Opinions may differ on this, but surely both should be
together in whatever grouping is decided on. In the main body of the book occasional spelling
mistakes — Hysterimorpha for Hystricomorpha, for example — and misinterpretations of references
read mar some entries. The largest octopuses, for instance, have an arm span, not arm length, of up
to 5m. Altogether, in spite of the bright promise of the cover, preface and pictures, this is a
disappointing book.
J. PoPE
Hardy Geraniums. P. F. Yeo. Pp. 192 with 44 colour plates and 121 figures (line and silhouette).
Croom Helm Ltd, London and Canberra. 1985. Price £25-00 (ISBN 0-7099-2907-2).
In this attractive book Peter Yeo has succeeded in combining botanical scholarship with accessibility
to the uninitiated (though keen) gardener or naturalist. It is true,as he says, that it is ““mainly about
identification’’, but it also contains a clear exposition of the current state of his work on the
classification of the genus Geranium as a whole.
The title indicates that it is about ‘Cranesbills’, not tender ‘Pot Geraniums’ (Pelargonium), while
the contents show that the author has a very thorough familiarity with the genus in cultivation in all
its aspects. Not only has he grown the majority of species and hybrids hardy in the British Isles, but
he and Dr H. Kiefer have carried out numerous crossing experiments. The result is an authoritative
account of Geranium in which personal horticultural experience is evident on almost every page.
The introductory chapters, especially, include much fascinating data that will be of value to
botanists as well as gardeners. The chapters on ‘Structure and terminology’ and ‘Structure in relation
to function’ show in a masterly way the characteristics of Geranium and how the variation in each
part of the plant is associated with environmental variations (habitat, altitude, pollination, seed
dispersal, etc.). The chapter on nomenclature is worth reading by itself as a clear, untechnical
exposition of the essentials of the International Code of Botanical Nomenclature, illustrated with
examples from Geranium. In addition to a well constructed dichotomous key to species and hybrids,
there is a multi-access key that is easy to use and should allow one to identify specimens in which all
the characters necessary for using the main key are not present.
In Peter Yeo’s new classification of Geranium, the genus is divided into three subgenera,
depending on the method of discharge of the seed. By far the majority of the species fall into
subgenus Geranium section Geranium, one of three sections of that subgenus. The author explains
206 BOOK REVIEWS
that he has not subdivided this unwieldy section into formal categories because, as yet, he is unable to
define the groups that can be recognized. These groups have, however, been given informal names
and descriptions. Although only 95 species varieties and cultivars are described in detail (as well as 11
hybrids), these include nearly all the British ones. Only G. purpureum is omitted, and it is briefly
differentiated from G. robertianum.
The illustrations are both excellent and helpful. The author’s well known abilities with a camera
are demonstrated by the 44 beautiful colour plates, and Frances Hibberd’s detailed drawings are
excellent. The silhouette or outline of a leaf of each species and hybrid provides a very useful
discriminating feature.
In sum, this is a delightful book, taxonomically exact, that will be invaluable for anyone wishing to
obtain information about Geranium in general, as well as on its cultivation. It is not yet, however, the
complete monograph that is badly needed to replace that of Knuth (1912).
~
N. K. B. Rosson
Ouvrages botaniques anciens. Catalogue des ouvrages prélinnéens de la Bibliothéque des Conserva-
toire et Jardin botaniques de la Ville de Genéve. Dressé et publié sous la direction de Hervé M.
Burdet. Pp. xxviii + 599, with coloured frontispiece and numerous illustrations in the text.
Conservatoire Botanique de Genéve. 1985. Price DM 250 (ISBN 2-8277-0105-—7).
This lavish catalogue of the ‘pre Linnean’ books in the library of the Geneva Botanic Garden will
prove a delight to every lover of old botanical literature and the student of the history of botanical
illustration. It is a handsome, large quarto volume, strongly bound in gold tooled green cloth and
finely printed on laid paper as befits a major contribution to the literature of botanical bibliography.
The order of presentation is chronological, but a comprehensive author index enables easy
reference to a particular title. The library’s holdings of very early herbals is surprisingly poor, with
only one incunabula being noted and only 27 entries for works published prior to 1550. In all just over
600 titles are listed. The bibliographical entry for each item is on familiar lines, with a transcription of
the title page, note of the imprint, pagination and register of signatures. Some minor errors in the
title transcriptions have been noted; for instance entries number 213 and 473 both transcribe colons
which are not in fact present on the title page, and pagination details under entries 390 and 448
appear to be slightly awry. Whilst such minor imperfections may cause irritation when using the
catalogue to compare and collate another copy of a particular volume, they do not significantly
detract from a work which does contain some detailed collations not easily obtainable from other
printed sources. It is a pity however that the opportunity was not taken to give details of the
provenance of important individual items and to include details of any unusual features. For example
the coloured frontispiece is a reproduction of a page from what appears to be a contemporary
coloured copy of Brunfel’s herbal of 1539, yet no mention is made in the catalogue entry of what is
surely an important attribute of this particular copy.
A principle attraction of the catalogue is in the generous number of really fine illustrations; the
great majority of entries are accompanied by a facsimile reproduction of a significant text page, title
page or portrait of the author, and it is this feature that gives so much pleasure and enables one to
browse with continual enjoyment through what could have so easily been a ‘dry as dust’ production.
The choice of illustrations is excellent, mirroring the art of botanical illustration over a period of
some 250 years, with a leavening of maps, butterflies, mammals, etc. (plus the odd ‘indelicate’
woodcut) to break the monotony which is sometimes inherent in works of this kind. Either the
library is fortunate enough to possess exceptionally ‘clean’ copies of the books described or the
illustrations have been very skilfully prepared, for nowhere is there much evidence of unsightly
library stamps or paper imperfections which so often mar reproduced illustrations of this kind.
It is to be hoped that this publication will receive sufficient support to encourage the publication of
a ‘post-Linnean’ volume in the not to distant future.
M. WALPOLE
BOOK REVIEWS 207
Mr. Marshal’s Flower Album from The Royal Library at Windsor Castle. Introduction and
commentary by John Fisher, Preface by Jane Roberts. Pp. 128, including 36 coloured plates.
Victor Gollancz, London. 1985. Price £20-00 (ISBN 0—5S75—03536-6).
The mid-seventeenth century Florilegium of Alexander Marshall is now in the Royal Collection at
Windsor Castle. It is of particular interest as Marshall was renowned as one of the earliest water-
colourists and he spent a good deal of time and ingenuity in deriving pigments from various plant
and vegetable sources to provide a unique freshness to his coloured drawings. How well he
succeeded it is not possible to judge from this album of reproductions for, despite the use of
modern colour printing techniques, one is left with a feeling of lack of authenticity.
Thirty plates (from a total of 164) are reproduced, all with a distinct charm and each portraying
several species of plant (many being novelties of the time); and although the general impression is
of an artist with considerable botanical skill, the reproductions are such that it is really impossible
to assess the fine detail which was the hallmark of so many of the fine botanical drawings of this
period. Certainly the unique vibrancy of colour Marshall is said to have created is not apparent. I
am left with the distinct impression that justice has not been done to the originals and the hope that
the opportunity of seeing the original work might sometime be possible.
To supply suitable text for a rather random selection of drawings presents a challenge to any
writer. John Fisher has chosen to quote quite liberally from the old herbal writers, Gerard,
Culpeper, Turner and the like, but also weaves into his writing an additional mixture of folklore
and nomenclature plus the odd note of taxonomic and historical interest which results in very
readable and at times entertaining narratives. The scholarly preface by Jane Roberts pieces
together some details of the 17th century background of Marshall’s work and is a valuable
contribution to a publication which brings to our attention a unique item of early English botanical
literature.
M. WALPOLE
God’s Acre: the flowers and animals of the parish churchyard. F. Greenoak. Pp. 192, with
numerous black & white and water-colour illustrations. Orbis Publishing Ltd, London. 1985. Price
£12-95 (ISBN 0-85613-800-2).
By the time this review is printed, I feel confident that this beautiful book will be familiar to most
naturalists and possessed by many. On almost every page, Clare Roberts’ delightful illustrations
grace the text; it is indeed the perfect ‘coffee-table book’. It is also a particularly timely
publication, for all over the country there are groups of people who quietly lavish care upon our
20,000 churchyards, and treasure them as oases of peace and beauty in an increasingly harsh and
strident world. To these faithful servants — and to the very many others who appreciate their
devotion — this book is especially aimed.
The care of churchyards is a complicated subject which until recently was hardly ever the
concern of naturalists — unless, of course, they were also the incumbents, and even then the natural
history of the churchyard was often taken for granted. In this self-conscious age, however, it is no
longer possible for us to enjoy nature without realizing how fragile are the wild plant and animal
communities our ancestors took for granted. So, in the last decade, even churchyards have come
under a new scrutiny. Francesca Greenoak’s book is designed to do two things: to open the eyes of
ordinary readers to the wealth of interest in the wild life of the churchyard, and to convert those
who already enjoy the serenity and beauty of particular churchyards to a middle way of
management when, as so often is the case, the local community has to decide how the churchyard
must be cared for. In the first of these aims this book will surely succeed. In the second aim,
however, I must confess to a slight sense of disappointment for, though the author herself clearly
appreciates the principles of ecological management, the book presents no very clear practical
guidance to the enthusiastic naturalist who may wish to help in the conservation of his or her
favourite churchyard. Should these practical guidelines not have appeared as another Appendix,
along with the (very valuable) Appendices devoted to relevant literature and organizations? I have
in mind a condensed form of Arthur Chater’s excellent paper in the Church of England periodical
208 BOOK REVIEWS
Churchscape (vol. 3, 1984, pp. 21-7) entitled ‘God’s Acre: the conservation of consecrated
vegetation’, which gives clear advice on, for example, ivy control, the use of ‘strimmers’ for cutting
rank grass, and the control of Manpower Services Commission working parties!
The title of this book is oddly misleading in one important respect. Francesca Greenoak is too
good a botanist to restrict her text to ‘flowers’, or even to the Flowering Plants. Indeed, she is
clearly a convert to the study of lichens, as pages 87-92 amply demonstrate, and the trees of
churchyards also find their place in the book. But what, incidentally, has happened to the
bryophytes? Some of the most remarkable rarities of churchyard tombs are mosses, and the
specialist ecology of moss communities on gravestones is full of interest for the naturalist. If the
index is correct, mosses receive mention or illustration on only four pages, and not one of the three
illustrated species is said to be drawn in a churchyard.
On churchyard animals, the book ranges widely, though inevitably the birds and insects get the
lion’s share of publicity. (Incidentally the list of the ‘20 commonest churchyard birds’ supplied as
Appendix V by the British Trust for Ornithology, is surprisingly accurate for my own local
churchyard, and I can imagine many readers happily making their own comparisons.) The new-
comers to the naturalist’s ‘quarry’ are the bats, and churchyards are, of course, especially interes-
ting as habitats to be studied. ‘Call in your local Bat Group’ is Francesca Greenoak’s advice —
advice which I can strongly recommend from personal experience.
Minor errors seem to be commendably few, though many B.S.B.I. members’ eyebrows may be
raised when they read (p. 125): ‘the true Date Palm of the Holy Land grows in Britain . . .’ and
‘has been planted in a number of churchyards’ in the south-west! (Is this the familiar confusion
between Phoenix and Cordyline?). The index has passed a small random test reasonably well,
though the important churchyard fern genus Asplenium, mentioned at least three times in the text,
seems to have slipped through the net.
In her ‘Author’s Note and Acknowledgements’, Francesca Greenoak explains how much of the
idea for the book stemmed from a survey, organized by Sue Parish and conducted by Women’s
Institutes, of 1400 churchyards in England and Wales, and acknowledges her indebtedness also to
several specialist societies, amongst them the B.S.B.I., who have conducted churchyard surveys in
recent years. Among the many people whose help is acknowledged individually I was pleased to
see Mary Briggs, Arthur Chater and Frank Perring. The preparation of the book was much
affected by the death in 1983 of John Talbot White — there is a very apposite quotation from his
writings given pride of place in the beginning of the first chapter on ‘History and Heritage’ — and
Francesca Greenoak herself took over only at this late stage. We all owe her a great debt for so
attractively marrying together much separate advice, and wish the book all the success it so richly
deserves.
S. M. WALTERS
Insects and flowers, the biology of a partnership. Friedrich G. Barth (translated from the German
by M. A. Biedermann-Thorson). Pp. ix + 279, with 37 plates, many in colour, and 97 text-figures.
George Allen & Unwin, London & Sydney. 1985. Price £30-00 (ISBN 0-04-574029-1).
The time has passed when it was possible to deal reasonably adequately with all the aspects of
pollination biology in one book. Now one has to consult several, and they vary in their assumptions
about the reader’s background. This one assumes little and is written in everyday language. It puts
our knowledge of insect-pollination into a wide historical framework, relating the facts where
possible both to everyday life and to scientific theory of the past and present. The author is an
entomological neuro-physiologist, and his main concern is to tell us about the structure, sensory
equipment, sensory perception and behaviour of insects as they relate to pollination. In doing so,
he has accomplished an outstanding piece of popularization, carrying us to at least a basic level of
understanding in a long series of complex enquiries (but I make exception to the last two sections
of Chapter 25). All of it is fascinating, and much of it astounding. At the beginning, the life-history
of vascular plants and the main types of flower, their various attributes and the means by which
they secure the services of insects in pollination are briefly outlined in chapters that are inter-
spersed with others about insects, but in the later part of the book the insects take over entirely.
BOOK REVIEWS 209
While the book is accessible to the layman it will be of value to students, and if it is not deep
enough for them the extensive bibliography will take them further. The book is excellently and
generously illustrated with line-drawings, well-chosen colour photographs and scanning electron
micrographs. (The picture of a bee-fly on a bugle flower on plate 11 has apparently been rotated
anti-clockwise through a right angle.)
Two points on which explanation is inadequate seem worth mentioning. On p. 115 the colour-
change in the guide-marks on the flower of Aesculus, which occurs when nectar-production ceases,
is discussed, and it is mentioned that flowers of most species simply wither at this stage. The point
not made is that the distant signal of the large many-flowered inflorescence can remain unimpaired
if the pollinated flowers remain fresh. On pp. 179-180 it is stated that, of the perfumes that are
collected by certain tropical American bees, two compounds are specially attractive, and then that
“the attractiveness of an odor component may be diminished when a second component is added
to it’’. The point is that the number of species of bee attracted is diminished; the mix of compounds
actually found in the orchids varies from one species to another, and the different ‘perfumes’
resulting attract bee species selectively, “‘thereby” as the author says “‘increasing the specificity of
pollination”.
The German edition of this book was published in 1982; it is now almost faultlessly translated
and reads very well (but Pteridium aquilinum gets the English name ‘eagle fern’; ‘palm ferns’ (p.
259) are presumably cycads). However, in the bibliography, English editions of other translated
works ought to have been cited instead of the originals, and the list of books on pollination ecology
for Chapter 3 should have been up-dated (most of the missing titles will be found under ‘Further
Reading’ in The sex life of flowers by Meeuse & Morris (1984), reviewed Watsonia 15: 416). Book
production is good but there are two irritating faults: (1) when plates are mentioned in the text
there is no page reference; (2) the chapter number should have been included in the running heads
to facilitate reference to the bibliography which is at the end of the book and divided into chapters.
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Obituaries
JOHN PATRICK MICKLETHWAIT BRENAN
(1917—1985)
With the death of Pat Brenan in London on 26th September 1985 the world of botany lost one of its
giants. This, however, is not the place to dwell upon his numerous and important contributions to
African floristics or his administrative accomplishments at Kew, but rather to review his work and
achievements in the study of the flora of our islands.
Born at Chiselhurst, Kent, on 19th June 1917, the younger son of Dr A. R. M. Brenan and his
wife Jill, Pat was attracted to the study of plants from early childhood, an interest encouraged by
his father and brother, both of whom were knowledgeable on British wild flowers. He received his
early education at Tonbridge, and it was from here in 1933 that he joined the B.S.B.I. (then the
Botanical Society and Exchange Club of the British Isles). This was a period of great transition in
the Society’s fortunes, for the autocratic G. C. Druce who had dominated its activities for so many
years had died in 1932, and a wind of change was blowing. The Society’s report for 1934 included a
spate of Kentish records from the enthusiastic new member, as well as some contributed by his
father and brother. A report of Bupleurum subovatum (as B. lancifolium) as a spontaneous garden
weed at Tonbridge indicated the high standard reached by the young botanist, for descriptions of
that bird-seed adventive did not then appear in British Floras and such finds were invariably
referred to the rare B. rotundifolium; furthermore, records of several Chenopodium species hinted
at an awakening interest in a critical genus on which he was to be eventually acknowledged an
international authority.
In the summer of 1935 Pat made contact with the skilled alien plant specialist Mrs C. I.
Sandwith, and together they visited Avonmouth Dock and other important exotic plant sites in the
Bristol area; later in the year they extended their forays to similar habitats at Barry and Cardiff
Docks. During this year too he paid one of his early visits to the Royal Botanic Gardens, Kew,
little anticipating that some 40 years later he would become its Director. Attention was also given
to interesting adventives growing by the Thames between Kew and Mortlake where, with his
maternal uncle, R. N. Parker, a botanist who had spent much of his life in the Indian Forestry
Service, he saw Ailanthus altissima, Ficus carica and Vitis vinifera. Late summer and early autumn
days were spent in visits to those happy hunting grounds for the alien plant enthusiast, the famous
Dagenham waste in Essex and the vast metropolitan rubbish-tips at Yiewsley, Middlesex and Iver,
Bucks. In 1936 he won a scholarship to the University of Oxford, and proceeded to Brasenose
College, where it had been his intention to read modern languages; a growing love of plants, a
natural inclination to taxonomy and influence by his brother persuaded him, however, to change to
natural sciences. Work now filled most of his waking hours, but such leisure time that was available
was spent in the field in the rural countryside of Oxfordshire and Berkshire, often in the company
of John Chapple who had liased closely with Druce, and who was at this time Honorary Secretary
of the B.E.C., P. G. Beak, the Populus specialist, R. Burn and N. E. G. Cruttwell. During the
long vacation further visits were made to Bristol and Cardiff with Mrs Sandwith and her son Noel,
who became one of Pat’s closest friends, an association which lasted until the sudden and untimely
death of Sandwith in 1965. Never was there a more knowledgeable partnership on British plants
than these two; a day in the field with them was both a privilege and awe-inspiring, and one
returned home realizing sadly how little one knew of the British flora. It was in 1936 also, in the
company of Ted Lousley, that Carex montana was refound between Tunbridge Wells and Eridge,
FE. Sussex, in a locality where it had first been noted as British by William Mitten in 1845; later he
and Pat saw Bromus interruptus growing in a field of Sainfoin at Upper Halling, W. Kent.
Alien studies with the Sandwiths at Bristol and Cardiff were resumed in the summer and autumn
of 1937, and a search for brewery malting adventives was undertaken at Burton-on-Trent, Staffs. in
the company of John Chapple. During term time field work continued in the upper Thames Valley.
2 OBITUARIES
In August 1938 and again in 1939 Pat toured Ireland with N. Douglas Simpson, making many new
and important plant finds. In 1939 too, while staying with Simpson at his Bournemouth home, a
joint foray to Poole Harbour, Dorset resulted in the discovery of Carex x tornabenii (C. distans X
extensa), a hybrid new to the British flora.
During 1940 Pat graduated with first class honours in botany and accepted a post at the Imperial
Forestry Institute, Oxford. By the end of his student days he had compiled a personal herbarium of
over 5000 British and Irish specimens, with an emphasis on critical material, but he showed little
regard for the Exchange Club activities of the Society, contributing but a few sheets of a single
gathering of Senecio squalidus x vulgaris collected near Oxford in 1946 by himself and John
Chapple.
His duties at the Imperial Forestry Institute consisted of teaching and research, and saw the
beginning of his African studies; and, although work occupied much of his time, some leisure hours
were still allocated to investigating the Oxfordshire and Berkshire flora, sometimes with F. B.
Hora, the mycologist, and J. N. Mills, who later specialized in Hieracium. In March 1945 he was
elected to the B.E.C. Committee (now Council) and to the Excursions and Field-work sub-
committee (now Meetings Committee), on which he served for a number of years; he also
organized and led a field meeting in the vicinity of Oxford. The results of his extensive field work in
Oxfordshire and Berkshire were brought together in a long paper in the Report of the Botanical
Society & Exchange Club for 1943-44; and in the same periodical appeared a joint account with J.
E. Lousley on floral variations in Stellaria holostea.
The year 1947 was an eventful one: Pat joined Peter Greenway and Ronald ‘Keay on a plant
collecting expedition to Northern Rhodesia (Zambia) and Tanganyika (Tanzania), and shortly
after his return to England again visited Africa with the Cambridge University Botanical
Expedition to Nigeria and the Cameroons under the leadership of Paul Richards. Despite this
increased activity, British botany was not forgotten; he briefly became B.S.B.I. Local Secretary for
Oxfordshire, v.c. 23, and joined the Society’s panel of specialists as the referee for Amaranthus
and Chenopodium, a function that he maintained to the end. His continued interest in alien plants
was confirmed by the publication of a paper entitled “‘A contribution to the adventive flora of
Southampton”’.
Pat’s reputation as an extremely competent taxonomist with an increasing knowledge of African
plants both in the field and the herbarium was now established, and in 1948 he transferred to the
Tropical African Section of the herbarium, Royal Botanic Gardens, Kew, as Senior Scientific
Officer, to work on the new Flora of Tropical East Africa. Notwithstanding a growing work load,
time was found to pen a few short notes on British plants. In April 1949 he was elected to the
B.S.B.I. Council, on which he served for three years; and in collaboration with John Chapple he
produced an account of the Australian Myriophyllum verrucosum in Britain, and followed it with
one of his most important papers, involving much detailed and careful research, “‘Artemisia
verlotorum Lamotte and its occurrence in Britain”. During this period he was still expanding his
alien studies — visiting the ‘shoddy’ fields of Bedfordshire for wool adventives and the soya bean
site near Harefield, Middlesex, for the wide range of N. American ephemerals.
In 1950 Pat married Jean Helen Edwardes and settled down to family life at his Kew home. He
was elected a Fellow of the Linnean Society of London in 1952 and in the following year joined the
B.S.B.I. Publications Committee where, apart from a few short breaks, he was a prominent figure
almost to the time of his death. His family and his work on African plants now filled his time as he
steadily built on his reputation as one of the world’s leading taxonomists with a special interest in
the families Leguminosae, Chenopodiaceae and Commelinaceae. During 1959 he became head of
the Tropical African Section of Kew Herbarium, and in 1961 produced his major paper on British
botany — a revision of the critical genus Amaranthus in these islands. Three years later he wrote the
account of Chenopodium for the first volume of Flora Europaea.
On the retirement of Dr C. E. Hubbard in 1965, Pat was a natural successor to the post of
Deputy Director and Keeper of the Herbarium and Library at Kew, and was urged, somewhat
against his will, to apply for the position (successfully). The move to an administrative situation at
the zenith of his scientific life doubtless imposed a burden on him, but he overcame it, and with
renewed vigour became Botanical Secretary of the Linnean Society from 1965 to 1972, President of
the Association of Tropical Biology (1970-1971) and President of the Kew Guild (1972-1973).
With the early and unexpected resignation of Professor J. Heslop-Harrison from the Directorship
OBITUARIES 213
of the Gardens in 1976, Pat was again the obvious, if somewhat reluctant, candidate to follow him,
which he did. In 1977 he was appointed a Visiting Professor to the University of Reading, and in
1978 was honoured by Portuguese botanists in being elected an Honorary Member of the
Sociedade Broteriana; he also received the highly coveted Victoria Medal of Honour from the
Royal Horticultural Society. To his many advisory posts he added that of Honorary Botanical
Adviser to the Commonwealth War Graves Commission, serving until 1982. In 1979 he was given
one of the Willdenow commemorative medals struck by the Berlin Botanic Garden, and in May
1981 he was elected President of our Society; he also became Director of the International Council
for the Development of Under-utilized Plants.
October 1981 saw Pat’s retirement from the Directorship of Kew, and in the months that
followed he travelled abroad with his wife, renewing old friendships and visiting S. Africa,
Australia and Hawaii. While in S. Africa in February 1982 he was one of three botanists awarded a
special medal issued by the South African Association of Botanists. On his return to England he
joined the Council of the National Trust.
Botanical interests were pursued almost to the end, and his large, loose-limbed figure was
frequently to be seen in the herbarium and library at Kew, where no matter how busy he was he
always found time for a friendly word. Time allotted to various committees was still willingly and
freely given, and at the time of his death he was serving over twenty organizations.
Pat Brenan was a brilliant scholar, fluent in several European languages and in Latin; a kindly,
modest man who showed great consideration to all; a private individual blessed with a remarkable
memory and a great curiosity, while beneath a somewhat serious exterior lurked a fine sense of
humour. His botanical life was aptly summed up by Dr Roger Polhill in his address at the memorial
service held at St Anne’s Church, Kew Green, on 23rd October, in the words — “To Pat all plants
were wondrous’’.
THE BRITISH AND IRISH WRITINGS OF J. P. M. BRENAN
1939
Galinsoga quadriradiata Ruiz & Pav. in Britain. Rep. botl Soc. Exch. Club Br. Isl. 12: 93-94.
1945
(With Simpson, N. D.) A hybrid sedge new to Britain. N. West Nat. 20: 202-206. [Carex distans X extensa = C.
x torbenii Chiov. |
1946
(With Mitts, J. N.) 308/5. Scabiosa arvensis L. Rep. botl Soc. Exch. Club Br. Isl. 12: 679-680. [Note on
aberrant form].
396/4 x 2. X Cirsium sabaudum Lohr (C. acaule X vulgare = lanceolatum). Rep. botl Soc. Exch. Club Br. Isl.
12: 682-683.
641/1. Myrica gale L. Rep. botl Soc. Exch. Club Br. Isl. 12: 686-687. [Note on monoecious bushes with
androgynous catkins}.
Notes on the flora of Oxfordshire and Berkshire. Rep. botl Soc. Exch. Club Br. Isl. 12: 781-802.
(With Lous-ey, J. E.) Floral variations in Stellaria holostea L. Rep. botl Soc. Exch. Club Br. Isl. 12: 840- 846.
1947
354/2b. Galinsoga quadriradiata Ruiz & Pav. var. hispida (DC.) Thell. Rep. botl Soc. Exch. Club Br. Isl. 13:
30.
A contribution to the adventive flora of Southampton. Rep. botl Soc. Exch. Club Br. Isl. 13: 106-112.
1948
196/11. Crataegus monogyna Jacq. var. xanthocarpa Lange. Rep. botl Soc. Exch. Club Br. Isl. 13: 260-262.
+435/13. Campanula alliariifolia (‘‘alliariaefolia’’) Willd. Rep. botl Soc. Exch. Club Br. Isl. 13: 263-265.
7596/26(2). Amaranthus blitoides S. Wats. Rep. botl Soc. Exch. Club Br. Isl. 13: 269-271.
+600/26(2). Chenopodium pumilio R. Br. Rep. botl Soc. Exch. Club Br. Isl. 13: 272-273.
Senecio squalidus X vulgaris L. Rep. botl Soc. Exch. Club Br. Isl. 13: 364.
214 OBITUARIES
1949
(With CuHappLe, J. F. G.) The Australian Myriophyllum verrucosum Lindley in Britain. Watsonia 1: 63-70.
(With Simpson, N. D.) The results of two botanical journeys in Ireland in 1938-9. Proc. R. Ir. Acad. 52B: 57-
84.
1950
Artemisia verlotorum Lamotte and its occurrence in Britain. Watsonia 1: 209-223.
1952
ALLEN, G. O. British Stoneworts (Charophyta). Watsonia 2: 213. [Review]
1953
*206/13. Galium parisiense L. Watsonia 2: 413.
1956
Notes on the flora of Oxfordshire and Berkshire: 2. Proc. bot. Soc. Br. Isl. 2: 105-114.
1961
Amaranthus in Britain. Watsonia 4: 261-280.
1964
Chenopodium, in TuTin, T. G., et al., eds. Fl. Europaea 1: 92-95. Cambridge.
1966
Noel Yvri Sandwith 1901-1965. Taxon 15: 254-255. [Obituary]
1967
Obituary: Noel Yvri Sandwith (1901-1965). Proc. bot. Soc. Br. Isl. 6: 418-422.
1968
The relevance of the national herbaria to modern taxonomic research, in HEywoop, V. H., ed. Modern
Methods in Plant Taxonomy, pp. 23-32. London.
1971
The role of the herbarium in the work of Kew. J. Kew Guild 8: 1127-1134.
1972
Profile. Professor John Heslop-Harrison. B.S.B.I. News 1(2): 30-31.
Edgar Milne-Redhead. Kew Bull. 26: 1-3.
1973
The value of herbaria for cultivated plants, in GREEN, P. S., ed. Plants wild and cultivated, pp. 38-47. London.
1975
Mr V. S. Summerhayes. The Times, 7th January 1975. [Obituary]
Obituary: Victor Summerhayes. Bull. Brit. Ecol. Soc. 6(2): 5-6.
1976
Gopwin, H. History of the British flora — A factual basis for phytogeography, 2nd ed. Kew Bull. 31: 411-412.
[Review]
1980
Obituary: Edward James Salisbury (1886-1978). Watsonia 13: 68-70.
1981
President’s message. B.S.B.I. News 2(28): 3.
1983
Presidential address, 1982. The British flora — A changing picture. Watsonia, 14: 237-242.
D. H. KENT
OBITUARIES 215
URSULA KATHERINE DUNCAN
(1910—1984)
Ursula Duncan M.A. LL.D., F.L.S., L.R.A.M. was born in Kensington, London in 1910, the
elder daughter of the late Commander J. A. Duncan and his wife, the former Dorothy Weston.
During her infancy, the Duncan family, whose origins in Angus can be traced back to 1705,
returned to Parkhill, Arbroath, their ancestral home since 1799. Dr Duncan’s childhood education
was entrusted to a governess, Miss Isobel Leslie, and under her tutelage she passed the Entrance
School Certificate to Cambridge at the age of 15, obtaining a distinction in Greek. Of studious and
intellectual bent, Dr Duncan registered as an external student of London University, obtaining a
BA in 1952 and a MA in 1956 in classics, a period during which she also visited Greece. She also
became a pianist of some distinction, excelling in musical theory and professing a preference for
Bach and Mozart; she became a Licentiate of the Royal Academy of Music in 1948.
During the Second World War she served for a time in the Censorship Department, based on
Inverness, dealing with the censorship of forces mail, an experience which she confided “‘brought
her face to face for the first time with harsher realities of personal life’’ and from which she derived
an understanding and experience of people which she valued in later life. The death of her father in
1943 curtailed her work in Inverness, and she returned to Parkhill to take over the supervision of
the family estate including 600 acres of farmland, an undertaking which she proceeded to carry out
with financial astuteness, foresight and considerable success for the rest of her life.
Her interest in botany commenced at the age of ten, and she was much encouraged in the pursuit
by her father after his retirement from the Royal Navy at the end of the First World War. What
was destined to be her major recreation throughout life started with vascular plants and extended
in 1939 to the bryophytes as well as the lichens.
She was a member of the Wild Flower Society from her youth and joined the Botanical Society
and Exchange Club of the British Isles in 1931, becoming the recorder for Easter Ross and serving
on the Committee for the study of the Scottish Flora for many years. As with her studies of
cryptogams she delighted in those genera which presented a taxonomic challege, such as
Potamogeton, Hieracium and Taraxacum; she always considered the grasses as the most intractable
and she became a considerable expert in this group. With these ‘little difficulties’ she kept up a
life-long correspondence with a wide range of specialists who, in turn, benefitted greatly from her
many new records, collections and valuable field data, mostly from botanically under-explored
areas of Scotland. Her personal generosity and kindness to her botanical friends, young or old,
amateur or professional, became one of the hallmarks of her considerable botanical flair and her
great success as a field botanist.
The publication of the Flora of East Ross-shire in 1980 marked the successful culmination of
many years of intrepid and tireless tramping to all areas of the vice-county, often alone, in
inhospitable terrain, and in all types of weather, always, characteristically, in wellington boots! She
also had a particular interest in the floras of Angus and the Island of Mull; indeed her preliminary
studies on the vegetation of Mull were largely instrumental in the initiation of the Flora of Mull
Project carried out by the Botany Department, British Museum (Natural History), to which she
subsequently gave generous and wholehearted support.
She was introduced to lichenology by Mr R. H. Brown of Oxford and Dr W. Watson of Taunton
and soon became one of the select few who kept lichenology alive during the period of its greatest
decline in Britain between 1945 and 1955. Even more importantly, by unstintingly putting her
time, knowledge and collections at the service of young aspiring lichenologists she undoubtedly
played a premier role in the renaissance of the subject in Britain. Of her three books on lichens,
the last, An Introduction to British Lichens, published in 1970, reflects the resurgence of interest in
the group and the major part she played in its revival. She was a founder member of the British
Lichen Society. As with other groups of plants, she contributed records continually to mapping
schemes and census catalogues, providing, for instance, over 25 per cent of the Scottish records in
W. Watson’s Census Catalogue of British Lichens (1954).
Her early studies on mosses and hepatics were with her namesake Mr J. B. Duncan, and she
soon became an active member of the British Bryological Society. It was characteristic that her
interest turned in particular to the challenge offered by the Sphagnales, for which she acted as a
referee and published an illustrated key. As with vascular plants and lichens, she also acted as a
216 OBITUARIES
referee for beginners, ecology students, mapping recorders, etc., attending to their many letters
and specimens with prompt and meticulous thoroughness.
She eschewed formal representation on committees and was happiest when in the field, either
gently and patiently encouraging ‘her beginners in smalls and greens’ or, with botanical friends,
amateur or professional, exchanging specimens and expertise, or leading one of the many field
meetings to various parts of the British Isles. She had a rare intuitive flair which made her an
outstanding field botanist, in addition to being a naturally gifted teacher with an instinctive ability
for conveying her enthusiasm and knowledge whatever the level of experience of the recipient.
Reticent about her many achievements and naturally modest by nature, she never sought the
limelight and was most content in the surroundings of her own home. Friends who visited her were
always welcome and were able to share with her her enthusiasm for gardening, particularly her
interest in vegetable growing, rhododendrons, Meconopsis, Cardiocrinum and, in the quarry which
furnished the building stone for Parkhill, a fine range of Primula species.
She amassed a large and important herbarium of vascular plants, rich in carefully collected
material of critical and difficult taxa and much of it identified or checked by specialists; she donated
this to Dundee Museum in 1983. Failing eyesight and ill-health caused her to give her cryptogams
to the herbarium of the Royal Botanic Garden, Edinburgh later in that year.
Dr Duncan was honoured by the Linnean Society with the Bloomer Medal, by the British Lichen
Society with an honorary membership and, in 1969, by the University of Dundee with the
Honorary Degree of Doctor of Laws, public recognition and expressions of_gratitude for her
private generosity and kindness to all whom she helped and who shared her enthusiasm for plants
and their habitats, as well as recognition of her unique contribution to British Botany.
P. W. JAMES
Watsonia, 16, 217-236 (1986) 217
Reports
CONFERENCE REPORT
RECORDING CRITICAL GROUPS IN THE FLORA OF THE BRITISH ISLES,
LIVERPOOL, 13th-14th SEPTEMBER 1985
INTRODUCTION
A total of 98 members of the B.S.B.I. and the Biology Curators’ Group attended this meeting
based at the Merseyside County Museums and the University of Liverpool. A feature of the
conference was the workshop sessions where small groups were guided through the difficulties of
identification in critical groups by session leaders, each of whom was particularly knowledgeable in
his or her field. To help these sessions, the Museum’s herbarium was available and a block of four
rooms enabled simultaneous sessions to be held, yet with the flexibility for members to change
from one session to another as they wished. Most of the sessions were held at the County Museums
but other venues included the Department of Botany at the University of Liverpool (Computer
Workshop) and the Halls of Residence of the University of Liverpool (Recorders’ Forum), whilst a
Reception by the Merseyside County Council and an evening talk on ‘Operation Groundwork’
were held at the Merseyside Maritime Museum.
The Society is greatly indebted to all those who helped to make the conference a success, and is
especially grateful to the generosity of the speakers and workshop leaders who helped to keep the
costs of the conference to affordable levels.
E. F. GREENWooD & J. R. EDMONDSON
ABSTRACTS OF PAPERS
LOCAL FLORAS: THE CRITICAL ELEMENT
THE CRITICAL ELEMENT — WITH SPECIAL REFERENCE TO THE ISLE OF MAN
The coverage of critical groups is disappointingly meagre in many modern local Floras. Three
reasons can be identified: printing costs, the trend towards shorter-term projects, and lack of
taxonomic self-sufficiency. That the sheer size of certain groups compels a certain economy of
treatment has to be accepted: sacrificing the coverage of critical taxa in the interest of producing a
Flora speedily is more open to challenge. Short-term projects have become the norm in those parts
of Britain where field botanists are sufficiently numerous to give rise to substantial teams and
where habitat alteration is sufficiently rapid and severe to cause Floras to date quickly. The
increasingly heavy emphasis in local Flora work since 1948 on dot-distribution mapping has tended
to militate against adequate recording of critical taxa, despite the special value of these in
indicating regional and sub-regional affinities and the greater need for their existence to be notified
for conservation purposes. By undertaking the study of these taxa themselves, instead of
delegating the task to a series of specialists, some or all of whom may be unfamiliar with the local
environment and with the local flora more generally, Flora writers can provide a more integrated
treatment and use their resulting breadth of view to make illuminating comparisons. The size of
this task, however, must vary from area to area. The author’s experience in the Isle of Man,
discussed by way of example, may have been by no means typical in this respect.
D. E. ALLEN
218 REPORTS
THE ALIEN FLORA OF GUERNSEY
Ten years ago Wild Flowers of Guernsey (1975) put the recorded wild and naturalized plants
there at 1340. Since then the total has risen to some 1400. Over half of these are aliens. They
come from the four corners of the globe, some are transient, but many have settled down and are
a welcome enrichment of the island’s tally, only very few endangering native vegetation. Naming
new ones calls for familiarity with foreign Floras, consultation of herbaria and a generous amount
of help from professional botanists, plantsmen and other specialists. One or two in Guernsey are
not yet satisfactorily named through the absence of type specimens or other reliable material.
But these newcomers add extra fascination to a fascinating island, not least in pondering when
some will become similarly established in England.
D. McCLintock
RECORDING IN IRELAND: PROBLEMS AND PRIORITIES
The organization of recording in Ireland was described and discussed, together with remarks on
problems encountered in the field. These include the distances recorders must travel from home
to their area of study, and problems of species identification. New vice-county recorders’ plans
were outlined, and reference was made to contributions by visiting botanists. ~
D. DooGuE
B.S.B.I. HANDBOOKS: PROGRESS REPORTS
CRUCIFERAE
The Cruciferae are well represented in Britain and Ireland, with about 300 species recorded to
date. They can be considered a critical group because there are a large number of superficially
similar species (especially in the ‘Yellow Crucifer’ aggregate) that cause problems of identifica-
tion at both the generic and specific level. The genera, despite forming natural groups, are
especially hard to define clearly. Within a genus, the species are usually easier to distinguish
though there are a few critical genera. Many of the characters require careful examination and a
combination often has to be used. It is usually necessary to have both mature flowers and ripe
fruit. Some characters are more easily observed on fresh rather than herbarium material, and
vice versa. The aim of the Cruciferae Handbook is, therefore, to provide an aid to identification
of species likely to be found in the British Isles and Ireland. It will follow the general style and
format of the other B.S.B.I. Handbooks. Taxonomy and nomenclature will follow Flora Euro-
paea but incorporate recent updates. Descriptions are being prepared from live and herbarium
material. Distribution maps will be provided for many species by the Biological Records Centre.
Ecological, cytological and other general information will be included where relevant. It is hoped
the Handbook will be illustrated by amateur artists. It is planned for publication in 1989.
li C.uG. Rick
LEGUMES
A publication on legumes is being prepared for the B.S.B.I. Handbook Series, and information
and help is sought from members. The Handbook will differ from its predecessors in a number of
ways. For each species the description, which will be comparatively brief, will be accompanied by
illustrations and notes on habitat and use. Distribution maps may also be included. Approxi-
mately 100 native and well established alien species will be treated in full but thumbnail sketches
will be included for many other aliens. The Handbook is anticipated to be of interest to foreign
REPORTS 219
botanists, particularly those from areas in which the weed legume flora includes many of our species.
A variety of identification aids, including multi-access keys are being considered as alternatives to
dichotomous keys. Separate chapters will deal with dispersal, legume/insect interactions, pollination
biology and pollen and honey sources. An innovative aspect of Handbook production is that, for the
tribe Vicieae (approximately one third of the species), morphological and other information will
come initially from the Vicieae Database in Southampton.
R. ALLKIN
WEEDS
Weeds constitute a familiar group of plants that affects the lives of all of us. Many definitions have
been proposed for the word ‘weed’, but most agree that a weed is an opportunist plant that causes
problems of one sort or another to agriculture, horticulture and other human industries. Weeds have
evolved alongside man and have become adapted to the various niches provided by his activities. In
order to be able to control weeds with any degree of efficiency, it is important to be able to identify
them correctly and to know at least a little of their biology, notably aspects of their life-history,
dispersal and ability to withstand destruction by man. Thus, a handbook that brings together
information on this group of plants is now in preparation, as there are surprisingly few authoritative
texts on the subject. It should be noted that weeds are intrinsically interesting, and they are perfect
material for the study of evolutionary processes — easily grown plants, adapted to a variety of
catastrophic habitats! The proposed Handbook is being designed for use principally in arable and
garden habitats, and other disturbed communities created and maintained by human activities.
Aquatic ecosystems are not to be included (although some species typical of wetter soils will be
treated), as this would introduce an unwieldy, extra element into the text. Grasslands, with the
exception of lawns, will not be given a full treatment. The systematic section of the book will contain
accounts of individual species and, in certain cases, groups of species within a genus. There will be
detailed descriptions, notes and ecology, distribution and history of particular species, and any
appropriate comments on pharmacology, folklore and more anecdotal material. It is thus hoped to
introduce weeds to a wider public.
J. R. AKEROYD
TAXONOMIC WORKSHOPS
A LOCALIZED APPROACH TO THE HAWKWEED PROBLEM
The emphasis of the workshop was to show how aspiring field botanists can gain practical knowledge
of the genus. It sought to cover the collection of specimens, the main sources of written information,
the major relevant herbaria, and the subdivisions of the genus in Britain.
Examples of the main series/sections were available for study. These were recently determined
specimens mostly from southern Scotland and northern England.
D. J. McCosH
VARIATION AND HYBRIDITY IN SOME BRITISH SPECIES OF ASPLENIUM AND POLYSTICHUM
Characters were given for the separation of (a) Asplenium adiantum-nigrum, A. billotii and the
hybrid A. X sarniense, and (b) the serpentine and non-serpentine forms of A. adiantum-
nigrum. A range of material was available for comparison, together with specimens of the
European A. cuneifolium. Help was also available with other difficult species in both Asplenium and
Polystichum, and in the latter genus particular attention was paid to the identification of hybrids by
spore examination.
A. SLEEP
220 REPORTS
EUPHRASIA
The genus Euphrasia in Europe consists of a successful group of hemiparasites of a diverse range of
grassy, herb-rich habitats, including marshes, saltmarshes, heather moorland, base-poor to base-
rich grassland, sea-cliffs and mountain ledges. Numerous morphologically distinct ecotypes exist,
some of which have a wide European range. It is convenient to recognize at least the major taxa as
separate species, though breeding barriers are often weak or non-existent and the distinctness of
the taxa is maintained largely by habitat separation. The British Isles have 21 of the currently
accepted 48 European species and with ever-changing patterns of land-use and disruption of
habitat-boundaries, the genus shows a complex pattern of hybridization and evolution of local
taxa. A flexible approach to the genus was encouraged and the workshop took an ecological view
of the major taxa and attempt to define the morphological characters and habitat requirements of
the recognized species. The problems of the local recorder, who may be faced with numerous
populations that do not readily fit any single species, were discussed, as was the treatment of
Euphrasia in recording schemes.
A. J. SILVERSIDE
INTERPRETING THE CHARACTERS OF THE CHARACEAE
Students of the Characeae have long indulged in discussion as to the validity of many of the
supposed species. Since the publication of the standard work for the British Isles (J. Groves & G.
R. Bullock-Webster, The British Charophyta (1920 & 1924)), the group has been the subject of a
major, world-wide revision (R. D. Wood, Monograph of the Characeae, (1965)) which has reduced
the number of species reported from these islands by half. Wood questioned the taxonomic
significance of many of the characters previously considered important in separating species, but
his views have not been accepted by the majority of the European workers, who remain
traditionalists. A further dimension has been added to the debate by experimental workers
(notably V. W. Proctor) whose success with laboratory cultures and breeding experiments has both
clarified and confused the situation. In preparing a Handbook to the Characeae of the British Isles,
the author has re-evaluated the 40 or so taxa recorded, adopting a mainly traditional approach in
interpreting the characters examined but tempering this with a regard to the work of Wood and the
experimentalists.
J. A. Moore
INFORMAL EVENING SESSION
OPERATION GROUNDWORK: WILDLIFE IN THE URBAN FRINGE
A detailed botanical survey has been carried out in the boroughs of St Helens and Knowsley,
Merseyside. A brief description of the methods used was given, especially of a new system tested
on the urban areas. The rural area is dominated by intensive barley production, while in the urban
areas much open space is ‘ryegrass desert’ or species-poor rough grassland. There are some
remnants of semi-natural vegetation including acidic grassland, brook communities and a fragment
of ‘heath’. Secondary habitats include some rich meadows, a number of 19th century plantation
woods and many marl pits, which together hold much aquatic life. However, many of the most
interesting plants are found on derelict industrial land. St Helens has a large and varied collection
of such sites, e.g. maritime species on drainage from waste heaps, calcareous grassland on Leblanc
process waste, orchids on sand waste, acidic flora on colliery shale, marshes, abandoned dams,
railways and the canal. A number of the species found on such sites are locally uncommon or rare —
or would be if there wasn’t so much derelict land in north-western England.
H. J. ASH
REPORTS 221
CURRENT RESEARCH INTO CRITICAL GROUPS
THE STATUS OF EPIPACTIS LEPTOCHILA AND E. DUNENSIS
Epipactis leptochila and E. dunensis are two very closely related, largely autogamous helleborines.
They were formerly thought to be ecologically, distributionally and morphologically distinct; E.
leptochila is a woodland plant of southern England and from northern France to Denmark, with a
narrow, acute, patent epichile. E. dunensis is endemic to sand-dunes in England and Wales, and
has a broad, short, subobtuse, recurved epichile. The only other morphological feature by which
they differ statistically is sepal length, which usually exceeds 10 mm in E. leptochila, but is shorter
in E. dunensis. E. muelleri, from central Europe is a woodland plant which is virtually
indistinguishable from E. dunensis. E. leptochila var. cordata is probably the same plant.
Since 1974, about 15 inland populations of Epipactis have been discovered, chiefly on lead and
zinc polluted sites under birch in Lincs., Yorkshire and Northumberland. These vary within and
between populations in epichile posture and dimensions and sepal length, and contain individuals
which are indistinguishable from southern E. leptochila, and those which cannot be separated from
E. dunensis/muelleri. It is concluded that E. dunensis cannot be maintained as a species, and it is
suggested that E. dunensis is returned to varietal rank as E. leptochila (Godf.) Godf. var. dunensis
ieand IT: A; Stephenson (J. Bot. 59: 205, 1921).
A. J. RICHARDS
DEFINING AND DELIMITING DACTYLORCHID TAXA
Classical taxonomic methods have proved inadequate for dealing with the extensive morphological
variation exhibited by dactylorchids (genus Dactylorhiza) within the British Isles. The resulting
controversies surrounding the classification and nomenclature of these plants have been
compounded by: 1) major differences of opinion over the meaning of taxonomic ranks, 2)
erroneous descriptions based on unrepresentative data, and 3) a failure to place British
populations within a European perspective. Multivariate analyses based on c. 50 morphological
characters sampled from 87 dactylorchid populations have enabled the variation within this genus
to be quantitatively expressed, and have led to a more objective delimitation of taxa based on
morphometric, distributional and ecological criteria.
R. M. BATEMAN & I. DENHOLM
IDENTIFYING BRITISH BROME-GRASSES
Although only a fragment of the genus, 22 species of Bromus are likely, in varying degrees, to be
encountered in Britain. Many of them present puzzling variation or are downright awkward. The
lecture gave delegates the opportunity to consider some of these taxa, including some of the most
‘difficult’ species pairs and groups, side by side. Some of the differential characters are hard to
communicate by mere words and delegates were asked to test out a new, draft key to British
material. New characters from spikelet morphology were illustrated. There was a brief analysis of
the reasons for the critical nature of some of these species. They include the breeding system,
developmental variation, environmentally induced plasticity, hybridization and the sheer smallness
of some of the differences. Inevitably, what ca:: be done by way of circumscription has to take into
account the variation seen as well as the reasons so far established (by experiment for instance) for
at least some ot it.
P. M. SMITH
222 REPORTS
GETTING TO GRIPS WITH RUBUS
The large number of local variants, named and unnamed, that exists in this group calls for a chain
of people versed in the Rubi of limited areas. Many of these variants will be unknown to the
national referees, and some means needs to be devised of enabling local specialists to examine each
other’s unnamed material. The matching of the recently-described R. norvicensis of the Norwich
and Ipswich districts with a bramble long known round Romsey in Hampshire suggests that even
floristically very different areas may prove to have species in common. More locally-based work
might be forthcoming if the genus were not seen as prohibitively difficult. The difficulties,
however, tend to be exaggerated. It is mainly in south-eastern England that the number of species
becomes bewildering: over much of the British Isles reasonably widespread ones are comparatively
few and their distribution patterns are more clear-cut. Given a good visual memory and a necessary
period of intensive field study, the principal local forms can normally be mastered inside a season
in these latter areas. Complications are caused by the frequency with which a number of species
hybridize and by the variability of many under different environmental conditions. Unfortunately
the only recent monograph, W. C. R. Watson’s Handbook of the Rubi of Great Britain and Ireland
(1958), confuses almost as much as it helps.
D. E. ALLEN
HYBRIDIZATION: CRITICAL GROUPS
IDENTIFYING HYBRIDS
Problems encountered in identifying hybrids are mostly the result of the nature of the hybrids
themselves, or of the would-be identifier’s poor understanding of it. The rule-of-thumb that
hybrids are intermediate and sterile works some of the time, but not all of it. With regard to
sterility, most hybrids are quite highly sterile, but most are fertile to some extent, and many are
quite highly and a few fully fertile. Equally, sterility exists commonly in nature from a wide range
of causes other than hybridity, e.g. adverse weather, lack of pollinators, lack of compatible pollen.
With regard to intermediacy, most hybrids are intermediate to varying degrees, but many are much
closer to one of the parents, and some are identical to one or other and some look unlike either.
The results of crosses between two individual plants often show a wide range of variation, only
some of which might conform with what we recognize as a hybrid. As a result, our determination of
hybrids will sometimes be accurate, sometimes an underestimate and sometimes an overestimate.
The only certainty is that we can never be 100% sure, because we can never re-create a particular
hybridization event.
C. A. STACE*
IDENTIFICATION IN RANUNCULUS SUBGENUS BATRACHIUM AND CURRENT WORK ON RANUNCULUS
PENICILLATUS
A new tabular key to species in Ranunculus subgenus Batrachium was distributed, with a request
for help with testing the key in the field. A tabular key is particularly useful in this group because it
enables identification to be based on all available features and incomplete material can be
identified as accurately as possible. Characters used in the key were discussed and illustrated.
Hybridization within the group not only presents difficulties for identification, but also plays an
important evolutionary role. The Ranunculus penicillatus group is thought to have arisen as
amphidiploid hybrids between R. fluitans and R. peltatus, R. aquatilis or R. trichophyllus. Within
R. penicillatus, three varieties — var. penicillatus, var. calcareus and var. vertumnus — are currently
recognized. Var. penicillatus differs from the other two varieties in having the capacity to form
* Unfortunately Dr Stace was unable to present his paper at the Conference and, at very short notice, Mr D. E.
Allen volunteered to contribute his paper on Rubus.
REPORTS 223
laminar leaves, whilst vars calcareus and vertumnus, which produce only capillary leaves,
intergrade morphologically. In addition, var. penicillatus is ecologically distinct, occupying base-
poor rivers in Ireland and western Britain, whereas both var. calcareus and var. vertumnus
generally occupy base-rich rivers and occasionally share a habitat. Var. penicillatus also appears to
have a different, if overlapping geographical distribution from the other two varieties, although
this difference may be determined largely by ecological preferences. These results suggest that var.
penicillatus merits separation from the other two varieties at subspecific rank, whilst both var.
calcareus and var. vertumnus should be retained within a second subspecies.
S. D. WEBSTER
RECENT ADVANCES IN THE STUDY OF TARAXACUM
Since the publication of The Taraxacum Flora of the British Isles (Richards 1972), considerable
inroads have been made into our understanding of the British status of this problematic genus. The
sectional taxonomy has recently been revised and this has been of considerable value to the
student. In addition, our knowledge of the Taraxacum flora has doubled the number of species
since that of Richards’ publication.
The workshop concentrated on the following aspects, supported by the use of appropriate
material:
—Hints on the collection, preservation, presentation and identification of dandelions.
—An outline of the ‘problem’ nature of the genus.
—Description and demonstration of the diagnostic features of the nine sections found in the British
Isles and some discussion of their inter-relationships.
—A review of our current understanding of the composition and distribution of the species known
in Britain.
C. C. HawortH & A. J. RICHARDS
MENTHA SPECIES AND HYBRIDS IN BRITAIN
In common with other perennial cultivated groups, the mints are difficult to treat taxonomically
and difficult to identify. Of the six species occurring in Britain, four are native in part of their
range. These are M. aquatica, M. arvensis, M. suaveolens and M. pulegium. The last, and also the
introduced Corsican M. requienii, do not normally cause identification problems. Most problems
are caused by M. spicata, the Garden Spearmint, belonging to sect. Mentha as do the first three
species mentioned above and among which hybrids occur frequently. The relationships of these
species were discussed and the importance of careful observations in the field and, if possible, in
cultivation, was underlined. The value of traditional characters for identification was considered,
together with others such as scent, epidermal hairs, sex and fertility. It was pointed out that a range
of characters should be assessed together when dealing with possible hybrids between genetically
very variable species, as often no hard-and-fast rules can be drawn up for their identification. As
with many critical, polymorphic groups, local variants are worth recognizing at least informally, as
they can provide evidence of means of vegetative spread and dispersal, and their history and origin
may sometimes be possible to unravel. In many cases these variants are impossible to treat in
formal taxonomic terms, when considered against the total variation encountered in the species or
hybrid as a whole.
R. M. HARLEY
DACTYLOID SAXIFRAGES
The Dactyloid Saxifrages of the British Isles are currently classified into four species, Saxifraga
cespitosa, S. rosacea, S. hartii and S. kypnoides. Recent research has both questioned and
224 REPORTS
confirmed certain aspects of this classification. An example of this is that cytological work has
shown the group to have a chromosome base number of x = 13. S. cespitosa is hexaploid (2n = 78)
and S. rosacea and S. hartii are tetraploid (2n = 52). S. hypnoides has been found to consist,
throughout its range (Iceland, Faeroes, British Isles, north-eastern France), of two geographically
separate chromosome races, a tetraploid (2n = 52) and a diploid (2n = 26). This total separation of
chromosome races suggests that they are biologically separate species. In proposing this taxonomic
change, a problem arises because the morphological differences between the two cytotypes are
slight. The only possib!e definitive characters are based on cell size (and chromosome number) and
thus a microscope needs to be used for identification. The most effective character was found to be
stomatal guard cell diameter. The demonstration illustrated the situation with S. hypnoides and
presented information on geographical distribution which is currently the best method of
separating the two cytotypes. S. rosacea and S. hartii, although having a consistent chromosome
number in the populations that have been studied, show characters that particularly question the
specific rank currently afforded S. hartii. This species was described from material collected on
Arranmore Island, Co. Donegal. Recent collections of plant material from this island proved to be
a further variant of S. rosacea, with some affinity to the variant found on Clare Island to the south.
It was concluded that S. hartii should be included within the variable S. rosacea but that fieldwork
was necessary on Arranmore Island in order to confirm this. The demonstration showed the
variability of S. rosacea, including the population from the Burren and Aran Islands, Co. Clare,
which has the most claim to be treated as a separate subspecies or species.
D. M. PARKER
ROSA
As a critical group of plants, roses are more akin to willows than say to brambles or hawkweeds.
The taxonomic solution to their diversity appears to lie in recognizing hybrids between a few
species rather than in characterizing many microspecies. This is often difficult and complicated by
the unique genetic performance of at least the British members of this genus during sexual
reproduction, a process which leaves the genetic factor of the female parent predominant.
Unfortunately, in the past, varietal and even species names have been allocated to many taxa
which we should now recognize as hybrids. Flora Europaea adds to the confusion, and the
treatment of roses in this work was compared with that of Wolley-Dod (A revision of the British
roses, in J. Bot., Lond., 1930-31, supplement, 1931), Clapham, Tutin & Warburg (Flora of the
British Isles, 2nd ed., 1962), and Melville (in Stace, C. A. (ed.), Hybridization and the flora of the
British Isles 1975). The main British species as well as some common hybrids were discussed and
characters of use in recognizing hybrids were pointed out. Recording from herbarium specimens is
often dubious. Field observation is paramount in the study of this genus, and fresh fruiting material
was used to illustrate the session.
G. G. GRAHAM
COMPUTER WORKSHOP
CARMARTHENSHIRE FLORA PROJECT, DOT-MAP PLOTTING PROGRAM
The use of a Sinclair Spectrum linked to a suitable dot-matrix printer to produce good quality
species distribution maps was demonstrated. The operator manually input each record to be
plotted and selected the appropriate map symbol to signify the particular status (e.g. post-1970,
native). Maps so constructed were saved to tape for subsequent recall and update and were of
sufficient quality to be ready for publication. The major disadvantage with the method was that
records needed to be abstracted manually from the master card-index before plotting, a very time
consuming business.
|
REPORTS 225
Future advances were demonstrated at informal evening sessions. Using a B.B.C. Model B
computer an innovative database has been developed which allows rapid and easy input by
displaying on the screen species lists derived from the Nature Conservancy Council’s Wales field
card. Plants recorded are selected by the cursor, a method which also allows status and habitat
information to be appended to each species. The economic use of the machine’s memory enables the
complete vice-county database of nearly 700 tetrads to be stored within the 1-3 megabytes available
on two double-sided floppy discs.
Interrogation of the database is very flexible. The present hardware takes about ten minutes to
search the whole and plot a distribution map for the selected species or combination of species.
Other enquiries can, for example, produce a list of species for a given tetrad or group of tetrads, sum
the number of species recorded in a tetrad, plot recorded habitats, list species by status, etc. The use
of hard disc or sideways RAM would significantly increase the speed of interrogation.
The system is adaptable to any county or area and to any floral or faunal group. A database to store
higher-plant data for the whole of Wales by 10-km-squares is already under development.
R. D. PrYcE
FLORA OF CUMBRIA DATABASE
A series of interactive computer programs (in FORTRAN 77) has been written to maintain and
utilize a database which contains species records catalogued by the tetrads in which the species
occurs. Program FILESPP is responsible for the input of new records into the database and has
facilities for checking, as far as possible, the accuracy of the data entered.
Program SELECTSPP interrogates the database to extract the following information:
i) Compile a list of tetrads in which a given species occurs and produce a distribution map.
ii) Compile a list of tetrads in which a given combination of species occurs, and produce a
distribution map for that combination.
ii) Compile a list of tetrads which have a total number of species lying between given limits, e.g.
tetrads with 150—200 species, and produce a distribution map.
iv) Compile a list of species in a given tetrad.
v) Compile a list of species in a given group of tetrads.
vi) Compile a list of species found in a given number of tetrads, e.g. a list of species in 10-15 tetrads.
vii) Compile a list of species-totals in each tetrad in the database. The maps produced are of
sufficient quality to be ready for publication.
A. J. C. MALLOCH
W. LANCASTER FLORA: USE OF A B.B.C. COMPUTER
A survey of the flora of West Lancaster (v.c.60) was started in 1964. At that time and for many years
afterwards the use of computers was not contemplated. As publication of a new Flora approaches,
the B.B.C. microcomputer has been utilized for capturing dot-map distributional data for onward
transmission and map production using the Flora of Cumbria Database at Lancaster University. The
B.B.C. microcomputer is also used as a word-processor for the production of text.
E. F. GREENWOOD, P. W. PHILLIPS & J. RAVEST
IDENTIFICATION BY COMPUTER
Computer programs for interactive identification on the BBC-B and TORCH microcomputers were
demonstrated. Examples were shown of sets of data relating to the critical genera Rubus,
Taraxacum and Euphrasia, as well as a set for grasses with vegetative characters only.
R. J. PANKHURST
226 REPORTS
THE NORTH WEST BIOLOGICAL FIELD DATA BANK’S CATALOGUING SYSTEM
The North West Biological Field Data Bank holds information on nearly 2,000 sites in north-
western England. The workshop demonstrated the use of the commercial programme dBase II to
construct the catalogue of these sites. On-line retrieval of the data was demonstrated and the use of
command files which allow the database to be used by the computerphobic person was explained.
Further expansion possibilities were described and some other commercially available database
programmes for micro-computers were also mentioned.
A. S. GUNN
P THE USES OF CRITICAL RECORDS
FLORA RECORDING AND THE NATIONAL VEGETATION CLASSIFICATION
The National Vegetation Classification is providing a systematic and comprehensive description of
all the vegetation in natural, semi-natural and major artificial habitats in Great Britain. From its
results it will be possible to produce a phytosociological profile for a large proportion of the British
vascular flora as well as for many bryophytes and lichens. It is hoped that the National Vegetation
Classification will encourage flora-recorders to set their observations in an ecological context and
serve as a framework for research on the conservation of individual taxa.
J. RODWELL
THE ROLE OF CRITICAL SPECIES RECORDING IN EVALUATING SITES OF SPECIAL SCIENTIFIC INTEREST
Biological Sites of Special Scientific Interest are usually notified on grounds of habitat or plant
communities for which the Nature Conservancy Council uses guidelines based on the concepts of
the Nature Conservation Review (1977): naturalness, diversity, size, rarity and fragility. This picks
up most good botanical sites, but there is also provision for S.S.S.I.s to be notified on grounds of
individual species or assemblages of species alone, both for animals and plants. For plants on
Schedule 8 of the Wildlife and Countryside Act 1981, that is ‘Red Data Book’ (British Red Data
Book 1. Vascular Plants, 2nd ed. (1983)) species which require special protection, the policy is to
notify all localities. For all other vascular plants, an assemblage score of 200 or more is used as a
guideline, which includes use of critical species. The remaining ‘Red Data Book’ species score 100,
species occuring in 16-30 10-km-squares in Atlas of the British Flora (1962) and Critical Supplement
to the Atlas of the British Flora (1968) score 50, 31-100 squares score 40. The Council is not bound
by the score and single species localities (including critical species) of particular note: British
endemics, species rapidly declining, those with fewer than six locations in Britain, and species at
the edge of their range receive special attention and may merit notification. The Council’s capacity
to use critical groups in evaluating S.S.S.I.s is limited by the information available about their
distribution. It would like to have better information to facilitate this process.
N. A. ROBINSON
CRITICAL SPECIES AND NATURE CONSERVATION
The objective of plant conservation in Britain must be the protection of our native taxa in their
original habitats. This is of particular importance for critical species groups or infraspecific taxa
where endemic variants have evolved since the separation of these islands from the Continent.
These taxa present several difficulties because they are recognized by a very small number of
specialists:
1. What are the taxa? Which specialists do we follow?
REPORTS 227
2. Where are the taxa? Who can collect distribution information over a short enough period for an
assessment of their status to be made?
3. What are the management requirements? Can site managers recognize the taxa and have
adequate knowledge to ensure their survival?
4. How can the Law be enforced? Should critical taxa be scheduled under the Wildlife and
Countryside Act 1981? Would ignorance of identity be a defence for picking a critical taxon?
5. In the light of these difficulties and the scientific importance of these taxa, decisions have to be
made on the number of sites for each we should be seeking to protect; we may find out, with the
help of experts, how many sites for each are already protected; and consider the role of botanic
gardens/universities in gathering and monitoring living collections of taxa within nominated
groups.
The relevance of collecting data for further editions of the Atlas of the British Flora (1962) and the
Critical Supplement to the Atlas of the British Flora (1968) and the monitoring of change in the
distribution of critical taxa must also be considered.
F. H. PERRING
FIELD EXCURSIONS TO NESS GARDENS AND RED ROCKS, 14TH SEPTEMBER 1985
The field excursion on Saturday, 14th September visited two quite different areas. In the morning,
Dr H. McAllister, Deputy Director of the University of Liverpool’s Botanic Gardens at Ness,
South Wirral, Cheshire welcomed delegates and took them on a guided tour of the Gardens. There
was far too much for delegates to see on one trip and certain features were selected for special
comment. These included the national collection of willows recently transferred from Long Ashton
Research Station, Bristol, taxonomic studies on Orobanche, and the native plant garden where a
number of endangered British species are grown.
In the afternoon, delegates drove a few miles north-westwards to the coast of the Wirral
peninsula at West Kirby. Here the Red Rocks S.S.S.I., managed as a nature reserve by the
Cheshire Conservation Trust, was visited. It is a small area of sand dunes with a brackish slack and
reedbed, and developing saltmarshes adjoining the site. In continuing the critical groups theme of
the conference, Atriplex littoralis, A. prostrata, A. glabriuscula and A. laciniata were distinguished.
In a new developing slack Juncus ranarius was noted and distinguished from J. bufonius sensu
stricto growing further inland. A feature of Red Rocks is the presence of Equisetum X trachyodon,
and considerable discussion took place when distinguishing this hybrid from E. variegatum, which
it was thought was growing as a robust upright variant in a nearby dune slack. The other parent, E.
hyemale, formerly occurred here but has not been seen for many years. The meeting finally ended
after a developing Salicornia marsh was examined, where S. ramosissima, S. fragilis and S.
dolichostachya were identified.
E. F. GREENWoop & H. MCALLISTER
EXHIBITION MEETING, 1985
The Annual Exhibition Meeting was held in the Department of Botany, British Museum (Natural
History), London, on Saturday, 30th November 1985, from 12.00 to 17.30 hours. The following
exhibits were shown.
SOME ADDITIONS TO THE FLORA OF EUROPE
The exhibit illustrated some of the new information that is being assembled for the revised edition
of Flora Europaea Volume 1, currently being prepared at Reading University. Herbarium material
‘and maps were displayed of taxa that have been reported as native to Europe, or
228 REPORTS
naturalized aliens, since the publication of Flora Europaea Volume 1 in 1964. The examples shown
were:
—Arenaria pomeli Munby, an annual species formerly thought to be endemic to Morocco, but now
known from southern Spain;
—Chenopodium pumilio R.Br., an alien from Australia that is naturalized in western and central
Europe;
—Crassula alata (Viv.) Berger, similar to C. tillaea, recently found on Crete and in the Cyclades,
which has its main distribution in N. Africa and the Middle East;
—Polygonum lanigerum R. Br., a species of the Old World Tropics that extends to the Middle East,
reported in southern Crete;
—Rumex vesicarius L., an annual species that has been refound at its only European station at
Nafplion in southern Greece;
—Tellima grandiflora (Pursh) Douglas ex Lindley and Tolmeia menziesii (Pursh) Torrey &
A. Gray, saxifragaceous species from Pacific N. America, which are becoming established in shady
places in the British Isles.
J. R. AKEROYD
SOME RECENT RUBUS DISCOVERIES IN HAMPSHIRE q
By 1983, 115 Rubus species were reliably on record from v.c. 11 and/or v.c. 12. In the following
year, five more species were added, their Hampshire occurrences representing in each case
startling extensions of their known British ranges.
—-R. hastiformis W. C. R. Wats., locally abundant in Cornwall and Devon, with rare outliers in
Glamorgan, Cardiganshire and Sussex;
—R. leptothyrsus G. Br. (R. danicus (Focke) Focke), a common Scottish species, not previously
known south of Suffolk and Herefordshire;
—-R. norvicensis A. L. Bull & Edees, known since 1972 in abundance in woods and hedges around
Romsey, extending to Winchester and Southampton, and in copses on Hayling Island. In 1984, a
chance look at the BM holotype of this recently described species of the Norwich and Ipswich
districts suggested the two might be identical, a hypothesis confirmed by A. L. Bull;
—R. tumulorum Rilst. (not exhibited), formerly believed to be confined to the Looe district of
south-eastern Cornwall and one or two places in Devon;
—R. winteri P. J. Muell. ex Focke, believed to be restricted to the central and east Midlands of
England. In 1984-85 it was found in copses on chalk in three widely separated parts of Hampshire,
one of them by the border with S. Wilts., v.c. 8, into which it probably continues.
Also exhibited was the holotype (BM) of R. hantonensis D. E. Allen, described in Watsonia, 15:
387 (1985), so far known only in Hampshire and the Isle of Wight.
D. E. ALLEN
DISTINGUISHING BETWEEN BETULA PENDULA, B. PUBESCENS AND THEIR HYBRIDS
Betula pendula Roth (2n=28) and B. pubescens Ehrh. (2n=56) can be separated reliably using |
three leaf characters: leaf tip width, ‘leaf tooth factor’, and distance to the first tooth on the leaf
base. This identification is carried out by using a discriminant function. Hybrids are identified as B.
pubescens by this method. Artificial hybrids can also be distinguished from both parent species by
using a pair of discriminant functions involving nine leaf characters. Stomatal length varies
between B. pendula, B. pubescens and the hybrids, those of B. pendula being the smallest and
those of the hybrids being intermediate.
M. D. ATKINSON & A. N. CODLING |
REPORTS 229
SEED-SET IN BRITISH REYNOUTRIA POPULATIONS
Members of the rhizomatous, perennial genus Reynoutria, giant knotweeds, are now established
members of the flora of the British Isles. As part of a study carrying on from A. P. Connolly’s work
on the genus, an examination of the reproductive biology of these plants has been undertaken.
Reynoutria japonica was found to have 88 chromosomes, and R. japonica var. compactum and R.
sachalinensis 44 chromosomes. Two groups of hybrids were found to have 44 and 66 chromosomes.
In order to examine the role played by seed set, seeds and parental rhizomes were collected from a
number of localities in the British Isles, and chromosome counts performed on the plants that were
raised. A most surprising result is that the majority of seedlings from parents with 88 chromosomes
had 54 chromosomes. These plants are thought to be hybrids with Fallopia aubertii, Russian Vine
(2n=20), as the male parent.
J. P. BAILEY
X ORCHIACERAS BERGONII (NANTEUIL) CAMUS: A BIGENERIC HYBRID NEW TO BRITAIN
Bigeneric hybrids between Aceras anthropophorum (L.) R. Brown and species of the genus Orchis
subgenus Militares are often recorded on the Continent, especially x Orchiaceras bergonii (Nan-
teuil) Camus (A. anthropophorum xX Orchis simia Lamarck). The first British specimen of this
hybrid (also the first involving Aceras) was discovered during 1985 in Kent, at the only British locality
where its parental species co-exist. Its identity was confirmed by detailed morphometric comparison
with its parents, using multivariate analyses, but its origin remains speculative. Photographs and
sketches of the hybrid and its parents were exhibited, and implications of the morphometric study
were outlined.
R. M. BATEMAN & O. S. FARRINGTON
BRITISH PLANTS AS ALIENS IN MALAYSIA
Specimens were shown of a number of plants from peninsular Malaysia which also occur in Britain.
Most of these are weedy species of disturbed soils above 100 m altitude. The following 29 species
were noted in 1976 and/or 1985; only those marked * are mentioned in current Floras and the rest
may be new records: Alopecurus aequalis, Artemisia vulgaris, Briza minor, Cardamine hirsuta, C.
flexuosa, Cerastium fontanum, C. glomeratum, Chrysanthemum segetum, *Cynodon dactylon,
?Fragaria vesca, Galinsoga parviflora, *Gnaphalium luteo-album, Hypochoeris radicata, Juncus
bulbosus, *Oxalis corniculata, *Plantago major, *Poa annua, *Pteridium aquilinum, Raphanus
sativus, Rumex crispus, Sagina procumbens, Sambucus nigra, * Setaria geniculata, *Solanum nigrum,
Sonchus oleraceus, *Stellaria media, Trifolium dubium, ?Vulpia bromoides, V. myuros.
H. J. M. BOWEN
HAMPSHIRE HIGHLIGHTS, 1985
This exhibit reported notable records within the county during 1985 of flowering plants & ferns. It
was illustrated with herbarium material kindly loaned by the British Museum (Natural History) & A.
R. G. Mundell, and with specimens provided by the exhibitors.
Plants reported from the geographical county for the first time (or which had apparently become
extinct from all known sites) were Allium oleraceum L., Berberis aggregata Schneid., Carex elongata
L., Cotoneaster dielsianus Pritz., Cotoneaster frigidus Wall., Cotoneaster perpusillus (Schneid.)
Flinck & Hylm6, Cotoneaster salicifolius Franch., Cotoneaster X watereri Exell, Papaver dubium L.
xX P. rhoeas L., Potamogeton coloratus Hornem., Ruscus hypoglossum L., Salvia reflexa Hornem.,
Setaria pumila (Poiret) Schultes. and Sisyrinchium iridifolium subsp. valdiviense.
230 REPORTS
New vice-county records were Equisetum hyemale L. (v.c. 12), Phacelia tanacetifolia Bentham
(v.c. 11), and Typha x glauca Godr. (v.c. 12). Other rare plants that were seen to thrive in the county
during 1985 were Briza minor L., Gastridium ventricosum (Gouan) Schinz & Thell., Lycopodiella —
inundata (L.) Holub, Polypogon monspeliensis (L.) Desf. and Himantoglossum hircinum (L.)
Sprengel (seen for the first time since 1948).
A. Brewis, J. D. Fryer, B. A. GALE & F. RosE
SOME MORE BRITISH SPECIES FROM THE WESTERN HIMALAYA
Herbarium sheets of the following species, found both in Britain and the western Himalaya, were
displayed: ;
Campanula latifolia (Ludlow & Sheriff, 1940), widespread in Pakistan and Kashmir, 2700-3700 m;
Lotus corniculatus (Polunin, 1956), widespread, 1200-3200 m; Dactylis glomerata (Polunin, 1956),
very common and variable, from Chitral eastwards, 2-400 m; Veronica beccabunga (Robson, 1971),
found at springs and stream margins, 1800-3500 m; Achillea millefolium (Polunin, 1956), very
common in the temperate and lower alpine zones, 1600-4000 m; Oxyria digyna (Southampton
University Botanical Expedition, 1981), very common in wet soil at high altitudes, 2700-5400 m;
Hippuris vulgaris (Zino, 1978), cool waters of the northern hemisphere, 1200-4500 m; Adoxa
moschatellina (Duthie, 1892), forest humus, 2700-3500 m; Convolvulus arvensis (Southampton
University Botanical Expedition, 1981), a common weed of fields.
C. CHADWELL
THE HAY-MEADOWS OF THE PICOS DE EUROPA, ASTURIAS, SPAIN
The Picos de Europa in northern Spain contain a large number of species-rich hay meadows which
are managed in a traditional manner. In May 1985 the students and staff of the Conservation Course
M.Sc. at University College London visited the area to study the meadows. The rich plant
communities were recorded and the specific management carried out to produce the meadow flora
was determined from the local farmers. Spain’s entry to the E.E.C. in 1986 may cause the
management of these meadows to be altered. Four scenarios were proposed for the future of the
meadows. Some of the options available which would lead to the conservation of the haymeadows
were discussed. The text of the exhibit was a condensed version of ‘““The Hay Meadows of the Picos
de Europa’’, Discussion Paper in Conservation, no. 43, Conservation Course, 1985, U.C.L.,
augmented by photographs and herbarium specimens.
A. COLSTON
THE GENUS OROBANCHE IN THE BRITISH ISLES
The broomrape genus Orobanche is represented in the British Isles by a number of local or rare
species with very restricted host ranges, and by a more frequent species, O. minor Sm., that
parasitizes several dicotyledonous hosts of diverse phylogenetic origin. However, due to the lack of
precise morphometric data, the characters that best separate Orobanche species are still
inadequately defined; as a result the specific identity of some broomrape populations parasitizing
Daucus carota, Picris hieracioides and Eryngium maritimum in southern England and Wales is
presently in doubt. This photographic exhibit depicted the native Orobanche taxa recorded from the
British Isles, summarized their known distributions and host ranges in the country, and served to
highlight existing areas of controversy within this taxonomically difficult group. Additional records
of broomrapes on unusual hosts or outside their accepted geographical ranges were requested.
I. DENHOLM & M. JONES
REPORTS 231
QUINQUENNIAL REVIEW 1985
During 1985 a field survey of most of the 62 species scheduled under the Wildlife and Countryside
Act 1981 has been carried out. Many other species were considered on request from B.S.B.I.
members, N.C.C. staff and other botanists. A list of the species researched, and the criteria used
was displayed, together with the recommendations to be given to N.C.C. Council and then to the
Secretary of State. It was proposed to add 15 species and to delete one, as it has become extinct.
Members were asked for their comments. Distribution maps showing the decline of Mentha
pulegium, Pulicaria vulgaris and. Teucrium botrys were displayed along with colour pictures of
many of the species surveyed.
L. FARRELL & P. WILSON
TOLYPELLA PROLIFERA (ZIZ & BRAUN) LEONH. AT AMBERLEY WILD BROOKS, WEST SUSSEX (v.c. 13)
On 19th August 1985, while assisting Sussex botanists who record regularly at Amberley, I found
Tolypella prolifera growing with Groenlandia densa in a ditch on the south side of the brooks. This
species has been recorded from recently cleared ditches and ponds in about a dozen vice-counties,
notably Cambridge, Huntingdon, Lincoln and Northants. Always considered rare, its large size
and spectacular appearancé were apparent in the photographs shown. Specimens from Amberley
and other parts of the Arun Valley (BM) show that the plant has been recorded in this area since
1900 but, like other stoneworts, its appearances are spasmodic and unpredictable. Most stoneworts
require unpolluted water and the periods of low competition provided by regular ditch cleaning
management, so it is particularly pleasing to find this interesting plant at a time when so much
wetland habitat is being lost or degraded.
R. FirzGERALD
THE ECOLOGY OF ABNEY PARK CEMETERY, HACKNEY, AND ITS EDUCATIONAL POTENTIAL
Cemeteries in urban areas provide a unique habitat which is extremely valuable both for wildlife
and as an educational resource. The 9 ha of Abney Park Cemetery are the only woodland in
the London Borough of Hackney. The Cemetery was designed by William Hosking in 1840, and
under the direction of George Loddiges developed into one of Europe’s most varied arboretums.
At present densely overgrown in places, it is in need of management, and the Borough Landscape
Architect invited the Urban Spaces Scheme (Polytechnic of North London) to advise on the
enhancement of the area’s ecological and educational potential. The exhibit showed the early
botanical history, current floristic composition, plans for future management and conservation, the
geology of the gravestones, an Interpretive Guide, and the cemetery’s grasses. A computerized
“Town Grass Key” has been developed for children on a BBC microcomputer.
M. HALE
ECOLOGICAL SURVEY OF HAM RIVERSIDE LANDS, RICHMOND
This area of some 74 ha has long been of botanical interest, with a diversity of habitats
including river bank, water meadows, gravel excavations and dumps. It is now a metropolitan open
space within Richmond-upon-Thames, and various individuals and organizations are co-operating
with the Parks Department to manage it as a recreational ‘mini-countryside’ with an important
natural history content. The exhibit showed a series of photographs taken during the past 23 years
and a recent detailed ecological study undertaken by students of the School of Horticulture at the
Royal Botanic Gardens, Kew.
F. N. HEPPER
232 REPORTS
GURREHOLM BJERGE BOTANICAL EXPEDITION, 1985
The Gurreholm Bjerge Botanical Expedition was a joint British and Irish four-man venture which
took place during July and August 1985. The aim of the expedition was to carry out floristic surveys
of a number of previously unbotanized valleys and mountain ranges in the @rsted Dal region of
north-eastern Greenland. The flora of this region has comparatively few species (a total of 143
were recorded by the group). However those which do occur are of great interest, particularly as
many of the common species (e.g. Saxifraga oppositifolia L. and Betula nana L.) are local alpine
plants in the British Isles.
The exhibit illustrated a range of plants seen during the course of the expedition. Amongst the
more notable finds were Potentilla stipularis L., an Asiatic species only found rarely in north-
eastern Greenland, the sedge hybrid Carex parallela (Laest.) Sommerf. x C. lachenalii Schkuhr..,
and Saxifraga nathorstii (Dusén) Hayek which is endemic to this area of Greenland.
J. HotpicH & D. A. Simpson
STAGES IN THE LIFE CYCLE OF OROBANCHE CRENATA FORSSKAL
The genus Orobanche consists of some 150 or so species of obligate parasites. Much of their
development takes place underground since their site of attachment is through the host root, and it
is only when they flower (which for perennial species such as Orobanche rapum-genistae L. may be
several years after seed germination) that they become apparent. For this reason the underground
development of British broomrapes has received little attention from most botanists. The exhibit
demonstrated the stages in the life cycle of a member of the genus Orobanche as shown by O.
crenata. The stages were shown using preserved specimens, diagrams, and photographs. The role
of O. crenata as an agricultural pest was also discussed.
M. JONES
DIOECY AND POLLINATION IN VISCUM ALBUM
Viscum album L. is dioecious, with marked differences in size and appearance between male and
female flowers. Female plants have been found to predominate, with approximately twice as many
females as males among flowering plants both in wild populations and among plants raised from
seed.
Since 1762, when Koelreuter suggested that V. album was insect-pollinated, opinions have
differed but there has been little evidence about the mode of pollination of its inconspicuous
flowers. Recent workers have stated that it is wind-pollinated. However, observations in
Herefordshire, where V. album is abundant, during March 1985 showed that both male and female
flowers were visited by large numbers of Diptera, especially the common greenbottle, Dasyphora
cyanella, apparently seeking nectar in female flowers (in which 0-02-0-04 ul of nectar with 48%-
55% s.e. concentration was found) but seeking pollen in the nectarless male flowers. Flies caught
on female flowers up to 6 m from the nearest male plant carried many V. album pollen grains. It
was clear that the flowers of V. album are well adapted for insect pollination, which was fully
adequate in the study area.
Qv Ou Nv Kay
HAWKWEEDS IN SOUTHERN SCOTLAND
Recent fieldwork has added considerably to the number of records of Hieracium species from
southern Scotland (v.cc. 72-84), and 74 species have been recorded from the region on good
authority. Most species are rare or local, although some are widespread and locally common. The
|
|
1
REPORTS 233
usual habitat for the majority of species is cliffs and rocks, often by streams. Others may be
found in grassland or on the edges of woods and a few in urban situations or on waste ground. It
is likely that hawkweeds occur more widely in the region than is suggested by currently available
records. Extra records from v.cc. 74-77 and 81-84, in particular, would go far towards producing
a comprehensive picture of their distribution in southern Scotland.
D. J. McCosuH
CONSERVATION SECTION, CAMBRIDGE UNIVERSITY BOTANIC GARDEN
The Nature Conservancy Council funds the propagation, display, seed-storage and autecological
study of rare British plants in the Garden. Low temperature and moisture content provide seed-
storage conditions under which viability can be maintained for years. A list of the contents of the
seed-bank is available from the Conservation Propagator. As our populations are under such
pressure, only those people specifically requested to do so by the N.C.C. should collect seed of
wild Red Data Book species. Rare plants in cultivation can be used to inform visitors about
conservation and the autecology of these species, and provide quality material for research
scientists or re-introduction. These functions require rigorously careful documentation and labell-
ing. Exhibited were notes and slides, also live and herbarium specimens of some of the species
forming the Conservation Collection. Annual changes in the underground morphology of
Himantoglossum hircinum, Lizard Orchid, were also illustrated.
V.M. MorGAn
THE EAST HAMPSHIRE HANGERS PROJECT
Sponsored by the Countryside Commission, E. Hants. District and Hampshire County Councils,
the Project aims to bring back into positive management the 680 ha of Hanger woodlands on
the Chalk and Greensand scarps between Petersfield and Alton. Most of these are ancient
woodlands, half are S.S.S.I.s.
Due to loss of traditional markets, the inaccessibility of the steep slopes, age and disease, the
Hanger structure and landscape is deteriorating. The Project Officer aims to persuade owners to
join management schemes, and he will organize grants and help to market produce. Because of
the Hangers’ unique and diverse biological richness, all are being surveyed initially; small-scale
felling will be followed by recoppicing and natural regeneration to maintain stand-types and
genetic stock, planting when necessary with native species. Visitors to the Project are very
welcome: more information is available from the Project Officer, whose telephone number is
Petersfield 66551.
J. OCKENDEN
THE GUERNSEY BAILIWICK, 1985
Herbarium sheets were shown of the more important finds of the year, viz: GUERNSEY: Davallia
canariense — new to the Channel Islands (C.I.); Ononis cf. alopecuroides — new to the C.I.;
Symphytum ‘Hidcote Blue’ — new to the C.I.; Juncus bulbosus f. viviparus — first find since the
18th century; Bromus ramosus — new to the C.I.; Brachypodium pinnatum — in two separate
places, known otherwise in the C.I. only from Alderney; Apera spica-venta — last seen 1958.
SARK: Myosoton aquaticum —again as a garden weed; Stellaria graminea; Fragaria vesca — unac-
countably rare in the bailiwick; Mentha xX villosa ‘Alopecuroides’; Luzula forsteri; Poa angustifo-
lia; Phleum pratense. HERM: Conyza sumatrensis — new to the Herm archipelago; Chamaemelum
nobile — first find for 148 years; Allium subhirsutum — known otherwise in the C.I. only in Sark;
Alopecurus pratensis. JETHOU: Portulaca oleracea; Phytolacca cf. polyandra — new to the C.L.;
234 REPORTS
Polygonum persicaria; Agrostis stolonifera — first record for 95 years; Vulpia bromoides — ditto;
Alopecurus myosuroides — new to the Herm archipelago.
Also shown were sheets of Silene vulgaris subsp. maritima with double flowers, found at
Jerbourg in 1984; and of this plant with a pallid calyx collected in 1910. Fuller accounts of both,
with names for them will appear in the Transactions of La Société Guernésiaise for 1985. In this
there will also be the first Supplement to the “Wild Flowers of Guernsey”’, and the new Check-List
for Herm, both available separately.
J. PAGE
ORIGINAL DRAWINGS FOR FRYER’S MONOGRAPH OF POTAMOGETON
Alfred Fryer began to study Potamogeton during the 1880s. He became an acknowledged authority
on the genus, and the first parts of his monograph The Potamogetons (Pond Weeds) of the British
Isles were published in 1898. He died before finishing the work, which was completed by A.
Bennett. Fryer’s section of the monograph was illustrated by Robert Morgan (1863-1900).
Morgan’s colour plates delighted Fryer and have been praised by later critics. Eight of his original
drawings were exhibited, by kind permission of the British Museum (Natural History).
C. D. PRESTON
THE CARMARTHENSHIRE FLORA DATABASE: A COMPUTER DATA STORAGE AND RETRIEVAL SYSTEM
ADAPTABLE FOR ANY HOME-BASED VICE-COUNTY RECORDER
Software to run on the B.B.C. Model B computer, written especially for vice-county recorders,
was demonstrated. The database as set up for the Carmathenshire Flora Project will store the
records for each of the 698 tetrads on only two floppy discs. The intended primary use of the
system is as a tool to enable fast interrogation of the floral data and not to entirely replace the
master card index. When inputting records, the program displays a series of screens of species lists,
based on the Biological Record Centre’s field card. Plants recorded are selected with the cursor
and habitat and status information is appended to each. Interrogation of the database enables a
variety of parameters to be extracted including, for example, the compilations of a list and plotting
of a distribution map of tetrads in which a selected species or group of species occurs by date, status
or habitat.
S. RHopEs & R. D. PRYCE
CAPSELLA AND CARDARIA IN THE BRITISH ISLES
In a recent study of Capsella in Greece, Svensson (Willdenowia 13: 267, 1983) showed that no
morphological or cytological character could be used to distinguish C. rubella Reuter from C.
bursa-pastoris (L.) Medicus. He concluded that C. rubella did not merit specific status and that it
should be included within C. bursa-pastoris sensu lato. A less detailed survey of British Capsella
shows many of the same features of variation observed in Greece. Specimens showing this
variation were exhibited and the characters traditionally used to separate the species were listed
and discussed.
Two subspecies of Cardaria draba (L.) Desv. have been introduced to the British Isles.
Subspecies chalepensis (L.) D. E. Schulz is much rarer than subsp. draba and has been recorded
from v.cc. 2, 39 and 70 (J. E. Lousley in Proc. bot. Soc. Br. Isl. 1: 577, 1955). It is distinguished by
its ovate siliculae which have cuneate to narrowly rounded bases. Specimens of the two subspecies
were exhibited, and details of further sites and live or dried material were requested.
T, GoGo Rich
REPORTS 235
ISOZYME VARIATION IN SEXUAL AND ASEXUAL TARAXACUM
In northern Europe, all dandelions are asexual, being polyploid agamosperms with diplosporous
parthenogenesis. In southern Europe, many dandelions are self-incompatible, diploid, sexual
species. It is to be expected that sexuals are more genetically variable than asexuals, and possibly
able to fill more environmental niches. This hypothesis was tested for 16 loci in eleven enzyme
systems for an asexual population of 7. pseudohamatum from Newcastle-upon-Tyne, and a sexual
population, probably referable to T. alacre, from the Auvergne, France. The Mendelian inheritance
of all varying loci was tested by experimental crosses in the sexual population.
In the sexual population, eight loci were polymorphic (50%), but the mean level of heterozygosity
at each locus was only 12%. In the asexual population, only one locus was polymorphic (6%), but the
mean level of heterozygosity at each locus was 43%. At five of the loci which were polymorphic in the
sexual, only invariable fixed heterozygotes were found in the asexual population. One monomorphic
locus in the sexual was also heterozygously fixed in the asexual. It is concluded that theoretical
predictions of low genetic variability but high levels of heterozygous fixation (which might result in
vigour) in asexuals are confirmed, at least in these populations.
A. J. RICHARDS & J. HUGHES
SPIRANTHES SPIRALIS (L.) CHEVAL. — FROM SEED TO FLOWERING PLANT IN 5 YEARS
The development of terrestrial orchids from seed to a flowering plant is generally considered to be a
lengthy process, involving a long period as an underground protocorm. Summerhayes (1951) states
that the first leaf is produced after 11 years in Spiranthes spiralis, with another 2-3 years before the
plant flowers. In asymbiotic culture, we produced plants with leaves within 6 months after sowing,
with a single plant producing an inflorescence within 5 years after transfer to a non-sterile shell-sand
soil. While we accept that conditions in a culture tube are different from those encountered in the
field, the exhibit suggested that the times for development from seed to flowering plant quoted in
most standard works on orchids are excessively long and require re-assessment.
T. C. E. WELLS & R. KRETZ
The following also exhibited:
BOTANY LIBRARY, BRITISH MUSEUM (NATURAL HISTORY). (a) Frank Kingdon-Ward, b. 6th November
1885; a centenary exhibition. (b) History of Botanic Gardens.
D. BurNHAM. Mutation in orchids.
A. J. ByFIELD. Nitella mucronata var. gracillima J. Groves & Bullock-Webster, a rare charophyte
from Co. Avon.
J. H. FREMLIN. Stereophotographs of some Irish plants, including Minuartia recurva.
A. N. Gipsy. Botanical postage stamps.
A. C. JERMy, J. M. Camus & A. M. Paut. Pilularia in the British Isles.
A. C. JeRMy & A. G. KENNETH. Diphasiastrum X issleri (Rouy) J. Holub in Scotland.
V. A. JOHNSTONE. Photographs of wild flowers of southern England.
L. M. KArLey. (a) Help! (b) Galls.
J. M. MuLiin & A. MELpeERts. Polypogon pumilus Clarke in Britain.
. PARKER. The dactyloid saxifrages of the British Isles.
. F. & M. E. Procror. Scanning electron micrographs of Carex leaf-surfaces.
. & M. C. F. Proctor. Fruit characters of Devon Sorbus species.
M. SAUNDERS & ST CHRISTOPHER’S SCHOOL, BURNHAM-ON-SEA. Operation Orchid.
Sisson. A selection of botanical illustrations.
R. StincsBy. The Keen of Hamar — a Shetland National Nature Reserve in its North Atlantic
ntext.
F. Stewart & S. J. LEACH. Fife’s disappearing pondweeds.
F. Stewart & H. E. Stace. Interesting records from West Perthshire.
M
c
E
e)}
a)
S.
D
M
M
c.
P.
D
c
N.
N.
236 REPORTS
O. M. Stewart. (a) Aira armoricana Albers. (b) Senecio squalidus L. x S. sylvaticus L. on
Arthur’s Seat? (c) Senecio x cardinensis J. E. Lousley and Vicia tetrasperma (L.) Schreb. in v.c.
73. (d) Flower paintings.
H. S. THompson. Works in process.
J. Tusss. Flora by a photographic botanist.
In the lecture-hall, the following members gave short talks illustrated by colour slides:
P. J. Horton. B.S.B.I. visit to Hungary.
L. FARRELL. Farrell’s travels in Scotland.
J. D. Hooper. Some autumn flowers in Mallorca.
E. MILNE-REDHEAD. A Black Poplar site of special biological interest.
P. F. Yeo. British Euphrasia species.
M. Jones. Coco-de-Mer, the world’s largest seed.
N3
N2
Vice-
counties
1
N
f.
W. Cornwall
1b. Scilly
E. Cornwall
S. Devon
N. Devon
S. Somerset
N. Somerset
. N. Wilts.
. S. Wilts.
. Dorset
. Wight
. Su Hants. F
i Ne Baats=
. W. Sussex
=. SUSSER:
. E. Kent
. W. Kent
. Surrey
_ is. BSsex
. N. Essex
. Herts.
. Middlesex
. Berks.
. Oxon
. Bucks.
. E. Suffolk
. W. Suffolk
. E. Norfolk
. W. Norfolk
. Cambs.
. Beds.
. Hunts.
. Northants.
. E. Gloucs.
. W. Gloucs.
. Mons.
. Herefs.
. Worcs.
. Warks.
S. Kerry
N. Kerry
W. Cork
Mid Cork
E. Cork
Co. Waterford
S. Tipperary
Co. Limerick
Co. Clare
. N. Tipperary
. Co. Kilkenny
. Co. Wexford
Co. Carlow
H14. Laois
39. Staffs.
40. Salop
41. Glam.
42. Brecs.
43. Rads.
44. Carms.
45. Pembs.
46. Cards.
47. Monts.
48. Merioneth
49. Caerns.
50. Denbs.
51. Flints.
52. Anglesey
53.. S. Lines:
54. N. Lincs.
35. CIES.
55b. Rutland
56. Notts.
57. Derbys.
58. Cheshire
59-°S; Lanes:
60. W. Lancs.
61. S.E: Yorke:
62.. N-E>Y orks.
63. S:W>Yarks:.
64. Mid-W. Yorks.
65. N.W. Yorks.
66. Co. Durham
67. S. Northumb.
68. Cheviot
69. Westmorland
69b. Furness
70. Cumberland
71. Man
72. Dumfriess.
73. Kirkcudbrights.
74. Wigtowns.
75. Ayrs.
H15.
H16.
Hid 7:
H18.
tbo.
H20.
H21.
H22.
H23.
H24.
HZ):
H26.
5 FW
H28.
IRELAND
S.E. Galway
W. Galway
N.E. Galway
Offaly
Co. Kildare
Co. Wicklow
Co. Dublin
Meath
Westmeath
Co. Longford
Co. Roscommon
E. Mayo
W. Mayo
Co. Sligo
NAMES OF VICE-COUNTIES IN WATSONIA
ENGLAND, WALES AND SCOTLAND
. Renfrews.
. Lanarks.
. Peebless.
. Selkirks.
. Roxburghs.
. Berwicks.
. E. Lothian
. Midlothian
. W. Lothian
. Fife
. Sturlings.
. W. Perth
. Mid Perth
. E. Perth
. Angus
. Kincardines.
. S. Aberdeen
. N. Aberdeen
. Banffs.
. Moray
. Easterness >
96b. Nairns.
H29.
H30.
H31.
HZ,
H33.
H34.
H35.
H36.
Hy
H38.
H 39.
H40.
. Westerness
. Main Argyll
. Dunbarton
. Clyde Is.
. Kintyre
. S. Ebudes
. Mid Ebudes
. N. Ebudes
. W. Ross
. E. Ross
. E. Sutherland
. W. Sutherland
. Caithness
. Outer Hebrides
. Orkney
. Shetland
Co. Leitrim
Co. Cavan
Co. Louth
Co. Monaghan
Fermanagh
E. Donegal
W. Donegal
Tyrone
Co. Armagh
Co. Down
Co. Antrim
Co. Londonderry
Botanical Society of the British Isles
BSBI HANDBOOKS FOR FIELD IDENTIFICATION
Salix lanata Each handbook deals in depth with one
of the more difficult groups of British 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
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No. 2 UMBELLIFERS OF THE BRITISH ISLES
T. G. Tutin. 1980. 197 pages, fully illustrated with line
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So
No. 3 DOCKS AND KNOTWEEDS OF THE BRITISH
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J. E. Lousley and D. H. Kent. 1981. 205 pages, with many
line drawings of British native and alien taxa. Wrappers.
ISBN 0 901158 04 6 £5575
No. 4 WILLOWS AND POPLARS OF GREAT BRITAIN
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R. D. Meikle. 1984. 198 pages with 63 line drawings of
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and hybrids. Wrappers. ISBN 0 901158 07 0 £6.75
No. 5 CHAROPHYTES OF GREAT BRITAIN AND
IRELAND
J. A. Moore. 1986. 144 pages with line drawings and
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Ed. A. J. Wilmott, 1948. 104 pages, 18 plates. Wrappers. £4.25
2. THE STUDY OF THE DISTRIBUTION OF BRITISH PLANTS
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Ed. J. E. Lousley, 1955. 189 pages, 2 plates and 23 text figs. £5.25
5. PROGRESS IN THE STUDY OF THE BRITISH FLORA
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16. THE POLLINATION OF FLOWERS BY INSECTS
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17. THE BIOLOGICAL ASPECTS OF RARE PLANT CONSERVATION
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18. PLANT LORE STUDIES
Ed. R. Vickery, 1984. 260 pages. Proceedings of a joint conference of BSBI and Folklore Society in
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19. ARCHAEOLOGY AND THE FLORA OF THE BRITISH ISLES
Ed. Martin Jones, 1986. Proceedings of a joint conference held in 1984.
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ENGLISH NAMES OF WILD FLOWERS
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INSTRUCTIONS TO CONTRIBUTORS
Papers and Short Notes concerning the systematics and distribution of British and European
vascular plants as well as topics of a more general character are invited.
Manuscripts must be submitted in duplicate, typewritten on one side of the paper only, with wide
margins and double-spaced throughout. They should follow recent issues of Watsonia in all matters
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Tables, appendices and captions to figures should be typed on separate sheets and attached at the
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drawings should be in Indian ink on good quality white card, blue-lined graph paper or tracing paper.
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Contributors are strongly advised to consult the editors before submission in any cases of doubt. 25
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Papers and Short Notes should be sent to Dr R. J. Gornall, Botany Department, The University,
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CAMBRIDGE
The Names of Plants
D. GLEDHILL
Mountain Flora of Greece
Volume 1 ARNE STRID
This, the first of two volumes, is an up-to-date and
This book will prove fascinating reading both for
those interested in translating the scientific
names of plants into everyday English, and those
who wish to learn how and why the
nomenclature has changed over the years.
159 pp. 1985 0521 305497 £20.00 net
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A Dictionary of Flowering
Plants and Ferns
Eighth Edition The Late J.C. WILLIS
Revised by H.K. AIRY SHAW
This famous publication is a comprehensive and
indispensable reference to the generic and family
names of flowering plants and ferns.
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The European
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The first in a series of six, this volume contains
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and 16 families of Monocotyledons including
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c.430 pp. 1986 0521 248590 £55.00 net
For further details please write to Annette Duggan at the address below.
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WATSONIA
_ GRAY HERBARIUM
* MAR 3 0 1987
Botanical Society of the British Isles
Patron: Her Majesty Queen Elizabeth the Queen Mother
Applications for membership should be addressed to the Hon. General Secretary,
c/o Department of Botany, British Museum (Natural History), Cromwell Road, London,
SW7 SBD, frem whom copies of the Society’s Prospectus may be obtained.
Officers for 1986-87
Elected at the Annual General Meeting, 10th May 1986
President, Mr D. E. Allen
Vice-Presidents, Dr N. K. B. Robson, Mr E. C. Wallace, Professor D. A. Webb
Honorary General Secretary, Mrs M. Briggs
Honorary Treasurer, Mr M. Walpole
Back issues of Watsonia are handled by Messrs Wm Dawson & Sons Limited,
Cannon House, Folkestone, Kent, to whom orders for all issues prior to Volume 16
part 1 should be sent.
Recent issues (Vol. 16 part 1 onwards) are available from the Hon. Treasurer of the
B.S.B.1., 68 Outwoods Road, Loughborough, Leicestershire.
Watsonia, 16, 237-247 (1987) 237
Presidential Address, 1986
DE ALLEN
PART ONE: ON BEING A HUNDRED AND FIFTY
Back in the early 1950s anyone who happened to work in the library of the Department of Botany
at the British Museum would be startled every now and again by a fierce volley of expletives that
emanated from somewhere on the upper floor. Traced to their source, these would be found to
have come from a little, cross-looking, very old man bent over an array of open books and scraps of
paper in what appeared at first sight to be a state of total confusion. He was a one-time member of
the staff, long since retired, the more curious visitors would be gently informed, who was toiling
away at a polyglot dictionary of botanical terms, which, alas, there was no likelihood of his
completing before he was taken from this world. And that, sad to say, did in due course turn out to
be the case.
The old man was a certain Antony Gepp, who in his time had done distinguished work as the
Department’s specialist in algology. And unless I am seriously mistaken, a Miss Branfill whose
name features in the earliest list of members of our ancestor, the Botanical Society of London, was
none other than his mother. A girl of sixteen then, she was to marry at twenty-nine, give birth to
him when in her fortieth year — and leave him to live on until he was well into his nineties. Thus
some of us here today have been privileged to touch hands, as it were, with the founders of the
Society at just a single remove. A hundred and fifty years is not so long as all that: a couple of
lifetimes only, at a rather extended stretch.
Yet in other ways 1836 does seem a very long time ago indeed. It was only two years before that
that slavery had been abolished — and then only within the British Empire: other Western nations
would continue to cling to it for a long time yet. At home meanwhile the penal laws were still
horrific. Transportation remained in vogue. The gibbet had only just been abolished. It was only
sixteen years since the last execution by beheading.
Politically, too, the scene was hardly a familiar one. William IV was on the throne and
Melbourne was in the second year of what was proving to be an agonising premiership. After a
period of acute unrest, of rick-burning in the countryside and rioting in the streets, the Reform Bill
had eventually been passed, only to leave the radicals as noisy as ever, their appetites merely
whetted. As they bayed at Melbourne’s heels, the King and the House of Lords growled and
snapped at him from above. Two years earlier, as if symbolically, Parliament had been in flames.
Now the Prime Minister himself was the subject of a squalid trial, accused of adultery with a
leading society beauty. The very next year the economy would collapse, a long depression would
start and the Chartist agitation would set in. In another two years there would be a serious uprising
in Wales and a chain of conspiracies in Yorkshire. It can hardly have seemed an auspicious time to
be founding a national society.
Yet a much more familiar world was also just starting to appear. Already, six years earlier, the
first passenger train service had been brought into being — and, soon after that, the first plant-
spotting out of train windows. In June 1832, on a journey across Lancashire by this:new form of
travel, the young William Christy made several interesting records while moving, in his awestruck
words, “‘at a speed which almost precluded any botanical observation” (Christy 1833). Within two
years of the Society’s founding the first excursion trains had even been introduced and Daniel
Cooper, our founder, was leading a group of the members to “‘a spot . . . called Woking’’, chosen
because it had become easy to reach by the newly-opened London and Southampton Railway
(Cooper 1838). Only one year later Mr Bradshaw was to bring out the first of his national
timetables.
It would be another four years yet, though, before the members would enjoy the penny post.
238 D. E. ALEEN
Even so, in the Society’s inaugural year the crippling stamp duty on paper had been brought down
to only a penny a sheet, one of the major spurts forward on the road to the abolition of the much-
decried ‘taxes on knowledge’. In that same year, too, the first provincial daily came out, and the
University of London was established as a national degree-conferring institution. 1836 also
witnessed, believe it or not, the first hospital training scheme, the earliest screw-driven steamboat
and the foundation of the gas appliance industry. More ominously, the first fatal train derailment
occurred, to be followed in the very next year by the earliest death from a parachute jump.
Already, too, by 1835 Fox Talbot (a botanist, be it noted) had taken the earliest surviving
photograph. Within four years of that even microphotography had been invented. A tremendous
transformation in the everyday environment was in progress.
By comparison, science lagged. 1836 was the year of Darwin’s return in the Beagle: over twenty
more would have to elapse before the society in whose rooms we meet today would be the first to
be told of its shocking outcome. 1836 was the year too in which Henslow held the last of his famous
Friday evening ‘at homes’ for the science students at Cambridge. Now he was off to be a country
vicar — and an absentee occupant of the Professorial Chair. The previous year the Royal College of
Physicians had rescinded its rule that only graduates of Oxford and Cambridge could be elected as
its Fellows, and the marvellous high summer which the sciences had enjoyed since the 1820s at
those two universities as a result was abruptly brought to an end. Within a decade the number of
medical students at Cambridge was down to a mere five or six yearly and empty classrooms were
greeting the science professors to their embarrassment (Becher 1986). By 1846 Babington would
be having to write: ‘“Never was botany at so low an ebb as now in this place” (Babington 1897). At
the embryonic London University the Chair of Natural History at King’s had been abolished
because of the shortage of students; and although the Chair of Botany there survived, the
emolument that it carried was so disgracefully low that Edward Forbes would have to hold it in
tandem with the Geological Society’s Curatorship — which paid half as much again — in order to
enjoy more than an artisan’s existence (Wilson & Geikie 1861).
But if botany in English Academia was in sad disarray (and matters were very different up in
Scotland, we must be careful to remember), it was a halcyon period for the subject elsewhere.
Field botany, fashionable since the Linnaean 1770s, was now borne along on a second and much
more massive wave, which penetrated far more sweepingly. Of its innumerable catches, almost
every One was engaged in forming a personal herbarium, and wore on their roamings through the
countryside that by now conventional emblem of the fraternity, the roomier, eighteen-inch
students’ vasculum (Allen 1959). Before long a member of that ancestral society of ours, Thomas
Bentall, would have succeeded in interesting his family firm in producing a special drying-paper to
accompany it.
Field botany was well into its great nineteenth-century expansion, brimming over with
confidence, tingling with excitement. Additions to the flora were continuing to be turned up in the
less well-worked corners of these islands at a not noticeably declining rate; and even if by then
most of the more obvious and spectacular species had been discovered, keener discrimination
seemed likely to ensure that the stream of novelties would not dry up for a long time yet. In that
very year of 1836, indeed, there had again been an impressive haul: Carex punctata Gaud.,
Cystopteris montana (Lam.) Desv., Calamagrostis stricta (Timm) Koel. and Spartina alterniflora
Lois. The previous one had yielded Ononis reclinata L., Erica mackaiana Bab. and Valerianella
carinata Lois., the next would see the first of Babington’s two successive forays to the Channel Isles
and two more prizes as a result: Ornithopus pinnatus (Mill.) Druce and, most glittering of all,
Spiranthes aestivalis (Poir.) Rich.
These were the years when the still-youthful Babington was busily making his mark,
crisscrossing the British Isles one long summer after another and by his efforts alone considerably
augmenting the total of species they were known to possess. Never to be a member of the
Botanical Society of London (though he did join the later Botanical Exchange Club), he was
twenty-seven when it was born. Watson, his resentful rival, and presently to be its mainstay, was
five years older at thirty-two. Babington’s Manual, the first really critical field handbook, was still
nine years off; but Watson’s New Botanist’s Guide, that updated register of what each county held,
was in the middle of coming out. Detailed treatments had appeared already of Mentha, Salix and
Rosa; Rubus had been marginally nibbled; but Hieracium was still effectively virgin territory. It
was obvious that there was a great deal waiting to be done.
PRESIDENTIAL ADDRESS, 1986 239
AN ABNORMAL DEVELOPMENT
Having set the scene, I must now reluctantly drag you away from it. For if I continue, I can only
recapitulate in brief what I have already set out at length in the book (Allen 1986) whose
publication we are also celebrating this day. For that same reason this is debarred from being the
kind of presidential address in which the president looks back down the road along which his
society has travelled and points out the milestones. I know that view too well now, in any case, to
be capable of describing it with an appetising freshness. I must leave you to discover it for
yourselves under the guidance of the written page.
You will learn that the Society has been lucky to survive for as long as it has. There were two
awkward junctures, indeed, at which its continuation was seriously in doubt. It has had to endure
periods of autocracy and neglect, civil war, bankruptcy and the loss twice over of all its records.
Only a body which met a very insistent need would have displayed such resilience.
All the same we must bear in mind that it has been a normal learned society for only just over a
third of its life — that is, for fifteen years or so at its start and for the forty which have passed since
the end of the Second World War. For much of the time in between it had a disembodied
existence, so disembodied indeed that when the time came for Charles Bailey to lay down the
Honorary Secretaryship in 1903 after twenty-four years, there was scarcely a single member who
had met him in the flesh. Even after that period, during the years when that postal club grew
steadily more accustomed to describing itself as a ‘society’, it existed as a physical entity more or
less only through its annual reports. It never came together for lectures and not until latterly even
for field meetings that were other than few, irregular and doubtfully official. Until the late 1940s
there was not very much to be proud of — and a history chiefly notable for its heavy punctuation of
rows and sniping.
So rather than give way to a Schwdrmerei of self-congratulation of the kind so tediously
traditional to commemorative occasions, it seems more in keeping to consider what the Society has
missed. By its failure to enjoy an ordinary existence what has it been deprived of? What gaps in our
activities are there still that might usefully be closed?
SOME DEPRIVATIONS
The most obvious of the deprivations is the lack of a library, a lecture-room, somewhere to call our
own. Had the Botanical Society of London only continued, instead of dissolving and selling off all
its possessions in 1857, there is every likelihood that we would now be in such richly-propertied
circumstances as the Royal Entomological Society, a body which was founded only three years
before us, took off from very much the same starting-point and in its infancy passed through some
parallel vicissitudes.
From one point of view, of course, this is a deprivation that has turned into an advantage. All
too many of the old-established metropolitan societies have found themselves under siege in recent
years from spiralling wage bills and rocketing rents. In all too many cases now the marvellous
collections of journal runs and books that they have accumulated through the years are
distressingly little-used, largely duplicated as they have come to be by the libraries of the
universities and of the major government institutions. Not a few societies, indeed, have found this
inherited burden too heavy to sustain and have been forced, or at any rate have opted, to transfer it
to some better-endowed pair of shoulders. Even so they have normally been able to arrange for
their members to continue to have access, and can thus go on offering this valuable extra privilege
to that limited proportion who enjoy no alternative facilities.
While we may have been spared an incubus, therefore, and along with it the expense of housing
a staff, we have probably suffered a deprivation on balance. Certainly we have suffered one by
having no place of our own in which to forgather before and after meetings of Council and
committees. Since the sad demise of ‘Domus Linnaei’, of hallowed memory, there has not even
been a handy establishment ‘just around the corner’ to serve as the recognized assembly-point at
which the formal business of the evening can be rehearsed or digested. Without a convivial
warming-up beforehand, or a no less convivial winding-down afterwards, committee meetings tend
to be too clinical and angular. Proceedings begin and end with an almost brusque abruptness;
240 DSEVALERN
newcomers are left at sea, unfamiliar with the background to the agenda (or with the identity of
who they are sitting opposite or next to); the full inside of issues remains too thinly known. A
society without an ‘ante-chamber’ life is a society without a valuable extra dimension.
Another by-product of having no premises is our lack of a programme of regular winter
meetings. For if we were the possessors of a lecture-room of our own, we would surely wish to
make use of it frequently. Certainly our ancestor could not have conceived of dispensing with such
an activity: it was, after all, of the very essence of a metropolitan society of the kind it aspired to
be. On the other hand since that ancestor of ours collapsed we have ceased to be metropolitan,
albeit latterly London-based. This has left us without the metropolitan reflexes and so without the
urge to restore the status quo ante and repair the deficiency. Had we those reflexes, we would
surely long since have followed the example of, most notably, the London Natural History Society
in not allowing the mere lack of facilities of our own to stand in the way of such a step.
But maybe we have done injury to ourselves by keeping to this non-metropolitan stance.
Without a regular forum in which new findings can be critically debated as well as disclosed a
scientific society is prone to a certain stunting. It is true that we have our symposia and
conferences, but these are but occasional affairs and often on very restricted topics. It is true, too,
that the conferences of Vice-county Recorders every other year give rise to ample and sometimes
vigorous debate; but, again, these are specialized in their focus and in any case cater for only a
limited section of the membership. There is no outlet for papers on non-prescribed topics, for the
reporting of work that is still at an early stage, for the airing of general critiques or the mere
floating of ideas. And because we do not have one, it has regularly fallen to the Linnean Society,
instead of to ourselves, to play host to those one-day gatherings of doctoral students who have
theses in train on the biosystematics of British vascular plants. It is excellent that such occasions
should occur and excellent that a national society should sponsor them; but it would surely be even
better if those engaged in these studies had an opportunity to outline their investigations to an
audience that is best placed of all to assist with material, observations and records from all over the
British Isles.
Filling this gap need not mean the instituting of a full-scale winter programme. It is doubtful if
the supply of suitable material would warrant that in any case; moreover, there are a sizeable
number of botanical topics on offer from other societies with just such programmes already. But
even just one or two meetings might help to brighten the dark nights and make the annual
Exhibition less of an oasis in that long, bleak desert between the end of one plant-hunting season
and the opening of another.
There is one further deprivation which I suspect it may not even have occurred to the Society
that it suffers. It arises from the fact that we confine ourselves to the flora of the British Isles — just
as we do to only certain sections of that flora and to only certain aspects of its study (though it is not
with those other limitations that Iam here concerned). Had only the Botanical Society of London
managed to carry on, the whole of the world — and, for that matter, the whole of the science of
botany — might still be our oyster, on the pattern of the metropolitan societies in general. When it
folded, the field of vision contracted abruptly to that of its surviving rump of members: a handful of
collectors, of very modest means, mostly out in the provinces, whose private herbaria were
restricted, more or less of necessity, to the flora of these islands and, by choice, to their vascular
plants alone. That new cement set hard, so hard in fact that it has remained in place ever since.
And who is to say whether the Society would have done any better had it taken a less insular
course?
All the same we should recognize that in settling upon these particular geographical bounds,
indeed by putting ourselves in thrall more generally to taxonomy’s ‘territorial imperative’, we have
settled our fate in a wider and subtler way. For because there are, unfortunately, so relatively few
taxa that the flora of these islands shares with the flora of Canada and the United States, we have
deprived ourselves of any chance of participating in what has been termed ‘tthe common culture of
the North Atlantic triangle” (Berger 1982). In almost every other field of scholarship this English-
speaking triangle houses what amounts to a single community, a community in which a specialist
based in Britain is as likely to find his closest kindred spirits in Berkeley or Toronto as in Glasgow
or Leicester. From both sides of the Atlantic contributors subscribe to, and write for, shared
journals; there are frequent transatlantic conferences and symposia; libraries and archives in the
British Isles are thick for large stretches of the year with researchers from North America. The
PRESIDENTIAL ADDRESS, 1986 241
social interpenetration is as wide as it is deep and a source of great enrichment, personally as well
as intellectually, for those with the good fortune to be able to partake of it. Alas, though, by the
nature of the course we have taken, it can never be for us. All that we can do is to eye it with
eternal envy and regret what we have had to miss.
So much for the shortfalls in our ways of functioning that we happen to suffer from by virtue of
the particular kind of society that we are. There is little enough that we can do about them, but I
have pointed to ways in which that little might be done.
PART TWO: WORK ON INFRASPECIFIC VARIATION
But what about the scientific content of our work? Are there any glaring gaps in that still waiting
to be filled? This is something that we shall be discussing this afternoon and I must be careful
therefore not to encroach on the area overmuch. There is one suggestion, however, that I would
like to put forward which I feel fairly confident no one else plans to propose at that later
juncture. It amounts to taking up where one of my predecessors in this office, Professor
Valentine, in effect left off in the stimulating Address that he delivered on this same occasion
eight years ago. In the course of a wide-ranging review of experimental work on the British flora
he remarked upon the fact that “interest in variation below the level of the species and in the
naming of taxa at the level of variety and form has tended to decrease in the writing of local
Floras”’ (Valentine 1979a). This is indeed only too true. Someone picking up and leafing through
the average local Flora published in recent years might well suppose himself back in the era of
the fixity of species, so largely absent is any hint that the entities listed are subject to variation —
and to variation that at least in the past has come in for taxonomic recognition.
The reason for the silence, of course, is that writers of local Floras tend to take their cue from
the national Floras currently in use, and the national Floras have yet to grow out of the general
revulsion between the Wars against the excess to which the earlier ‘stamp-collecting’ school had
taken the naming of minor variants. The experimental studies which came into fashion in the
1930s showed that some of these taxa were quite without substance, reverting to normal when
tested in cultivation, while others were no more worthy of a name than many analogous variants
that had merely happened to escape notice. More basically, they also showed that the variation
_ In many species is so complicated or so extensive that it is hardly feasible to attempt to do justice
to it taxonomically. In any case the established hierarchy of categories — subspecies, variety and
form — had come to seem artificially restrictive, even supposing that any broad measure of
agreement could be reached on how they were interpreted.
Unnerved by the strictures of the new experimentalists, Flora-writers responded by dropping
more or less all mention of infraspecific variants from their pages, doing so indeed with consider-
able relief, for they secured as a result a less cluttered text and useful savings in space. Left
without names and descriptions, recorders in the field in their turn gradually lost the habit of
noting the variants that they met with. The widespread use of mapping cards with room only for
the names of species (and only for a proportion of those) has merely intensified the trend.
It is a trend, though, that has surely now been allowed to go much too far. The study of
micro-evolution is no less important than ever it was and it is a study in which field botanists
ought to be contributing their due. For despite the salutary correctives to past simple-mindedness
administered by the genecologists and their like, there is in fact a great deal of value that could
still be contributed by straightforward observation. Many of the taxa lodged in the past literature
have too readily come to be dismissed as no longer worth recording merely because they are
based on a departure from the norm in just a single character. Yet it does not need much
searching through the older Floras and back numbers of journals to turn up evidence that
differences even as slight as this are by no means without interest in the pattern of their occurr-
ence geographically. Thanks to the fact that many of the ablest and most assiduous field botanists
of the second half of the nineteenth century were marooned in isolated country parishes or
confined to the area they could cover in the course of a Sunday ramble on foot, their explorations
tended to become ever more minutely intensive and the many described minor variants, no less
than the microspecies, were the beneficiaries of this. There was a rich harvest as a result, which
has still been only very partly cut.
242 DD: EB. ALEEN
There is a form of Medicago lupulina L., for example, in which the pods are tubercled instead of
smooth. S. F. Gray christened it var. scabra as early as 1822. According to workers in the period
around the turn of this century this is the prevailing form of the species round Malvern, in
Herefordshire and Worcestershire (Towndrow 1911), whereas in Oxfordshire it is scarce (Druce
1890) and in Derbyshire even scarcer (Drabble & Drabble 1911). Should that pattern prove to be
confirmed, it is odd enough to need explaining. Similarly there is a form of Filipendula ulmaria (L.)
Maxim. in which the normal tomentum is missing on the under-surface of the leaves. The character
is known to breed true (Druce 1911) and to remain unaltered by differential cultivation (Beeby
1888; Yapp 1912). Yet again, oddly, while rare in some districts it is the prevailing version of the
species in others. If we are to believe the old records, a puzzling alternation of this kind is also
displayed by the variant of Leontodon taraxacoides (Vill.) Mérat with a hairy involucre.
Rarely in such cases have any ecological differences been reported that might provide a clue to
the selective factors at work — always assuming there are any and that the patterns are not merely
random. One exception is the claim by Druce (1897) that the form of Polygonum convolvulus L.
which has the calyx segments winged, var. subalatum Le}. & Court., is the representative of that
species in rich garden ground, bushy places and hedges, while the ‘normal’ form is characteristic of
cornfields. That observation, however, seems hard to square with the finding by Drabble & Long
(1932) that the variant is the commoner of the two by far in the Isle of Wight. It may be, though,
that Polygonum convolvulus is one of those species in which the native, or at least old-established,
populations are conveniently distinct in some external way from more recently-arrived ones or
from mere casual specimens. Examples of this are to be found in Papaver rhoeas L., Sisymbrium
officinale (L.) Scop., Silene vulgaris (Moench) Garcke, Lapsana communis L. and Cirsium arvense
(L.) Scop., in each of which the plants prevailing in more southerly parts of Europe differ in one
way or another in their hairiness. Even more conveniently, populations from further south or east
in Europe may have different flower colours: in Raphanus raphanistrum L., for example, purple as
opposed to yellow or white, in Anagallis arvensis L. blue as opposed to scarlet. One of the
strongest reasons for studying infraspecific variation, it has always seemed to me, is that it can so
often help in clarifying status. Indeed it is only thanks to enthusiasts for this line of work, and to the
taxonomic trail that they have left behind them, that we know of seemingly native ecotypes of
species that otherwise tend to be dismissed as non-indigenous: the condensed, Erophila-like form
of Arabidopsis thaliana (L.) Heynh., for instance, on rocks high up in the central Pennines for
which Druce (1924) coined the epithet brevicaulis, or the prostrate plants of Spergula arvensis L. in
short close turf in the Channel Isles which E. F. Linton (1907) first brought to notice and described
as var. nana.
Spergula arvensis, as it happens, has also been the subject of a model study in recent years (New
1958, 1959) in which the nature of the two main ways in which that species has long been known to
vary has now been definitively elucidated. In one of these variants, known as var. sativa (Boenn.)
Mert. & Koch, the seed-coat lacks the usual papillae; in the other the plant as a whole is densely
hairy. It turns out that, although there is no link between the two genetically, both exhibit much of
the same geographical trend, increasing proportionately from south-south-east to north-north-west
and with increasing altitude, not only within the British Isles but across Europe more generally. It
would seem logical to infer from this that both characters are adaptations to a climate combining a
greater coldness with a greater wetness, and experiments have indeed shown that the plants with
non-papillose seed-coats do germinate more readily at low temperatures.
Many other variants probably occur with increasing frequency in a particular direction like this,
in response to a gradual crescendo in the environmental tendency that favours them. In some cases
the variation takes the form of a more or less steady shift in the proportion of one character at the
expense of another sharply contrasting with it — a polymorph-ratio cline in the technical
terminology. An example is the gradual outnumbering north-westwards across Europe of blue-
flowered Anagallis arvensis by the scarlet type. Typically, as in this case, just a single pair of genes
is involved; but sometimes the situation is more complex. In Raphanus raphanistrum there are four
different colour forms, purple, white, pale yellow and deep yellow, which replace one another
across Europe in a series of broad but widely overlapping belts. In the south of Britain the white
and pale yellow forms commonly grow intermingled and there discrimination between these by
insect pollinators has recently been discovered to be extremely marked (Kay 1976). Presumably in
this case the influence of climate is at one remove.
PRESIDENTIAL ADDRESS, 1986 243
After flower colour, in which frequency differences are so much more obvious, the most
numerous reported examples of ratio clines seem to be in leaf-marking. The best-known of these is
the north-south one in Arum maculatum L., the populations of which in the north and west of the
British Isles have unspotted leaves almost exclusively (Prime 1955), although even as far south as
Wiltshire spotted plants constitute no more than a fifth of the total (Grose 1957). By contrast, in
Dactylorhiza praetermissa (Druce) So the cline is an east-west one, with a lowering in the
incidence of leaf-marking across southern England as one travels towards Kent (Heslop-Harrison
1958). Another east-west example, this time in a fruit character, would appear to occur in Stellaria
neglecta Weihe; for according to Marshall (1914) the form of that with the seeds bluntly instead of
acutely tubercled, his var. decipiens, is scarce in Somerset but easily predominates in Sussex and
Surrey.
We do not know what degree of stability these ratio clines possess, for no measuring of
populations has yet taken place over a sufficiently lengthy period. Certainly in butterflies short-
term fluctuations of a sometimes startling magnitude have been reported. That they may occur in
flowering plants as well might appear to be indicated by the marked increase that has taken place in
recent years in the rayless form of Aster tripolium L. (var. discoideus Reichb.) on many East Coast
salt-marshes, at the expense of the normal rayed one. Gray (1966) has proved statistically that the
rayless plants are more characteristic of the lower zones of the marshes and believes their greater
ability to withstand tidal submergence gives them a selective advantage there. The spread of
Spartina may have been responsible for changes in the ecology which have provided the necessary
opening. On the other hand it seems that there is more than just raylessness to this taxon:
according to another author (Burtt 1970) it also has a different habit, thicker and more brittle
leaves and a shorter pappus. So it is perhaps better classed as an ecotype than as the kind of single-
character variant typical of ratio clines.
But not all clinal variation involves clear-cut entities of the sort just described. Very often the
graduated response to a graduated environmental influence is expressed developmentally, in
stature, say, or leaf shape; and because the genes controlling growth tend to be more numerous
and their interaction complex, the variation in that takes the form of a continuous dimensional
trend in a particular character or character-cluster. In these cases simple percentage counts are
clearly out: measurements have to be made and biometrics grappled with. For that reason the
record of the earlier botanists in discriminating such variation is less impressive. In many species
indeed they overlooked its existence entirely, simply because of the very gradualness of the
changes in appearance. Alternatively, they dismissed it as of no taxonomic worth, with the
standard verdict of ‘grades into type’. When they did give it recognition, the solution adopted
depended on how narrowly or widely the variation ranged. If it was sufficiently narrow, as in the
east-west cline in the length of the outer calyx teeth in Lythrum portula (L.) D. A. Webb (Druce
1911; Allen 1954b), it was feasible not only to give a varietal name to what was regarded as the
extreme (that is, the furthest the character departed in shape or size from whatever was taken to be
the norm), but also to capture the whole of the variation in between with a single ‘omnibus’ varietal
name as well. If the variation was much wider, however, as in the leaf-cutting of Anthriscus
sylvestris (L.) Hoffm. (Druce 1917), no one name could be made to do service for so long a bridge
of intermediates and, rather than coin a whole series of names for these, just the two extremes
were treated to taxonomic recognition. The futility of doing anything more than this was well
illustrated by a study made in Denmark and part of Sweden of the extent of the variation that
needed to be accounted for in that same species: it transpired that a total of sixteen different taxa
would have had to be created if the taxonomic potential had been pursued to its logical conclusion
(Petersen 1915). In a similar study of the leaf-cutting in Pimpinella saxifraga L. the artificiality of
the practice of giving a name, in this case the so-called ‘var. dissecta (Retz.) Spreng.’, to merely
one part of what was shown to be a continuum was reasonably called into question (Petersen 1921).
But just how such variation is to be handled taxonomically, if indeed it can be accommodated
within the established system at all, is something that still remains unresolved.
A clear distinction is often not easy to make out, at any rate from the literature, between truly
continuous geographical trends and cases where a variant merely becomes more common in a
particular climatic region. For example, is the form of Sagina subulata (Sw.) C. Presl without the
usual glandular pubescence, var. glabrata Gillot, mainly in the north and west of the British Isles
(Harrold 1978) as one end of a ratio cline — as the golden yellow form of Melampyrum pratense L.,
244 DIET AEVEN
var. hians Druce, so clearly is — or is it a mutant which has been able to colonize those parts thanks
to a selective advantage there but scarcely to penetrate the more southern and eastern populations
of the species? Exactly the same question has to be asked of the form of Angelica sylvestris L. with
the upper leaf segments more or less decurrent, var. decurrens Fisch., Mey. & Lallem., which is
said to prevail in the north and west similarly (N. D. Simpson, pers. comm. 1951). On the other
hand it is hard to see why the form of Gnaphalium uliginosum L. with hairy fruits, var.
pseudopilulare Scholtz, should apparently be commoner than the glabrous-fruited form in France
(Corbiére 1894; Hariot 1895) whereas the paucity of records appears to suggest that the opposite is
the case on this side of the Channel. Under-recording is the stumbling-block at present in seeking
to interpret such patterns.
All the same some variants are so conspicuous that it is hard to believe that their distribution is
not known with tolerable completeness. My favourite in this connection is the colour form of Jris
foetidissima L. in which the normal livid purple is replaced by clear lemon yellow with merely
purple lines. It was named var. citrina by Bromfield early last century. There is a strong
concentration of it in eastern Dorset, in Purbeck, where five localities had been discovered by
1895. It has also been recorded from two places in the Isle of Wight, from one in central Dorset and
from near Newton Abbot in Devon; a single plant was also found near Bath in 1924. Though
scattered over quite a wide stretch of country, the finds are sufficiently centred on Purbeck to
suggest very strongly that that is where this mutant arose. It is a great pity it escaped being mapped
in the Critical Supplement to the Atlas (Perring & Sell 1968).
Hardly less conspicuous is the form of Bidens cernua L. with ray florets, var. radiata (Roth)
Lindl. Of this there is a similar clustering of records, this time in Cheshire and South Lancashire;
but it has also long been known in a locality in as far from there as Somerset, so in this case
independent mutations, perhaps on many occasions (for the variant is known on the Continent),
seem likely. The same may be true of the rich magnolia purple form of Trifolium repens L., var.
rubescens Sér., so prevalent in Scilly but also (McClintock 1975) in several places in Guernsey.
Variants less obvious than these, however, must be presumed to have been at best only patchily
recorded. Only when there has been cause for a species to be investigated intensively, as in the
accounts prepared for the Biological Flora of the British Isles, have there been exceptions. It is only
thanks to the unusually thorough study carried out for that work by Lewin (1948) of the variation
of the species of Sonchus that we know that there is a form of Sonchus asper (L.) Hill with white
achenes which is common in the Orkneys and on the north and west Scottish mainland. It is
apparently without a name. Many may think, indeed, that variants as slight as this, probably the
product of just a single gene, are too trivial to warrant lumbering the literature with yet further
taxa. On the other hand if they are left unnamed, they are likely to go unrecorded (unless they
should happen to be the subject of someone’s special interest) and the surely not unimportant fact
that they have achieved a distinctive, reasonably compact range may consequently be overlooked.
It is not unimportant, of course, for the reason that any mutant capable of carving out a range for
itself must be presumed to have some selective influence in its favour and may thus be an incipient
local race, perhaps in the course of time to accumulate a further measure of distinctiveness
sufficient to entitle it to subspecific rank. Just such a process seems to be in the course of happening
in the case of the suberect ecotype of Anagallis arvensis which occurs mainly on sand dunes in the
west of Britain and Ireland. Some of the populations of that, but by no means all, have acquired
the additional distinction of flesh-coloured flowers — the so-called var. carnea (Schrank) Boenn.
(Rilstone 1938; Allen 1954a). The similarly western coastal race of Calystegia sepium (L.) R.Br.,
subsp. roseata Brummitt, is one step further on: as well as its rosy corolla it has more acute leaf tips
and more pubescent stems (Brummitt 1967).
That brings me to the vexed question of how the three infraspecific categories acceptable under
the international rules are most appropriately interpreted. Clearly, hard-and-fast dividing-lines are
not to be expected, as we have learned not to expect them between subspecies and species. At the
same time it is surely not utopian to look for a reasonable measure of consensus about which level
in the hierarchy of taxonomy is appropriate to which level of the hierarchy observable in nature.
Until the 1930s we seemed to be making good progress in that direction. Then along came
genecology and the advocating of alternative courses. The more extreme of these was the
abandonment of the traditional terminology in favour of an entirely new one freed from the
shackles of taxonomy altogether. That ‘deme’ concept, however, despite the appeal of its goods-
PRESIDENTIAL ADDRESS, 1986 245
train versatility, seems to have proved but a passing vogue. The other course was the compromise
one of taking one of the traditional categories and fashioning it anew. The chosen victim was the
subspecies. The term ‘variety’, it was held, had become too irretrievably ambiguous to be usable
any longer — and any variation below that level was too trivial to be worth bothering with. But
whereas a subspecies had previously been thought of as more or less synonymous with a
geographical race, the genecologists took the view that no sustainable distinction could be drawn
between the subdivisions of a species created by lengthy isolation or adaptation to a different
climatic zone and those subdivisions, typically much smaller and much more restricted in their
occurrence, which result from adaptation to particular specialized habitats (‘ecotypes’, as they
have come to be known). This was despite the fact that these latter are, by their very nature,
capable of arising independently in more than one place and are thus the very antithesis of what
plant geographers have been accustomed to think of subspecies as denoting — namely, broken-off
bits of a species which, by virtue of their uniqueness, can serve as valuable historical evidence. An
essential part of the coinage, in other words, was damagingly debased. The subspecies became
“the dumping ground for many sorts of situations’’, used for the accommodating of entities as
disparate as cryptic or semi-cryptic polyploids, maritime or alpine ecotypes, physiological races
and other morphs differing in relatively minor characteristics — in addition to the traditional
geographical races (Stace 1976).
This broadening of the subspecies has introduced a further layer of confusion into what was
already a more than sufficiently confused state of affairs. What is more, followers of this approach
who have made revisionary incursions into our flora in the post-war years have tended to leave
behind them an irritating lopsidedness. In the Shetlands, for example, the dwarf ecotype there of
Silene dioica (L.) Clairv. has been raised to subspecific rank (Baker 1947, 1948) — maybe rightly, in
view of its distinctiveness geographically as well — but the no less distinctive race of Senecio
aquaticus Hill that occurs in such profusion in those islands remains unpromoted from the varietal
level at which Druce (1921) described it. More seriously, the heritable forms of Geranium
robertianum L. that occur on shingle round our coasts have been grouped all together, though
according to Yeo (1973: 339) quite unjustifiably, and elevated similarly to a subspecies (Baker
1956) — receiving as a result, ironically, their proper due at last from recorders — but, again, the
analogous versions of Galium aparine L. and Solanum dulcamara L. have been allowed to languish
in mere varietal obscurity in contrast. The same goes for the ‘cushion’ ecotypes of maritime
habitats that have been described in a number of species: alone among these that of Valerianella
locusta (L.) Betcke has been dignified with the higher rank (Sell 1967).
On top of this inconsistency — or rather, lurking beneath it — there is a terrible chaos in the
nomenclature. Continental names have long been taken over for similar-sounding British variants
without any comparison of specimens, let alone consulting of type material; there has been no
thorough searching of even the British literature to check whether earlier valid names exist;
descriptions are very often inadequate. Yet better wrong names than no names at all, surely: we
should not allow such problems to serve as an excuse for total inaction. For I hope I have said
enough by now to have convinced you that there is a major gap here in our knowledge of the flora
of these islands, which it is high time that this Society led the way in seeking to close.
An essential preliminary, though, is for us to know what has been discovered already. As I have
attempted to demonstrate, there is a vast store of data on the subject buried in the past literature:
what we require in the immediate instance, and urgently, is a compendium bringing it all between
two covers. When Professor Valentine addressed you on this same topic in 1979, he spoke in the
knowledge that the projected five-volume ‘Flora of Great Britain and Ireland’ would be taking care
of that deficiency (Valentine 1979b). Since then, alas, that project has foundered, and the Society
now remains the only hope. But it has successfully produced already a bible on British hybrids
(Stace 1975): I see no reason why it cannot follow suit now with a companion volume on British
infraspecific variation.
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Baker, H. G. (1956). Geranium purpureum Vill. and G. robertianum L. in the British flora — II. Geranium
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Burtt, B. L. (1970). Infraspecific categories in flowering plants. Biol. J. Linn. Soc. , 2: 233-238.
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VALENTINE, D. H. (1979b). Flora of Great Britain and Ireland. Watsonia, 12: 279.
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Witson, G. & Gerke, A. (1861). Memoir of Professor Edward Forbes, p. 324. Edinburgh.
Yapp, R. H. (1912). Spiraea Ulmaria, L., and its bearing on the problem of xeromorphy in marsh plants. Ann.
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Bot. J. Linn. Soc. , 67: 285-346.
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Watsonia, 16, 249-267 (1987) 249
Variation in Jasione montana L. (Campanulaceae) and related
species in Europe and North Africa
J. PARNELL
School of Botany, Trinity College, Dublin 2, Ireland
ABSTRACT
This paper describes the variation of European and North African Jasione montana L. North African material
is assigned to J. montana subsp. cornuta Greuter & Burdet. All other J. montana belongs to J. montana subsp.
montana within which six varieties are recognized. These are J. montana subsp. montana var. montana, J.
montana subsp. montana var. litoralis Fr., J. montana subsp. montana var. latifolia Pugsl., J. montana subsp.
montana var. echinata Boiss. & Reuter, J. montana subsp. montana var. bracteosa Willk. and J. montana
subsp. montana var. imbricans J. Parnell, var. nov. Only the first three of these varieties occur in Britain. A
number of Jasione species are relegated to synonymy with these taxa though the closely related J. penicillata
Boiss., J. corymbosa Poiret ex Schultes and J. heldreichii Boiss. & Orph. are maintained as species. The two
latter species are each recognized as containing two varieties; these are J. corymbosa var. corymbosa, J.
corymbosa var. glabra (Durieu ex Boiss. & Reuter) J. Parnell, stat. et comb. nov., J. heldreichii var. heldreichii
and J. heldreichii var. papillosa J. Parnell, var. nov.
INTRODUCTION
The genus Jasione L. (Campanulaceae) contains twelve species. It can be conveniently split into an
annual/biennial section and a perennial section. Although two of the perennial members, Jasione
bulgarica Stoj. & Stefanov and Jasione foliosa Cav., have, on occasion, been placed in separate
genera, I feel that Jasione clearly forms a single natural entity. In this paper I have applied a
combination of a detailed, experimental, biometric study of Jasione montana L. from the British
Isles and herbarium observations of material from elsewhere to re-assess the infraspecific
classification and relationship of all non-perennial Jasione species.
J. montana is the most widespread species in the genus, occurs commonly throughout Europe,
and has an ill-defined pattern of infraspecific variation. Details of its autecology, morphology and
distribution are given in Parnell (1985). Plants of this species may be either annual or biennial, may
vary from almost glabrous to hirsute, may be branched in the lower half or unbranched and be
from 1 to 65 cm tall. Recent revisions vary considerably in their treatment of this infraspecific
variation. In a continuing series of papers, Rivas-Martinez (1967, 1976, 1979) and Greuter (1981)
recognize a number of subspecies of J. montana, whereas Tutin (1976) recognizes none.
The only complete revision of Jasione is that of Schmeja (1931) but his work has, with good
reason, been laid aside and it is the earlier work of Pugsley (1921) which most clearly outlines the
current understanding of British Isles J. montana. Pugsley (1921) distinguished five, rather ill-
defined, varieties; these are a decumbent sand-dune taxon J. montana var. litoralis Fr., a gross
cliff-top taxon J. montana var. latifolia, a dwarf cliff-top taxon J. montana var. maritima Bréb., a
glabrous variant J. montana var. laevis Duby and a tall variant J. montana var. major Mert. &
Koch. The major problems with these taxa are that Pugsley’s own annotated specimens (BM) do
not correspond with his descriptions and that they were described in the absence of information on
phenotypic variation in J. montana. Additionally, continental workers have described many largely
undistinguished taxa, which may or may not occur in the British Isles.
250 J. PARNELL
3
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FicureE 1. The location of populations sampled for morphological analysis.
VARIATION IN J. MONTANA WITHIN THE BRITISH ISLES
SAMPLING, SCORING, CHARACTER SELECTION AND PRELIMINARY ANALYSIS
A total of 59 populations from a wide geographical and ecological range were sampled during the
summers of 1977 and 1978. Fig. 1 shows their general location; grid references are given in Parnell
(1980).
At each site 25 flowering specimens were collected at random, pressed and taken back to the
laboratory for measurement. At some localities, seeds from another 25 randomly selected
individuals were collected. These were sown individually in potting compost in 10-cm flower-pots
and grown to maturity in a cold frame. Representative specimens of the wild populations are
deposited in ABD and/or TCD. No population of less than 100 individuals was sampled.
39 morphological measurements, including four derived ratios, were made on all Scottish
populations. Of these 39 measurements, 20 proved redundant due either to high correlation with
another measurement or lack of variation and it was found that the remaining 19 adequately
described a plant’s morphology. These are listed in Table 1 and some are illustrated in Fig. 2.
VARIATION IN JASIONE MONTANA 251
TABLE 1. CHARACTERS USED IN THE ANALYSIS OF BRITISH ISLES JASIONE MONTANA
. Plant height, measured from the base of the leaf-rosette to the top of the plant
. Plant width, the widest diameter of the leaf-rosette or the total spread of any decumbent stems
. Leaf length, measured on the fourth leaf down from the top of the main peduncle, is taken along the midrib
from the base
4. Leaf breadth (the maximum breadth of the same leaf used in 3)
5. The maximum diameter of the peduncle, taken from just under the involucral bracts (Fig. 2, No. 5)
6. The maximum diameter of the largest flowering head (Fig. 2, No. 6)
:
8
WN
. The maximum diameter of the involucre (Fig. 2, No. 7)
3. The maximum length of the outermost involucral bract, from the base to its widest point (Fig. 2, No. 8)
9. The maximum length of the pedicel (Fig. 2, No. 9)
10. The maximum length of the calyx (Fig. 2, No. 10)
11. The maximum width of the calyx-teeth (Fig. 2, No. 11)
12. The number of flowers in the terminal inflorescence
13. The number of flowering stems
14. The ratio of plant height to width
15. The ratio of leaf length to breadth
16. The ratio of involucre to inflorescence diameter
17. The hairiness of the hypanthium. Recorded as present or absent
18. The hairiness of the plant. Recorded on a 0-5 subjective scale with O=No hairs, 1=very few scattered
hairs, 2=sparsely hairy, 3=moderately hairy, 4=hairy, 5=very hairy
19. The habit of the plant. Recorded as either decumbent or upright
Three characters, numbers 2, 12 and 13 (Table 1) were found to be non-normally distributed.
These data were normalized using either a square root or a Log), transformation. The transformed
values were used in all further calculations. Three other characters, numbers 17, 18 and 19, were
not measured on a continuous scale and were therefore not used in the multivariate analysis.
The size and shape of the involucral bracts has been a commonly used diagnostic feature both of
British varieties and continental subspecies of J. montana and also of other Jasione species.
However, analysis indicated that involucral bracts showed considerable variation in size, shape,
toothing and colour both within a single plant and also between plant* within a single population.
The bracts illustrated in Fig. 3 are all taken from the main inflorescence of four adjacent
individuals from a single Irish population of J. montana. Small variations in bract characters are,
therefore, unlikely to be of taxonomic importance though large differences in the outer involucral
bracts, e.g. ovate vs lanceolate, may still be of use.
Preliminary analysis also showed that there was continuous variation in height and leaf
dimensions between and within populations. Therefore plants formally distinguished as J. montana
var. major Mert. & Koch (mostly on the basis of their large size (Pugsley 1921)) do not warrant
recognition. Equally, dwarf plants formally distinguished as J. montana var. maritima Bréb.
(Pugsley 1921) were mostly found to occur intermingled with plants of larger size. A single
population (C10) consisting entirely of dwarf plants and only separable on size characteristics from
J. montana var. montana was found. Continuous variation also occurred in the density of the
indumentum between plants within a population, some being almost glabrous whilst others were
hirsute.
NUMERICAL ANALYSIS OF POPULATIONS
The main numerical analysis was carried out on 1025 plants representing 41 of the 59 sampled
populations. Additional analysis, using means of population characters, was carried out on the 32
populations of J. montana sensu stricto.
I used two fundamentally different multivariate techniques to analyse the data. The first,
Principal Components Analysis (PCA), I used to examine patterns of variation or groupings of
plants present in both meaned and raw data. Technically, PCA aims to produce axes which
summarize the raw data and against which combinations of plants or other entities can be plotted.
Its interpretation is often difficult because simple biaxial plots, which are the type most often used,
are limited views of any variation pattern, and their interpretation depends on the percentage of
the variance accounted for by each axis. The programme used was BMDP4M (Frane & Jennrich
202 J. PARNELL
Figure 2. Morphological measurements made on the flowers and inflorescences of J. montana. The numbers
refer to the corresponding characters described in Table 1.
AAAQOOOM OOO 406
OASOOSDSOVO
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FiGureE 3. Bract outlines, from the outermost (left-hand) to the innermost (right-hand) bract, for four adjacent
individuals from a single population of J. montana at Carnsore point, Ireland (GR 31/12.04).
1977). The second technique, Discriminant Analysis (DSC), was used to test whether groups of
individuals assigned to different varieties of J. montana could be separated on the basis of a
particular set of morphological characters. It was also used to determine if populations retained
their distinctive characters (i.e. group membership) on cultivation. DSC allows any change in
morphology to be reflected as a change in group membership. Technically DSC can, as Marriot
(1974) states, be thought of as a multivariate extension of analysis of variance. The computer
analysis used was published in the SPSS series (Nie et al. 1975).
In addition to the multivariate statistics used, mean values and confidence limits were calculated
for each of the characters measured. These univariate statistics were used to quantify and express
differences in the morphology of the groups seen on PCA and discriminated by DSC.
VARIATION IN JASIONE MONTANA 253
RESULTS OF PCA, BASED ON ANALYSIS OF INDIVIDUAL PLANTS
Altogether, 16 axes were required to explain fully all the variance in the sample. Marriot (1974)
points out that axes with eigenvalues less than unity account for less of the variance than a single
individual variable and do not therefore represent a condensation of the data and should not be
plotted. In this analysis only four eigenvalues had values greater than unity and only three
substantially so. Axes one, two and three accounted for 40%, 12% and 10% of the total variance
respectively. Axes one and two are plotted in Fig. 4.
Membership of previously accepted groups is indicated on the PCA plot (Fig. 4). The groups are
defined partly on groupings noted in the field and partly on the basis of previous descriptions. The
groups used were:
1. J. montana var. litoralis Fr., defined on the basis of its decumbent habit, short stature (<20 cm),
small lanceolate leaves (0.5—1 X0.2 cm) with unthickened margins, and involucral bracts which are
rhomboidal, acute or obtuse and sub-entire and which do not exceed the inflorescence;
2. J. montana var. latifolia Pugsley, of upright habit, 10-30 cm tall and distinguished by its large
sub-spathulate leaves (1—2.5 x0.4—0.9 cm) which often have thickened margins, and its involucral
bracts which are broadly ovate, acute or obtuse with obscure teeth and which exceed the
inflorescence;
3. J. montana var. montana, containing all other plants of J. montana.
In Fig. 4, those plants which showed maximum overlap between the varieties, thereby
emphasising any overlap between groups, have been plotted. Despite this, it is evident that Fig. 4
offers at least moderate support for there being a separation between the varieties recognized.
RESULTS OF PCA, BASED ON ANALYSIS OF POPULATIONS OF J. MONTANA VAR. MONTANA
PCA of J. montana var. montana populations (Fig. 5) showed that there are morphological
differences between populations of this variety from different parts of Britain. For example, plants
from Scotland tend to have more flower stalks, smaller bracts and to be taller in relation to their
basal spread than plants from the rest of the country. Some individual morphological
characteristics also showed variation between different geographical regions, e.g. more than 55%
of flowers of plants from Cornwall, Devon and Dorset had a glabrous hypanthium, decreasing to
38% from south Wales and only 18% of those from north Wales and southern Scotland. Such
differences are relatively minor in comparison to those that exist between the different varieties of
J. montana detailed above and seem to be largely eliminated on cultivation under standard
environmental conditions, but they are evident in herbarium collections.
RESULTS OF DSC, BASED ON ANALYSIS OF INDIVIDUAL PLANTS
An objective measure of the separation between the groups recognized in the PCA of individual
plants was obtained by discriminant analysis. Two significant discriminant axes were extracted.
The first and second discriminant functions respectively accounted for 67% and 33% of the total
variance, were highly significant (77=1519, p<0.01 and 7°=569, p<0.01) and therefore gave a
good summary of group differences. The F-values obtained for differences between the groups
were all highly significant at p<0.01 (i.e. between J. montana var. montana and J. montana var.
latifolia, between J. montana var. montana and J. montana var. litoralis, and between J. montana
var. litoralis and J. montana var. latifolia). Thus the groups tentatively identified on PCA are very
different from each other.
Fig. 6 shows the mean values for each character, together with the 99.9% confidence limits for
all characters measured for all three varieties. The most important difference between J. montana
var. /atifolia plants and those of the other two varieties lies in the possession of bracts which are
significantly longer to their widest point (Fig. 6A), in the larger size of the inflorescence and
diameter of the peduncle (Fig. 6B, 6C & 6D), in their taller, narrower growth-form (Fig. 6E, 6K &
6L) and generally larger dimensions throughout (Fig. 6F, 6G, 6H, 61 & 6J). Plants of J. montana
var. litoralis differ from plants of the other two varieties in their greater breadth, which is almost
equal to their height (Fig. 6E & 6L), and their smaller flowering heads (Fig. 6B & 6C).
Thus these multivariate analyses have supported the separation of British Isles J. montana into
three varieties; J. montana var. montana, J. montana var. litoralis and J. montana var. latifolia and
have shown that morphological differences exist between populations of J. montana s.s. from
different areas of Britain.
254 J. PARNELL
Axis twa®20
FiGure 4. Principal components analysis on all plants sampled. The 500 plants giving the greatest degree of
overlap are plotted. Axis One and Axis Two account for 40% and for 12% of the total variance respectively.
©O=J. montana var. litoralis, \=J. montana var. latifolia, @=J. montana var. montana.
VARIATION IN JASIONE MONTANA 259
. Axis two
Axis three
Ficure 5. Principal components analysis of J. montana var. montana populations. Axes One, Two and Three
account for 52%, 15% and 11% of the total variance respectively. @=Scottish material, O=Welsh material,
A=Dorset material, @=Cornish and Devon material.
J. PARNELL
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VARIATION IN JASIONE MONTANA 257
STABILITY OF THE MORPHOLOGICAL GROUPINGS RECOGNIZED
A number of populations representing all the varieties of J. montana were grown under uniform
environmental conditions. Plants were inspected and morphological measurements made on
them at maturity. These measurements were plotted in the form of polygraphs and were also
used in conjunction with the DSC functions already obtained from the wild populations (see
above) to estimate the genetic component of the variation pattern of the varieties.
Representative polygraphs of the populations before and after cultivation are shown in Fig. 7.
Wild populations are designated S16 (J. montana var. montana), 105 (J. montana var. latifolia),
S05 and $14 (J. montana var. litoralis), and C10 (J. montana var. maritima). After cultivation
under experimental conditions, the populations bear the prefix E, e.g. ES16.
J. montana var. litoralis populations (SOS &S14) vary in their response to cultivation (ESOS &
ES14) (Fig. 7). ESOS plants maintain their diagnostic characteristics on cultivation, e.g. low
height to width ratio but ES14 plants do not and come to resemble the experimental population
of J. montana var. montana (ES16). Thus it seems that there are genetically determined plants
assignable to J. montana var. litoralis but that there are also plants which, though similar in
morphology, are in reality ecads or phenotypic copies of that taxon. This pattern of results was
repeated in other tested populations of J. montana var. litoralis. Wild populations of J. montana
var. maritima (C10) changed greatly on cultivation (EC10, Fig. 7) and came to resemble J.
montana var. montana (ES16). J. montana var. latifolia (105) remained distinct on cultivation
(EI0S, Fig. 7).
DSC analysis was used to test whether population means had shifted under cultivation, i.e.
whether wild populations and the cultivated populations derived from them would be classified
into the same group by DSC. DSC supported the conclusions drawn from the polygraphs by
assigning all plants of EIO5 to J. montana var. latifolia, ESOS to J. montana var. litoralis and
ES16, ES14 & EC10 to J. montana var. montana.
Thus these experiments have shown firstly that plants grown from seed taken from natural
populations of J. montana var. litoralis and J. montana var. latifolia maintain their distinctive
morphologies on cultivation in uniform conditions and that the characteristics used to distinguish
them are therefore genetically determined; secondly that phenotypic copies of J. montana var.
litoralis exist and finally that plants of J. montana var. maritima do not maintain their dwarf
stature on cultivation.
CROSSING EXPERIMENTS AND OBSERVATIONS ON POLLEN FERTILITY
Crossing experiments and observations on pollen fertility (Parnell 1980, 1982d, 1985) revealed no
evidence of breeding barriers between or within any of the varieties. Artificial geitonogamous
pollination, i.e. pollination between different flowers on the same inflorescence, produced a
moderate amount of seed whereas strictly autogamous pollination did not. The distance seeds are
likely to be dispersed in natural populations was likely to be less than 1.5 m, serving to restrict
gene flow. Pollen infertility was variable both within an inflorescence and between different
plants within a population and was also in some populations surprisingly high (c.36%), the
infertility possibly acting as an insurance against excessive inbreeding (Parnell 1982d). So, despite
the obvious physical adaptations of J. montana flowers to outcrossing (Knuth 1909; Parnell
1982d), successful inbreeding can and probably does occur at a fairly high frequency in natural
populations.
THE VARIATION OF BRITISH ISLES MATERIAL
Experimental analysis has shown that it is possible to recognise taxonomically distinct, genetically
determined, fully interfertile varieties of J. montana in the British Isles — J. montana var. litoralis
and J. montana var. latifolia. The former is an ecotype confined to sand-dunes (Parnell 1980)
whilst the latter grows on cliff-tops and roadside banks near the sea. Scattered populations of
both occur throughout the British Isles. In general, Parnell (1982a) showed that J. montana is
cytologically very uniform with n=6 or 2n=12 but that J. montana var. litoralis is unique in the
occasional possession of either one or two 6 chromosomes. Neither ecotype has a geographically
circumscribed distribution and therefore neither warrants subspecific status.
Plants formerly recognized as J. montana var. major and J. montana var. laevis cannot be
separated from J. montana var. montana and should not be maintained at varietal rank. Equally,
258 J. PARNELL
$e
= *< E105
Ficure 7. Polygraphs illustrating the morphological differences between and within populations of J. montana
s.1. after growth under experimental conditions (prefixed by E) as compared with wild material. Wild
populations are J. montana var. montana (S16), J. montana var. latifolia (105), J. montana var. litoralis (SOS &
$14) and J. montana var. maritima (C10).
> ae
VARIATION IN JASIONE MONTANA 259
plants of J. montana var. maritima fail to maintain their distinctive features on cultivation and
therefore do not warrant recognition.
The geographical differences noted between northern and southern J. montana are small in
comparison to the genotypical differences noted above and do not warrant formal taxonomic
recognition.
The possibility that J. montana can act as an inbreeder has, surprisingly, not been suggested
before. Undoubtedly geitonogamy must be of fairly common occurrence in any plant which bears
more than a single flower and is not an obligate inbreeder. Why, in J. montana, such pollination
should result in successful seed set whereas autogamous pollination does not is unknown. What is
obvious though is that the morphology of J. montana belies its ability to self-fertilize. Such self-
fertilization can be, at least to some degree, balanced by genetic recombination. As J. montana has
a low chromosome number (2n=12) and rather small chromosomes with mostly few, terminal
chiasmata (c. 1.4 per bivalent (Parnell 1982a)), it is clear that its chromosomal system is also acting
to restrict rather than increase genetic variability. The occurrence of such physical and genetic
inbreeding mechanisms allows successful recombinants to be conserved and together with
restricted gene flow may allow the establishment of small morphological differences in plants from
northern and southern Britain or northern and southern Europe.
VARIATION IN J. MONTANA THROUGHOUT THE REMAINDER OF ITS RANGE
CONTINENTAL MATERIAL
Analysis of herbarium specimens and natural plant populations of J. montana showed that the
small morphological differences seen between J. montana from northern Scotland and south-
western England (see above) were paralleled in material from continental Europe. For example, J.
montana from southern Scandinavia is similar to Scottish material whereas that from central and
southern Spain differs in its larger, often more deeply toothed bracts and pale green leaves.
However, numerous intermediates connect northern and southern European specimens and there
is no sharp discontinuity between them.
Spanish J. montana is very variable and two subspecies have been recognized, viz. J. montana
subsp. echinata (Boiss. & Reuter) Rivas-Martinez and J. montana subsp. blepharodon (Boiss. &
Reuter) Rivas-Martinez. The former subspecies is distinguished by its usually pale green, often
hispid, leaves, purple, very deeply toothed and often strongly papillose outer involucral bracts and
hispid calyx. However numerous intermediates between this and J. montana s.s. occur in Spain and
also in Sicily, e.g. plants from the Sierra Marenna, near Cordoba (Spain), collected by Wilmott in
BM have all of the above features except for the toothed purple bracts. The combination of
characters defining J. montana subsp. echinata also breaks down because in the rest of the range of
J. montana, e.g. in Britain, rare individuals occur which have the bract characteristics of J.
montana subsp. echinata but none of the other features (Parnell in ABD). No chromosome counts
are available for J. montana subsp. echinata. J. montana subsp. blepharodon is distinguished on the
basis of its long (>2.7 mm), often basally ciliate calyx teeth, its generally sub-sessile flowers and
large, ovate, more or less entire bracts. This combination of characters is unique. However, as
both the number and position of the ciliate hairs on the calyx teeth is very variable (a situation
similar to the variation in pubescence of the hypanthium of British J. montana) as is also the
relative size and shape of the involucral bracts, it is clear that this combination of characters also
breaks down. Finally, intermediates between these two subspecies also occur, e.g. Reverchon 195
from Andalucia in E has ciliate calyx teeth and sub-sessile flowers but deeply toothed, purplish,
strongly papillose outer involucral bracts. Bjorkqvist et al. (1969) showed that J. montana subsp.
blepharodon had chromosomes which were numerically and morphologically identical with J.
montana S.s.
Specimens of J. montana from the cliffs and dunes of the Atlantic coasts of Finisterre (Spain)
and the Gironde (France) have often been distinguished as J. montana var. sabularia Cout.
Typically, these decumbent plants (Fig. 8) bear short, imbricate, strigose leaves which have a
markedly revolute margin. This combination of characters is unique and seems to occur only in
specimens from coastal habitats in this region. Unfortunately the lectotype (in BM) and original
description of this taxon (Coutinho 1913) neither correspond to the description given above nor in
260 Js PARNBEL
[ 1¢m
Jasione sontana L. subsp. @ontana var.
umbruans TF Crenars
Novcory re
DET. J. PARNELL 18/.9/1984
FicureE 8. Holotype of J. montana L. subsp. montana var. imbricans J. Parnell. Finisterre, 1928, Lacaita 32283
(BM).
fact refer to a distinguishable taxon. Thus J. montana var. sabularia Cout. has been consistently
misinterpreted. J. montana var. sabularia auct. non Cout. has the same habit as J. montana var.
litoralis and is probably closely related to it.
Thus, though there is considerable morphological variation in continental European J. montana,
the differences between specimens are often dependent on their distance apart with seemingly
acute differences breaking down when plants from intermediate geographical localities are
considered. J. montana var. sabularia Cout. is indistinguishable from J. montana s.s. However, J.
montana var. sabularia auct. non Cout. is distinct, and bears a close relationship to J. montana var.
litoralis.
NORTH AFRICAN MATERIAL
Jasione cornuta Ball is endemic to north-western Africa and differs from typical J. montana in its
long, apiculate, large involucral bracts, relatively broad, papillose leaves (Fig. 9) and occasionally
fimbriate calyx teeth. Some J. cornuta is different from any southern Spanish material
notwithstanding an obviously close relationship between them. It would seem that the apiculate
bracts and fimbriate calyx teeth of J. cornuta are a yet more extreme example of the great variation
in bract and calyx characters that occurs throughout the range of J. montana, being especially
noticeable in southern Spain and exemplified there by J. montana subsp. blepharodon. The broad,
papillose leaves of J. cornuta also indicate a close relationship to J. montana subsp. echinata where
these features occur in a less extreme form. Some specimens of J. cornuta are also obviously closely
related to J. corymbosa Poiret and intermediates between the two species occur.
J. cornuta and southern Spanish J. montana share a number of morphological features. However
the morphological differences between J. cornuta and J. montana s.\., combined with the
geographical isolation of the former, mean that J. cornuta is best treated as a subspecies of J.
montana, as suggested by Greuter (1981), though no reasons are given.
RELATED SPECIES
Both J. corymbosa Poiret ex Schultes and J. penicillata Boiss. have been treated as subspecies of J.
montana by Greuter (1981) and Rivas Martinez (1976) respectively. J. penicillata is strongly
VARIATION IN JASIONE MONTANA 261
Ficure 9. Plants of J. montana L. subsp. cornuta (Ball) Greuter & Burdet, collected from Morocco illustrating
its long apiculate bracts and broad leaves. Morocco, 13/4/1969, P. & J. Davis D 49337 (E).
pubescent, probably annual, endemic to the Sierra Tejeda and surrounding mountains and differs
from all non-teratological J. montana in being usually unbranched, though the lectotype in G is
highly branched, in bearing leaves all the way up the peduncle (Fig. 10), and in its linear-
spathulate, apically villous calyx teeth. Its relatively isolated geographical location and unusual
morphology indicate that it is best maintained at specific rank. J. corymbosa is typically a stout
glabrous or hispid annual which is again leafy almost to the apex of the peduncle (Fig. 11), has
subulate, villous calyx teeth and differs from all J. montana in usually bearing short, fastigiate
inflorescence branches. It occurs in southern Spain and Morocco. Cytologically it is similar to J.
montana s.s. with 2n=12 and has a similar karyotype (Parnell unpublished). In its common state it
is so distinct from J. montana that it would seem best to retain it at specific rank.
J. heldreichii Boiss. & Orph. is morphologically similar to J. montana, differing largely in its
lanceolate, aristate, very deeply toothed, involucral bracts. The bract shape and the degree of
toothing are so different from that of J. montana that J. heldreichii plants are always easily
distinguishable. Some plants of J. heldreichii, from north-western Turkey, can be separated from
the rest because of their short, triangular calyx teeth and very strongly papillose and marginally
thickened leaves and involucral bracts. Such plants only occur in the same region of northern
Turkey as J. heldreichii s.s. and so only warrant varietal rank (Fig. 12). J. heldreichii has 2n=12
and a similar karyotype to J. montana s.s. (Contandriopoulos 1966).
[1 om
FicurE 10. Plants of J. penicillata collected from Andalucia. Andalucia, 1889, Reverchon 194 (BM).
262 J. PARNELL
Figure 11. Plants of J. corymbosa collected from Morocco illustrating their generally fastigiate branching.
Morocco, 29/6/1974 Reading University/B.M. Exped. 1112 (BM).
Figure 12. Holotype of J. heldreichii Boiss. & Orph. in Boiss. var. papillosa J. Parnell. Istanbul, 8/8/72, Uslu
2070 (E).
VARIATION IN JASIONE MONTANA 263
ENUMERATION OF TAXA
Only the synonyms of taxa recognized at the varietal level or above are cited. Sources for
chromosome counts are given in Parnell (1982c) except where indicated.
1. JASIONE MONTANA L., Sp.Pl., 928 (1753).
Jasione vulgaris Gaterau, Descr. Pl. Montauban, 153 (1789).
Jasione undulata Lam., Encycl. Méth. Bot., 215 (1789), pro parte, excl. var. B.
Jasione appressifolia Pau in Not. Bot. Fl. Esp., 1: 19 (1887).
Jasione espadanae Pau in Not. Bot. Fl. Esp., 1: 19 (1887).
Jasione adpressifolia Pau, orth. mut. ap. Willk., Suppl. Prodr. Hisp., 126 (1893).
Jasione mediterranea Rouy, Fl. Fr., 10: 92 (1908).
Jasione hungarica Simonkai, nom. nud.
a. Subsp. MONTANA
1. Var. MONTANA
Jasione montana var. ramosa Latourette, Chloris Ludgensis, 25 (1785-1787), nom. nud.
Jasione montana var. prolifera Latourette, Chloris Ludgensis, 25 (1785-1787), nom. nud.
Jasione montana var. major Mert. & Koch in Rohling, Deutschl. Fl., 147 (1826).
Jasione montana var. laevis Duby, Bot. Gall. , 311 (1828).
Jasione montana var. hirsuta Duby, Bot. Gall. , 311 (1828).
Jasione montana var. prolifera A.DC., Monogr. Campan., 102 (1830).
Jasione montana var. maritima Bréb., Fl. Normand., 179 (1835).
Jasione montana var. glabra Petermann, Fl. Lips. Excurs., 168 (1838).
Jasione montana var. stolonifera DC., Prodr., 415 (1839).
Jasione montana var. gracilis Lange, Pug. Pl. Hisp., 155 (1861).
Jasione montana var. genuina Willk. in Willk. & Lange, Prodr. Fl. Hisp., 1: 126 (1870).
Jasione montana vat. gracilis Timbal-Légrave in Bull. Soc. Sci. Phys. Nat. Toulouse, 3: 419 (1875-
1876).
Jasione montana var. typica Trautvetter in Acta Horti Petrop., 6: 44 (1879).
Jasione montana var. hispida G. Beck., Fl. Nieder-Osterr., 1110 (1890).
Jasione montana var. boraei Rouy, Fl. Fr., 10: 91 (1908).
Jasione montana var. timbali Rouy, Fl. Fr., 10: 91 (1908).
Jasione montana var. sabularia Coutinho, Fl. Port., 603 (1913).
Jasione pyrenaica var. semiglabra Sennen, Plantes de Pyrenées-Orientales , 5726 (1927), nom. nud.
Jasione montana var. glaberrima Podpéra in Polvika, Domin & Podpéra, Klié k uplné kvétené
republiky C.S.R., 542 (1928).
More or less erect, sparingly branched (c. 2 flowering stems), usually villous biennials (rarely
annual or short lived perennial) 2-60 cm tall. Stems usually leafless in their upper and leafy in their
lower halves. Aerial leaves (0.90.2 cm) linear-oblong to lanceolate, always entire and often
undulate (rarely crenate), the margin, unthickened, often weakly papillose and sometimes ciliate.
Peduncles thin (<1 mm), outer involucral bracts, dark green, occasionally weakly papillose, ovate,
parabolic, inner bracts, green or colourless, linear to lanceolate sometimes ovate, sometimes entire
but more often crenate, serrate or laciniate, shorter than the flowers. Calyx teeth usually c. 2 mm
long, subulate, green and often pubescent but never ciliate or fimbriate. Corolla usually blue,
sometimes white or pink, <40 flowers in the primary inflorescence. Various abnormal forms of this
variety occur and are described in Parnell (1982b, 1982c) and Scannell (1977). Distribution:
throughout the range of the species excluding North Africa. n=6, 2n=12.
ii. Var. ECHINATA (Boiss. & Reuter) Willk. & Lange, Prodr. Fl. Hisp., 2: 282 (1870).
Jasione echinata Boiss. & Reuter, Pugillus, 73 (1852). Type: Gauchin, 1837, Boissier (Lectotype:
G! fide Burdet, Charpin & Jacquemoud, in Candollea, 38: 414). Jasione montana subsp. echinata
(Boiss. & Reuter) Rivas-Martinez in Publ. Inst. Biol. Appl. Barcelona, 42: 122 (1967).
Jasione montana var. dentata sensu Boissier, Fl. Orient., 3: 885 (1875) et auct.
264 J. PARNELL
Jasione montana var. arenaria Boiss. in sched.
Jasione stricta Pomel, Nouv. Mat. Fl. Atl., 1 (1874).
Jasione montana var. cartiliginea Sennen, Plantes d’ Espagne, No. 3461 (1918), nom. nud.
Plants are erect and sparingly branched 2-15 (—30) cm tall. Differs from var. montana in the
usually pale green leaves, and pale green, often hispid, often generally purplish, long, thin, very
deeply toothed and strongly papillose outer involucral bracts. Distribution: mainly in south-
western Europe (Spain) but also in North Africa, Sicily, Italy and Corsica.
iii. Var. BRACTEOSA Willk. in Bot. Zeit., 5: 863 (1847).
Jasione blepharodon Boiss. & Reuter, Pugillus, 172 (1852). Type: Gibraltar, 1849, Boiss. & Reuter
(Lectotype: G! fide Burdet, Charpin & Jacquemoud, in Candollea, 38: 412 (1983)). Jasione
montana subsp. blepharodon (Boiss. & Reuter) Rivas-Martinez in Candollea, 31: 113 (1976);
Jasione corymbosa Poiret ex Schultes in Roemer & Schultes var. blepharodon (Boiss. & Reuter)
Batt., Fl. Alger. Dicot., 571 (1889).
Jasione montana var. littoralis Boiss., Voy» Bot. Midi. Esp., 2: 396 (1839) non J. montana var.
litoralis Fr., Nov. Fl. Suec., 269 (1814). (I have been unable to find any specimens of this taxon
and Burdet et al. (1983) have been unable to trace any in G. However, Boissier’s original
description and subsequent synonymy (Boissier & Reuter (1852)) makes this the most likely
resting place for it.) N
Jasione baetica Rodrig. in sched.
Differs from var. montana in being usually annual with papillose leaf-margins, ovate outer
involucral bracts, mostly subsessile flowers and long (>2.7 mm), ciliate calyx-teeth. Distribution:
mainly in south-western Europe (Spain). 2n=12.
iv. Var. LATIFOLIA Pugsl. in J. Bot., Lond., 59: 215-216 (1921). Type: Bray Head, Co.
Wicklow, 1881, Britten & Nicholson (Lectotype: BM!, designated here).
Jasione montana var. megaphylla Vicioso in Anal. Jardin Botanico de Madrid, 6: 79 (1965).
Differs from var. montana in its lanceolate-spathulate leaves with thickened but not strongly
papillose leaf-margins, thickened peduncles (c. 1.5 mm), large inflorescences with ovate involucral
bracts projecting beyond the edge of the inflorescence and with usually >60 flowers in the main
inflorescence. Distribution: scattered throughout Europe. n=6.
v. Var. LITORALIS Fr., Nov. Fl. Suec., 269 (1814). Type: Halmstad par., Halmstad Holland,
1814, E. Fries (Lectotype: UPS! Lower specimen, designated here).
Jasione montana var. littoralis Koch, Syn. Fl. Ger., 463 (1837).
Jasione montana var. tenella Petermann, Fl. Lips. Excurs., 168 (1838).
Jasione montana var. nana Boreau, Fl. Centre Fr., 2: 286-287 (1840).
Jasione montana var. nana Gren. & Godron, FI. Fr., 2: 398 (1850).
Differs from var. montana in having decumbent to ascending flowering stems. Plants are usually as
tall as they are wide with >8 flowering stems and small (0.5—2 cm), narrow (c. 2 mm wide) leaves.
Distribution: mainly on the coasts of Europe and Scandinavia (on sand) but also inland (on sand or
granitic rocks). n=6 or n=6+1f or n=6+2f.
vi. Var. IMBRICANS J. Parnell, var. nov.
Typus: Finisterre, 1928, Lacaita 33283 (Holotypus: BM). Fig. 8.
Jasione montana var. sabularia auct. non Cout.
A varietate montana habitu decumbent et a varietate montana et varietate litorali caulibus floriferis
paucis (1-3), foliis brevibus strigosis imbricatis marginibus saepissime revolutis, differt.
Differs from var. montana in its decumbent habit and from both var. montana and var. litoralis in
VARIATION IN JASIONE MONTANA 265
its few (<4) flowering stems with short, imbricate, strigose leaves which usually have a strongly
revolute margin. Distribution: coasts of Finisterre (Spain), Gironde (France).
b. Subsp. CORNUTA (Ball) Greuter & Burdet in Willdenowia, 11: 40 (1981).
Jasione cornuta Ball in J. Bot., Lond., 11: 373 (1873). Jasione corymbosa subsp. cornuta (Ball)
Murbeck in Jahandiez & Maire, Cat. Pl. Maroc., 737 (1934).
Differs from subsp. montana in being usually annual with pale green leaves with strongly papillose
margins, ovate, long acuminate outer involucral bracts, and linear/linear-spathulate aristate and
often weakly fimbriate calyx teeth. Distribution: North Africa only.
2. JASIONE PENICILLATA Boiss., Elenchus, 63 (1838). Type: Sierra Tejeda, 1837, Boiss.
(Lectotype: G! fide Burdet, Charpin & Jacquemoud in Candollea, 38: 416 (1983)).
Jasione blepharodon subsp. penicillata (Boiss.) Rivas Goday in Bol. Soc. Brot., ser. 2, 47: suppl.
168 (1973). Jasione montana subsp. penicillata (Boiss.) Rivas Martinez in Candollea, 31: 113
(1976).
Differs from J. montana in being an annual or short-lived biennial 1-10 cm tall without a distinct
leaf rosette but with lanceolate, entire aerial leaves 0.9 xX 0.2 (—0.3) cm with a thin, neither
undulate nor papillose margin and with the peduncle usually leafy up to the capitulum.
Distribution: Sierra Tejeda and surrounding mountains, Spain.
3. JASIONE CORYMBOSA Poiret ex Schultes in Roemer & Schultes, Syst. Veg., 5: 474 (1819).
Type: Tangier, 1794-1799, Broussonet (Lectotype: M!, designated here).
Jasione corymbosa Poiret in Lam., Encycl. Méth. Bot. Suppl., 3: 130 (1813), nom. prov.; Jasione
montana subsp. corymbosa (Poiret) Greuter & Burdet in Willdenowia, 11: 40 (1981) nom.
illegit.
Differs from J. montana in being usually 3-10 (—15) cm tall, often fastigiately branched and with
spathulate to ovate or obovate aerial leaves (6 x 2-4 mm) generally present up to the capitulum.
2n=12 (Parnell, unpublished). Distribution: southern Spain and Portugal and north-western
Africa.
1. Var. CORYMBOSA.
As above and also usually villous.
ii. Var. GLABRA (Durieu ex Boiss. & Reuter) J. Parnell, stat. et comb. nov.
Jasione glabra Durieu ex Boiss. & Reuter, Pug. Plant. Nov., 72 (1852). Type: Oran, 1840-1842,
Durieu (Lectotype: G!, designated here). Jasione corymbosa subsp. glabra (Durieu ex Boiss. et
Reuter) Batt. in Batt. et Trabut, Flore de l’Algerie, 570-571 (1888-1897).
Jasione arenaria Salzm. in sched.
Differs from var. corymbosa in being spindly and more or less glabrous with ovate involucral
bracts. Confined to sand-dunes and connected by intermediates to J. cornuta.
4. JASIONE HELDREICHI Boiss. & Orph. in Boiss., Diag. Pl. Nov., 3(6): 120 (1859). Type:
Macedonia, 1857, Orph. (Lectotype: G!, designated here).
Jasione jankae Neilr., Aufz. Ungarn. Slavon. Gefasspfl. Nachtrage und Verbesserungen, 43-44
(1870).
Jasione glabra Vel. in Ost. Bot. Z., 34: 424-425 (1884).
Jasione heldreichii var. microcephala Vel., Fl. Bulg., 374 (1891).
Differs from J. montana in having lanceolate, aristate, deeply toothed or cut/slashed involucral
bracts. Biennial or perennial. Distribution: Balkan Peninsula and south-eastern Europe. 2n=12
(Contandriopoulos 1966).
266 J.. PARNELL
i. Var. HELDREICHII
Jasione montana var. dentata DC., Prodr., 415 (1839), non Boissier, Fl. Orient., 3: 885 (1875) et
auct.
As above and with weakly papillose, slightly thickened leaf margins and green involucral bracts
with long triangular calyx teeth.
ii. Var. PAPILLOSA J. Parnell, var. nov.
Typus: Istanbul, 1972, Uslu 2070 (Holotypus: E!). Fig. 12.
A var. heldreichii foliis bracteisque involucri fortissime papillosis, marginibus conspicue
incrassatis, bracteisque pagina interiore purpureis et dentibus calycis breviter (nec longe et
anguste) triangularibus, differt.
Differs from var. heldreichii in its very strongly papillose and marginally thickened leaves, and
involucral bracts which are also distinctly purple on their inner surface and in having short
triangular calyx teeth. Distribution: north-western Turkey.
ACKNOWLEDGMENTS
I thank Dr C. C. Wilcock, for general help and encouragement in the biometric part of this work
and for reading an early draft of this manuscript, as did Professor D. A. Webb who also provided
the Latin descriptions; Professor D. H. S. Richardson and Dr D. L. Kelly who also commented on
the draft and the curators of the following herbaria who kindly lent me specimens: ABD, BM,
DBN, E, G, K, M, P, PAD, RNG, UPS. Some of this work was undertaken during the tenure of a
Science Research Council Studentship.
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GREUTER, W., ed. (1981). Med-checklist Notulae, 3. Willdenowia, 11: 23-43.
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Marriot, F. H. C. (1974). The interpretation of multiple observations. London.
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Barcelona, 42: 107-126.
VARIATION IN JASIONE MONTANA 267
Rivas-MarTINEZ, S. (1976). De plantis Hispaniae notulae systematicae, chorologie et ecologicae, 1. Candollea,
$1: 111-117;
Rivas-MarTINEZ, S. (1979). Brejoales y jarales de Europa occidental (Revisi6n Fitosociolégica de las clases
Calluno-Ulicetea y Cisto-Lavanduletea). Lazoroa, 1: 5-127.
SCANNELL, M. J. P. (1977). Dwarf Jasione montana. Watsonia, 11: 431.
ScHMEJA, O. (1931). Beitrage zur Kenntnis der Gattung Jasione. Beih. bot. Centralbl., 48: 1-51.
Tutin, T. G. (1976). Jasione L., in Tutin, T. G. et al., eds. Flora Europaea, 4: 100-102. Cambridge.
(Accepted July 1985)
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Watsonia, 16, 269-281 (1987) 269
The pattern of morphological variation in the Salicornia
europaea L. aggregate (Chenopodiaceae)
M. J. INGROUILLE and J. PEARSON
Department of Biology, Birkbeck College, Malet Street, London, WCIE 7HX
ABSTRACT
A numerical taxonomic analysis of diploid Salicornia L. (Chenopodiaceae) plants from four salt marshes in W.
Sussex (v.c. 13) and Essex (v.cc. 18 & 19) was carried out by minimum variance clustering (Ward’s Method)
and principal components analysis. The pattern of morphological variation both between and within marshes
provides no evidence for the separate recognition of Salicornia europaea L. and S. ramosissima J. Woods.
INTRODUCTION
In the British flora, the distinction between perennial and annual species, and between diploid
(2n=18) and tetraploid (2n=36) species of Salicornia L. (Chenopodiaceae) is well established. The
woodiness of Salicornia perennis Miller (=Arthrocnemum perenne (Miller) Moss) clearly identifies
it. Diploid and tetraploid annual species may be distinguished by a number of characters. The
number of stamens, the shape of the spike-segment, the length of the terminal spike, the angle of
the upper branches to the main stem and the size of the lateral flowers in relation to the central
flowers are all useful characters in the field (Ball & Tutin 1959; Ball 1964; Ball & Brown 1970). It is
within each ploidy level where difficulties arise, especially for the non-expert.
A major difficulty in Salicornia taxonomy emerged in the course of an ecophysiological study on
salt marshes in Norfolk, Essex, W. Sussex and Pembrokeshire. In a few locations it proved difficult
to distinguish taxa, especially the diploids, even when typical representatives of each species were
present within a single marsh.
The first edition of Flora of the British Isles (Clapham et al. 1952) records three diploid species in
the S. stricta Dumort aggregate (=S. europaea L.), and three diploid species in the S. prostrata
Pallas aggregate, as well as S. disarticulata Moss (=S. pusilla J. Woods). The second edition
(Clapham et al. 1962) reduces these species to four: $. europaea, S. ramosissima J. Woods
(including the S. prostrata agg.), S. pusilla and S. obscura P. W. Ball & Tutin. This trend in the
reduction of numbers of species recognized was outlined by Ball (1964) in Flora Europaea, where
S. obscura is recorded as a probable variant of S$. europaea. Of the three British diploid species
presently recognized, S. pusilla is distinctive in having a single floret in each cyme rather than the
normal three florets of S. europaea and S. ramosissima. This study is confined to the last two
species as these are often the most difficult to distinguish in the field.
Of the distinguishing features between S. europaea and S. ramosissima, emphasis is placed on
the width of the lower fertile segments of the terminal spike, the width of the scarious border and
the colour of the segments and cymes (Ball & Tutin 1959; Clapham et al. 1962; Ball 1964).
However, there is a range of overlap and variation in these characters that make their use difficult.
Furthermore, any segment colouring serves to highlight the scarious border and perhaps over-
emphasize its importance. It was found that identification of the two species at any one location
was generally reliable if the habitat and colour differences were distinct. For example, at Itchenor
in W. Sussex (v.c. 13), S. europaea is found on the lower marsh associated with bare ground and
the tetraploid S. dolichostachya Moss, and here its colour is generally dark green with some red
round the cyme. S. ramosissima is found on the middle and upper parts of the marsh, largely
associated with Puccinellia maritima (Hudson) Parl., with the segments and cymes coloured
purple. The two populations are separated by a broad, almost pure zone of Spartina anglica C. E.
270 M. J. INGROUILLE AND J. PEARSON
Hubbard. Less well-defined marshes, or marshes heavily dissected with creeks, often make such
identification more difficult because the habitats are less clearly defined. However, phenological and
genotypic studies have shown that differences between the two species at these levels do exist and
have emphasized the habitat differences (Jeffries et al. 1981; Jeffries & Gottlieb 1982). Jeffries &
Gottlieb (1982), using electrophoresis, found that six out of 30 isozymes tested in diploid species at
several locations were consistent with two distinct homozygous genotypes, representing S. europaea
and S. ramosissima.
The aim of this study was to carry out a numerical taxonomic study on the morphology of several
populations of both species and to determine whether inter-location and intra-location differences
could be detected. Any such investigation suffers from the difficulty of maintaining in cultivation, on
a regular basis, Salicornia plants. Although germination of seeds is easy, maintaining representative
plants in terms.of succulence and growth form has proved impossible so far. As a result this work has
been carried out entirely on collections of wild material. The lack of any testing of genotype —
environment interaction for any of the morphological characters measured is a severe drawback.
Of necessity, a classification of Salicornia must relate to the pattern of variation in nature since one
of the most important requirements of a general purpose classification is that species can be
identified by the field botanist. Although classifications can be erected for many purposes, a
classification in which it is impossible to identify named taxa except by specialist techniques has a
very restricted value.
~“
MATERIALS AND METHODS
SAMPLED POPULATIONS
Both S. europaea and S. ramosissima are recorded as being common and locally abundant or
dominant in the parts of Essex (Jermyn 1974) and W. Sussex (Hall 1980) that have been sampled.
Four salt marshes were sampled (Table 1), three in Essex (v.cc. 18 and 19) and one in W. Sussex
(v.c. 13). Marshes were chosen that include typical representatives of S. europaea and S.
ramosissima. At each locality the marsh was sampled at two sites at different levels. Plants were
selected from the lower marsh sites from the most seaward populations of the mudflats at St Peter’s,
Canvey and Itchenor. At Tollesbury, the lower marsh is foreshortened by erosion and at this location
plants were collected from the bare mud next to the eroded cliffs. Middle or upper marsh sites were
those areas with the most landward group of plants of a reasonable size.
Ateach site on the marsh a relatively small area was sampled, keeping to the same altitude as much
as possible. Underdeveloped or damaged plants and those outside the S. europaea group were
ignored. For the purpose of this study the S. europaea group was defined as in Flora Europaea (Ball
1964), consisting of all 3-flowered diploid plants.
Plants were collected over a two week period in late September and kept in polythene bags at 4°C
until being scored. It proved possible to keep plants fresh for 2—3 weeks in this way without any signs
of shrinkage or distortion. Scoring took place within this period.
TABLE 1. SITE DETAILS OF SAMPLED POPULATIONS
Number of Grid
Sampled site plants reference
N. Essex, v.c. 19
Tollesbury lower marsh 50 52/97.10
middle marsh 50
S. Essex, v.c. 18
St Peter’s lower marsh 50 62/03.08
middle marsh 50
Canvey lower marsh 24 51/82.83
middle marsh 50
W. Sussex, v.c. 13
Itchenor lower marsh 50 41/78.01
upper marsh 49
VARIATION IN SALICORNIA EUROPAEA 271
penultimate
fertile segment
middie floret
lateral floret 3rd fertile segment
i margin
scarious g 2nd fertile segment
sterile segment
5mm
TERMINAL SPIKE
FicurE |. Spike characters.
SCORING OF CHARACTERS
Plants were scored for 52 characters consisting of 47 metric characters covering all parts of the
plant (including the branching pattern, spike and floret size) and five multistate characters
assessing the coloration of the sterile and fertile segments and of the florets of the terminal spike
(Fig. 1). An additional 28 characters were derived as ratios of some of the initial 52 characters in
order to assess the shape of the whole plant and parts of the terminal spike (Table 2).
In choosing characters, no attempt was made to identify ‘useful’ characters, nor to examine
correlations between characters other than logically correlated characters, e.g. height of the spike
and height of each of the segments making up the spike, which were rejected.
DATA PREPARATION AND NUMERICAL ANALYSIS
Initial data preparation was carried out on a Vax — 11/750 computer. Data sets consisting of
matrices of individuals as Operational Taxonomic Units (OTUs) against characters were compiled.
Characters were encoded as scored without adjustment, with ratio characters calculated and added
to the sets.
Subsequent analysis was carried out on the following data sets:
a) all sites and populations consisting of 373 individuals and all 80 characters;
b) eight sets, one for each sampled site of all the individuals from a marsh and all 80 characters;
c) a reduced data set consisting of all 373 individuals and 36 characters — these being the ones
relating only to the terminal spike (characters 30-52 and 64-76).
The analysis was carried out on the reduced data set as well as the whole data set in order to
identify the effects of environmentally induced variability. One of the most obvious characteristics
of Salicornia plants is the variation in height and branching pattern of plants on different parts of a
marsh. This variability may be environmentally induced and leads to difficulty in the strict
comparison of branch characteristics between a plant with and without branches. Terminal spike
characters are strictly comparable (being always present) and may be the least environmentally
labile characteristics. Subsequent data preparation was carried out by using procedures of the
CLUSTAN package (Wishart 1978) release 2 version 1C on an Amdahl 470 computer. All further
analysis was carried out using this package of clustering and ordination procedures.
272 M. J. INGROUILLE AND J. PEARSON
TABLE 2. CHARACTERS SCORED FOR TAXOMETRIC ANALYSIS
GENERAL MORPHOLOGY
. Height of plant from rooting point to apex
. Height from rooting point to 1st branching point
. Number of internodes
. Length of Ist internode
. Length of 2nd internode
. Length of penultimate internode
. Length of ultimate internode
. Length of longest 1st (basal) primary branch
. Number of fertile segments in 1st primary branch
10. Number of sterile segments in Ist primary branch
11. Length of longest 2nd primary branch
12. Number of fertile segments in 2nd primary branch
13. Number of sterile segments in 2nd primary branch
14. Length of the longest penultimate branch
15. Number of fertile segments in penultimate branch
16. Number of sterile segments in penultimate branch
17. Length of ultimate branch
18. Number of fertile segments in ultimate branch
19. Number of sterile segments in ultimate branch
20. Distance from apex to apex of ultimate branch
21. Distance from apex to apex of lst primary branch
22. Number of secondary branches in Ist primary branch
23. Number of secondary branches in 2nd primary branch
24. Branch node with the most secondary branches
25. Maximum number of secondaries on a primary branch
26. Length of longest secondary branch
27. Number of fertile segments on the longest secondary
28. Number of sterile segments on the longest secondary
29. Length of the longest tertiary branch
OOnNNNKRWN PR
TERMINAL SPIKE CHARACTERS
30. Length
31. Number of fertile segments
32. Number of sterile segments
33. Maximum width of 3rd fertile segment
34. Width of middle floret
35. Width of 3 florets
36. Width across apex of 2nd fertile segment
37. Minimum width of 2nd fertile segment
38. Maximum width of 2nd fertile segment
39. Distance between florets on 2nd fertile segment
40. Distance from tip of 3rd fertile segment to apex of middle floret
41. Height of middle floret of 3rd fertile segment
42. Height of side floret of 3rd fertile segment
43. Height of triangular apex of 2nd fertile segment
44. Width of the scarious margin of 2nd fertile segment
45. Length of last sterile segment
46. Maximum diameter of penultimate fertile segment
47. Minimum diameter of penultimate fertile segment
48. Colour of sterile segment*
49. Colour of florets”
50. Colour of fertile segments*
51. Distribution of coloration**
52. Sterile segments***
*Green or yellow=0, diffuse pink=1, red=2. **Basal or even=0, apical=1. ***Not yellow=0, yellow=1.
VARIATION IN SALICORNIA EUROPAEA 273
TABLE 2 cont’d. CHARACTERS SCORED FOR TAXOMETRIC ANALYSIS
RATIO CHARACTERS
53. 1/11
54. (1-2-8)/3
55. 54/(445)
56. 54/(6+7)
57. 23/8
58. 24/(8+1—2)
59. 11/14
60. 11/20
61. 17/20
62. 29/11
63. 29/32
64. 33/(40+41)
65. 34/41
66. 35/42
67. 37/43
68. 37/38
69. 35/39
70. 45/(40+41)
71. 38/46
72. 37/47
73. 46/47
74. 36/43
75. 33/37
76. 33/38
77. 45/7
78. 38/1
79. 8/20
80. 9/20
Before carrying out any statistical procedures, the data were standardized by converting to
standard scores to give each measured character equal weighting. A similarity matrix with Squared
Euclidean Distance as the measure of similarity between individuals (OTUs) was calculated.
Subsequent ordination and clustering methods attempted to produce a low dimensional, but
undistorted, simplification of this matrix.
Initially a principal components analysis was carried out and scatter diagrams of the first few
components constructed.
Cluster analysis was performed using Ward’s Method as the criterion for the fusion of clusters.
Ward’s Method attempts to find a set of clusters with the minimum total within cluster variance.
Dendrograms were constructed to illustrate the cluster patterns produced.
The inclusion of so many characters, some of which are highly correlated, poses a number of
problems. The inclusion of correlated characters may be justified theoretically on the basis that
they each represent the phenotypic expression of a more basic but unmeasured pleiotropic gene or
genes. The inclusion of different measures imposes a kind of objective weighting; the more times
the expression of the pleiotropic gene is measured the more accurately it is assessed.
It is because of the problem of correlated characters that principal components analysis is so
useful. Here the basis of the method is to seek to simplify the data matrix by finding character
correlations. The results of a cluster analysis and a principal components analysis can validate each
other.
The inclusion of many characters increases the possibility that numerical noise will conceal any
patterns which exist but there is no way of choosing the best characters before the analysis is
carried out except in an arbitrary, subjective manner. The alternative, as reported here, is to carry
out the analysis, detect the best characters a posteriori, and then use these alone to see what
pattern of variation exists. The best characters can be chosen a posteriori on the basis of having low
intra-cluster variability and high inter-cluster variability, i.e. they are good cluster diagnostics.
274 M. J. INGROUILLE AND J. PEARSON
TABLE 3. PERCENTAGE VARIABILITY ACCOUNTED FOR BY THE COMPONENTS OF THE
PRINCIPAL COMPONENTS ANALYSIS
Full data set (80 characters) Spike data set (36 characters)
Component % variability cumulative % % variability cumulative %
1 18.26 18.26 23.77 23377
Z 13253 S179 P2325) 36.01
3 6.05 37.84 9.05 45.06
4 5.86 43.70 6.46 51152
{ { { { {
15 1.87 90.76
29 ; 0.76 90.43
RESULTS
ORDINATION
Principal components analysis, both of the whole data set and the reduced (spike) data set, failed
to identify any useful (i.e. discriminatory) components. The data were not summarized adequately
by any of the first few components although the first four components accounted for about half of
the total variability (Table 3). For example, the principal component scatter plots shown in Figs. 2
& 3 did not reveal any distinct groups or clusters. Rather there was a broad spread of individuals
with no obviously distinct modes.
CLUSTER ANALYSIS
Clustering by Ward’s method produced well defined clusters in both the whole and reduced data
sets (Figs. 4 & 5) but it is necessary to validate the clustering procedure. In part this can be
achieved by a comparison of different methods of analysis and by comparing the results of the one
COMPONENT 2
PRINCIPAL COMPONENT 1
FiGuRE 2. Principal component scatter diagram the first 2 components from an analysis of the whole data set.
Clusters labelled as in Fig. 4 and in Table 4.
VARIATION IN SALICORNIA EUROPAEA we ke)
COMPONENT 2
PRINCIPAL COMPONENT 1
Ficure 3. Principal component scatter diagram of the first 2 components from an analysis of the spike data set.
Clusters labelled as in Fig. 5 and Table 5.
Cluster
variance
73.2
56.1
39.0
Cluster
membership y y ] y y y ] Ty 9
Tollesbury
lower marsh = = 4 A = 1 1 216 10
middie marsh — = - 26 2 4 9 ——- 9
Saint Peter’s phy os
lower marsh = 46 1 i 1 1 ‘ oe Z
middle marsh ~— = 2 = - 15 29
Canvey =
14 = 3 - 4 2 ~ -
lower marsh ay
middie marsh 190 1 8 2 - 15 1 13
Itchenor
lower marsh 2 _ _ 1 44 7 - sg ae
upper marsh 1 _ 29 6 = 8 - 2- 3
Ficure 4. Phenogram of cluster analysis of whole data set. Clusters A, B and C defined in Table 4. Source of
cluster members at the 10 cluster phenon level indicated.
276 M. J. INGROUILLE AND J. PEARSON
Cluster
variance
105.0
80.4
55.9
31.3
6.8
1 2 3 4 5 6 7
Cluster
membership
Tollesbury V y
lower marsh 13 6 8 10 2 3 8
middie marsh 7 2 4 1 2 29 7
Saint Peter’s
lower marsh 2 1 = 47 =
middie marsh 28 2 14 1 - 3 2
Canvey
lower marsh 9 1 2 = 1 10 1
middle marsh 4 21 4 1 10 3 Tf
Itchenor
lower marsh = - = 1 Zs 5 44
upper marsh a 12 2 2 29 1 3
FiGurE 5. Phenogram of cluster analysis of spike data set. Cluster D and E defined in Table 5. Source of cluster
members at the 7 cluster phenon level indicated.
clustering method (Ward’s Method) on the different data sets. Since the clusters isolate different
portions of the principal components scatter plots (Figs. 2 & 3), there is a correspondence between
different methods of analysis. The phenograms based on the complete data set and the reduced
(spike) data set showed a good correspondence especially at the lower levels of clustering, and
therefore the clustering procedure is fairly stable.
More important than this kind of internal validation is biological validation to determine
whether the clusters produced have any biological significance. The following points suggest that
they do.
In the analysis of the whole data set of all the individuals, small clusters regularly identify single
sampling sites. 65% of OTUs are most closely grouped to a cluster which is made up only of
individuals from the same site. Most of these pure clusters are small ranging, with from two to eight
individuals, but four clusters, two from the lower marsh at St Peter’s and one each from Itchenor
lower marsh and Tollesbury middle marsh, have 13-16 individuals. There are, in addition, a
number of other clusters as large or larger which would be from a single site if not for a few mis-
matched individuals (Fig. 4).
The best diagnostic characters which distinguish the large clusters of both phenograms (Figs. 4 &
5) are listed in Tables 4 & 5. Included is character no. 44, width of the scarious margin of fertile
VARIATION IN SALICORNIA EUROPAEA 277
segments, which has been described as being important (Ball & Tutin 1959). Here it has proved to
be a very poor diagnostic. The other characters, however, do act as good diagnostics and relate
very well to those used in traditional classification.
The large clusters identify variants very similar to the accepted species concept (Ball 1964)
especially clusters A & C of the cluster analysis of the full data set (Fig. 4).
TABLE 4. CHARACTER DISTRIBUTION OF THE THREE CLUSTER STAGE OF THE WHOLE
DATA ANALYSIS
S. europaea Cluster Means S. ramosissima
Character sensu Ball (1964) A (n=157) B (n=142) C (n=74) sensu Ball (1964)
30. Spike length 10-50 mm 34.5 mm 22.8 mm 23.4 mm 5—30(40) mm
SD=8.3 SD=6.2 SD=8.7
37. Minimum width | 3-5 mm 3.6 mm 3.0 mm 3.1 mm 2.5-4 mm
of 2nd fertile segment SD=0.3 SD=0.4 SD=0.5
44. Width of 0.1 mm 0.18 mm 0.20 mm 0.24 mm 0.1-0.2 mm
scarious margin SD=0.11 SD=0.12 SD=0.09
64. Fertile segment 1.6 ils) 1.9
width/length SD=0.3 SD=0.3 SD=0.4
49. Colour of Diffuse Red 16s) 2.0 I) Dark Red
florets SD=0.6 SD=0.8 SD=0.8
2. Height to Ist 41.8 mm 42.9 mm 11.3 mm
branch SD=26.1 SD=30.1 SD=6.2
55. Internode 0.6 0.6 1 |
unevenness SD=0.2 SD=0.4 SD=0.5
8. Length of 52.0 mm 55.0 mm 98.9 mm
basal branch SD=28.2 SD=41.4 SD=39.8
25. Max. no. of Simple — 0.4 0.5 19 Typically
secondary branches much branched SD=0.5 SD=0.6 SD=0.5 much branched
29. Length of longest 0.8 mm 0.1 mm 7.5 mm
tertiary branch SD=3.0 SD=0.6 SD=7.4
1. Height of plant (100-)150- 195 mm 179 mm 161 mm 30-400 mm
300(—350) mm SD=43 SD=50 SD=40
Source of
cluster members
Tollesbury Lower marsh 39 11 0
Middle marsh 20 30 0
St Peter’s Lower marsh 3 Z 45
Middle marsh 47 3 0
Canvey Lower marsh 4 6 14
Middle marsh 30 10 10
Itchenor Lower marsh 3 43 4
Upper marsh 11 37 1
SD=standard deviation.
TABLE 5. CHARACTER DISTRIBUTION AT THE TWO CLUSTER STAGE OF THE SPIKE DATA
ANALYSIS
S. europaea Cluster Means S. ramosissima
Character sensu Ball (1964) D (n=168) E (n=205) — sensu Ball (1964)
30. Spike length 10-50 mm 34.5 mm 21.6 mm 5—30(-40) mm
SD=8.0 SD=6.0
37. Minimum width 3-5 mm 3.6 mm 2.9 mm 2.5-4 mm
of 2nd fertile segment SD=0.3 SD=0.4
44. Width of 0.1 mm 0.18 mm 0.22 mm 0.1-0.2 mm
scarious margin SD=0.10 SD=0.12
SD=standard deviation.
278 M. J. INGROUILLE AND J. PEARSON
DISCUSSION
The failure of the principal components analysis to discriminate any groups is remarkable because
the first four components encompass a large part of the total variation. This failure may reflect the
absence of identifiable variants or that the taxonomic structure is too complex, with very many
variants, to be easily simplified.
A comparison of the contribution of spike characters and vegetative characters to the clustering
is interesting. For example, at the three-cluster stage of the whole data set (Fig. 4 & Table 4), the
smallest and most distinct cluster (C) exhibits many of the characteristics of S. ramosissima, such as
being well branched and having a short spike. Cluster A exhibits the character of S. europaea,
having a long spike and being only moderately branched. Cluster B has the spike characteristics of
cluster C and the branching characteristics of cluster A.
In this cluster analysis, the vegetative characters are of equal importance to spike characters.
This is not surprising, despite the wide range of phenotypic plasticity reported by Ball & Tutin
(1959) in vegetative characters, because many are correlated. As a result much emphasis has been
placed on spike characters. Here the 20 best diagnostic characters of the 80 recorded (derived from
the cluster analysis) include eleven vegetative measurements. All of the ten best diagnostics of
cluster C are vegetative.
Cluster B may, however, represent an ecophene of variant C — a poorly branched variant
growing in suboptimal conditions. Tutin (in Clapham ef al. 1962) notes that ‘‘S. ramosissima,
though typically much branched, bushy and erect” is ‘‘often quite unbranched when growing in
crowded pure stands or in competition with other plants. Crowding reduces the degree of
branching and this is often accompanied by an increase in the length of the terminal spike.”
This last point may explain the failure of the principal components analysis to separate the two
taxa when it is carried out on the spike data alone.
There is little correlation between clustering and the level on the marsh of the sampling site. At
St Peter’s there is a separation of individuals of lower from middle marsh sites (Table 4). At
Tollesbury this is only on the basis of spike characters. Notably here, it is the well-branched plants
which grow in the lower marsh. Itchenor and Canvey show little clustering of sites within each
marsh. At Itchenor this is due to the existence of a large proportion of intermediate individuals
(Cluster B). At Canvey the clusters are more equally represented.
The lack of stratification of variants as shown by clusters A, B and Cis not due to the inclusion of
environmentally induced variation in vegetative characters. An examination of the clusters from
the analysis of the spike data shows that 45% of cluster D and 48% of cluster E plants come from
lower marsh sites.
The cluster analysis of both spike and full data sets does show a remarkable ability to identify
local variants. At the seven-cluster stage of the spike data analysis (Fig. 5), between 44% and 75%
of individuals of each cluster come from a single site. The only sites not identified are those of
Canvey lower marsh, where only 24 individuals were sampled, and Tollesbury lower marsh, where
the lower marsh is foreshortened by erosion. In particular, St Peter’s lower marsh and Itchenor
lower marsh have very distinct local variants. Such local variants are probably the result of the
regular self-pollination reported by Dalby (1962). This would encourage the evolution of distinct
inbred lines. The hierarchy of clusters at each site may relate to a pattern of familial relationships.
Means and standard deviations of the best cluster diagnostics for each site are shown in Table 6.
As can be seen, there is little that can be said in a general way about the variation in particular
characteristics between sites. Each site is defined in a different way. The degree of variation
present between individuals within each site is assessed by the clustering method which finds
‘spherical’ minimum variance clusters. The clustering level of the final clustering procedure
provides an estimate of the error sum of squares. Tollesbury and Canvey sites which are probably
the least well stratified marshes have the greatest range of variant individuals and the least well
distinguished variant groups.
VARIATION IN SALICORNIA EUROPAEA 279
TABLE 6. MEANS AND STANDARD DEVIATIONS IN PARENTHESES OF DIAGNOSTIC
CHARACTERS FROM SITES WITHIN MARSHES. MEASUREMENTS IN MM
Tollesbury St'Peter’s Canvey Itchenor
Lower Middle Lower Middle Lower Middle Lower Upper
Character marsh marsh marsh marsh marsh marsh marsh marsh
30. Spike length Ps | 34.7 Lig .7 24.2 30.3 23.0 37.8 27.8
(5.6) (8.4) (5.9) (7.8) (8.9) (8.0) (6.6) 7 1)
37. Minimum width 3.0 S25) 2a 331 3.4 S.2 3.7 33
of 2nd fertile segment (0.3) (0.4) (0.5) (0.4) (0.4) (0.5) (0.3) (0.4)
44. Width of 0.16 0.11 0.28 0.29 0.16 0.20 0.14 0.25
scarious margin (0.12) (0.13) (0.18) (0.10) (0.07) (0.08) (0.06) (0.10)
64. Fertile segment 1.4 1.4 1.9 es) 1.8 7 15 17
width/length (0.2) (0.3) (0.4) (0.3) (0.4) (0.4) (0.2) (0.3)
2. Height to Ist 30.2 = il a) 9.7 TFS 1537, 22.9 31:6 40.7
branch (19.1) (27.0) 6:3) (27.4) (6.2) (16.7) (17:0): ‘QY5)
55. Internode Te3 3.2 1257 2.9 7.9 136 Sf) 5.6
unevenness (3:1) (3.3) 61) (1-5) (1:9) (3:2) (2.4) (2.2)
8. Length of basal 60.4 41.4 76.8 38.6 91.8 80.7 63.6 59.9
branch (6r2) (36.2) (30.9) (27.7) (46.4) (5225) (36.0) (34.5)
25. Max. no. of 4.1 aE 16.7 aS) 14.3 10.4 4.5 7.4
secondary branches (5.4) (4.6) (6.0) (3-5) (8.5) (9.0) (5.8) (6.0)
29. Length of longest 0.08 0.32 6.80 0.0 6.00 2.20 0.16 0.90
tertiary branch (0.57) (2.26) (7.0) — (7.4) (5.6) Chaise. (3:14)
49. Colour of florets 2.40 1.42 2.70 1.94 Jey 1.74 1.00 1.76
(0.53) (0.70) (0.50) (0.84) (0.78) (0.69) — (0.66)
Overall variability 7.45 pes, 6.75 6.62 3.5 9.90 6.73 6.62
Error Sum of Squares Nee pre
from cluster analysis 11.90 17.93 15 By he ie Sie)
Sample size 50 50 50 50 24 50 50 49
CONCLUSION
The evidence presented here suggests that by far the most important kind of variation in the
Salicornia europaea agg. is small scale. Where sites are compared, local variants can be identified,
but the variation between sites is not consistent, and thus it is not possible to find a lower marsh
and a middle/upper marsh variant in every marsh. In addition, taking the plants as a whole, there is
little evidence for the recognition of distinct taxa. Cluster analysis may define clusters which are
very similar to the recognized taxa but this is the result of a rather arbitrary division of a more or
less continuous spectrum of variation. Such clusters fulfill only one of the four criteria Stace (1980)
and Davis & Heywood (1973) cite for the recognition of plant species, i.e. the first of the following:
1. The individuals should bear a close resemblance to one another such that they are always
recognisable as members of that group.
2. There are gaps between the spectra of variation exhibited by related taxa.
3. Each taxon occupies a definable geographical area or ecological niche.
4. Sexual taxa should have individuals capable of interbreeding with little or no loss of fertility and
there should be some reduction in the level or success (measured in terms of hybrid fertility) on
crossing with other taxa.
That the second criterion is not fulfilled is shown by the principal components scatter diagrams.
The inability to detect breaks in the spectrum of variation is not due to the inclusion in the analysis
of extraneous characters that are very sensitive to environmental variation. Scatter plots of two of
the best diagnostic characters show the same complete spectrum of variation (Fig. 6).
These scatter plots also illustrate the failure to fulfill criterion 3. Variants that do exist are not
consistently associated with a level on a saltmarsh.
280 M. J. INGROUILLE AND J. PEARSON
Itchenor Tollesbury Canvey Saint Peters
Spike length
2nd segment minimum width
Ficure 6. Scatter plots of the best cluster diagnostics; spike length and minimum width of 2nd fertile segment
(mm). X=upper or middle marsh plants, O=lower marsh plants.
The information presented here provides no evidence for the fourth criterion. No such
information is available but the work of Jeffries & Gottlieb (1982) is very suggestive and
contradicts our findings. They were able to detect just two electrophoretic morphs which differed
consistently in their isozyme pattern for six different enzyme systems. Each ‘electromorph’ was
consistently associated with either lower or upper marsh. More important was the fact that no
heterozygotes were detected indicating that the electromorphs do not normally interbreed.
Jeffries & Gottlieb (1982) identify the upper marsh electromorph with S. ramosissima and the
lower marsh electromorph with S. europaea. They further suggest that these two species form, in
the British Isles, two distinct homozygous lineages and that the inability to detect any genetic
variability within the species throws doubt on previous reports of genetic variants for
morphological characters.
If the electrophoretic evidence is taken to its limits all the morphological variation within each
species is environmentally induced. The many local variants are then the result of local conditions.
Until a proper genetic analysis is carried out this must remain a conjecture. What our work shows
is that morphological recognition of the two species is not possible and that it would be better to
group them all under S. europaea L. The alternative of recognising ‘chemical species’ is not a
practical possibility.
ACKNOWLEDGMENTS
We would like to acknowledge receipt of a Natural Environmental Research Council grant to J.
Pearson. We would like to thank Dr D. C. Havill for his advice and Graham King for his technical
assistance.
REFERENCES
BALL, P. W. (1964). Salicornia L. in Turin, T. G. et al., eds. Flora Europaea, 1: 101-102. Cambridge.
BALL, P. W. & Brown, K. G. (1970). A biosystematic and ecological study of Salicornia in the Dee Estuary.
Watsonia, 8: 27-40.
VARIATION IN SALICORNIA EUROPAEA 281
BALL, P. W. & TuTin, T. G. (1959). Notes on annual species of Salicornia in Britain. Watsonia, 4: 193-205.
CLAPHAM, A. R., Tutin, T. G. & WarBurG, E. F. (1952). Flora of the British Isles. Cambridge.
CLapHaM, A. R., Tutin, T. G. & WaARBURG, E. F. (1962). Flora of the British Isles, 2nd ed. Cambridge.
Da.sy, D. H. (1962). Chromosome number, morphology and breeding behaviour in the British Salicorniae.
Watsonia, 5: 150-162.
Davis, P. H. & HeEywoop, V. H. (1973). Principles of angiosperm taxonomy. New York.
HA t, P. C. (1980). Sussex plant atlas. Brighton.
JEFFRIES, R. L., Davy, A. J. & Rupmik, T. (1981). Population biology of the saltmarsh annual Salicornia
europaea agg. J. Ecol., 69: 1-15.
JEFFRIES, R. L. & Gorties, L. D. (1982). Genetic differentiation of the microspecies Salicornia europaea L.
(sensu stricto) and S. ramosissima J. Woods. New Phytol., 92: 123-9.
JERMYN, S. T. (1974). Flora of Essex. Colchester.
Stace, C. A. (1980). Plant taxonomy and biosystematics. London.
Wisuart, D. (1978). CLUSTAN, user manual, 3rd ed. University of Edinburgh.
(Accepted November 1985)
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Watsonia, 16, 283-286 (1987) 283
Hieracium westii, sp. nov.
POD SELL
Botany School, Downing Street, Cambridge, CB2 3EA
ABSTRACT
A new species, Hieracium westii P. D. Sell, sp. nov., is described and the account of its discovery set out.
A NEW HIERACIUM
Hieracium westii P. D. Sell, sp. nov.
TYPUS: By rocky stream, Allt nan Giubhas, Argyll, v.c. 98, GR 27/265.523, c.380 m alt., P. D.
Sell 78/138 & A. G. Kenneth (Holotypus: CGE). Fig. 1.
Ab H. submuroro Lindeb. (quo nomen plerumque false cognovitur) caulis foliisque pilosioribus,
foliis caulorum numerosioribus, squamis involucri latioribus obtusioribus, ligulis effectis differt,
praeterea ab H. duricipiti F. J. Hanb. (quid facie simulat) foliis paucioribus erectioribus
glaucoviridibus, pedunculis rectis suberectis, capitulis magnioribus, ligulis aureis distinguitur.
Planta phyllopoda. Caulis 28-60 cm altus, gracilis, pilis stellatis dispersis praecipue superne, aliis
simplicibus eglanduliferis paucis brevibus et mediocribus pallidis interdum basi nigris ubique, altis
glanduliferis minutis paucis flaviusculis ubique, aliis glanduliferis brevibus paucis obscuris superne
vestitus. Folia glaucoviridia, interdum leviter purpureotincta, utrinque et margine pilis simplicibus
eglanduliferis brevibus mediocribusque pallidis numerosis, inferne in mediocosto pilis simplicibus
eglanduliferis longis paucis, ubique pilis glanduliferis brevissimis flaviusculis dispersis vestita. Folia
basalia plerumque pauca, plusminusve erecta; primigena 0.7—2.0 cm longa, 0.5—1.5 cm lata,
subrotunda vel late elliptica, obtuso-mucronata, denticulata vel dentata, basi rotundata vel
abrupte contracta; cetera 3-10 cm longa, 1.5—5 cm lata, ovata, lanceolata vel elliptica, rotundo-
obtusa vel acuta, ordinate dentata, dentibus saepe plusminusve mammiformibus, basi plerumque
asymmetrica cuneataque interdum truncata; petioli breves vel mediocres pilis simplicibus
eglanduliferis mediocribus vel longis pallidus numerosis vestiti. Folia caulina 1-2 (—3), 1.5-5.5 cm
longa, 0.2—2.5 cm lata; inferiora lanceolata anguste acuta vel acuminata, dentata dentibus angustis
saepe cuspidatis, sessilia vel petiolis brevibus vel mediocribus semiamplexicaulibus; superiora
plerumque bractiformia. Capitula 2-10, 20-30 mm in diametro, bases rotundata; pedunculi recti,
suberecti, pilis stellatis numerosis vel aliquantum densis, aliis glanduliferis brevibus et brevissimis
obscuris numerosis, altis simplicibus eglanduliferis brevibus mediocribusque obscuris paucis vel
numerosis vestiti; acladium interdum brevissimum. Involucri squaamae 10-13 mm longae, 1-1.5
mm latae, atriusculo-virides, interiores marginibus pallidioribus, ante anthesin incumbentes,
anguste lineari-lanceolatae, longe contractae ad anguste subacutum vel acutum apicem, interiores
fere filamentosae, pilis glanduliferis inaequalibus brevissimis brevibus mediocribusque obscuris
numerosis, aliis simplicibus eglanduliferis brevibus mediocribusque obscuris paucis vel numerosis,
aliis stellatis interdum paucis versus basem vestitae. Ligulae aureae vel fere aurantiaco-aureae,
leviter dentatae, apice pilis simplicibus eglanduliferis brevissimis pallidis numerosis obsitae.
Stigmata obscuri. Receptaculi alveoli margine breviter dentati. Cypselae 3.5-4.0 mm longae,
obscurae.
Phyllopodous. Stem 28-60 cm, slender, with scattered stellate hairs mainly in the upper part; few,
short and medium, pale, sometimes dark-based simple eglandular hairs throughout; few,
yellowish, minute glandular hairs throughout and a few, short, dark glandular hairs in the upper
284 P: DU SELLE
Wf
NA oay
Figure 1. Hieracium westii P. D. Sell; based on the holotype from Argyll, P. D. Sell 78/138 & A. G. Kenneth
(CGE): a) whole plant, b) involucre showing indumentum, c) inner (left) and median (right) involucral bracts,
d) ligule. Scale bar = a) 30 mm; b, c) 5 mm; d) 7 mm
HIERACIUM WESTII 285
part. Leaves bluish-green, sometimes slightly purple-tinted, with numerous short and medium,
pale simple eglandular hairs on both surfaces and the margins with a few long ones on the midrib
beneath, and scattered, very short, yellowish glandular hairs throughout. Basal leaves usually few,
more or less erect; primordial 0.7—2.00.5-1.5 cm, subrotund or broadly elliptical, obtuse-
mucronate at apex, denticulate or dentate, rounded or abruptly contracted at base; later
3-10 1.5—5 cm, ovate, lanceolate or elliptical, rounded-obtuse or acute at apex, regularly dentate,
the teeth often more or less mammiform, usually asymmetrical and cuneate, sometimes truncate at
base; petioles short or medium, with numerous medium or long, pale simple eglandular hairs.
Cauline leaves 1-2 (—3), 1.5-5.5X0.2—2.5 cm, lower lanceolate, narrowly acute or acuminate at
apex, dentate, the teeth usually rather narrow and often cusped, with short or medium, semiam-
plexicaul petioles, or sessile, upper leaf usually bract-like. Capitula 2-10, 20-30 mm in diameter,
base rounded; peduncles straight and suberect, acladium sometimes very short, with numerous or
rather dense stellate hairs, numerous short and very short, dark glandular hairs, and few to
numerous, short and medium, dark simple eglandular hairs. Involucral bracts 10-13 1-1.5 mm,
blackish green, the inner with paler margins, incumbent in bud, narrowly linear-lanceolate, long
tapering to a narrow subacute to acute apex, the inner almost filamentous, with numerous,
unequal, very short, short and medium, dark glandular hairs, few or numerous, short and medium,
dark simple eglandular hairs and sometimes a few stellate hairs towards the base. Ligules golden or
nearly orange-yellow, rather shallowly dentate, with numerous, very short, pale simple eglandular
hairs at apex. Stigmas discoloured. Margins of receptacle pits shortly dentate. Achenes 3.5—-4.0
mm, dark.
I had hoped to celebrate Dr Cyril West’s hundredth birthday by describing this new species after
him, but he died on 25th March 1986, aged 98. Almost the whole of his thirty-odd years of
retirement had been devoted to the study of the genus Hieracium. Our last piece of work together
was to compare in detail the British plants that had been called H. submurorum Lindeb. with those
from Scandinavia. We decided the British plant was quite distinct and wrote a detailed description,
but never published it. It seems appropriate that this species should bear his name.
Hieracium westii was first discovered by E. S. Marshall by a streamlet in Cam Glen, Glen Etive,
Argyll, v.c. 98, on 27th June 1888. The sheet in Marshall’s herbarium (CGE) bears the note in his
handwriting “The first discovery of this good ‘species’ in Britain’. Two more sheets with the same
date are labelled “stream above ‘Tighe Craige Duibhe’ Clach Leathad (say 1800 feet) near
Kingshouse, Argyll’. They had been determined by Lindeberg as ‘‘Proximum H. submurorum” in
1890 and were confirmed by Elfstrand in 1894. One of these is also labelled ‘“‘First known British
specimens”. Seeds from near Kingshouse were grown by Marshall and there are cultivated
specimens in his herbarium collected in 1890 and 1891. He collected it again from a mountain
stream near Kingshouse on 17th July 1889. A cultivated specimen of Marshall’s in the Bickham
herbarium (CGE) is labelled ‘‘root from Allt Giubhas’’. Hanbury (1892) writes ‘“‘This was collected
in some quantity by the Rev. E. S. Marshall and myself, on the mountains round Kingshouse,
Argyll, our specimens agreeing well with the type No. 112, Hierac. Scand. Exsicc.”’. In Hanbury’s
Illustrated monograph of the British Hieracia it is beautifully depicted on plate 21, but only the
Argyll plant in the list of localities given is referable to it. The illustration was based on a Clach
Leathad plant supplemented by a cultivated specimen. A cultivated specimen in herb. Hanbury
(BM), said to have been used for this purpose, could not possibly have supplied the inflorescence,
but some of the leaves may possibly have been used. It is not absolutely clear how many different
localities these early botanists saw it in, but there seem to have been at least two.
It does not appear to have been seen again until rediscovered by John Raven on 14th July 1953 at
Allt nan Giubhas where it has since been examined by Cyril West, Beverley Miles, Archie
Kenneth and myself (all in CGE). It grows in crevices on rocks and boulders, and beside the stream
in association with Anthoxanthum odoratum, Calluna vulgaris, Deschampsia cespitosa, Luzula
sylvatica, Potentilla erecta, Ranunculus acris, Solidago virgaurea, Sphagnum sp., Succisa pratensis
and Vaccinium myrtillus. The stream of Allt nan Giubhas is one of the finest localities for alpine
and subalpine hawkweeds in the whole of Scotland, containing in some abundance H. anfracti-
forme, H. callistophyllum, H. dasythrix, H. dewari, H. duriceps, H. eximium, H. gracilifolium, H.
holosericeum, H. lingulatum, H. marshallii, H. memorabile, H. pictorum, H. pseudanglicum, H.
subhirtum, H. tenuifrons and H. vulgatum as well as at least one other undescribed species.
286 P2/ Di SEEL
Hieracium westii is characterized by its bluish-green, suberect, toothed leaves, few capitula on
straight suberect peduncles, dark involucral bracts clothed with numerous, unequal, dark,
glandular hairs and few to numerous, dark, simple eglandular hairs and golden to nearly orange
ligules (Fig. 1).
ACKNOWLEDGMENTS
I am grateful to N. K. B. Robson for checking the Latin description, A. O. Chater for reading and
commenting on the text, and Tim Sell for drawing the illustration.
~
REFERENCE
Hansury, F. J. (1892). Further notes on Hieracia new to Britain. J. Bot., Lond., 30: 129-134.
(Accepted July 1986)
Watsonia, 16, 287-289 (1987) 287
Hieracium pellucidum Laest., H. lucidulum (A. Ley) Roffey and
H. asteridiophyllum P. D. Sell & C. West
PDS See.
Botany School, Downing Street, Cambridge, CB2 3EA
ABSTRACT
The taxonomic history and distribution of Hieracium pellucidum Laest. and H. asteridiophyllum P. D. Sell and
C. West (Asteraceae) are discussed and their nomenclature is set out.
INTRODUCTION
When Dr C. West and I described H. asteridiophyllum (Sell & West 1955) we gave no explanation
of our treatment of, and its relationship to, the allied H. pellucidum. H. asteridiophyllum can be
distinguished from H. pellucidum by its having simple eglandular as well as glandular hairs on the
involucral bracts and stellate hairs on the underside of the leaves. This paper discusses the
taxonomic history of these two species, outlines their distribution in Britain, and sets out their
nomenclature.
HISTORY
H. pellucidum was described by Laestadius (1824) and there is a specimen in the Stockholm
herbarium (S) labelled ‘““Angermanland ad Tasjo, 1824, Laestadius’, which appears to be the
holotype. Whether the date is that of the actual collection is unsure, but in those days it was
possible to get a new species published in the same year that it was collected. The volume in which
it was published appeared in two parts, both of which are dated 1824 but without a precise day or
month. H. pellucidum was in the first part. In my opinion, the Scandinavian plants described as H.
pellucidum are identical with those found in the British Isles. There is a good description in Pugsley
(1948); the Scandinavian distribution is given by Samuelsson (1954), and the distribution in the
British Isles by Perring (1968).
In both Scandinavia and the British Isles. any sizeable population of H. pellucidum seems to
contain plants with two different leaf shapes. In one the leaves are large, ovate in outline and
somewhat pointed. In the other they are smaller, more elliptical and rounded at the apex. Plants
occur also that are more or less intermediate. Scandinavian hieraciologists include both these
variants in H. pellucidum and I would agree with them. The Laestadius specimen is of the pointed
leaved variant.
The Rev. Augustin Ley grew two variants (but perhaps not the two mentioned above) side by
side for six or seven years, and considered that they remained distinct. As well as the leaf
difference, he thought that the round-leaved plant had smaller capitula, which formed a somewhat
umbellate inflorescence, and named them H. murorum var. lucidulum (Ley 1900). H. pellucidum
was regarded as another variety of H. murorum. Although it is quite clear from the description
what Ley intended, there is some difficulty in typifying var. /ucidulum. All the specimens in his
herbarium (CGE) cited under var. /ucidulum are still labelled ‘‘var. pellucidum”, “‘H. murorum
var.’’, or in one case “H. murorum lucentius var. nov.”’. Later specimens, labelled H. murorum
var. /ucidulum or H. lucidulum, are the same plant. Fortunately Ley says ‘‘Sent out by me through
the London Bot. Exchange Club in 1896, from Llangadoc, as var. pellucidum Laestad.”’ In his
herbarium is a sheet of this gathering bearing two complete plants, a stem without basal leaves, and
288 P.D*SSEEL
a stemless basal rosette. The plants are in accord with the original description and show well the
subumbellate inflorescence. I designate this sheet as the lectotype of H. murorum var. lucidulum
A. Ley. In Ley’s herbarium there are no specimens labelled either /ucidulum or pellucidum that
are H. asteridiophyllum.
W. R. Linton (1905) transferred var. Jucidulum to H. pellucidum. His description does not differ
much from Ley’s, but specimens collected by Ley in 1900, and others by E. F. and W. R. Linton in
1897 from Craig Cille [Craig y Cilau], Brecon, were both distributed as H. murorum var. lucidulum
in their “Set of British Hieracia no. 139” and are H. asteridiophyllum.
Williams (1902) transferred var. Jucidulum to H. silvaticum. His description is more or less that
of Ley, but he cites the Lintons’ ‘Set no. 139”, which is H. asteridiophyllum. Ley (1909) says “‘It
appears that the plants we have been accustomed to call H. pellucidum Laestad. in Britain, and
some which have been so named by great Scandinavian authorities, are now mostly placed under
H. serratifrons Almq., and that the very distinct plant we have been naming /ucidulum Ley is the
true H. pellucidum Laestad.” This is true in that many of Ley’s specimens of pellucidum belong to
what I call the H. exotericum agg. It is not known what the plants were which Ley grew to compare
with var. /ucidulum. Zahn (1921) raised var. /ucidulum to a subspecies of H. murorum, but his
description, clearly based on the Lintons’ “Set no. 139”, is of H. asteridiophyllum. Roffey (1925)
raised var. /ucidulum to the rank of species, not, I think, because he knew anything about it, but
because Zahn had called it a subspecies. Druce (1928) also had it as a species. Pugsley (1948) kept
separate what he cited as H. pellucidum var. lucidulum W. R. Linton emend., with the references
*H. pellucidum var. lucidulum W. R. Lint. Brit. Hier. 43 (1905) pro min. parte; H. pellucidum Ley
in Jour. Bot. xlvii 13 (1909), pro min. parte; and H. murorum subsp. lucidulum Zahn, |.c. 318
(1921). Pugsley’s description of H. pellucidum var. lucidulum is clearly that of H. aster-
idiophyllum. Of the specimens cited by Pugsley (1948), those from Craig y Cilau are H.
asteridiophyllum, the Pwll Byfre plant is H. discophyllum P. D. Sell & C. West, and the Ingleton
specimen I have not seen.
DISTRIBUTION
H. pellucidum is one of several Hieracium species which are widespread in Scandinavia and absent
from Scotland and Ireland, but which appear in some quantity on the limestone cliffs of northern
England, Derbyshire and Wales.
H. asteridiophyllum is endemic to the cliffs of Craig y Cilau just south-west of Crickhowell,
Brecon (v.c. 42) (GR 32/181.166 to GR 32/190.156), incorrectly called Craig Cille by A. Ley. They
are mostly in a National Nature Reserve and consist of a shallow north-east-facing scarp of tiered
cliffs and steep slopes, with much scree, at an altitude of 275-460 m. It is one of the largest
exposures of upland limestone in south Wales. In 1975 Lynne Farrell and I did a census of
Hieracium (and Sorbus) on this cliff, and saw 460 plants of H. asteridiophyllum. We examined
three levels of cliff; 387 plants were in the lower section, only one was seen in the middle part, and
72 in the upper part. Although occurring along the whole length of the cliff, most plants were at the
northern end. H. pellucidum is also on the cliff, but we saw only 24 plants, again mainly at the
northern end.
NOMENCLATURE
HIERACIUM PELLUCIDUM Laest. in Kungl. Svenska Vet.-Akad. Handl., 1824: 172 (1824).
TYPE: Angermanland ad Tasjo, Sweden, 1824, L. L. Laestadius (Holotype: S).
H. murorum var. lucidulum A. Ley in J. Bot., Lond., 38: 3 (1900) (TYPE: River side rocks,
Llangadoc, Carmarthenshire, lst June 1896, A. Ley (Lectotype: herb. Ley in CGE).
H. silvaticum var. lucidulum (A. Ley) Williams, Prodr. Fl. Brit., 137 (1902).
H. pellucidum var. lucidulum (A. Ley) W. R. Linton, Brit. Hier. , 43 (1905).
H. murorum subsp. lucidulum (A. Ley) Zahn in Engler, Pflanzenreich, 76 (1V. 280): 318 (1921)
quoad basionymum exclud. descript.
H. lucidulum (A. Ley) Roffey in F. J. Hanb., London Cat. Brit. Pl., 11th ed., 27 (1925).
THREE HIERACIUM SPECIES 289
HIERACIUM ASTERIDIOPHYLLUM P. D. Sell & C. West in Watsonia, 3: 233 (1955). TYPE:
Daren Cilau (Craig Cille), Brecon, 1952, C. West (Holotype: CGE).
H. murorum subsp. lucidulum sensu Zahn in Engler, Pflanzenreich, 76 (IV. 280): 318 (1921)
quoad descript. exclud. basionymum.
H. pellucidum var. lucidulum sensu Pugsley in J. Linn. Soc. Lond. (Bot.), 54: 141 (1948) quoad
descript. et pl. Craig Cille.
ACKNOWLEDGMENTS
J. R. Akeroyd and A. O. Chater kindly read through and commented on the manuscript of this
paper.
REFERENCES
Lagstapius, L. L. (1824). Besktifning ofver nagra sallsyntare Vaxter fran norra delarne af Sverige jemte
anmarkningar i Vaxtgeografien. Kungl. Svenska Vet.-Akad. Handl., 1824: 172.
Ley, A. (1900). Some Welsh hawkweeds. J. Bot., Lond., 38: 3-7.
Ley, A. (1909). Brecon and West Yorkshire hawkweeds. J. Bot., Lond., 47: 8-16.
Linton, W. R. (1905). An account of the British Hieracia. London.
PERRING, F. H. (1968). Critical supplement to the atlas of the British flora. London.
Pusey, H. W. (1948). A prodromus of the British Hieracia. J. Linn. Soc. Lond. (Bot.), 54: 1-356.
Rorrey, J. (1925). Hieracium L. in Hansury, F. J., The London catalogue of British plants, 11th ed. London.
SAMUELSSON, G. (1954). Maps of a selection of Scandinavian Hieracium species. Kungl. Svenska Vet.-Akad.
Hand Ser. 4, 5.():
SELL, P. D. & West, C. (1955). Notes on British Hieracia, I. Watsonia, 3: 233-236.
WILLIAMS, F. N. (1902). Prodromus florae Britannicae. Brentford.
ZAHN, K. H. (1921-1923). Compositae-Hieracium in ENGLER, H. G. A. (ed.), Das Pflanzenreich 76 (IV. 280).
Leipzig.
(Accepted July 1986)
Watsonia, 16, 291-297 (1987) 291
The history and nomenclature of Thomas Edmondston’s
endemic Shetland Cerastium
R. K. BRUMMITT
The Herbarium, Royal Botanic Gardens, Kew, Surrey, TW9 3AB
DH kent
75 Adelaide Road, Ealing, London, W13 9ED
Poo, WUSBY
Department of Plant Sciences, The University, St Machar Drive, Aberdeen, AB9 2UD
and
R. C. PALMER
11 Fleet Way, Didcot, Oxfordshire, OX11 8BZ
ABSTRACT
A bibliographic history of the endemic Shetland Cerastium currently known as C. arcticum Lange subsp.
edmondstonii (H. C. Watson) A. & D. Léve is given, together with some biographical notes on Thomas
Edmondston. The correct name for the species is shown to be C. nigrescens (H. C. Watson) Edmondston ex H.
C. Watson, which includes subsp. nigrescens endemic in Shetland and subsp. arcticum (Lange) Lusby, comb.
nov., occurring in Wales and mainland Scotland and widespread in northern Europe.
INTRODUCTION
During work on a forthcoming Flora of the Shetland Islands, one of the present authors (R. C.
Palmer) found cause to query the correct name and authorship of the Cerastium, endemic to the
Shetlands, variously known by the epithets edmondstonii and nigrescens. On further investigation
with two others (R. K. Brummitt & D. H. Kent) it became obvious that the citation in Flora
Europaea (Jalas 1964) of C. nigrescens Edmondston ex H. C. Watson as a synonym of C. arcticum
Lange subsp. edmondstonii (H. C. Watson) A. & D. Love was suspect. The specific name C.
arcticum was not published until 1880, and since H. C. Watson died in 1881 at the age of 77 the
likelihood of his having validated C. nigrescens after the publication of C. arcticum seemed remote.
Further enquiries revealed a recent unpublished thesis by the fourth present author (Lusby 1984)
investigating the same problem and reaching similar, but not identical, conclusions. Our joint
findings are here merged into one paper.
Examination of the literature has revealed several incorrect nomenclatural assumptions and
numerous Overlooked important bibliographical details in accounts by earlier authors, despite
several papers having been devoted to the taxonomy and nomenclature of this group of species.
We have here tried to bring together the historical details and recall a little of the remarkable
Edmondston, as well as to resolve the nomenclature of the plant. The several bibliographic
references given in the text below instead of in the References at the end are to notes published
anonymously.
Mr P. D. Sell (Cambridge) has very generously made available his own extensive notes on the
292 R. K. BRUMMITT, D. H. KENT, P. S. LUSBY AND R. C. PALMER
species concerned, and has encouraged us to publish our findings while reserving his own judgment
on the taxonomy of the group.
EDMONDSTON AND THE DISCOVERY OF THE SHETLAND CERASTIUM
Thomas Edmondston was born in the Shetland Islands on 20th September 1825, the eldest son of
the medical practitioner on the island of Unst. Although delicate early in life, he soon showed
himself to have an extraordinarily quick and retentive memory, and a predilection for natural
history from the age of four (Seeman 1853: 67-78). In 1837, at the age of eleven, he compiled a
remarkably competent catalogue of the plants of Unst, listing 174 species by their Latin names. On
September 3rd that year he had the good fortune to meet William Dawson Hooker, elder son of
W. J. Hooker and brother of J. D. Hooker, who was returning from a naturalists’ expedition to
northern Norway when his ship sought shelter from bad weather at Baltasound in Shetland.
Hooker was entertained there by the Edmondstons, and noted Thomas as “a particularly
intelligent boy, and passionately fond of Natural History” (W. D. Hooker 1839: 110-111). Later
Dr Edmondston sent a copy of Thomas’s manuscript catalogue to W. J. Hooker, noting that the
only assistance he had had was from two books, Willdenow’s /ntroduction and McGillivray’s
edition of Withering, and it was eventually printed by W. D. Hooker in the second edition of his
Notes on Norway (Edmondston 1839). This included a reference (p. 114) to ““Cerustium alpinum;
very rare, on serpentine; June.”
In 1837, Dr Gilbert McNab visited Unst and was shown the Cerastium by the young
Edmondston. McNab sent material to the Botanical Society of Edinburgh, where it was exhibited
and commented on by the President, Dr Robert Graham, and in a report of the meeting in Annals
of Natural History, 1: 406 (1838), Graham’s determination of the plant as ‘Cerastium latifolium
var.’ is recorded.
At the age of fifteen, Edmondston published a list of plants observed in the Shetland Islands
(Edmondston 1841), including the first record of Arenaria norvegica Gunn. from the British Isles
and recording ‘Cerastium latifolium’ from near Baltasound. In the same year, he visited the
Scottish mainland, where he saw full-grown trees for the first time, and was appointed Assistant
Secretary of the Edinburgh Botanical Society (Jackson 1888). By March 1841, Edmondston was
also already corresponding with C. C. Babington of Cambridge about his Shetland Cerastium,
commenting in May the same year that ‘‘a reference to the Linnaean herbarium would settle any
doubts” (Mrs Edmondston 1868: 117). In March 1843, at the age of seventeen, with the support of
Babington he published a paper in The Phytologist (Edmondston 1843a) on the taxonomy of alpine
and arctic species of Cerastium, drawing attention particularly to the Shetland plant, which he
referred to C. latifolium L. He followed Bentham’s opinion, quoted in Lindley (1829: 51), by
referring the Scottish and Welsh mountain plants to C. alpinum L., in which a var. piloso-
pubescens Benth. was also recognized.
Shortly afterwards, H. C. Watson, one of the leading amateur botanists of the day, replied in the
May issue of the same journal, disagreeing with Edmondston’s taxonomy and considering the
Shetland plant “‘in all likelihood a mere form or variety of the same species as the C. /atifolium (of
British authors) found in many of the Highland mountains” (Watson 1843a). In the August issue
Edmondston responded spiritedly, maintaining that the Shetland plant was distinct (Edmondston
1843b), but in the following issue Watson replied again saying that after consulting the Linnean
herbarium he considered that Edmondston had confused the characters of the taxa (Watson
1843b). Watson here emphasized seed characters as the best means of distinguishing the species,
but the following year (Watson 1844) he noted that the names of his packets had been switched.
In 1844 Edmondston sent good herbarium specimens bearing live seeds to Watson, and to other
members of the Botanical Society of London, under the name ‘“‘Cerastium nigrescens Edmond.”’.
Specimens are still preserved in the Watson herbarium at Kew, together with some prepared in
1845 and 1846 from plants grown by Watson from the seeds sent by Edmondston. However, in the
first edition of the London Catalogue of British Plants, published by the Botanical Society of
London in 1844, the plant appears under the name C. J/atifolium L. b. edmondstonii, a nomen
nudum, without any authority given but almost certainly at Watson’s instigation (W. J. Hooker
1844: 294; Allen 1983). This seems to be a sign of weakening in Watson’s opposition to
THOMAS EDMONDSTON’S CERASTIUM 293
Edmondston’s view that the Shetland plant was taxonomically distinct from mainland plants, and
in The Phytologist of March the next year (Watson 1845) he grudgingly acknowledged that “‘Mr
Edmondston’s specimens are distinguishable from my other wild specimens by their shorter
capsules and usually (not constantly) broader leaves; which are the only characters to distinguish
them, even as a mere variety, from the species of the Highland mountains”. He adopted the
name ‘“‘Cerastium latifolium (Linn.) var. Edmondstonii (Lond. Cat.)” in the title of the paper,
and this brief description would be sufficient to validate this name. The note was dated 12th
February 1845 and, judging by the average time of several weeks between submission of articles
and their appearance in The Phytologist, it is likely that publication was towards the end of
March.
Strangely enough, however, two to three pages later in the same journal (The Phytologist, 2:
95-96, 1845), in a report of the proceedings of the Botanical Society of London for 7th February
1845 published by G. E. Dennes, reference was made to the specimens of the Shetland Cerastium
presented by Edmondston to the Society, noting that they were labelled “‘Cerastium nigrescens
Edmond. in Shetland FI. ined.” and that “It would thus seem that Mr Edmondston has changed
his opinion regarding its specific identity with the Linnaean species’’. This was considered by
Druce (1908, 1911, 1922) to constitute validation of the name, but we agree with Marshall (1911)
that it is not validly published here. The author, Dennes, did not accept the name (Jnternational
Code of Botanical Nomenclature, Art. 34.1a) and went on to say that it may be doubted whether
Edmondston’s specimens can be distinguished from the Highland and Linnaean C. Jatifolium.
However, in 1845 Edmondston’s Flora of Shetland was also published (Edmondston 1845).
Jackson (1888) referred to it as published in Spring 1845, and it seems likely that it was published
in February or the first week of March. Neither of the notes in the March issue of The Phytologist
referred to above (both written in mid February) made any mention of it, but a letter by
Edmondston himself to his uncle dated 8th March 1845, reproduced in the biography published
by his mother (Mrs Edmondston 1868: 222), refers to his book as “‘published at last”. On p. 29 of
his Flora, Edmondston included ‘‘Cerastium latifolium B Edmondstonii. (London Botanical
Society’s Catalogue of British Plants)’’, with “‘C. latifolium Edmondston in Phytologist p. 498”
and ‘“‘C. nigrescens Edmonds. MSS.” in synonymy followed by a detailed description of the
Shetland plant. This constitutes validation of the name C. latifolium var. edmondstonii (I.C.B.N.
Art. 35.3), and apparently precedes Watson’s publication of the same name (see above) by about
a month. In the preface to his Flora, however, on p. xv Edmondston wrote ‘‘On the Serpentine
hills near Baltasound grows Arenaria Norvegica and Cerastium nigrescens .. .”. Druce (1922)
suggested that the preface was written after the text, but since Edmondston wrote of the plant in
question on p. 30 “‘Mr Watson, after an investigation of numerous specimens, still considers this
plant as not truly distinct from C. /atifolium, and from deference to his authority I give up the
point . . .” this seems questionable. Since the name C. nigrescens was clearly not accepted in the
Flora it cannot now be considered to have been validly published there. Thus two different
epithets, edmondstonii and nigrescens, became established in the literature almost simultaneously
for the same plant between 1844 and 1845, but in neither case was a specific binomial under
Cerastium validly published. Furthermore, Edmondston himself seemed unable to make up his
mind which to use.
Having gained the friendship of the leading Scottish professionals such as W. J. Hooker and J.
Hutton Balfour and established for himself a reputation of being one of the most brilliant young
botanists in the country, Edmondston was appointed Professor of Botany at Glasgow in January
1845 at the age of nineteen. A few months later, however, he took the fateful decision to take
part in an expedition to the Pacific and California as naturalist aboard H.M.S. Herald (see The
Phytologist, 2: 185, 1845) which he joined on 21st May 1845. After sailing via Buenos Aires and
the Falkland Islands, the ship rounded Cape Horn and visited Chile and the Galapagos Islands
before calling at Sua Bay in Ecuador. Here, on the evening of 24th January, as Edmondston was
boarding a small boat to re-join ship, a man in front of him caught the lock of a rifle in his
trouser leg and accidentally fired it, shooting Edmondston in the head and killing him instantly
(Seemann 1853: 66). His death was announced in, The Phytologist, 2: 580 (1846). This tragic loss
at the age of twenty of a man surely otherwise destined to become one of the great botanists of
the nineteenth century meant that the Shetland Cerastium was left in a position of doubt and
uncertainty which has persisted to the present day.
294 R. K. BRUMMITT, D. H. KENT, P. S. LUSBY AND R. C. PALMER
LATER TAXONOMIC HISTORY
The subsequent taxonomic history of the Shetland plant reveals many varying opinions. Although
the name C. /atifolium L. had been commonly applied to British plants, Lange (1880) regarded this
as a species of the southern Alps only, and described the widespread northern taxon as C. arcticum
Lange. He did not actually mention the Shetland plant, but his circumscription of his new species
was such that it could well have included it. Beeby (1887) accepted Lange’s species and referred
the Shetland plant to it as C. arcticum var. edmondstonii (H. C. Watson) Beeby. Murbeck (1898)
apparently adopted a similar species concept but called the species C. edmondstonii (Wats.) Murb.
& Ostenfeld, with C. arcticum Lange in synonymy, though failing to quote any specific name
earlier than C. arcticum or to explain why the epithet edmondstonii was considered to have
priority. Asplund (1918) discussed the status of C. edmondstonii but regarded it as a synonym of C.
arcticum Lange. Ostenfeld (1920) adopted a specific concept similar to that of Murbeck, but
changed the name of the species to C. nigrescens Edmondston on the false assumption that
Edmondston had validated the specific name in his Flora of Shetland in 1845. Hylander (1945:
148-149), however, in a detailed nomenclatural analysis, accepted the name C. edmondstonii to
cover the widespread species.
In the Flora of the British Isles, Clapham (1952) called-the species C. edmondstonii, with ‘“‘C.
arcticum auct., vix Lange” in synonymy, distinguishing the Shetland plant as “var. nigrescens
Edmondston’’; this must be incorrect since the type of both the specific and varietal names must be
the Shetland plant. A note by Brett (1953) argued for C. arcticum as the correct name, but failed to
consider any of the bibliographical details. Love & Léve (1956) also reverted to the specific name
C. arcticum and recognized within it a widespread subsp. edmondstonii, including not only the
Shetland plant but also plants from the Faeroes and eastern Iceland, and then distinguished the
Shetland plant within this as var. nigrescens. Hultén (1956) with some reservation regarded the
Shetland plant as specifically distinct, calling it C. edmondstonii (Watson) Murb. & Ostenf., and
referred the Scottish mainland plants to C. arcticum Lange var. arcticum and var. alpinopilosum
Hultén. In the second edition of the Flora of the British Isles, Clapham (1962) also regarded the
Shetland plant as a species endemic there, but under the name C. nigrescens Edmondst. ex H. C.
Watson, with C. edmondstonii in synonymy, and called the widespread mainland and arctic plant
C. arcticum Lange. Jalas (1964), in Flora Europaea, adopted the same nomenclature as Léve &
Love but restricted subsp. edmondstonii to the Shetland plant, an opinion with which we
tentatively agree. Bocher (1977), however, while dividing C. arcticum into three subspecies, sank
C. edmondstonii without trace into C. arcticum subsp. arcticum, commenting that the Greenland
plant is inseparable from that from Iceland and the Faer6es, which ignores the fact that the type of
C. edmondstonii must be from Shetland.
NOMENCLATURE
The question which now remains is that of the correct name for the species when the Shetland plant
is included in the widespread species. The name C. arcticum dates only from 1880, while the
epithets nigrescens and edmondstonii have been in the literature considerably longer. Was neither
adopted at specific rank in a valid name before 1880?
The binomial C. edmondstonii is given in Index Kewensis, Suppl. 5, 1921, as first validated by
Murbeck & Ostenfeld in Murbeck (1898) and this appears to be correct. The original volumes of
Index Kewensis in 1893 gave C. nigrescens as having been validated by Syme in English Botany, 3rd
ed., 2: 87 (1864), but this was corrected in Supplement 12 in 1959 to H. C. Watson, Cybele
Britannica, 1: 233 (1847). However, although Watson here gave a binomial C. nigrescens, he
numbered it 197b under his 197 C. Jatifolium, commenting that “I therefore describe its
distribution apart from that of C. latifolium, though still inclining to hold it a variety of the Linnean
species’’. It is clear that Watson did not recognize it at specific rank and so cannot be held to have
validated the specific name here. However, despite the fact that he used a binomial form, we may
accept that he validated here the name C. latifolium var. nigrescens (see I.C.B.N., Art. 24.4). The
publication by Syme in 1864, referred to above, clearly accepted C. latifolium var. nigrescens, and
THOMAS EDMONDSTON’S CERASTIUM 295
although the binomial ““C. nigrescens Edmondston mss” was given beneath, it is not valid since it is in
synonymy only (.C.B.N. Art. 23.1 (d)).
However, examination of Watson’s many other works reveals that he did in fact publish the
binomial C. nigrescens in his first supplement to the Cybele Britannica (Watson 1860). It appears on
p. 81 in a list of species simply as Cerastium nigrescens, and, despite the fact that no author citation
was given, it is clearly referable back through the title of the book to the name used in his original
work of 1847 in which he validated the name C-. latifolium var. nigrescens. Strangely, there might be
some doubt whether the book is effectively published under the requirements of the /nternational
Code, for at the foot of the title page it reads “Printed for private distribution’. A notice on p. 2,
however, says that it ‘‘will be sent to all the provincial botanists of Britain whose present addresses
are known to the author. ... As the author cannot bind himself certainly to proceed with the
Supplement .. . he feels that it cannot properly be made a published work’’. ‘Publication’ in
Watson’s sense meant commercial publication, whereas in a botanical sense ‘‘Publication is effected
.. . by distribution of printed matter (through sale, exchange or gift) to the general public. . .”
(.C.B.N. Art. 29.1). The latter is clearly fulfilled by Watson’s Supplement, and this publication must
be taken as validating the specific combination.
Watson did not produce any more supplements to his Cybele Britannica. However, in his later
Compendium of the Cybele Britannica, he first (Watson 1868: 126) sank C. nigrescens completely
into C. latifolium, but later (1870: 492) gave it as a segregate taxon of his aggregate species latifolium,
in the form ‘“‘Cerastium (latifolium?) nigrescens, Edm.”’. His comments in his introduction (Watson
1870: 462-463) that “segregates ... can be distinguished from almost all the rest, the non-
indigenous plants, by their names being given in triplets, the name of the aggregate species being
inserted between the generic name and the distinctive name of the segregate. . . Itis at the reader’s
own choice to receive that distinctive name as varietal or as specific, in accordance with his own views
in each separate instance” make it clear that he did not clearly adopt specific rank for nigrescens here.
This cannot, however, detract in any way from his unambiguous publication of a specific binomial in
his 1860 supplement, and it is clear that the name C. nigrescens (H. C. Watson) Edmondston ex H.
C. Watson predates C. arcticum Lange by 20 years and must be taken up as the correct name for the
latter. Unfortunately then, following Flora Europaea taxonomy, the widespread taxon hitherto
known as C. arcticum subsp. arcticum requires a new combination.
Correct names of British taxa and synonymy relevant to Britain are given below. Since it is obvious
from the above that nigrescens and edmondstonii were essentially alternative epithets for the same
taxon, we have deliberately chosen the same lectotype for both, making all names involving either
epithet homotypic.
CERASTIUM NIGRESCENS (H. C. Watson) Edmondston [Fl. Shetland, xv, 29 (1845), in synon. |
ex H. C. Watson, Part First Suppl. Cyb. Brit., 1: 81 (1860). TYPE: Shetland, Baltasound, Aug.
1844, Edmondston (Lectotype: K, in herb. H. C. Watson; chosen here).
C. latifolium var. edmondstonii [anon. in Lond. Cat. Br. Pl., 2 (1844) nom. nud.| Edmondston, FI.
Shetland, 29 (Feb. or early March 1845); H. C. Watson in Phytologist, 2: 93 (later March 1845).
TYPE: as for C. nigrescens above, chosen here as lectotype.
. latifolium var. nigrescens H. C. Watson, Cyb. Brit., 1: 233 (1847).
alpinum var. edmondstonii (Edmondston) Hook.f., Stud. Fl. Br. Isl., 3rd ed., 60 (1884).
. arcticum var. edmondstonii (Edmondston) Beeby in Scott. Nat., n.s., 14: 24 (1887).
. edmondstonii (Edmondston) Murb. & Ostenf. in Bot. Notiser, 1898: 246 (1898).
. arcticum forma nigrescens (H. C. Watson) Druce in Moss, Cambr. Br. Fl., 3: 48 (1920).
. edmondstonii var. nigrescens (H. C. Watson) Clapham in Clapham, Tutin & Warburg, Fl. Br.
Isl., 299 (1952). ‘
C. arcticum subsp. edmondstonii (Edmondston) A. & D. Léve in Acta Horti Gothoburg., 20: 110
(1956).
C. arcticum var. nigrescens (H. C. Watson) A. & D. Love loc. cit. (1956), nom. inval., without
proper basionym reference.
Oye views)
a. subsp. NIGRESCENS
C. latifolium var. acutifolium Edmondston, FI. Shetland, 30 (1845). Described from Shetland, no
specimen traced.
296 R. K. BRUMMITT, D. H. KENT, P. S. LUSBY AND R. C. PALMER
C. nigrescens var. acutifolium (Edmondston) Druce in Rep. botl Soc. Exch. Club Br. Isl. , 6: 476
(1922).
b. subsp. ARCTICUM (Lange) Lusby, comb. nov.
C. arcticum Lange, Fl. Danica, 17 (50): 7 (1880). TYPE: Greenland, Vahl (Lectotype: see Hultén
(1956: 459), not seen). :
C. arcticum subsp. arcticum A. & D. Léve in Acta Horti Gothoburg., 20: 110 (1956).
C. latifolium var. compactum Syme in Sowerby, English Botany, 3rd ed. 2: 87 (1864). Type not
traced.
C. arcticum var. alpinopilosum Hultén in Svensk Bot. Tidskr., 50: 450 (1956). Type not stated.
C. latifolium var. smithii Syme in Sowerby, English Botany, 3rd ed., 2: 87 (1864) is a superfluous
illegitimate name for C. atratum Lap. var. piloso-pubescens Benth., Cat. Pl. Indig. Pyren. Bas
Languedoc, 70 (1826) and must be typified by a plant from the Pyrenees. The latter name was
applied to British plants, presumably in error, by Lindley (1829) apparently on the advice of
Bentham.
A question may arise concerning the authorship of the name C. nigrescens given above. As
noted, Edmondston first proposed this name but did not validate it. The first validly published
name including the epithet nigrescens was C. latifolium var. nigrescens published by H. C. Watson
in 1847, and because this combination was never proposed by Edmondston it séems desirable to
attribute it only to H. C. Watson (cf. Laundon 1985). The specific binomial was proposed by
Edmondston in manuscript, and was validly published by H. C. Watson in 1860. Hence the citation
given above of C. nigrescens (H. C. Watson) Edmondston ex H. C. Watson. It may well seem
paradoxical that according to the present rules of nomenclature the epithet nigrescens, which was
first proposed by Edmondston, has to be attributed to Watson as the first validating author, while
the epithet edmondstonii, which was applied to the same plant almost simultaneously by Watson,
has to be attributed to Edmondston himself! Such is the result of the bibliographic mix-up between
the two men.
In our discussion we have omitted reference to the var. acutifolium described by Edmondston in
his Flora, though it is included in the synonymy above. The taxonomic status of this will be the
subject of further research by one of us (P. S. Lusby).
REFERENCES
ALLEN, D. E. (1983). London Catalogue publication dates and authors. Watsonia, 14: 408-409.
ASPLUND, E. (1918). Beitrage zur Kenntnis der Flora des Eisfjordgebietes. (Cerastium). Arkiv for Botanik, 15
(14): 25-28.
Beesy, W. H. (1887). On the flora of Shetland. Scott. Nat., 1887: 20-32.
Bocuer, T. W. (1977). Cerastium alpinum and C. arcticum, a mature polyploid complex. Bot. Notiser, 130.
303-309.
Brett, O. E. (1953). Cerastium arcticum Lange. Nature, 171: 527-528.
CLAPHAM, A. R. (1952). Cerastium edmondstonii, in CLAPHAM, A: R., TuTin, T. G. & WARBURG, E. F. Flora of
the British Isles, p. 299. Cambridge.
CLAPHAM, A. R. (1962). Cerastium arcticum and C. nigrescens, in CLAPHAM, A. R., Tutin, T. G. & WARBURG,
E. F. Flora of the British Isles, 2nd ed., pp. 236-237. Cambridge.
Druce, G. C. (1908). British plant lists and their discrepancies: Cerastium arcticum Lange. Ann. Scot. nat.
Hist., 1908: 239-240.
Druce, G. C. (1911). The alpine Cerastia of Britain. Ann. Scot. nat. Hist., 1911: 38-44.
Druce, G. C. (1922). Flora Zetlandica. Rep. botl Soc. Exch. Club Br. Isl., 6: 457-546.
EDMONDSTON, Mrs. (1868). The young Shetlander, or Shadow over the sunshine, being life and letters of
Thomas Edmondston. Edinburgh.
Epmonpston, T. (1839). List of plants observed in the Island of Unst, Shetland, during the summer of 1837, in
Hooker, W. D. Notes on Norway, 2nd ed., pp. 111-117. Glasgow.
EpMONDSTON, T. (1841). List of phanerogamous plants, together with the cryptogamic orders Filices,
Equisetaceae, and Lycopodiaceae, observed in the Shetland Islands. Ann. Mag. nat. Hist., 7: 287-295.
EpMONDSTON, T. (1843a). Notice of a new British Cerastium. Phytologist, 1: 497-500.
EDMONDSTON, T. (1843b). Note on Cerastium latifolium. Phytologist, 1: 677-678.
THOMAS EDMONDSTON’S CERASTIUM 297
Epmonpston, T. (1845). A Flora of Shetland. Aberdeen.
Hooker, W. D. (1839). Notes on Norway, 2nd ed. Glasgow.
Hooker, W. J. (1844). The London Catalogue of British Plants. Lond. J. Bot. , 3: 288-295.
Hu ten, E. (1956). The Cerastium arcticum complex. Svensk bot. Tidskr., 50: 411-495.
Hy Lanper, N. (1945). Nomenklatorische und systematische Studien tber nordische Gefasspflanzen. Uppsala
Univ. Arsskr., 7: 1-337.
Jackson, B. D. (1888). Edmondston, Thomas. Dictionary of National Biography, 16: 397-398.
JALAS, J. (1964). Cerastium arcticum, in TuTin, T. G. et al., eds. Flora Europaea, 1: 141. Cambridge.
LANGE, J. (1880). Cerastium arcticum. Flora Danica, 17 (50): 7.
Launpbon, J. R. (1985). Proposal to emend recommendation 46E.1. Taxon, 34: 333.
LINDLEY, J. (1829). A synopsis of the British flora. London.
Love, A. & Love, D. (1956). Conspectus of the Icelandic flora. Acta Horti Gothoburg., 20: 65-291.
Lussy, P. S. (1984). The history and taxonomy of the Shetland taxon of Cerastium arcticum Lange. B.Sc. thesis,
University of Aberdeen.
MarsHaLl, E. S. (1911). Cerastium nigrescens Edmondston. Ann. Scot. nat. Hist., 1911: 119.
Murseck, S. (1898). Studies 6fver kritiska karlvaxtformer, 3. De nordeuropeiska formerna af slagtet
Cerastium. Bot. Notiser, 1898: 241-268.
OsTENFELD, C. H. (1920). A list of arctic Caryophyllaceae with some synonyms. Meddr Grgnland, 37 (12):
221-227.
SEEMANN, B. (1853). Narrative of the voyage of H.M.S. Herald, 1. London.
Watson, H. C. (1843a). Note on the supposed new British Cerastium. Phytologist, 1: 586-587.
Watson, H. C. (1843b). Note on the Cerastium latifolium of the Linnean herbarium. Phytologist, 1: 717-718.
Watson, H. C. (1844). Notes on the specific characters and varieties of some British plants. Lond. J. Bot., 3:
63-81.
Watson, H. C. (1845). On the Cerastium latifolium (Linn.) var. Edmondstonii (London Cat.); and on the
seeds of Cerastium latifolium and C. alpinum. Phytologist, 2: 93-94.
Watson, H. C. (1860). Part first of a supplement to the Cybele Britannica. London.
Watson, H. C. (1868, 1870). A compendium of the Cybele Britannica. London.
(Accepted March 1986)
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Watsonia, 16, 299-302 (1987) 299
Typification of Festuca rubra L., F. ovina L.
and F. ovina var. vivipara L.
CE. JARVIS
Department of Botany, British Museum (Natural History), London SW7 5BD
C. A. STACE and M. J. WILKINSON
Department of Botany, University of Leicester, Leicester LE] 7RH
ABSTRACT
Festuca rubra L., F. ovina L. and F. ovina var. vivipara L. (F. vivipara (L.) Smith) are each lectotypified so as
to preserve the usual current application of the names. The lectotype of the first name is in GB, and those of the
latter two in LINN.
INTRODUCTION
Festuca ovina L. and F. rubra L. are two widely distributed and ecologically important grasses that
exemplify two large, very variable aggregates of taxa. In Flora Europaea (Markgraf-Dannenberg
1980) 92 and 23 species respectively are attributable to these two aggregates, so that the precise
typification of F. ovina and F. rubra is a matter of considerable taxonomic significance. F. ovina
var. vivipara can be conveniently dealt with at the same time.
FESTUCA OVINA L., SP. PL., 73 (1753)
Linnaeus took his diagnostic phrase-name unchanged from his earlier (1745) Flora Suecica account
and listed four polynomials in synonymy, adding “Habitat in Europae collibus apricis aridis
vulgatissimum’’. He also appended an unnamed variety f, later to be named as var. vivipara.
Linnaeus’ own herbarium in LINN contains surprisingly little Swedish material and it seems that
his Swedish herbarium was probably kept apart from his main collection and was lost or destroyed.
Accordingly, there is no material which can be unequivocally associated with the Flora Suecica
accounts.
The first synonym is from Linnaeus’ Samling of et hundrade wdxter upfundne pé Gotland, Oland
och Smaland (1741) but it is not associated with any known extant material. The second is from van
Royen (1740) but no associated material in Adriaan van Royen’s herbarium in L has been found
(J. F. Veldkamp pers. comm. 1986). A polynomial from Linnaeus’ Flora Lapponica (1737)
account is cited third and associated with this are two specimens, one in Linnaeus’ Lapland
herbarium in the library of the Institut de France, Paris (see Fries 1861; Stearn 1957, p. 115) and
another (92.1 in the Savage (1945) catalogue) in LINN bearing the entry number “‘55” and “‘e
Lapponia”’ in Linnaeus’ handwriting. These two specimens are possibly duplicates, and belong to
the F. ovina aggregate. The fourth synonym is from Bauhin’s Pinax Theatri Botanici (1623) also
cited through Scheuchzer’s Agrostographia (1719). The Burser herbarium in UPS does not in this
instance contain any material which Linnaeus might have used in association with the Bauhin
polynomial.
Specimen 92.2 in LINN also belongs to F. ovina agg. but it was sent to Linnaeus by Arduino in
1761 and hence has no relevance for the purposes of typification. There are four specimens in the
Linnaean herbarium in S which have in the past been referred to F. ovina, but examination shows
that two are entirely unannotated. However, one of the others (fiche 37.19) bears “1. ovina” and
300 C. E. JARVIS, C. A. STACE AND M. J. WILKINSON
on the reverse “‘e Lapponia”’ in Linnaeus’ handwriting. We have seen only a microfiche of this
specimen, and from this cannot draw any conclusions as to the precise identity of the plant, even
to the extent of determining it as F. ovina agg. rather than F. rubra agg. The last specimen (fiche
38.5) relates to F. ovina var. vivipara and is discussed under that name.
Specimen 92.1 in LINN does not bear the name or number of the Species Plantarum account,
but the presence of the Flora Lapponica number and the correspondence of the specimen with
that in Paris indicates that Linnaeus almost certainly collected it in Lapland in 1732 and it was
hence available to him in 1753. It agrees fully with the Flora Lapponica diagnosis ‘‘Poa spiculis
ovato-angustis aristato-acuminatis”’. This specimen was also examined by the late C. E. Hubbard
(Kew) and by the late P. Auquier (Liége) and considered by both of them to be a suitable choice
of lectotype and to be in accordance with the currently accepted concept of F. ovina, as well as
with the protologue. Accordingly we designate specimen 92.1 in LINN as the lectotype of F.
ovina L. (Plate 2A).
In the 1970s the concept emerged of ‘typical’ F. ovina sensu strictissimo as a diploid taxon
characteristic of many regions of northern and central Europe, including northern Britain, but
one of uncertain occurrence in southern Britain and not occurring in France or Belgium. This
concept was developed mainly by Auquier, firstly in his doctoral thesis (Auquier 1974), where he
stated (p. 196) “il parait vraisemblable que le matériel original de son (Linnaeus’) F. ovina
corresponde aux populations diploides communes en Europe septentrionale et centrale; Linné
met d’ailleurs son espéce en synonymie avec des phrases descriptives tirées de flores de Scandi-
navie et de Suisse”. Later the same concept was used by Auquier in other works, notably in the
standard Belgian Flora (Auquier 1978), where (p. 757) he stated: “‘Festuca ovina L. s. str.,
espéce d’Europe sept., centr. et or., n’existe pas a l'état spontané dans le territoire [Belgium,
Luxembourg, and immediately adjacent parts of France, Germany and Holland] de la Flore’.
Another leading festucologist, M. Kerguélen (Guyancourt), has followed the same interpretation
(e.g. Kerguélen 1983, p. 16), and Markgraf-Dannenberg (1980, p. 145) also considered F. ovina
to be a diploid taxon (the basis of her citing Belgium in the distribution is not known). The view
that typical F. ovina is diploid and rare in western Europe has become generally accepted by the
leading Continental festucologists, and we also are of this opinion. At Auquier’s request, Hub-
bard had sent detailed measurements of specimen 92.1 to the former (letter of 2 May 1974), after
which Auquier expressed the view (letter to Hubbard of 8 May 1974) that ‘““Les mesures relevées
sur l’enchantillon 92.1 concordent bien avec celles que j’ai notées sur des recoltes de Scandinavie
et d'Europe centrale. Ces plantes diploides different par leur mensuration des populations tét-
raploides d'Europe occidentale’. The critical measurements of the proposed lectotype are:
spikelet length extrapolated to fourth floret 4.8-5.7 mm; second lemma length 3.0-3.5 mm;
second lemma awn length 1-1.3 mm. Auquier (1977, p. 100) stated in print that “le nom de F.
ovina doit étre réservé aux populations diploides largement répandues en Europe boréale, cen-
trale et orientale,” and after he finally examined specimen 92.1 in LINN on 1 November 1979 he
confirmed (orally to C.A.S.) that he considered this sheet to represent the diploid taxon. Hub-
bard’s views are expressed in a note at the Linnean Society dated 8-9 April 1974, and in greater
detail in the letter of 2 May 1974 mentioned above. Specimen 92.1 was also examined by M.J.W.
in 1983, and the same conclusions drawn. A paper typifying F. ovina by specimen 92.1 was
planned by Auquier and Hubbard (see Auquier 1977, p. 100), and at the time of their separate
deaths in 1980 a draft manuscript had been prepared by Auquier (oral communication to C.A.S.
1980).
In most of Britain, Belgium and northern France, as well as in some regions further east, F.
ovina sensu Strictissimo is replaced by various tetraploid taxa. Their degree and level of distinc-
tion from each other and from F. ovina are being examined in several laboratories; some
preliminary conclusions have been presented by Wilkinson & Stace (1985), but this is not the
subject of the present paper.
FESTUCA OVINA VAR. VIVIPARA. L., FL. SUEC., 2ND ED., 31 (1755)
Linnaeus recognized this proliferating grass as an unnamed variety in 1753 with the diagnosis
‘Festuca spiculis viviparis’’, but two years later gave it a formal varietal epithet. Four synonyms
were cited, together with “Habitat in alpibus Lapponicis, ubi nullum gramen magis frequens”’.
TYPIFICATION OF FESTUCA SPECIES 301
There are no relevant specimens in the Linnaean herbaria in H, MW, SBT or UPS. Specimen
92.5 in LINN (noted by Frederiksen 1981, p. 287) is labelled by Linnaeus “‘1f” (the number of the
taxon in Species Plantarum) and ‘“‘Lappo” (=Lapponia, the area of collection).
The first synonym is taken from Linnaeus’ Flora Lapponica account, no. 56, and there is a
specimen bearing this name and number in his Flora Lapponica herbarium in the library of the
Institut de France, Paris. There is also a further specimen bearing this number in Linnaeus’
handwriting in S (fiche no. 38.5), which is almost certainly a duplicate. Of the remaining three
synonyms, two are drawn from works by Scheuchzer and the third is from Ray. All three are
accompanied by illustrations, but none can be associated with specimens that Linnaeus would have
seen.
Specimen 92.5 in LINN was clearly used by Linnaeus when writing his Species Plantarum
account and we formally select it as the lectotype. Hubbard (ms. note in Linnean Society dated 8-9
April 1974) reached the same conclusion.
Although Linnaeus’ polynomial covers all proliferous Festuca taxa, which belong to many
species (most but not all of them within the F. ovina aggregate), the lectotype conforms precisely
with the modern concept of F. ovina var. vivipara, now usually known as F. vivipara (L.) Smith, Fi.
Brit., 1: 114 (1800). F. vivipara in northern Scandinavia exists as both triploids and tetraploids and
the lectotype could be either.
FESTUCA RUBRA L., SP. PL., 74 (1753)
As with F. ovina, Linnaeus used the polynomial name that he had earlier used in Flora Suecica
(1745, p. 93) as his diagnosis, and he cited one pre-Linnaean synonym from Scheuchzer. He also
noted ‘Habitat in Europae sterilibus siccis”. At the end of the entry Linnaeus added a sentence
with further information for distinguishing F. rubra from F. ovina, suggesting that he knew F.
rubra well: ““Magnitudine, colore maturitatis rubro, culmo tereti sed altero latere planiusculo,
distinguitur a F. ovina’’.
There are two specimens (92.9, 92.10) in LINN which are linked with this name. Specimen 92.9
is a member of the F. rubra aggregate and is labelled by Linnaeus “3 rubra” and, on the reverse,
‘“Lapponia 52”’. The former represents the name and number of the species in Species Plantarum,
and the latter would appear to be the origin of the specimen and its number in Linnaeus’ Flora
Lapponica account. However, entry number 52, Poa spiculis ovato-oblongis, foliis subulatis, is
nowhere cited in the synonymy of F. rubra, but is instead to be found along with the reference ‘FI.
lapp. 52” as a synonym of Poa angustifolia L. There is a specimen, numbered ‘‘52” by Linnaeus
and with the name “‘Poa spiculus ovato oblongis’’, in Linnaeus’ Lapland herbarium in the library of
the Institut de France, Paris. From a photograph it is clear that this specimen is a Festuca, not a
Poa, and probably belongs to F. rubra agg.; it is possibly a duplicate of specimen 92.9 in LINN.
However, since its name and number are referred in Species Plantarum to Poa angustifolia, not to
F. rubra, it cannot be regarded as a syntype of the latter, nor used to typify it.
Specimen 92.9 does not represent the taxon now generally known as F. rubra (sensu
strictissimo), but is an example of F. richardsonii Hooker (F. rubra ssp. arctica (Hackel)
Govoruchin). This conclusion was reached without reservation by P. Auquier when he examined
the specimen on 1 November 1979, and by S. M. Cunningham and C.A.S. when they examined it
in 1982 (see also Kerguélen 1983, p. 9).
Specimen 92.10 also bears the number “‘3” in Linnaeus’ handwriting, but Linnaeus additionally
wrote ‘‘2”’ on the sheet. This suggests some change of mind as to whether F. duriuscula L. (number
2 in Species Plantarum) or F. rubra was the identity of this specimen, which is therefore not
relevant for the typification of F. rubra.
There are three specimens in the Linnaean herbarium in S but none can be regarded as a
syntype. One is unannotated by Linnaeus, a second was received by Linnaeus from Arduino in
about 1761, and the third, although annotated by Linnaeus, lacks the figure ‘‘3”’ and was almost
certainly acquired by him after 1753. The last sheet bears a specimen each of F. rubra ssp. rubra
and F. rubra ssp. commutata Gaudin (F. nigrescens Lam.). No further relevant specimens are
known amongst the Linnaean herbaria in H, MW, SBI or UPS.
However, a further specimen has recently come to light in the Botanical Museum, Goteborg
(GB), which was not widely known to possess any Linnaean materials. During an informal
302 C. E. JARVIS, C. A. STACE AND M. J. WILKINSON
discussion with C.E.J., Dr Lennart Andersson, now Botanical Curator at GB, said that he thought
Linnaean material existed there. Dr Andersson’s predecessor, Dr Bo Peterson, kindly confirmed
that just one Linnaean specimen is in the herbarium, and that this is a sheet of Festuca rubra,
collected near Uppsala and evidently in Linnaeus’ possession by 1753. The sheet bears on the recto
“3 rubra’”’ and on the verso “in paludosis prati regii Upsalia’’, all in Linnaeus’ handwriting. The
presence of the “‘3”’ indicates that it is almost certain that the specimen was in Linnaeus’ possession
before 1753. Whilst it is true that the ecological information on the specimen does not agree with
that given in the protologue in 1753 (i.e. “in Europae sterilibis siccis’’), it is notable that the Flora
Suecica (1745) account rather unusually omits any habitat details. It may well be that the GB
specimen was collected prior to 1745 (and Linnaeus did state in Flora Suecica “habitat ubique in
Suecia, praesertim in Uplandia’’). Ecological descriptions for a species such as this were probably
rather unimportant to Linnaeus as he frequently omitted mention of them. In any case, it is clear
that Linnaeus possessed the specimen before 1753 and regarded it as belonging to his F. rubra, and
SO we accept it as a Syntype.
Examination of the specimen by C.A.S. and M.J.W. shows that the collection consists of only
the top parts of three flowering stems, but measurements of the spikelet parts (e.g. lemmas 5—6
mm, spikelets to fourth floret 7-8 mm) reveal that it is an example of the plant now commonly
known as F. rubra ssp. rubra. Hence we select it here as the lectotype (Plate 2B).
~
ACKNOWLEDGMENTS
We are greatly indebted to the late Dr P. Auquier (Liége) for many fruitful discussions with
C.A.S. on the taxonomy of the F. ovina aggregate, and similarly to Dr M. Kerguélen
(Guyancourt) for much valuable help concerning the taxonomy and nomenclature of both F. ovina
agg. and F. rubra agg. We are extremely grateful to Dr L. Anderson and Dr B. Peterson
(Goteborg) for drawing our attention to the existence of the Linnaean specimen in their care, and
for making it available for study. Our thanks are also due to the curators of the herbaria in L, LINN
and § for allowing access to material, and to the library of the Institut de France, Paris, for
permitting C.E.J. to study Linnaeus’ Lapland herbarium. Dr J. F. Veldkamp kindly checked the
existence and identity of van Royen material in L for us.
REFERENCES
AvuagquleR, P. (1974). Biosystématique, taxonomie et nomenclature du groupe de Festuca ovina L. s.l. (Poaceae)
en Belgique et dans quelques régions voisines. D.Sc. thesis, University of Liége.
AvuauleR, P. (1977). Taxonomie et nomenclature de quelques Festuca tétraploides du groupe de F. ovina L. s.1.
(Poaceae) en Europe moyenne. Bull. Jard. bot. nat. Belg., 47: 99-116.
AUvuQulIER, P. (1978). Festuca L., in DE LANGHE, J.-E. et al. Nouvelle flore de la Belgique, du Grand-Duché de
Luxembourg, du nord de la France et des régions voisines, 2nd ed., pp. 749-758. Meise.
FREDERIKSEN, S. (1981). Festuca vivipara (Poaceae) in the North Atlantic area. Nord. J. Bot., 1: 277-292.
Fries, T. M. (1861). Anteckningar rorande en i Paris befintlig Linneanska herbarium. Ofversigt Kongl.
Vetensk. Akad. Forh., 18: 255-272.
KERGUELEN, M. (1983). Les Graminées de France au travers de “Flora Europaea” et de la “Flore” du
C.N.R.S. Lejeunia, n. sér., 110.
MARKGRAF-DANNENBERG, I. (1980). Festuca L., in Tutin, T. G. et al., eds. Flora Europaea, 5: 125-153.
Cambridge.
SAVAGE, S. (1945). A catalogue of the Linnaean Herbarium. London.
STEARN, W. T. (1957). An introduction to the Species Plantarum and cognate botanical works of Carl Linnaeus,
in LINNAEUS, C. Species Plantarum. A facsimile of the first edition 1753, pp. 1-176. London.
WILKINSON, M. J. & Stace, C. A. (1985). The status of Festuca ophioliticola and related taxa. Soc. Echange PI.
vasc. Europe et Bassin médit., 20: 69-73.
(Accepted July 1986)
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Watsonia, 16, 303-309 (1987) 303
Typification and status of the mysterious Festuca guestfalica
Boenn. ex Reichb.
M. J. WILKINSON and C. A. STACE
Department of Botany, The University, Leicester, LEI] 7RH
ABSTRACT
The discovery of the type specimen of Festuca guestfalica Boenn. ex Reichb. is described and its diagnostic
features are presented. It is closely related to F. ovina L. and probably a tetraploid. It is concluded that there is
insufficient evidence to identify any plants from the British Isles as this central European taxon.
THE FESTUCA OVINA AGGREGATE
In its broadest sense F. ovina L. consists of a large number of taxa widely distributed in Eurasia
and North America. In Flora Europaea (Markgraf-Dannenberg 1980) 92 species (numbers 79-170)
belong to this aggregate. Hackel (1882), in his monograph of European fescues, had earlier used
almost exactly the same scope for his ‘F. ovina Linn. sens. ampliss.’; in fact only one of the 92
species recognized in the aggregate by Markgraf-Dannenberg (F. hystrix Boiss.) was excluded
from it by Hackel.
In British Floras a range of circumscription has been adopted (Table 1). Howarth (1925)
described the aggregate as ‘F. ovina L. sensu ampliss.’ and Tutin (1952) as ‘F. ovina agg.’
However, both these authors divided the group into separate species (five and four respectively).
Howarth (1948) reduced the number of species he recognized from five to four, F. supina Schur
having previously been recorded in error. Hubbard (1954) recognized five species. The later
editions of Tutin and Hubbard do not differ from the first, apart from the use of the name F.
longifolia Thuill. instead of F. trachyphylla (Hack.) Krajina by Tutin (1962).
Markgraf-Dannenberg (1980) recorded seven of her 92 species for Britain, and these were all
briefly treated by Tutin (1981). In addition F. armoricana Kerguélen and F. huonii Auquier are
known from the Channel Isles, but Markgraf-Dannenberg’s record of F. indigesta Boiss. from
Ireland is probably an error of identification. Since 1980 an eighth species, F. ophioliticola
Kerguélen, has been recorded from Britain (Wilkinson & Stace 1985); all eight are listed in
Table:1,
TABLE 1. TAXONOMIC TREATMENTS OF F. OVINA AGG. IN FIVE DIFFERENT WORKS
Markgraf-
Howarth 1925 Howarth 1948 Tutin 1952 Hubbard 1954 Dannenberg 1980
F. capillata F. tenuifolia F. ovina ssp. F. tenuifolia F. tenuifolia
tenuifolia
F. ovina F. ovina F. vivipara F. vivipara F. vivipara
F. ovina ssp. F. ovina F. ovina
ovina F. guestfalica
F. ophioliticola
F. glauca F. glauca F. glauca F. glauca F. longifolia
F. longifolia F. longifolia F. trachyphylla F. longifolia F. trachyphylla
F. lemanii
304 M. J. WILKINSON AND C. A. STACE
Our concept of F. ovina in Britain is covered by the descriptions of F. ovina, F. guestfalica and F.
ophioliticola given in Flora Europaea, but it excludes the other five species. It is possible to
subdivide taxonomically this concept of F. ovina (which may be called F. ovina sensu stricto), as
Markgraf-Dannenberg has done, but we prefer to recognize such segregates at infraspecific levels
rather than as species (Wilkinson & Stace 1985).
Festuca ovina has been typified (Jarvis, Stace & Wilkinson 1987) by a specimen in LINN which
represents a northern diploid plant (Festuca ovina sensu strictissimo) which is common in
Scandinavia and northern Britain but rare in southern England and apparently absent from
Belgium and France. In these more southern areas it is replaced by various tetraploid taxa.
Apparently the earliest name at specific level that could be used for any of these tetraploids is F. .
guestfalica Boenn. ex Reichb., and it is the application of this name that is the subject of the
present paper. <
TYPIFICATION OF FESTUCA GUESTFALICA
FESTUCA GUESTFALICA Boenn. ex Reichb., Flora Germanica Excursoria, p. 140 (3) (1831).
F. ovina ssp. eu-ovina var. vulgaris subvar. guestphalica (Boenn. ex Reichb.) Hackel, Monogr.
Fest. Europ., 87 (1882).
The name F. guestfalica has never been typified and was ignored by twentieth-céntury botanists
until Markgraf-Dannenberg (1980) adopted the name in Flora Europaea for a plant that she
claimed was distributed from Britain and France to Poland and Czechoslovakia. Reichenbach’s
description is too vague to give a good impression of the identity of the taxon, except that it
belongs to the F. ovina aggregate. If it is part of Festuca ovina sensu stricto it is, from its
distribution, probably a tetraploid. Markgraf-Dannenberg’s (1980) concept was apparently
intuitive, based upon her wide and long experience of the genus in central Europe. Although she
gave “Br” in the distribution, we have seen no British material so labelled by her. The few records
of F. guestfalica for Britain that have appeared in the literature recently are all errors, based on a
misinterpretation of Markgraf-Dannenberg’s concept of the species (e.g. Ellis 1983, p. 172).
Kerguélen (1982, 1983) has considered this problem, and gave a brief description of a Festuca
occurring in the Paris region that he thought came under Markgraf-Dannenberg’s concept of F.
guestfalica. However, he considered that this latter concept embraced several taxa, and he
surmised that the typical species of Reichenbach, the Parisian plant, another related taxon from Le
Mans, F. ophioliticola ssp. calaminaria Auquier, and at least three populations studied by Huon
(1970) in western France probably came under this name. However, he had no information on the
nature of the type material, identification of which he justifiably considered ‘‘indispensable”’.
Reichenbach’s “‘F. guestfalica a Bnngh.”’ (implying the name was supplied by Boenninghausen)
was based on “‘F. valesiaca var. B Weihe D. Gras. XI. no. 264.”’, said to come from “‘An Kalkfelsen
im Sauerlande in Westfahlen” (Westphalia, W. Germany, hence the epithet guestfalica). W.
Lippert (Munchen) kindly informed us (in litt. 1983) that ““D. Gras. XI” refers to Weihe’s
exsiccatum set Deutsche Graser ftir Botaniker und Oeconomen, XIte Sammlung, a set of which he
and other central European grass taxonomists had been seeking unsuccessfully for some time.
In January 1984 we distributed with the University of Leicester Botanic Garden annual Seedlist
an open request for information about Weihe’s exsiccatum. This resulted in the location of at least
two syntypes of F. guestfalica.
At Helsinki there is a set of fifteen bound volumes of Weihe’s Deutsche Grdser, volume XI of
which contains no. 264, which is indeed labelled “264. Festuca valesiaca var. 6B Am Felsen im
Sauerlande” (Plate 3A), with no other data. Volume XI was kindly lent to us by I. Kukkonen. It
contains 25 sheets (nos 251-275), as presumably do all the other volumes, but is undated (Fig. 1).
Dr Kukkonen tells us that Volume I is dated 1st October 1817, and Volume XII October 1824, and
that, since it contains a reference to Mertens & Koch’s third edition of R6hling’s Deutschlands
Flora (1823), Volume XI must have appeared in 1823 or 1824. Sheet no. 264 is clearly the plant
upon which Reichenbach’s F. guestfalica was based.
From Leningrad also we received anonymously on loan a single sheet of no. 264 from the same
volume. This has the same small printed label as that at Helsinki, under which someone has written
‘Weihe’. This sheet is obviously another syntype. Presumably the set of exsiccata at Leningrad has
305
TYPIFICATION OF FESTUCA GUESTFALICA
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306 M. J. WILKINSON AND C. A. STACE
been cut up and disposed into the herbarium in systematic sequence. No other examples of this
rare set of exsiccata have come to light.
We also received on loan from Edinburgh a sheet ex “Herb. Dérfler. Purchased 1912.” that
bears no other printed labels but two handwritten ones (Plate 3B). The upper was written by
Weihe (handwriting identified for us by B. Gries of Miinster, H. E. Weber of Osnabriick and G.
Wagenitz of Gottingen) and reads: ‘‘Festuca valesiaca B hirsuta . . . Weihe’’. The lower label starts
“543 b Festuca guestfalica . . .”’, followed by a description and ends “‘(Weihe D Graser 11 Heft)
. . . guestfalia montibus calcareis.’’ The handwriting has been identified for us by F. Kottwitz of
Nortrup as that of Boenninghausen. The absence of the number 264 on this sheet indicates that this
specimen cannot be considered a syntype, but in fact it might well be one. The description written
by Boenninghausen agrees with that in Reichenbach’s Flora, and the number 543 b might be an
error for 243 b; which is the number of F. guestfalica in the Flora. Moreover the reference to
Weihe’s exsiccata, Volume 11, by Boenninghausen, and use of the name F. valesiaca B by Weihe,
tie the specimen in with no. 264 in the exsiccata volumes. There remains the possibility that
Weihe’s and Boenninghausen’s labels were referring to the fact that the Edinburgh specimen was
taxonomically the same as F. guestfalica, rather than it being a type, even though the specimens
appear so similar that they might have come from the same gathering. The most important aspect
of the Edinburgh specimen, however, is that it bears a label by Boenninghausen with the name F.
guestfalica, which (uniquely) confirms Reichenbach’s attribution of the origin of. the name.
Accordingly, we designate the Helsinki exsiccatum as lectotype, the Leningrad specimen as
isolectotype, and the Edinburgh specimen as a possible isolectotype.
DESCRIPTION AND DISTRIBUTION OF FESTUCA GUESTFALICA
Our concept of F. guestfalica is based upon the three specimens mentioned above and upon a
number of other herbarium specimens in HAL, W and Z, from Germany and Switzerland, that
resemble the former closely and are obviously conspecific with it. Diagnostic measurements of the
type material are as follows: plant height 45-60 cm; leaf-blades Y-shaped to V-shaped in section,
with 2 grooves, 1 ridge, 6-7 veins and continuous or slightly broken, rather thin sclerenchyma;
panicles (6)8-12 cm; spikelets (to tip of 4th floret excl. awn) (6.1)6.5-7 mm; lower glume
(2.6)2.8-3.6(4.4) mm; upper glume (3.2)3.5-4.4(4.6) mm; lemmas 44.5 mm; lemma awns
(0.3)0.5-1(1.2) mm. The most notable feature of the types is the very lax panicle, setting the plants
well apart from most others in the Festuca ovina aggregate.
A more extended description based upon all the above material follows (Fig. 2) (measurements
are means of specimens):
Plants laxly tufted, with only intravaginal shoots, without rhizomes, retaining or shedding old
leaves. Culms (30)36—-58(68) cm, with 2-3 nodes (excl. inflorescence), the uppermost not pruinose,
usually visible beyond subtending sheath and reaching 11-26% up culm; stem below inflorescence
0.40.7 (0.8) mm wide, scabrid or occasionally smooth, strongly grooved when dry. Leaves green,
subpruinose or not; sheaths 2.9-7 cm, fused for 0-39% of length, smooth or occasionally
scabridulous in upper half, hairy or glabrous; auricles short, minutely ciliated or occasionally
glabrous; ligules <0.5 mm, minutely ciliated; blades 6.8-19.7 cm, very lank, not acutely pointed,
0-40(50)% on each plant curved at tip, scabrid at least in upper third, glabrous or hairy at base.
Inflorescences nodding or slightly so, 6-12.4 cm, very lax, with (8)10—14 nodes and 20—36(41)
spikelets; branches not pruinose or occasionally subpruinose, not narrowing below spikelets,
scabrid or scabridulous, the lowest two 1.7—-3.6 cm apart. Spikelets (6.6)6.7—7(7.1) mm, with 3-8
florets (the most apical one reduced and sterile), not pruinose or occasionally subpruinose; pedicel
1.4-2.3(2.7) mm, scabrid or scabridulous; lower glume narrowly triangular to narrowly lanceolate,
2.5-3.30.6-0.8 mm, scabrid at tip, ciliate at margin, with 1 vein; upper glume lanceolate to
narrowly so, 3.45—4.3 X0.9-1.15 mm, scabrid or hairy at tip, ciliate at margin, with 3 veins; lemmas
3.84.4 (excl. awn) X1.6-1.9 mm, lanceolate, hairy or scabrid in distal half, with 5 veins, with awn
0.6-1.1(1.9) mm; paleas linear-lanceolate, 4-4.8x0.65—0.85 mm; anthers yellow or purple,
(2.2)2.4-3.1 mm.
Leaf-blade in section: laterally compressed to oval, with margins not or slightly infolded,
(0.46)0.57-0.69(0.75) mm diameter/thickness ratio 1.9-2.5(2.7), with (5)7 veins, 2(4) grooves
TYPIFICATION OF FESTUCA GUESTFALICA 307
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FicurE 2. Spikelet (A) and transverse sections of leaf-blades of sterile shoots (B—G) of F. guestfalica. (A)
lectotype; (B) lectotype; (C) Edinburgh possible isolectotype; (D) Leningrad isolectotype; (E, F, G)
unlocalized German and Swiss specimens in Ziirich.
and 1(3) ridges on adaxial surface; sclerenchyma forming thin broken ring or sometimes
unbroken ring 1-2 cells thick; adaxial midrib 0.07-0.1(0.17) mm wide and 0.02-0.06 mm deep;
adaxial epidermal cells tuniform in size, lacking bulliform cells, with prickles (12.5)20—40(46.5)
um long.
Leaf-blade adaxial epidermis: stomata (33.5)35.5—39.75(44.5) um long, (20)40—80(85)%
solitary, 8-56(76)% with accompanying prickle-cell, 3-24% with accompanying silica-cell; prickles
63-93% solitary, 7-14(34)% forming rows of 2-few cells, 2-9(18)% with accompanying silica-cell.
Leaf-blade abaxial epidermis: stomata absent; long-cells with highly sinuous walls,
(105)140-185(250) um long; silica-cells and cork-cells present; prickles usually present.
We have encountered two British specimens that agree well with the lectotype in all critical
characters. One is a voucher (now in K) of a plant used by T. J. Jenkin in the hybridization
308 M. J. WILKINSON AND C. A. STACE
experiments carried out at Aberystwyth in the 1930s, 1940s and 1950s. The plant is that described
by him as F. ovina (Sheet 28, Experiment B143) (Jenkin 1955). He found it to be a tetraploid
(2n=28), and he obtained hybrids between it and F. rubra L. It originally came from an acid heath
in Pembrokeshire, Wales. The other is a plant collected fresh by T. C. G. Rich at Gaitbarrows, W.
Lancashire, England and cultivated at Leicester. Like the type of F. guestfalica, it came from a
limestone area. It is also a tetraploid (2n=28), with all the chromosomes metacentrics or
submetacentrics and the ratio of largest to smallest chromosomes 1.7.
We cannot confirm or deny the more extensive distribution of the species given by Markgraf-
Dannenberg (1980). We have seen only three specimens determined by her, and these do agree
with our concept of the species, but it must be remembered that her circumscription of the species
was drawn without having seen the type specimens. The plants from France thought of as possible
F. guestfalica by Kerguélen (1982, 1983) are not good matches for the type material, and we have
seen no material of F. guestfalica from France, Belgium or Holland.
TAXONOMIC STATUS OF F. GUESTFALICA
At first glance the type and similar specimens from Germany and Switzerland appear to represent a
very distinct taxon. By far the most distinctive character is the widely spreading, lax pan‘cle, quite
unlike that usually found in F. ovina sensu stricto or indeed within the F. ovina aggregate. The
most similar western European taxa appear to be F. ovina ssp. ophioliticola (Kerguélen) M.
Wilkinson and F. lemanii Bastard, both of which normally possess much more contracted
inflorescences, shorter, stiffer leaf-blades and shorter leaf-sheaths than F. guestfalica. Apart from
this F. lemanii differs mainly in its shorter pedicels, shorter lemma-awns, shorter anthers, 24
(rather than usually 2) grooves on the leaf-blades, and longer stomata (Table 2). These differences
seem consistent and within the F. ovina aggregate are of the order normally representative of
different species.
F. ovina ssp. ophioliticola differs additionally from F. guestfalica mainly in its shorter anthers
(more or less as in F. lemanii) and shorter upper glumes (Table 2). These differences are less
convincing.
It is possible that F. guestfalica is a distinct species, but equally possible that it is an extreme
variant of F. ovina ssp. ophioliticola. The two British plants closely resembling F. guestfalica throw
some light on this point. Both Jenkins (1955) and T. C. G. Rich (pers. comm. 1985) collected the
plants in the belief that they were representative of the segregate of F. ovina common in each of
the areas concerned. Yet no herbarium material of wild-collected F. ovina that we have seen from
those areas (or indeed from anywhere in Britain) closely resembles F. guestfalica. The two British
specimens that do seem identifiable with that taxon represent cultivated material, and it is likely
that they do not represent phenotypes normally encountered in the wild. Bidault (1968, p. 242)
listed panicle length and innovation leaf-length as two of the more plastic characters in Festuca.
The wide disjunction between the two British sites, and between those and the type locality of F.
guestfalica, suggest that it is unlikely that the former are outposts of a central European taxon.
We conclude that more investigations in Westfalia are needed to investigate the status of F.
TABLE 2. DIAGNOSTIC CHARACTERS OF FESTUCA LEMANII, F. GUESTFALICA AND
F. OVINA SSP. OPHIOLITICOLA
Character F. lemanii F. guestfalica F. ovina ssp. ophioliticola
Pedicel length (mm) 0.6-2(2.4) 1.4-2.9 (1.3)1.4-2.8(3.6)
Spikelet length (mm) (6.1)6.5-7.5(8.5) (6.6)6.7—7(7.1) (5.5)5.9-7(7.5)
Upper glume length (mm) (3.4)3.5—4.6(4.8) 3.45-4.3 3-3.9(4.9)
Lemma awn length (mm) (0.34)0.6-1.6(1.9) 0.6-1.1 (0.3)0.5—-1(1.6)
Anther length (mm) 1.8-2.5 (2.2)2.4-3.1 (1.65)1.85—2.2(2.85)
No. leaf-blade grooves 2-4 2(-4) 2(-4)
Stomatal length (um) (31)38-45.8 (33.5)35.5-39.75(44.5) 31-39.5
Chromosome number 42 728 28
TYPIFICATION OF FESTUCA GUESTFALICA 309
guestfalica, but that, whatever the results of such studies, it is probable that the British plants are
extreme ecophenes of F. ovina ssp. ophioliticola. If these two taxa are the same, it should be noted
that F. guestfalica is the correct name at the species level, but ssp. ophioliticola is correct at the
subspecies level.
ACKNOWLEDGMENTS
We are most grateful to the many herbarium curators who searched (mostly unsuccessfully) for
specimens on our behalf, especially to Dr I. Kukkonen (Helsinki) for loaning us Volume XI of the
Weihe set of exsiccata. We are also indebted to the following for much helpful advice on Weihe
and Boenninghausen and on the taxonomy of Festuca: B. Gries, M. Kerguélen, F. Kottwitz, W.
Lippert, F. Runge, G. Wagenitz and H. E. Weber; and to T. C. G. Rich for providing living
material of Festuca.
REFERENCES
Bipau Lt, M. (1968). Essai de taxonomie expérimentale et numérique sur Festuca ovina L. s.|. dans le sud-est
de la France. Rev. Cytol. et Biol. vég., 31: 217-356.
Exus, R. G. (1983). Flowering plants of Wales. Cardiff.
HackEL, E. (1882). Monographia Festucarum Europaearum. Kassel & Berlin.
HowartH, W. O. (1925). On the occurrence and distribution of Festuca ovina L., sensu ampliss., in Britain. J.
Linn. Soc. (Lond.), Bot. , 47: 29-39.
Howartn, W. O. (1948): A synopsis of the British Fescues. Rep. botl Soc. Exch. Club Br. Isl. , 13: 338-346.
Hussarp, C. E. (1954). Grasses. Harmondsworth.
Huon, A. (1970). Les fétuques de l’ouest de la France. Recherches de biosystématique et de biogéographie.
Thése Fac. Sciences Rennes, sér. C., 100/36 (1967). Rennes.
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ovina var. vivipara L. Watsonia, 16: 299-302.
JENKIN, T. J. (1955). Interspecific and intergeneric hybrids in herbage grasses, XIV. The breeding affinities of
Festuca ovina. J. Genet., 53: 118-124.
KERGUELEN, M. (1982). Les Festuca des groupes de F. ovina L. et de F. rubra L. s. lat. dans la région
parisienne. Cahiers des Naturalistes. Bulletin des Naturalistes parisienne, 38(1): 1-12.
KERGUELEN, M. (1983). Les graminées de France au travers de «Flora Europaea» et de la «Flore» du C.N.R.S.
Lejeunia, n. sér., 110.
MARKGRAF-DANNENBERG, I. (1980). Festuca L., in Tutin, T. G. et al., eds. Flora Europaea, 5: 125-153.
Cambridge.
TuTin, T. G. (1952). Festuca L., in CLAPHAM, A. R., TuTin, T. G. & Warbure, E. F. Flora of the British Isles,
pp. 1421-1427. Cambridge.
Tutin, T. G. (1962). Festuca L., in CLAPHAM, A. R., Tutin, T. G. & WARBURG, E. F. Flora of the British Isles,
2nd ed., pp. 1125-1130. Cambridge.
TuTIn, T. G. (1981). Festuca L., in CLAPHAM, A. R., TuTin, T. G. & WarsBure, E. F. Excursion Flora of the
British Isles, pp. 448-449. Cambridge.
WILKINSON, M. J. & Stace, C. A. (1985). The status of Festuca ophioliticola and related taxa. Soc. Echange PI.
vasc. Europe et Bassin médit., 20: 69-73.
(Accepted July 1986)
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Watsonia, 16, 311-327 (1987) 311
The distribution of Erica erigena R. Ross in Ireland
P. J. FOSS.and’G. J7 DOYLE
Botany Department, University College, Belfield, Dublin 4
and
E. C. NELSON
National Botanic Gardens, Glasnevin, Dublin 9
ABSTRACT
Erica erigena R. Ross (E. mediterranea auct. non L., E. hibernica (Hook. & Arn.) Syme non Utinet), Irish
Heath (Ericaceae), has a disjunct distribution in Europe. It occurs in the Iberian Peninsula, at Bordeaux in
western France, and in counties Mayo and Galway in the west of Ireland, but is absent from the remainder of
the European mainland. The distribution of E. erigena in Ireland is mapped and discussed in detail, and site
maps are presented for the major stations of the heather in Mayo and Galway. Relevant earlier accounts are
discussed. The Irish stations of this species are at present secure, but face a potential threat from further
exploitation of blanket peat and reclamation of former peatland.
INTRODUCTION
Erica erigena R. Ross (E. mediterranea auct. non L., E. hibernica (Hook. & Arn.) Syme non
Utinet), Irish Heath, has a markedly discontinuous distribution in western Europe. It occurs in the
Iberian Peninsula in southern Spain (see Cebrian 1948; McClintock 1971), along the coast of
Portugal from Serra de Monchique to Oporto (Malato-Beliz 1982), in north-western Spain (Fraga
1984) and in eastern Spain in the province of Valencia (Mansanet et al. 1980; Costa, Peris &
Figuerola 1983). There is also a small population north of Bordeaux in western France (Besancon
1978), but the heather does not occur elsewhere on the continental mainland. It is not a native
species in Britain, but was formerly naturalized in Cornwall (Williams 1911). In Ireland, E. erigena
is confined to counties Galway and Mayo, where its distribution is again disjunct. The most
southerly site is Errisbeg, near Roundstone in Connemara, Co. Galway; the most northerly
population is on the Mullet Peninsula, Co. Mayo. The western extremity of its Irish range is on the
shore of Lough Nakeeroge on Achill Island, and the most easterly site is on the eastern shore of
Lough Conn, both in Co. Mayo.
The first published account of the heather in Ireland was presented by Mackay (1830) who
reported the Errisbeg colony. However, herbarium specimens had been collected in western
Ireland by Edward Lhwyd as early as 1700 (Nelson 1979). These are lodged in the Morrisonian
Herbarium, University of Oxford (OXF). Since 1830 numerous papers have dealt with the
distribution of FE. erigena in Ireland, but examination of these reports shows that its Irish
distribution is not clearly defined. Many of the accounts are couched in general terms, others are
confused. In this paper we aim to present a comprehensive and updated description, with site maps
that clarify the present distribution of the plant in the west of Ireland. The distribution of E.
erigena in all its known sites in Galway and Mayo is discussed systematically, and pertinent earlier
descriptions are noted. This paper is based on the authors’ field studies between 1982 and 1984,
and voucher specimens for all the populations noted have been deposited in the Irish National
Herbarium, National Botanic Gardens, Glasnevin (DBN).
312 P. J. FOSS. G. J. DOYLE AND E. C. NELSON
eae ee Se
Ballina ¢
Lough ~@
Conn
Ao a GALWAY
AOE
| 20Okm
it untae ll ree 2%
Ficure 1. The distribution of Erica erigena in Co. Mayo and Co. Galway. Detailed distribution maps are
subsequently presented for those areas outlined on the map: the associated numbers refer to figure numbers in
the text. The colonies lying outside the major areas of distribution are shown by means of solid squares.
DISTRIBUTION OF ERICA ERIGENA 313
TABLE 1. THE GEOLOGY OF THE AREAS IN WHICH ERICA ERIGENA GROWS IN CO. MAYO
AND CO. GALWAY, LISTING SITE, IRISH NATIONAL GRID REFERENCE, GEOLOGY AND
LITERATURE SOURCE
Site Grid Ref. Geology Reference
Errisbeg L690.415 Base-rich metagabbros, especially Bremner & Leake 1980
black gabbros, epidiorites and
dolorites
Gowlaun L745.640 Silurian shale Whittow 1974
Mweelrea L780.655 Palaeozoic sediments, grits, brown Whittow 1974
sandstones, conglomerates &
sedimentary rocks of volcanic origin
Clare Island L705.870 Dalradian sandstones & slates, Cole et al. 1914
Carboniferous sandstones, shales
and conglomerates, and various
Silurian rocks
Lough Beltra M070.980 Old Red Sandstone on the eastern Whittow 1974
shore: Carboniferous sandstones on
the south and western shores
Lough Conn G145.130 Carboniferous limestones Whittow 1974
G195.147
Crossmolina G170.240 Carboniferous limestone Whittow 1974
Furnace L975.980 Old Red & Carboniferous Whittow 1974
sandstones
Mallaranny L825.980 Dalradian schists Whittow 1974
Curraun L770.970 Dalradian schists & quartzites: Old Whittow 1974
Red Sandstone on the southern
coast
Achill F697.085 Dalradian schists Whittow 1974
F585.080
Bangor-Erris Dalradian schists & Quartzites, Whittow 1974;
Lewisian gneiss & Carboniferous Linton 1964;
sandstone. These are overlaid by Orme 1969;
podzolized glacial drift, from the Synge 1968;
Munsterian Cold Stage, covered by Synge & Stephens 1960;
blanket peat Doyle 1982
DISTRIBUTION
The distribution of Erica erigena in Co. Mayo and Co. Galway is shown in Fig. 1. The major
centres of distribution, for which individual maps are presented later in the text, are indicated in
the diagram. The main localities are dealt with in detail in the text. The geology of each site is
presented in Table 1, along with the Irish National Grid reference and appropriate literature
references.
ERRISBEG, ROUNDSTONE, CO. GALWAY
Mackay (1830) noted finding E. erigena at Roundstone, and described its distribution as “a
declivity by a stream, in boggy ground, at the foot of Urrisbeg mountain, near Roundstone, on its
western side; occupying a space of above half a mile [0-75 km] in length, and covering between two
and three acres [c. 1 ha] of ground’’. Nelson (1982) noted that Mackay confused the story of the
rediscovery of the plant in later years. Mackay referred to visiting Connemara in 1829, while in the
paper, read on 30th November 1830, he stated that he had found it during an excursion “‘in the
latter end of the last month” (i.e. October 1830). Praeger (1934, 1939) gave a similar account of the
Errisbeg site but also indicated that the species occurred “‘around Lake Nalawney as well as on the
slope of Urrisbeg [sic] from 350 feet [106 m] down to Lough Bollard’’. Gay (1957) reported the
heather ‘‘around L. Nalawney on the south and east side, a little on the stream leaving the lough,
314 P. J. FOSS, G. J. DOYLE AND E. C. NELSON
i Lough
> Bollard ee
Errisbeg
Erica erigena colonies
isolated individuals
Ficure 2. The distribution of Erica erigena at Errisbeg, Roundstone, Co. Galway. The main streams running
down the north-west facing slope are marked A and B. The outlet stream from Lough Nalawney is marked C.
The contour lines are marked in metres.
but more on the stream entering it from Errisbeg’’. He also found some plants ‘‘on Errisbeg well
isolated from the main area”. Webb & Scannell (1983) defined the distribution of E. erigena at
Errisbeg as follows — “In fair quantity around L. Nalawney, spreading up the feeding streams for
some distance, and rather sparsely down the effluent stream and over the saddle to L. Bollard,
where there are a few plants on the S. shore; there is also an outlying colony on wet, level bog
about 1 km S.W. of L. Nalawney”’.
DISTRIBUTION OF ERICA ERIGENA 315
The population of FE. erigena at Errisbeg is more extensive than indicated in these earlier
accounts. The heather grows in a narrow belt, at its widest 1-5 m broad, on much of the shore of
Lough Nalawney (Fig. 2). There is a larger colony on the northern shore where the stream (A in
Fig. 2) that rises near the summit of Errisbeg enters the lough. Along this particular stream the
plant grows in abundance for a distance of 1-5 km up the slope of Errisbeg, to an elevation of 140
m. The heather has colonized the small tributaries of this stream but does not extend further than
200 m from the main channel. The stream passes through two large, quaking Schoenus nigricans L.
fens, where isolated E. erigena bushes are confined to elevated Schoenus tussocks around the
edges.
The heather is also plentiful along the second stream on Errisbeg (B in Fig. 2) but does not
extend much above an elevation of 100 m.
Where the streams form deep, steep-sided channels the heather grows on the banks in dense
stands often 1-2 m wide. Where the streams flow through level terrain and water spreads over a
wide area, the plants form stands up to 30 m across. There is considerable variation in the habit of
bushes growing by the streams, with a general decrease in size and vigour with increasing altitude.
The heather occurs along the outlet stream (C in Fig. 2) from L. Nalawney. It is abundant close
to the lough, but rapidly decreases until only isolated bushes occur on the stream bank.
About 20 small bushes form four discrete colonies in the area between Lough Nalawney and
Lough Bollard. A similar number of bushes grow along a 200 m stretch of the south-eastern shore
of Lough Bollard, and in a small flush running into that part of the lake. Another site has been
found (N. Kirby, D. Hogan & C. O’Connell pers. comm. 1984) on the eastern shore, to the north
of the previous site. Here the heather grows in a small flush flowing into the lake, and on well-
developed tussocks of Schoenus nigricans and Molinia caerulea (L.) Moench.
GOWLAUN, CO. GALWAY
The Gowlaun colony, described in detail by Scannell (1976), may be the remnant of a population
mentioned in general terms by J. MacKinnon as “‘around Salruck Pass’ (Colgan & Scully 1898).
Praeger (1934) reported that E. erigena was “‘not infrequent” in the Kylemore area, but this
species is not known at Kylemore today.
North-east of Gowlaun E. erigena grows on the headland east of Islandlyre, on the northern
shore of the Culfin River estuary (A in Fig. 3). At this coastal site, E. erigena grows in two distinct
habitats. A small colony of depauperate bushes is found on level reclaimed peaty ground, where a
hummock/hollow system has developed as a result of treading by domestic animals. The second
colony is located on a small, stony bluff, where the heather grows vigorously from cracks in the
rocks. A few isolated bushes occur along the edge of the Culfin River (B in Fig. 3), near the road.
These plants were first noted by Mrs. Willoughby (Magor 1981).
MWEELREA, CO. MAYO
E. erigena was first reported on Mweelrea, on the northern shore of Killary Harbour by Simon
Foot & Joseph Hooker in 1835 (Mackay 1836a). Ninian Niven (1836) visited Mayo and Galway,
including Mweelrea, in August 1836, and noted E. erigena, but it is not clear if he saw the plant at
Mweelrea or at Errisbeg, which he also visited. Hart (1883) found the plant along a stream near
Bunnaglass up to 30 m above sea level. Praeger (1911a) reported it from Killary, growing on a
rocky bluff at the base of Mweelrea Mountain. Gay (1957) found a few bushes by a stream on the
western part of the northern shore. He recorded large quantities further east, where it was
dominant over extensive areas and grew to an elevation of 250 m on the slopes of Mweelrea, and
also found it colonizing abandoned ‘lazy beds’ (potato drills) in some places. D. A. Webb stated
that the heather grew in great quantity immediately north of the harbour mouth, on the flanks of
Mweelrea, from 120 m down to sea level, and that this was the second largest station in Ireland (see
McClintock 1969).
The distribution of E. erigena on the northern side of Killary Harbour is shown in detail in Fig. 3.
The heather is first encountered on two small streams that run off the western flank of Mweelrea,
south of the Bunanakee River. It is generally confined to the stream edges and to adjacent level
flushes, but some isolated bushes grow beside a path linking the two streams. To the east of this
area E. erigena becomes more abundant, growing along numerous streams and in flushes, from sea
level to 150 m. The largest stand of the heather in this area spans a group of four streams about 3-5
316 P. J. FOSS, G. J. DOYLE AND E. C. NELSON
River
Q@
N os
| Mw e e | req Q
ss “3 Z 827
e
sete Soe foes,
A
499
Bunnaglass
lslandlyre
FicureE 3. The distribution of Erica erigena at Gowlaun, Co. Galway, and at Killary Harbour and Mweelrea,
Co. Mayo. The coastal site near Gowlaun is marked A. The station on the Culfin River is marked B. Mountain
peaks are indicated by solid triangles, the associated numbers are heights in metres. Extensive colonies and
isolated heather bushes are depicted by stippled areas and solid circles respectively.
km from the mouth of the harbour. FE. erigena dominates the lower slopes of the mountain,
growing on stream banks, in flushes and on small boulder clay banks above the high water mark.
The plant has colonized abandoned ‘lazy beds’ in the vicinity, where it is confined to the troughs.
The most easterly site is on the stream-bed at Bunnaglass, as originally reported by Hart (1883).
CLARE ISLAND, CO. MAYO
In an earlier botanical account of Clare Island, Praeger (1903) indicated that a “constant search”’
had failed to locate this or the other ‘‘western heaths”’ (i.e. Daboecia cantabrica (Huds.) C. Koch
and Erica mackaiana Bab.). In the botanical section of the report of the Clare Island Survey,
Praeger (1911b) remarked that one of the important additions to the island’s flora was E. erigena.
He reported E. erigena as “‘abundant on boulder-clay slopes (no trace of peat) facing north-east,
from storm level to about 70 feet [c. 20 m], between Portlea and Ooghcorragaun . . . A couple of
outlying plants on top of the low cliff at east end of Portlea’’. The heather was found in the same
spot by McClintock (1969) “‘still in fair quantity and condition”’.
The Clare Island site was visited in August 1984. The heather is confined to a steep, north-east
facing, deeply eroded boulder-clay bank in the area described by Praeger, about 200 m south of the
abandoned harbour at Portlea. It was growing vigorously, reaching a height of 75 cm.
LOUGH BELTRA, CO. MAYO
The descriptions of the populations of FE. erigena at Lough Beltra demonstrate most clearly the
need for a general restatement of this species’ distribution. Praeger (1934) noted that “this little-
DISTRIBUTION OF ERICA ERIGENA 317
known lake is remarkable for the fine fringe of Erica mediterranea [E. erigena| which, as at
Carrowmore Lake to the NW [north-west], covers the broad stony storm-beach of the eastern
side, half of it submerged during average floods’”’. Webb (1954) described the fringe of heather as
“very narrow and precise’. Gay (1957) mapped the colony on the eastern side of the lake and
other colonies along the edges of a stream entering the lough from the south and by
Derrynafreva and Derrynaloughan loughs. McClintock (1966, 1969) noted the Beltra colonies; in
the latter paper he wrote that ‘‘there is still a good colony covering a quarter of a mile [0-4 km]
or so towards the south end of the west side of this lake and also on the land side of the road in a
young forestry plantation’, and he also stated that ‘““Webb has seen it at the extreme south end”’.
McClintock’s reference to a colony on the western shore was an obvious printer’s error
(McClintock pers. comm.).
The distribution of the heather around Lough Beltra was mapped in the spring of 1984. It
grows in abundance on both the eastern and western shores. On the eastern shore, the fringe is
20 m wide and extends 1-5 km along the shore (B in Fig. 4). Throughout, the heather grows on a
thin layer of peaty sand overlying a rocky substrate, and is either the dominant shrub, or co-
dominant with Myrica gale L. There is a rocky storm-beach in front of the main heather zone,
where isolated bushes grow on eroding soil hummocks. On the landward side of the main fringe,
scrub dominated by Alnus glutinosa (L.) Gaertn. and Ulex europaeus L. contains a few isolated
heather plants. There are some bushes scattered to the east of the road, both in the forest
plantation as mentioned by McClintock (1969) (A in Fig. 4), and to the north of the main shore
colony (C in Fig. 4).
On the western side of the lough, E. erigena forms a sparse fringe along much of the shoreline,
at or just below the high water mark (D in Fig. 4). In some places there are large colonies that
penetrate about 30 m inland on cutover bog. The heather does not occur in some areas where the
shore is sandy. It also forms an encircling fringe on the two larger islands in the western part of
the lough.
On the southern shore (E in Fig. 4) there is a well-developed colony. Much of the area is
covered by a narrow band of heather 1-4 m wide, but in one place the heather dominates a
headland about 30 m across. It is found only on one of the two streams that enter this area from
the cutaway bog to the south (F in Fig. 4). Between the lough shore and the confluence of the
two streams there are isolated bushes on the steep banks. Along the remainder of the channel
the heather grows abundantly on the peaty edges and extends on to adjacent cutover bog at the
southern end of the stream (F in Fig. 4).
E. erigena is also found on the shores of two loughs that lie to the west of the stream. The
smaller Derrynaloughan Lough (G in Fig. 4) comprises two lakelets on a feeder stream to the
Newport River, the main outflow from Lough Beltra. E. erigena grows on the shores of both
lakelets, for a short distance along several drains flowing into them, and on the exit stream from
the more westerly of the two. The larger Derrynafreva Lough (H in Fig. 4) lies directly on the
Newport River, and has a discrete colony of E. erigena, forming a band 10-15 m wide along its
eastern shore.
LOUGH CONN, CO. MAYO
Specimens of EF. erigena, collected at Enniscoe House on the western side of Lough Conn, were
sent by J. T. Mackay to W. McNab at the Royal Botanic Gardens, Edinburgh on 24th October
1837 (McClintock 1966). Praeger (1900) reported six heather bushes on the eastern shore of L.
Conn “on a patch of bog on the shore opposite Annagh Island’’, and commented that this was
the most easterly station in Ireland. He also described its habitat on the western shore, again on
boggy ground, from the Errew peninsula northwards to Enniscoe House. Gay (1957) and
McClintock (1969) saw the colony on the western shore, but neither succeeded in finding the
eastern shore colony. McClintock (1969) also remarked that this area had been searched in vain
in 1956 by D. A. Webb and again in 1968.
E. erigena still grows on both the western and eastern shores of Lough Conn. On the western
shore (G145.130) the plant grows on cutover bog, particularly on the edges of drains, on
desiccated peat hags, and less frequently in wet, disturbed depressions. There are two separate
areas where the heather grows in abundance, one on the Errew Peninsula and the other on a
headland immediately south of Enniscoe House. The intervening land has been reclaimed for
318 P. J. FOSS, G. J. DOYLE AND E. C. NELSON
a
Cloondaff Beltra
Birreen
FicureE 4. The distribution of Erica erigena at Lough Beltra, Co. Mayo. A = the plantation site, B = the east
shore colony, C = the isolated bushes on the landward side of the road, D = the west shore fringe, E = the
southern shore colony, F = the colony on one of the inflow streams on the southern shore, G = the
Derrynaloughan Lake colony, H = the Derrynafreva Lake colony. Stippled areas are extensive colonies, solid
circles indicate isolated bushes. Mountain peaks are indicated by solid triangles, the associated numbers are
heights in metres.
agriculture, and here the heather has been reduced to a few isolated bushes along the shore,
forming a connection between the two large populations.
The heather was collected in March 1984 on cutover peat on the eastern shore (G195.147),
opposite Annagh Island as reported by Praeger (1900). Two small colonies were found which have
a total of seven mature plants, 50-70 cm tall, and an undetermined number of smaller plants. This
remains the most easterly station of EF. erigena in Ireland.
CROSSMOLINA, CO. MAYO
The Crossmolina station, discovered by P. J. O’Hare (McClintock 1969), lies 6 km north of Lough
Conn, beside the road from Crossmolina to Killala, near Cloonachoor (G 17.24). E. erigena grows
DISTRIBUTION OF ERICA ERIGENA 319
abundantly on cutover bog, extending about 100 m on either side of the road. It has colonized dry
peat hags, and the intervening wet depressions where the heather forms stout tussocks.
THE LAKE COMPLEX AT FURNACE, CO. MAYO
The precise distribution of E. erigena around Furnace (L 975.980) is not obvious from published
accounts. Marshall (1900), commenting on its distribution in west Mayo following a visit in 1899,
wrote that “it extends eastwards nearly to Newport, but apparently avoids the limestone’’,
presumably referring to the occurrence of the heather near Furnace (Fig. 5), 5 km north-west of
Newport. Praeger (1934) also suggested that the heather grew close to Newport. Webb (1952)
described a fringe of the heather around Furnace Lough, which he suggested (Webb 1954)
resembled the fringe on Loughs Beltra, Carrowmore and “other lakes”. Gay (1957) recorded E.
erigena around most of Furnace Lough, in small quantity at the south-west end of Lough Fadda,
and occasionally on the outlet stream between the lough and Carrowsallagh Bridge. He also
reported it on the most easterly tributary of the stream running southwards from Bengorm
Mountain, and along the abandoned railway embankment in the area. McClintock (1966, 1969)
described the distribution of the plant around Furnace Lough and stated that it was ‘‘abundant on
the fringes of much but not all of this lake and by smaller lakes and streams to the west’’.
The distribution of E. erigena around the loughs at Furnace is shown in Fig. 5. The heather
Bengorm
Za
7 Furnace f
/_,Lough _f as
a oie ee 0 eee RD a Se Oe ey.
Ae ar
On ~
Carrowsallagh
Bridge
Mallaranny- Newport road
|km
Clew
Bay
Figure 5. The distribution of Erica erigena around the lakes near Furnace, Co. Mayo. A, B, C = extensive
colonies on cutover peat, D = a small lake with a colony on its western shore, E = the colony on one of the
tributaries of the Yellow River, F and G = Loughs Navroony and Naprasky, H = the colony on an unnamed
lakelet and the stream entering Furnace Lough, I = Lough Pollagowly, J = Lough Skahaghadrantan, K =
Lough Fadda, L = Doontrusk Lough, M = Lough Stirkeen. Extensive colonies are stippled, isolated bushes
- are indicated by solid circles.
320 P; Js FOSS?.G?73: DOYLEAND:-Ey'Ce NELSON
generally forms a distinct fringe along the shore of Furnace Lough, 1-3 m wide, with bushes up to2 m
tall. The heather also covers more extensive areas of cutaway peat on the north-western shore (A in
Fig. 5), on the western shore (B in Fig. 5) and again on the southern shore (C in Fig. 5), where
colonies extend 30 m inland in places. On the eastern shore of the lough, it occurs as isolated bushes
and only forms the typical fringe in sheltered bays. It is absent where the shore is sandy, as at the
mouth of the Yellow River. To the east of the lough, south of the Yellow River the heather extends
along a small flush eastwards of the road for 40 m. The heather is found further up this flush system,
on the western shore of a small lake (D in Fig. 5). E. erigena also grows on one of the tributaries of the
Yellow River to the north of the road (E in Fig. 5), and on the Yellow River to the south.
Distinct fringes occur on two drains that enter the lough on the west side. In the north-western
corner, it grows along the edges of the drain that connects Furnace Lough with Lough Navroony (F)
and Lough Naprasky (G). There are isolated bushes and occasional small colonies on the southern
shore of L. Navrooney. The heather is found around the entire shore of L. Naprasky; the bushes are
small and unevenly distributed in some areas, but form a large colony on the south-western shore,
and grow around the edges of, and sparingly on a quaking Schoenus flush. The plant also grows
extensively on cutover peat to the west of L. Naprasky. The heather has colonized the sides of a
second drain, flowing into the south-western corner of Furnace Lough from an un-named lakelet (H)
on which the plant is found on the northern shore.
E. erigena grows around other small loughs to the west of Furnace Lough. Isolated bushes are
found along the entire shore of Lough Pollagowly (I). There is a distinct fringe, 1-2 m wide around
the whole shore of Lough Skahaghadrantan (J) and numerous bushes along the stream entering the
lake from the south-west. The heather forms a sparse fringe around Lough Fadda (K) and extends
onto cutover bog to the south, and along the drain to Doontrusk Lough (L), where a few isolated
bushes are found on the southern shore. The heather grows on the drainage channel connecting
Lough Fadda and Lough Stirkeen (M), where it fringes the eastern and northern shores. The plant
extends south-westwards along the outflow stream from L. Stirkeen as far as the abandoned railway
line.
E. erigena grows on the stream running southwards from Bengorm. It forms a continuous band on
the banks of the most easterly tributary and on the main stream as it passes through rough pasture.
Only isolated bushes are found where the channel passes through an area of relatively intense
agricultural activity, to just south of the abandoned railway line. The plant does not grow on the
wooded section of the river bank, north of Carrowsallagh Bridge as suggested by Gay (1957). A
small patch is found west of the stream on the steep north-facing railway embankment. It no longer
grows at Carrowsallagh Bridge as recorded by McClintock (1969).
MALLARANNY, CO. MAYO
The largest population of E. erigena in Ireland is on Bellacragher Bay and Claggan Mountain north
of Mallaranny in Mayo. It was first reported by Newman (1839) who stated that it grew in great
abundance at ‘‘Molyrhany, a cluster of some 12 or 18 cabins”. Moore (1852) erroneously stated that
in this area one could see “‘at least a quarter of a million acres in extent covered with the lovely
heath”. Praeger (1934) gave a more conservative account saying that the heather grew from
Mallaranny to the “head of Bellacragher Bay’. Webb (1954) rejected Moore’s estimate as
exaggerated and noted that the heather was common at Mallaranny and on the shores of
Bellacragher Bay as far as the northern end of Claggan Mountain. Gay (1957) stated that E. erigena
was well represented on both sides of the bay, from sea level to 80 m altitude, but was restricted to
stream sides and damp valleys higher up the slopes. He indicated that the heather persisted along the
river running northwards to Bellaveeny Lodge, until the river entered blanket bog on level ground.
He also mapped the plant just east of, and directly behind the village of Mallaranny.
The distribution of FE. erigena around Mallaranny is shown in Fig. 6. To the east of the village there
are colonies (1—2 m wide) along the banks of the Bunnahowna and Murrevagh rivers, from just south
of the old railway line to 150 m altitude on the southern flank of Claggan Mountain. These rivers
have steep sided valleys that afford protection for the heather which is vigorous and up to 2-75 m tall.
On the slope immediately north of Mallaranny, the heather forms separate colonies in flushes, on
the edges of small streams and on cutover peat. The plants are stunted, as a result of exposure and
grazing. The heather also occurs on the abandoned railway embankment on the level surface and
sloped banks.
DISTRIBUTION OF ERICA ERIGENA 321
Blacksod |
Bellaveeny
Achill
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FicureE 6. The distribution of Erica erigena at Mallaranny and Curraun, Co. Mayo. Bellacragher Bay runs
northwards from Mallaranny, between Claggan Mountain and the Curraun Peninsula, and connects with
Blacksod Bay to the west. Extensive colonies are stippled, isolated bushes are indicated by solid circles.
Mountain peaks are depicted by solid triangles, the associated numbers are heights in metres.
West of Mallaranny the plant covers an extensive area on the slopes of Claggan Mountain. This
colony, the largest in Ireland, extends for 7 km from the village westwards to the junction of the
roads from Achill Island and Bellaveeny, thence along the eastern shore of Bellacragher Bay, and
the base of the north-west facing slopes of Claggan Mountain. Colonies exist north-eastwards on
the stream flowing towards Bellaveeny, until this enters the deep peat-covered coastal plain north
of Claggan. On the eastern side of Bellacragher Bay the heather forms an almost continuous belt
from sea level to 50 m altitude, and occupies stream beds and flushes on the hillside up to 150 m
altitude.
CURRAUN PENINSULA, CO. MAYO
According to Hart (1883), Curraun was formerly known to botanists as the headquarters of E.
erigena in Ireland. He reported it on the north-west side of the peninsula, below 50 m altitude. This
322 P. J. FOSS, G. J. DOYLE AND E. C. NELSON
should probably be interpreted as the western side of Bellacragher Bay (Fig. 6) rather than all of
Curraun. Praeger (1934) stated that the heather was abundant on the lower ground of Curraun, but
did not indicate if this referred to the whole peninsula. Webb (1952, 1954) gave further
descriptions stating that the plant occurred on the western side of Bellacragher Bay and on the
southern side of Curraun, where it was abundant beside almost every stream but absent from the
intervening bog. Gay (1957) showed colonies on the south coast and on a stream flowing from the
lakes marked Ard Loughs on the Ordnance Survey map. He also noted a new station on the west
side of the peninsula at Belfarsad. White (1968) presented a detailed vegetation map for Curraun,
on which she showed colonies on the eastern side of Bellacragher Bay, three widely separated
colonies on the south coast, and another new colony north-east of Belfarsad. McClintock (1969)
and Vanden Berghen (1972) did not add to the detail of the earlier reports.
The distribution of E. erigena on Curraun is shown in Fig. 6. Along Bellacragher Bay, the
heather occurs sporadically, extending from north of Glennanean Bridge to the southern end of
the bay. The heather occurs on stream banks, in small flushes, along the abandoned limestone
railway embankment, and on abandoned ‘lazy beds’ above the high water mark. It is also found
west of the road, generally confined to the banks of streams and flushes.
On the south coast the heather is found near Cushlecka, on both sides of the road growing on the
banks of a stream, and behind a beach of Old Red Sandstone boulders where it has colonized a
partially drained peat bog. It also forms two discrete colonies on a stream at Dooghbeg on both
sides of the road. At Carrickteige it occurs near the confluence of two streams, one of which flows
from Lough Ard Beg. This colony extends to the sea shore, and spreads along the top of a storm
beach for about 150 m. It also grows sporadically along the road to the east of Carrickteige. At
Lough Beg the plant occurs on the banks of a stream, abundantly on a flushed area on the northern
side of the road, sparingly on heavily grazed level ground, and on stream banks between the road
and sea shore. It also occurs close to the road further west, on the stream bank which flows down
from Ard Lough.
On the western side of the Peninsula the heather grows along the Bunanioo River (south of
Glassillaun), at Glassillaun, Mweewillin and between these two villages, and south of Belfarsad. A
large colony of impoverished bushes is situated north-east of Belfarsad, on the edge of a stream
and on severely grazed and eroded blanket peat.
ACHILL ISLAND, CO. MAYO
More (1889) mentioned that he had seen E. erigena on Achill in 1872, and cited two new Achill
localities discovered by J. R. Sheridan. Colgan & Scully (1898) listed the sites discovered by
Sheridan as Ridge Point and near Dugort (see also Moffat 1898). Praeger (1904) stated that the
heather was rare on Achill Island, growing chiefly in the east of the island. He specifically referred
to the following locations: the stream behind Valley Strand; from Lough Doo to Bull’s Mouth (the
Ridge Point site); by the streamlet which rises near Black Lough, from its source to its mouth near
Salia; and finally in the west, seen only half way along the northern margin of Annagh Lough (the
more easterly of the two lakes named Lough Nakeeroge on Ordnance Survey maps). This
particular station is the most westerly in Ireland.
D. A. Webb (see McClintock 1969) recorded E. erigena at Mweelin Lough, north-east of
Dooega village in 1957; our recent attempts to find the plant in this locality have been unsuccessful.
McClintock (1966, 1969) found the plant in just two Achill sites, at Annagh Lough (Lough
Nakeeroge East) and at Doogort.
An earlier account appears to suggest that the plant was found at Achill Sound. Moore & More
(1866) said it grew “all along Achill Sound, on the shore of the mainland opposite Achill Island”.
Colgan & Scully (1898) omitted the comma between the two phrases, thereby suggesting that the
plant was not found on Achill Island, but was confined to the mainland side (i.e. Curraun).
We have refound only the two sites seen by McClintock, although the other areas were searched
thoroughly. At Annagh Lough in the north-west of the island, about 40 bushes were seen in a small
bay on the northern lakeshore, and at the western end ten plants were growing along a small
stream that flows into the lough.
East of Doogort, and south-west of Lough Nambrack, the heather grows in a distinct colony (200
x 50 m) on cutover bog on the northern side of the road. There are a few bushes in a similar
habitat south of the road.
DISTRIBUTION OF ERICA ERIGENA 323
BANGOR-ERRIS, CO. MAYO
The earliest extant collection of herbarium material of E. erigena from the Barony of Erris seems
to have been by C. Vernon who sent it to David Moore in 1849 (fide DBN). However, Mackay
(1836b) credited the discovery to John Wynne, and it is he who was acknowledged by Colgan &
Scully (1898). Moore (1860) stated that he had traced the plant from Carrowmore Lake in the
north-west as far south as Achill Sound, but noted that it was confined to a coastal belt, up to 3
miles [4-8 km] inland, and was most abundant close to the sea. According to Colgan and Scully
(1898) Hart found E. erigena in two localities on the Mullet Peninsula in 1887. Praeger (1905) gave
further information of the distribution on the Mullet, indicating that the plant “haunts the tract
west of Belmullet, between the northern hill-bogs and the sands and tillage which extend
southwards’. Praeger (1934) summarized the distribution of the plant in the Erris area, indicating
that it was ‘widely spread’’, and was found fringing Carrowmore Lake, and occurred on boggy
ground on the Belmullet Peninsula.
Webb (1954) discounted the notion of the plant’s widespread occurrence in Erris, stating that he
could find it in only two localities other than at Carrowmore, one at a pair of lakelets 3 km north of
Gweesalia, the other around Lough Nahelly 3 km further north-west. Gay (1957) mapped the plant
at L. Carrowmore; along the road from L. Nahelly to Tristia; at Derrynameel (west of Glencastle
Hill), and at the well on the road from Bangor to Belmullet. Gay was unable to find it on the
Mullet Peninsula.
P. J. O'Hare noted several new colonies, including two south of the Glenamoy Peatland
Experimental Station (O’Hare 1959). He found the species at Muingerroon Bridge (on the road
from Glenamoy to Barnatra), at Faulagh (east of Barnatra), both east and west of Derrynameel on
the road from Barnatra to Derrycorrib, on the road from Derrycorrib to Bunnahowen, and near
Munhin Bridge on the main Belmullet to Bangor Road. He also recorded a number of locations on
the Mullet Peninsula, which had been described in general terms by Praeger (1934). McClintock
(1966) found another colony above the saltmarsh at Barnatra in 1957.
The distribution of E. erigena on the Erris Peninsula is shown in Fig. 7. It covers extensive areas
north of Gweesalia, but is generally restricted to places where the peat has been disturbed,
including cutover bog, drains at the edge of forestry plantations, and on the edges of drains along
roads such as that running north-westwards from Tristia.
The plant is abundant in disturbed blanket bog to the west of the road from Gweesalia to
Doolough Lodge, and is confined to drains, the bases of peat banks, and less commonly to water-
filled depressions between cutover peat hags. In some of these areas E. erigena is the dominant
species.
E. erigena is also found to the east of the same section of road, growing on relatively undisturbed
deep blanket bog. In this situation the plants are isolated depauperate individuals, and the species
never becomes dominant.
At Lough Nahelly there are small plants around the western shore behind a 20 m wide sedge
zone. The heather extends onto cutover bog further west of the lough, occurring in drains, at the
base of peat banks and on troughs in the cutover bog. Plants are generally small and the species is
never dominant.
Further north, it grows in a ditch beside the road from Bunnahowen to Derrycorrib, where there
are a few isolated bushes. It is found on the road from Bunnahowen to Derrynameel, growing
along the roadside and spreading westwards onto an adjacent cutover bog. Two colonies occur
near Derrynameel. At the eastern site the heather has colonized a desiccated peat hag and a wet
depression along its edge; the colony extends 30 m from the road, and includes stout bushes up to
1 m tall. At the western site the plant occupies a large area of drained and partially reclaimed peat,
which is subjected to heavy grazing by cattle. In this situation there are numerous dead bushes, and
the living plants are depauperate.
At Barnatra, the heather occurs on cutover bog on the landward side of the road, and more
extensively along a 300 m stretch on the seaward side. Here it generally grows above the level of
the highest spring tides, though a few bushes clearly lie below the flotsam line near the upper
saltmarsh. Another colony, near Faulagh, covers an area some 20 X 5 m on flushed cutover peat.
This site is heavily grazed by sheep and cattle. Further east about 40 bushes grow on grassy,
cutover peat on the hillside north-east of Muingerroon Bridge.
The stations reported from south of Muingerroon Bridge, and south of the road opposite the
P. J. FOSS)'G.' J) DOYLE AND EJ) GUNELSON
324
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DISTRIBUTION OF ERICA ERIGENA 325
Glenamoy Research Station were not found during our survey. The two latter sites, described by
O’Hare (1959), are in areas now covered by mature conifer plantations.
The populations at Carrowmore Lake are depicted in Fig. 7. Many of the earlier accounts are
vague, but Gay (1957) mapped the well-known fringe on the western shore, the colony on the
stream entering the lough on the western side, and another in the south-eastern corner. The
extensive colony on the north-eastern shore has not been reported before. The western shore
colony stretches almost uninterrupted for 3 km from Derreens to Gortmore, and varies in width
from 1—10 m; the bushes are 50-100 cm tall. The plants grow on shallow sandy peat and rocky
mineral soil. O’Hare (in McClintock 1969) observed a marked expansion in this population over 25
years ago when the water-level was lowered and seedlings colonized the exposed lake bed. The
heather also extends up the edge of the stream entering the lough on the western shore to an
altitude of 90 m. The species covers extensive areas of soligenous peat on the northern and
southern corners of the large bay lying due east of Gortmore. There are isolated stunted bushes on
the shore between these two colonies, and on the northern headland of the bay. In the north-
eastern corner of the lough the plants form a sparse colony along much of the shore; here it is
generally small (15-40 cm) and grows below the obvious high-water drift mark.
The heather is found on the main Bangor to Belmullet road, just west of Munhin Bridge. It
grows along the edge of two small streams that flow in the depressions between cutover peat banks,
and extends about 250 m from the road. The heather also covers an area of 200 x 200 m on a large
area of flushed, intact peat to the west of the more westerly of the streams. At Fauleens, a few
plants grow along a stream north of the road and numerous bushes grow on cutover peat along the
base of the hill north of this stream.
On the Mullet EF. erigena is found in the general vicinity of Tonmore. It grows on cutover bog,
colonizing the elevated dry peat hags, drain edges, and embankments on the edge of roads. It
forms an extensive colony along a small road running westwards from a chapel (marked on the
Ordnance Survey map) to Tonmore, and then southwards to the junction of five roads. The other
colonies on the Mullet are small and isolated. In every case, plants are small and weak and the
species is rarely the dominant one in the vegetation.
CONCLUSIONS
The distribution of Erica erigena in Ireland must be regarded as very local, although it is more
widespread than has been previously reported. Several new stations for the species have been
located and a number of poorly known sites refound. The conservation status of the heather in
western Ireland would seem to be secure at the present time. The major sites do not appear to be
threatened, although some are overgrazed. The heather seems to be increasing in the cut-over peat
sites, as it grows vigorously along desiccated peat hags and flat peat disturbed during cutting.
However, the heather’s long term prospects are not so guaranteed. It will probably survive in its
lakeshore habitats where interference is unlikely, but it will soon be threatened where it grows at
the edge of extensive blanket peatland areas, and on the cutover peat sites. In recent years there
has been extensive exploitation of blanket peat in the west of Ireland (Doyle 1983), where there
has been extensive reclamation for forestry and some commercial exploitation for fuel. Forestry
plantation is a serious threat as it is carried out at a rate of some 5000 ha per year (Dillon et al.
1976), and once established tends to obliterate the native peatland vegetation entirely (Doyle &
Moore 1982). If this heather is to survive in these peatland edge and cutover sites a proper
conservation plan will be required.
ACKNOWLEDGMENTS
We wish to thank Miss M. J. P. Scannell, D. McClintock, D. Small and Professor D. A. Webb for
discussion on the distribution of E. erigena in western Ireland. We acknowledge the assistance
provided by P. J. O’Hare (Assistant Director, Agricultural Institute, Oak Park, Co. Carlow), who
allowed us to consult his records of FE. erigena on the Bangor-Erris peninsula. We thank Dr N.
Kirby and D. Hogan of the Connemara National Park for stimulating discussion and practical help
326 P. JUFOSS) GG; J; DOYLEANDIEHCANEESON
at Roundstone. This work forms part of a Ph.D. thesis carried out at the Botany Department,
University College Dublin by P.J.F. We acknowledge the financial support of the Botany
Department, the Irish Department of Education and the Office of Public Works.
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(Accepted November 1985)
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Watsonia, 16, 329-338 (1987) 329
Short Notes
THE HERBARIUM OF BOURNEMOUTH NATURAL SCIENCE SOCIETY
A recent visit to the museum of this Society resulted in the discovery that the returns on which the
entries for its herbarium (BMH) in Kent (1957) and Kent & Allen (1984) were based were
misleading in certain important respects.
The herbarium consists in fact of just two separate collections: a Dorset one formed in 1910-18
by Charles Baylis Green, and a much larger one representative of the British Isles as a whole. The
first of these, mounted on small and rather flimsy sheets, contains material from many parts of v.c.
9, not (as stated) from the Isle of Purbeck alone, and extends to certain critical groups (though not,
for example, Rubus), the sheets of which carry determinations by the respective national referees
of the inter-war period. Oddly, though, it would appear not to have been consulted by Good
(1948) when compiling his checklist of the flora of the vice-county.
The other collection, described simply as ‘formed by members’’, seems to be basically the
combined herbarium of T. A. & C. Cotton, which has hitherto been untraced. This doubtless came
to the Society in or around 1925, the year of death of both of these collectors, who latterly had
Bournemouth connections. Thomas Atkinson Cotton was Honorary Secretary for several years of
the Watson Botanical Exchange Club, of which his wife Charlotte was a long-standing member as
well, and many of their sheets, as expected, have labels that bear the stamp of that. Not all the ones
from other collectors, however, can be from that source and some of these the Cottons may have
bought, for in at least certain other botanical directions they are known to have spent generously.
Possibly that is how the Society comes to be the possessor of numerous sheets of W. H. Purchas,
the joint author of the Flora of Herefordshire (particularly valuable, as his main collection was
destroyed in the wartime bombing of Bristol City Museum). R. V. Sherring, on the other hand,
another collector who is substantially represented, was a Vice-President of the Society, so his
specimens must be presumed to have been acquired directly and independently. That was
definitely the case with two other sets of specimens that have been incorporated into this general
Society collection. One of these consists of various common Rubus species from the Bournemouth
district which the Rev. W. Moyle Rogers is on record as having donated to the Society in 1909-10,
the other of what remains of the presumably once much more extensive herbarium of John
Frederick Rayner, author of A supplement to Frederick Townsend’s Flora of Hampshire and the
Isle of Wight (1929). For, sad to say, the whole of the Society’s general collection was drastically
thinned at some time in the 1930s or 1940s — by a misguided honorary curator who deemed it
superfluous for any taxon to be represented by more than a single sheet (N. Douglas Simpson pers.
comm. 1951). Among the many specimens destroyed Rayner’s appear to have featured in
disproportionately large numbers, probably because the sheets on which they are mounted are an
awkwardly smaller size than the rest. That they were also collected at more recent dates may also
have been a factor. Unfortunately, as a result, most of the vouchers for Rayner’s more interesting
records have thereby been lost irretrievably.
REFERENCES
Goon, R. (1948). A geographical handbook of the Dorset flora. Dorchester.
KENT, D. H. (1957). British herbaria. London.
Kent, D. H. & ALLEN, D. E. (1984). British and Irish herbaria. London.
D. E. ALLEN
Lesney Cottage, Middle Road, Winchester, Hants., SO22 5EJ
330 SHORT NOTES
ATRIPLEX LITTORALIS NEW TO SHETLAND
During fieldwork in Shetland, v.c. 112, in August 1985, some plants of Atriplex littoralis L. were
discovered near Boddam (GR 411/397.155) in S. E. Mainland. This discovery extends the range of
this species by about 240 km north of its previously mapped limits in the British Isles (Taschereau
1985).
The plants were growing in tall, ungrazed and rather rank vegetation, mainly of Elymus repens
(L.) Gould and Juncus gerardii Loisel., at a low level just above the shore, and very close to the
lower limit of angiosperm cover. Four plants were seen; two were collected and pressed (one
deposited at BM and one with W. Scott, Scalloway for incorporation in ZCM, Lerwick). These
plants have very compact inflorescences at anthesis; low magnification stereo microscopic study
showed apparently normal pollen formation. In October, the site was revisited independently by
W. Scott and the writer, and further plants were found in the same limited area. At this time, the
plants were much taller and in fruit, some of which dropped easily from the stems when touched.
W. Scott examined some fruits and was able to confirm the earlier identification based on foliage
characters.
The very limited area occupied by A. Jittoralis near Boddam and its previous omission from
Shetland records (apart from the inconclusive and unsupported reference to this species by Hilliam
(1977)), suggest that it may be a recent newcomer to the islands.
The northern limit in Britain (Taschereau 1985) lies at approximately 57° 40’ N., corresponding
very closely with the northern limit for frequent occurrences as a native in the Skagerrak and Gulf
of Finland in the Baltic (Hulten 1971), suggesting that common climatic conditions limit extensive
establishment in the north, although there are scattered locations for A. littoralis up the west coast
almost to the extreme north of Norway, perhaps as a result of the effects of the Gulf Stream. The
species is absent from Faerée and Iceland (Aellen 1964).
If A. littoralis has indeed recently arrived in Shetland from a source in mainland Britain, then
one might expect it to be found in the south of the archipelago. For climatic reasons too, one might
expect a more thermophilous species to gain a foothold in the extreme south of Shetland. In this
connection, the comments by Bullard (1985) about Orkney are relevant. She writes that the
“apparently unlikely Atriplex littoralis turns up occasionally as strand-line seedlings which fail to
overwinter’, but she does not refer to the survival of any mature plants.
In conclusion, it is not at all easy to predict the future of the Boddam colony, bearing in mind the
spasmodic fluctuations in population size of this species at Milford Haven in S. Wales (Dalby 1969,
1971), the often temporary nature of shoreline habitats and the location of the site rather north of
the general limit for Atriplex littoralis, but its chances may be promising if future winters are not
too severe. W. Scott, however, is inclined to suspect that the Boddam colony has been there a long
time, overlooked, and is likely to persist unless its habitat is destroyed.
ACKNOWLEDGMENTS
Thanks are due to Dr M. G. Richardson and the Nature Conservancy Council for permission to
publish this report, which is based on observations made during contract work for N.C.C. in
Shetland; both Walter Scott and Mike Richardson made helpful comments on the manuscript.
REFERENCES
AELLEN, P. (1964). Atriplex, in Tuttn, T. G. et al., eds. Flora Europaea, 1: 95-97. Cambridge.
BuLLARD, E. R. (1985). In Berry, R. J. The natural history of Orkney, p. 250. London.
Datsy, D. H. (1969). A further note on Atriplex littoralis in Milford Haven. Nature in Wales, 11: 141-142.
Datpy, D. H. (1971). Grass-leaved orache at Waterloo. Nature in Wales, 12: 169-170.
Hittiam, J. (1977). Phytosociological studies in the southern isles of Shetland. Ph.D. thesis, University of
Durham.
Hutteén, E. (1971). Atlas of the distribution of vascular plants in northwestern Europe. Stockholm.
PERRING, F. H. & Watters, S. M., eds. (1982). Atlas of the British flora, 3rd ed. Wakefield.
SHORT NOTES 331
TASCHEREAU, P. M. (1985). Taxonomy of Atriplex species indigenous to the British Isles. Watsonia, 15: 183-
209.
D. H. DALBy
Department of Pure and Applied Biology, Imperial College, London, SW7 2BB
>)
———
a
—-
m
ie’.
x
ww
a
FESS =
oes en
a
100 zm
Figure 1. Material of PolystichumxXillyricum (Borbas) Hahne from Bundalloch, collected by the author
(Ferguson 1785) and housed in the University of Antwerp Herbarium (RUCA). A. Outline of frond; B. Detail
of three fertile pinnae; C. Mature pinnae (9th and 10th pinnae from base); D. Empty sporangia. Illustration by
A. Torfs.
332 SHORT NOTES
AN ADDITIONAL SITE FOR POLYSTICHUMXILLYRICUM (BORBAS) HAHNE IN
NORTH-WESTERN SCOTLAND
In the course of a field trip in August 1984, an unusual shield-fern was encountered in the Glennan
Valley, 1 km east of Bundalloch, v.c. 105 (GR 18/906.276). It was growing on a relatively dry site
about 40 m above sea-level. One frond, 275 cm, with thirty pairs of pinnae, was collected for
further examination (Fig. 1).
The presence of sori ruled out the possibility that one was dealing with an immature frond of
Polystichum aculeatum (L.) Roth which, while not encountered in the Glennan Valley, is recorded
from the same 10-km square (Jermy et al. 1978). Likewise, P. lonchitis (L.) Roth, while still
unrecorded from the Glennan Valley, is known from two of the neighbouring squares. However,
in this case the species is only simply pinnate. There seemed to be no alternative but to assume a
hybrid origin. This was confirmed by examining the sori. While the indusia and some of the
sporangia were well developed, the sporangia proved to be empty (Fig. 1). The spores had
aborted. Since the nearest site for Polystichum setiferum (Forsk.) Woynar is more than 25 km
away, the putative parents are probably P. aculeatum and P. lonchitis, i.e. one is dealing with the
hybrid P. xXillyricum (Borbas) Hahne. Since P. Xillyricum is practically indistinguishable from
P. Xlonchitiforme (P. lonchitis Xsetiferum) on morphological grounds, this identification must
await confirmation in the form of a chromosome count.
Although P. aculeatum and P. lonchitis are largely sympatric in Scotland (J ermy et al. 1978), the
hybrid has so far only been reported from a single site, i.e. Inchnadamph (Stirling 1974), although
it has also been recorded from Co. Leitrim, v.c. H29 (Page 1982). The present record confirms the
surmise (Sleep & Synnott 1972; Page 1982) that this hybrid could be found in other areas of the
British Isles.
REFERENCES
JerMy, A. C., ARNOLD, H. R., FARRELL, L. & PERRING, F. H. (1978). Atlas of ferns of the British Isles. London.
Ps 3E, C. N. (1982). The ferns of Britain and Ireland. Cambridge.
Steep, A. & Synnott, D. (1972). Polystichum Xillyricum: a hybrid new to the British Isles. Br. Fern Gaz., 10:
281-282.
STIRLING, A.McG. (1974). A fern new to Scotland — Polystichum Xillyricum in West Sutherland. Watsonia, 10:
25h.
D. K. FERGUSON
Department of Botany, University of Antwerp (RUCA), Groenenborgerlaan 171, B — 2020,
Antwerp, Belgium
FURTHER NOTES ON DACTYLORHIZA INCARNATA SUBSP. CRUENTA_ IN
SCOTLAND
In a recent paper (Kenneth & Tennant 1984), we announced the discovery in 1982 of Dactylorhiza
incarnata (L.) So6 subsp. cruenta (O. F. Mueller) P. D. Sell from West Ross, v.c. 105, at that time
new to mainland Britain, and gave some descriptive information about the plants and details of
their habitat. Since that time, further field work has been undertaken and the data obtained during
1984 have now made it desirable to provide some additions and modifications to the original
information published in that paper.
Flower measurements were not carried out in 1982, and 1984 measurements are therefore given
in Table 1. Studies carried out on the different populations in 1984 showed that the uppermost
population did not have larger flowers as suggested in our paper (Kenneth & Tennant 1984) and
also the general appearance and colour of the flowers in this population was barely distinguishable
from those of the lower populations, unlike the situation found in 1982. The flower colour and
markings were well within the range of D. incarnata subsp. cruenta from Europe, and probably
SHORT NOTES 333
TABLE 1. CHARACTERS OF D. INCARNATA SUBSP. CRUENTA IN WEST ROSS (1984
MEASUREMENTS)?*
Height (cm) 7.5-21 (14)
Length of longest leaf (cm) 3-9 (5.4)
Maximum width of longest leaf (cm) 0.8-2.0 (1.1)
Ratio leaf length to width 4.91
Length of inflorescence (cm) 2.5-5.8 (3.5)
Number of flowers in spike 5-20 (12)
Labellum length (mm) 5.5-7.5 (6.4)
Labellum width (mm) 5.0-8.0 (6.4)
Spur length (mm) 5.0-7.0 (6.0)
Spur width (mm) 2.0-3.0 (2.5)
Length of bract peripheral cells (um) 36-67 (52)
*Field measurements made on 26 plants in June 1984, being additions and modifications to the original, 1982
data (Kenneth & Tennant 1984). Mean values are shown in brackets.
also from Ireland (R. Bateman pers. comm.). The peak flowering period in 1984 was also earlier
than in 1982, being from mid- to late June, and in 1984 there was no significant difference in
flowering state between the uppermost and lowermost populations. In 1982, flowering in the
uppermost population extended well into the second half of July. It is therefore likely that the
differences observed between these populations in 1982 were due to climatic factors as we
previously suggested, such as late snow-lie affecting only the upper population in that particular
year.
The main features which distinguish the West Ross plants from other variants of D. incarnata in
Britain are the distinctive, largish, violet-brown spots on both sides of the expanded sheathing
leaves, the heavy streaking or flecks of anthocyanin on the ridges or veins of the upper stem, bracts
and ovaries, and the labellum shape and pattern (Fig. 1). The West Ross populations of subsp.
cruenta differ in general appearance in all these respects from D. incarnata (L.) So6 subsp.
pulchella (Druce) So6, which was present amongst one of the populations examined; subsp.
cruenta also differed from subsp. pulchella at the same locality because of the slightly less erect,
less channelled, sheathing leaves in the former and, in the case of few-flowered plants, because of
the more spreading ovaries, giving subsp. cruenta a laxer floral spike. Additionally, the labellum of
subsp. cruenta was more distinctly trilobed, with a more intense pattern of distinctive, violet-
coloured loops, rings and dots (Fig. 1).
The shape and dimensions of the bract peripheral cells agree closely with published values for
the taxon, being rounded and barrel-shaped in outline, and not at all angular as is typically found in
tetraploid taxa.
Approximately 7.5% of the West Ross plants of subsp. cruenta examined had unspotted leaves.
It was noted, however, that such plants deviated slightly in other characters from typical plants
with spotted leaves, and that they were present only in the single population where D. incarnata
subsp. pulchella also occurred. It is conceivable that such plants might have been inter-subspecific
hybrids with subsp. pulchella, rather than mere morphs.
In spite of the number of interesting plants listed in Table 2 of the original paper (Kenneth &
Tennant 1984), which described the habitat, several were in fact noted in the nearby vicinity,
whereas those species which were more closely associated with the dactylorchids showed that the
habitat seemed to differ little from other Schoenus nigricans - Molinia caerulea - Scirpus cespitosus
- dominant communities which are so widespread in the north-western Highlands. The majority of
the dactylorchids occurred in flushed grassland where the soil pH was nearly neutral to mildly
alkaline. Additional species found to be present but not listed in the original paper (Kenneth &
Tennant 1984) included Ranunculus flammula, Myrica gale, Dactylorhiza incarnata subsp.
pulchella, Sphagnum sp. and Breutelia chrysocoma.
Specimens and photographs of D. incarnata subsp. cruenta from West Ross have been placed
in E.
334 SHORT NOTES
FiGuRE 1. D. incarnata subsp. cruenta from West Ross.
(a) Whole plant; (b) sheathing-leaf illustrating leaf spotting on upper surface; (c) sheathing-leaf illustrating leaf
spotting on lower surface; (d) flower illustrating steaked ovary, spur, and markings; (e) labellum shape
(pressed) and markings, subsp. cruenta (right) compared with subsp. pulchella (left). Scale bar=10 mm (a-c),
3 mm (d, e).
ACKNOWLEDGMENT
We should like to thank R. H. Roberts for provision of data on the bract peripheral cells and
general comments on the text.
REFERENCE
KeNNETH, A. G. & TENNANT, D. J. (1984). Dactylorhiza incarnata (L.) S06 subsp. cruenta (O. F. Mueller) P.
D. Sell in Scotland. Watsonia, 15: 11—14.
A. G. KENNETH & D. J. TENNANT
Marhead Grange, Arkendale, Knaresborough, North Yorkshire, HGS ORG
SHORT NOTES 335
ALLIUM AMPELOPRASUM L. IN ANGLESEY
Allium ampeloprasum L. is a native of southern and western Europe, with its main distribution
centred around the Mediterranean basin. It is a variable species within which three
morphologically differentiated taxa are usually recognized. These are now treated as varieties: var.
ampeloprasum with a dense, globose umbel of flowers (up to 500); var. babingtonii (Borrer) Syme,
with an irregular umbel having few flowers and numerous large bulbils (usually 8-12 mm) and
often with small, secondary umbels borne on longer pedicels; and var. bulbiferum Syme, with
fewer, smaller bulbils (6-8 mm) in the umbel (Stearn 1980). However, there is still no general
agreement on the taxonomy. Although var. babingtonii was formerly treated as a distinct species,
A. babingtonii Borrer, it is now generally regarded as a variety of A. ampeloprasum. Opinions
differ regarding var. bulbiferum, which Clapham et al. (1981) now do not distinguish from var.
ampeloprasum.
On the Continent, where most records seem to refer to var. ampeloprasum and only a few to var.
bulbiferum, the plants are found on disturbed ground (Stearn 1980). In the British Isles, however,
it has been recorded from a variety of habitats, mostly around the coast from the Isles of Scilly,
Cornwall, Guernsey, Steep Holm (Somerset), Flat Holm and Porth Kerry (Glamorgan) to Tenby
(Pembrokeshire) (Clapham et al. 1962; Ellis 1983). The Channel Island plants appear to belong to
var. bulbiferum, which is also recorded from [le d’ Yeu, Vendée, on the western coast of France
(Stearn 1980). In Cornwall, where both var. ampeloprasum and var. babingtonii are recorded (the
latter being much the more frequent) the plants occur mainly on roadside verges, Cornish ‘hedges’,
waste ground and occasionally in more natural habitats such as sandy coastal marshes (Margetts,
pers. comm. 1986). In the Isles of Scilly, where only var. babingtonii occurs, it is found on
hedgebanks, field borders, cliffs, dunes and rough bushy places (Lousley 1971). In Ireland, where,
similarly, all populations belong to var. babingtonii (Scannell & Synnott 1972), they are found
mostly on the islands along the western and north-western coasts, usually growing in rock crevices
and sandy places (Praeger 1934).
Its status in the British Isles is uncertain: while some authorities consider it to be native (Praeger
1934; Scannell & Synnott 1972), others think it more probably an ancient introduction (Margetis &
David 1980; Ellis 1983).
All previous Welsh records of A. ampeloprasum refer to var. ampeloprasum. The habitat is
described as rocky sea coasts, but only on Flat Holm, where it was first recorded in 1837, has it
been confirmed recently (Ellis 1983).
The occurrence of this species near South Stack on Anglesey is therefore of some significance.
The exact date of its discovery is uncertain, but Mr D. B. Hornby (pers. comm. 1985) appears to
have been the first to notice it in the early 1970s. He recalls seeing about five or six plants in flower
then, which he took to be a kind of “‘wild onion’’. Records of the species first appeared in the Royal
Society for the Protection of Birds’ South Stack Warden’s Annual Report about 1976, but it was not
until 1983 that one of us (R.H.R.) became aware of the locality. This was visited in the autumn of
that year to confirm the identification and determine the size of the population, which was found to
be 43 flowering plants with a smaller number not flowering.
In August 1985, only 22 flowering plants could be seen, again with a smaller number not
flowering, but a number of broken stems suggested that some flower-heads had been collected.
Additionally a further small group of plants (four in all, but only two of them flowering) was found
about 50 m away from the original site.
The South Stack plants have bulbs about 2.5 to 3.5 cm across with several offsets inside the tunic.
Their flowering stems are about 120 cm tall, with broad, grey-green, linear leaves around 36 cm
long and 3 cm broad at the base. The globose umbels, about 5 to 6 cm across, have a large number
of densely-packed flowers, their corollas tinged with pale mauve, but no bulbils. The stamens are
only slightly exserted, and the antheriferous points of the filaments are almost as long as the
undivided part. It seems, therefore, that these plants, like those from all other Welsh localities, can
be referred to var. ampeloprasum.
Although not close to habitations, the plants have no appearance of being indigenous in this
locality. In the original site they grow within an area of a few square metres in a humus-filled
hollow at the edge of a field and close to its boundary wall, which provides the plants with some
degree of shelter from the wind. The habitat of the second group of plants is similar and the main
336 SHORT NOTES
associated species are indicative of uncultivated headlands and nutrient-rich soils — Achillea
millefolium, Anthriscus sylvestris, Arrhenatherum elatius, Cirsium arvense, Dactylis glomerata,
Festuca rubra, Galium aparine, Heracleum sphondylium, Holcus lanatus, Lathyrus pratensis,
Pteridium aquilinum, Rubus fruticosus agg., Rumex acetosa and Urtica dioica.
How this Allium came to South Stack is a matter of conjecture, for there is no obvious local
source of either seed or bulbs. It was once gathered from Bangor, Caernarvonshire, in 1876, but
has not been recorded there since and is now considered to have been an introduction (Ellis 1983).
The fact that Griffith (1895) does not mention this record in his Flora is perhaps significant.
Clearly the small population at the South Stack locality is a fragile one. The flower-heads are
often picked by local people and this can result in dislodgement of the loosely-rooted bulbs, and it
is hoped that anyone visiting this site will refrain from picking any of them.
Hopefully this population of A. ampeloprasum will continue to flourish, for its occurrence on
Anglesey, and in particular in the environs of South Stack, adds another interesting species to a
locality long noted for its rarities.
ACKNOWLEDGMENT
We wish to thank Dr J. S. Parker for his useful comments.
REFERENCES
CLAPHAM, A. R., TuTin, T. G. & WarBurRG, E. F. (1962). Flora of the British Isles, 2nd. ed. Cambridge.
CLAPHAM, A. R., Tutin, T. G. & Warsura, E. F. (1981). Excursion Flora of the British Isles, 3rd. ed.
Cambridge.
E.uis, R. G. (1983). Flowering plants of Wales. Cardiff.
GriFFiTH, J. E. (1895). Flora of Anglesey and Carnarvonshire. Bangor.
LousLey, J. E. (1971). The flora of the Isles of Scilly. Newton Abbot.
Marcetrs, L. J. & Davin, R. W. (1980). A review of the Cornish flora. Redruth.
PRAEGER, R. L. (1934). The botanist in Ireland. Dublin.
SCANNELL, M. J. P. & Synnott, D. M. (1972). Census catalogue of the flora of Ireland. Dublin.
STEARN, W. T. (1980). Allium, in TuTin, T. G. et al., eds. Flora Europaea, 5: 49-65. Cambridge.
R. H. RoBerts & P. DAy
51 Belmont Road, Bangor, Gwynedd, LL57 2HY
TYPIFICATION OF HIERACIUM AMPLEXICAULE L.
Hieracium amplexicaule was described by Linnaeus on page 803 of Species Plantarum in 1753. The
diagnosis Hieracium foliis amplexicaulibus cordatis subdentatis, pedunculis unifloris hirsutis caule
ramoso is taken from page 387 of the Hortus Cliffortianus (1738) with only the minor change of vix
dentatis to subdentatis. In the herbarium of Hortus Cliffortianus at the BM is a single sheet labelled
Hieracium folio caulina amplexico Triumfett. and in an unknown hand amplexicaule. This
specimen is in agreement with the diagnosis and I designate it as the lectotype of Hieracium
amplexicaule L. It belongs to the taxon described by Zahn (1921-1923) as H. amplexicaule subsp.
amplexicaule and by Pugsley (1948) as H. amplexicaule. The synonym Hieracium pyrenaicum
longifolium amplexicaule Tournef., Inst. 472, given by Linnaeus, is probably referable to the same
species. His B Hieracium pyrenaicum rotundifolium amplexicaule Tournef., Inst. 472, may well be
a different species, possibly H. speluncarum Arvet-Touvet. The specimen in the Linnaean
Herbarium (LINN), Savage Cat. 954.33, labelled amplexicaule in Linnaeus’ hand, belongs to the
same taxon as the lectotype in Hort. Cliff.
Hieracium amplexicaule is native on mountain rocks, mainly calcareous, from Spain and S.
France to Austria and Hungary. It is naturalized on old walls further north. [ts distribution as a
naturalized plant in the British Isles has been discussed by Bevan (1982, 1983).
SHORT NOTES = lll
REFERENCES
Bevan, J. (1982). Section Amplexicaulia Zahn. Hier. Group Notes, 2: 4-8.
Bevan, J. (1983). Section Amplexicaulia Zahn. Hier. Group Notes, 4: 3-4.
LInNAEus, C. (1738). Hortus Cliffortianus. Amstelaedami.
LINNAEuS, C. (1753). Species plantarum. Holmiae.
PucsLey, H. W. (1948). A prodromus of the British Hieracia. J. Linn. Soc. Lond. (Bot.), 54.
SAVAGE, S. (1945). A catalogue of the Linnaean Herbarium. London.
TourNeEForT, J. P. DE (1719). Institutiones rei herbariae, 3rd ed. Lugdunaeo.
ZAHN, K. H. (1921-1923). Compositae-Hieracium L. in ENGLER, H. G. A., ed. Das Pflanzenreich, 75 (IV,
280). Leipzig.
PDA SEEL
Botany School, Downing Street, Cambridge, CB2 3EA
PELORIC FLOWERS IN THE TRIBE ANTIRRHINEAE (SCROPHULARIACEAE)
Flowers of European species of the tribe Antirrhineae (including Anarrhinum Desf., Antirrhinum
L., Asarina Miller, Chaenorhinum (DC.) Reichenb., Cymbalaria Hill, Kickxia Dumort., Linaria
Miller and Misopates Rafin.) are normally highly zygomorphic (bilaterally symmetrical) but
occasional plants have a more or less actinomorphic (radially symmetrical) corolla. This
phenomenon has been given the name ‘peloria’, the subject of a dissertation published by Linnaeus
(1749), and was known to pre-Linnaean authors. Such plants were sometimes treated as belonging
to a different genus from those with zygomorphic flowers (Rafinesque-Schmaltz 1840), while other
authors regarded them as distinct species, varieties or insignificant forms. Most authors today give
them no formal taxonomic recognition but regard the plants as monstrosities or genetic mutants.
Peloric flowers occur in other Scrophulariaceae such as Digitalis L. (Arber 1932), and some
American species of the tribe Antirrhineae (particularly Lophospermum D. Don, Maurandya
Ortega and Rhodochiton Otto & A. Dietr.) have corollas that normally approach the
actinomorphic state.
Peloric flowers do not represent a single morphological state. In Linaria vulgaris Miller, variants
with no spur (‘‘anectaria’” of Linnaeus (1749)) occur in addition to those with five spurs
(‘“‘quinquenectaria” of Linnaeus (1749)). Indeed, in a single specimen of Linaria or Kickxia, it is
sometimes possible to find corollas with one, three, five or no spurs (cf. Chavannes 1833: plate 8).
Plants of Linaria purpurea (L.) Miller, both native in Sicily and naturalized in Britain, are
occasionally encountered with single inflorescences containing mostly normal zygomorphic flowers
but also scattered peloric flowers.
Considerable research on Antirrhinum majus L. (cf. Stubbe 1966) has resulted in one of the best
genetically documented species amongst flowering plants. The inheritance of peloric flowers in this
taxon is reasonably well understood, though other species of the tribe have scarcely been studied.
Development of peloric flowers was originally thought to be controlled by a single recessive gene
and it is interesting to note that Darwin (1868: 46) published a report of crossing true-breeding
peloric plants with normal Antirrhinum majus. Dominance of the zygomorphic form over the
peloric form, and the expected 3:1 ratio of zygomorphic to peloric plants in the second generation,
are apparent from the numbers of offspring recorded by Darwin, though this was years before the
classic genetic work of Mendel (1866) was widely known. It was later shown that linked multiple
alleles were involved in the production of peloric flowers, especially those of the Cycloidea (or
Radialis) series (Kuckuck 1936). It is not surprising that genes controlling zygomorphy of the
corolla should be linked, as even small differences in the structure of the Antirrhineae corolla leads
to reproductive isolation, owing to the behaviour of the normal pollinators (Mather 1947).
Through hybridization, progeny of other species of Antirrhinum have been obtained with peloric
flowers (Brieger 1935). The most familiar results of breeding plants with peloric flowers are the
““Penstemon-flowered”’ garden snapdragons.
A simple genetic basis for peloric flowers does not account for situations where only a few
flowers in an inflorescence are peloric. It is possible that the development of the corolla could be
influenced by external factors such as insect predation or viral infection. Viral infection has been
338 SHORT NOTES
attributed as the basis of plants known as Cymbalaria toutonii A. Chev. (cf. McClintock 1982),
generally considered to be referable to C. muralis P. Gaertner, B. Meyer & Scherb. though with
very different foliage and a smaller corolla than the normal species. It is also possible that genes
controlling the zygomorphic development of the flower might be labile, with random mutation
accounting for inflorescences with varying symmetry of the corollas. Perhaps the best known labile
genes in Antirrhinum majus belong to the Pallida series and control the flower colour. Back
mutation results in a corolla with ‘broken’ coloration which varies from one flower to another and
some strains of garden snapdragon have this patterning. There is a labile allele in the Cycloidea
series which results in the peloric corolla reverting to the normal zygomorphic form.
Much is still unknown about peloric flowers in the Antirrhineae, but with such a long history of
morphological observation and the detailed genetic investigation of Antirrhinum majus, species of
the tribe would provide ideal experimental subjects for investigation into the development and
control of zygomorphy in flowering plants.
REFERENCES
ArBER, A. (1932). Studies in flower structure. I. On a peloria of Digitalis purpurea, L. Ann. Bot., 46: 929-939.
BrIEGER, F. G. (1935). The inheritance of self-sterility and the peloric flower shape in Antirrhinum. Genetica,
17: 385-408. s
CHAVANNES, E. L. (1833). Monographie des Antirrhinées, plate 8. Paris & Lausanne.
Darwin, C. (1868). The variation of animals and plants under domestication, 2: 46-47. London.
Kuckuck, H. (1936). Uber vier neue Serien multipler Allele bei Antirrhinum majus. Z. indukt. Abstamm.-u.
VererbLehre, 71: 429-440.
LINNAEUS, C. (1749). Dissertatio botanica de Peloria. Amoenitates academicae, 1: 280-297. Holmiae.
MatTuer, K. (1947). Species crosses in Antirrhinum. I. Genetic isolation of the species majus, glutinosum and
orontium. Heredity, 1: 175-186.
McCuintock, D. (1982). The story of Cymbalaria toutonii A. Chev. Watsonia, 14: 182-183.
MENDEL, G. (1866). Versuche tiber Pflanzenhybriden. Verh. naturf. Ver. Briinn, 4: 3—44.
RAFINESQUE-SCHMALTZ, C. S. (1840). Autikon botanikon, p. 20. Philadelphia.
StuBBE, H. (1966). Genetik und Zytologie von Antirrhinum L. sect. Antirrhinum, p. 421. Jena.
D. A. SUTTON
British Museum (Natural History), Cromwell Road, London, SW7 5BD
Watsonia, 16, 339-352 (1987) 339
Book Reviews
Studies in plant demography. A Festschrift for John L. Harper. Edited by J. White. Pp. xxxiii+393.
Academic Press, London. 1985. Price £44.00 (hardback; ISBN 0~-12-746630-4); £21.00
(paperback; ISBN 0—12-746631-2).
This volume celebrates the 60th birthday of one of the most influential plant ecologists of the last
30 years. Professor John Harper has long been at the forefront of the integration of plants into the
science of population biology, previously the domain of the zoologist. He and his numerous
students and co-workers have investigated births, deaths, numbers, survivorship and growth
patterns in plant species, in order to provide quantitative assessments of evolution through natural
selection. The material in this book thus falls rather outside the systematics fields that concern
most B.S.B.I. members, but provides a valuable introduction to, and summary of, an important
area of botany. Harper’s philosophy has been strongly influenced by Charles Darwin, and here we
are certainly on common ground. This Festschrift includes contributions on life histories,
reproductive biology, effects of neighbours, plant-animal interactions, weed ecology, and aspects
of agriculture and conservation. Weeds, which Harper himself once described, in characteristically
impish manner, as “‘slightly improper material for biological studies”, are a major theme, not least
with reference to the need to study invasions of alien weeds — very “‘improper”’ material indeed,
but meat and drink (of one sort or another) to many of us! There are items of interest for all here,
from Eupatorium to emus, and there is also a useful bibliography of Harper’s publications.
The contributors emphasize the use of quantitative observations, experiments and analyses in
plant ecology, which Harper has done much to transform into an exact science. Nevertheless, they
and we should not forget that the naturalists’ ‘anecdotal’ (to use an unhappily somewhat
derogatory term of the Harperian school) field observations can be the basis for hypothesis,
discussion and answers to the biologist’s questions. The critical mind is ultimately the best analyst,
as Harper clearly demonstrates; may he long continue to stimulate us in his retirement.
J. R. AKEROYD
The Oxford dictionary of natural history. Edited by M. Allaby. Pp. 688. Oxford University Press,
Oxford. 1985. Price £20.00 (ISBN 0-19-217720-6).
This Dictionary was written by a team of 25 specialists in different aspects of natural history and
contains over 12,000 entries. There are no illustrations. Unlike the Penguin Dictionary of British
Natural History, one of the few other specialist dictionaries of the subject, this book attempts a
worldwide coverage. The Preface explains the criteria which were applied in allocating space to the
various aspects of the subject. On the whole the editor has achieved a very reasonable balance.
However, botanists may detect a distinct zoological bias in entries such as ‘“‘nomen conservandum”’
or “nomen nudum’’, where the /nternational Code of Zoological Nomenclature is referred to
without mention of its botanical counterpart. A further entry describes the ICZN but does not
mention that there are other codes. This omission reflects the fact that whilst the Dictionary is
excellent in its detail it is lacking in careful definitions of the generalities. This is unfortunate,
because often it is broad concepts that exclude the amateur from the domain of the professional
and preserve the barriers this Dictionary hopes to remove. There are no entries for
‘nomenclature’, ‘‘systematics’’, ‘“code’’, or ‘‘rank’’, and the single line entry on “‘taxonomy’”’ is not
cross-referenced to the entries which describe methods of classification. There are also inaccuracies
resulting from omissions in the definitions of some fundamental terms. The fact that taxa above the
species level have types is not apparent from the entry for ‘“‘type specimen” and neither is it clear
that ‘‘phenetic classification”? can use data other than physical similarities or that a ““synonym”’ is
not simply a different name for a species or variety.
340 BOOK REVIEWS
A large proportion of the Dictionary comprises entries relating to plant and animal taxa of
various ranks, particularly families. The selection of taxa included is well balanced but could
have been improved botanically by including the alternatives to such conserved family names as
Compositae, Palmae or Gramineae especially since Aster, Areca, and Poa all have entries.
Cross-referencing between the taxonomic entries is generally good but there are exceptions. The
entry for ““Palmae” could usefully have referred to that on “‘rattan” which itself does not mention
palms but provides only a list of cross-referenced genera. The entries for two of these,
Plectocomia and Plectocomiopsis, are missing whilst others (e.g. Bejaudia, Calospatha) provide
more detail for these monotypic genera than is provided for some families.
Despite these criticisms the Dictionary will be useful as a source of information for those
interested in ecology, conservation and in the biology of tropical organisms in particular. In hard
back form it does not seem likely to match the volume of sales achieved by cheaper, soft back
specialist dictionaries on other subjects.
S. BLACKMORE
No ordinary gardener, Thomas Knowlton, 1691-1781. B. Henrey, edited by A. O. Chater. Pp.
324, with 32 black & white illustrations. British Museum (Nat. Hist.), London. 1986. Price
£17.50 (ISBN 0-565-00976-1).
A scholarly work, this is more than a well researched biography. Much valuable authoritative
information is given concerning the gardener’s employers, correspondents and other associates.
The introduction is a masterly and highly readable summary of Knowlton’s career and various
activities. Part 1 deals with the period 1691-1726, during which Thomas Knowlton’s most
important position was as head gardener to James Sherard at the time when the famous garden at
Eltham, Kent was being established. Part 2 covers the rest of Knowlton’s life, during which he
was head gardener at the Londesborough estate in Yorkshire. It consists mainly of transcriptions
from Knowlton’s letters to eminent figures of the day, notably Hans Sloane and the botanists
Samuel Brewer and Dr Richard Richardson, together with detailed commentaries on the topics
mentioned. The abstracts selected not only illustrate Knowlton’s wide interests, but also
demonstrate how 18th century conditions affected his life.
Records for some 50 species of British native plants are included in the text; ten are in
abstracts from the diary Knowlton kept during a visit to Guernsey in 1726 to study the Guernsey
Lily, and the rest in letters written at Londesborough. Most of the latter records are for
Yorkshire, but some are for Hertfordshire and Guernsey. Some records point to a very different
past. Observations concerning the occurrence of Stratiotes aloides add to our knowledge of the
distribution at that time of this now very rare S.E. Yorks. species. It occurred for instance in
plenty between Hull and Beverley, in dykes on both sides of the road. Primula farinosa was
found by the R. Wharfe at Otley, much further down the valley than it is now known. There are
several records for Fountains Abbey, including Cardamine impatiens, Paris quadrifolia, Primula
farinosa and Cephalanthera damasonium. The record for the last species: ‘‘Hellesborine flore albo
ger wild att fountain abby” is puzzling as the locality is so far north of its otherwise recorded
distribution.
Knowlton’s use of pre-Linnaean nomenclature and his difficult handwriting presented a
challenge to the authoress, as did his unconventional spelling. One error arose from the latter.
Marsh Gentian was described as growing “plentifully in our Sawampis comons at Home &
Everingham neare me in yorkeshie . . .”. Home is Holme (upon Spalding Moor) with adjacent
wet heathland and not Londesborough, Knowlton’s home village on the chalk wolds, as
suggested by Blanche Henrey.
Knowlton built up a living collection of uncommon wild plants, and there was clearly much
exchanging of plants and seed of both native and imported species amongst gardeners and
landowners. He had experience in growing exotic species at Eltham, and his skill in cultivating
such species became widely recognized. He wrote a treatise on the cultivation of coffee trees
which is reproduced. After a stove was erected at Londesborough he became increasingly
interested in growing such plants and was notably successful in growing pineapples.
BOOK REVIEWS 341
There are appendices giving information concerning Knowlton as tree planter and garden
planner, bibliophile, subscriber to publications and as a man of property. There is also an abstract
from his will and a classified list of plants mentioned in the book. I find the separate plant index of
particular value. There are numerous references to source material in foot notes, but I would like
to have seen a full bibliography.
The book is a good quality production. It is a fascinating social and historical document which,
whilst of the strongest appeal to the garden historian, will I believe be enjoyed by a very wide
readership.
It is sad that Blanche Henrey did not live to see the book published; the reader is indebted to A.
O. Chater for ably carrying out the exacting task of preparing the final typescript.
F. E. CRACKLES
Rubi Westfalici. Die Brombeeren Westfalens und des Raumes Osnabriick (Rubus L., subgenus
Rubus). H. E. Weber. Pp. 452, with 67 black & white photographs and 123 distribution maps.
Westfalisches Museum fiir Naturkunde, Minster. 1985. Price DM 68.00 (ISBN 3—924590-07-9).
This is another major work by the leading European batologist of our day. It is carefully printed on
firm glossy paper and strongly bound with a hard cover. There is a coloured illustration of ripe
blackberries as large as life on the outside of the cover to sweeten the appetite. Westphalia is a
province of western Germany bordering Holland and therefore near enough to the British Isles for
a comparison of its flora with ours to be interesting. Indeed, of the 123 species of Rubus recorded
for Westphalia one third are also found in the British Isles.
The book consists of a full explanatory and historical introduction, two keys to help in the
identification of the species, detailed descriptions, maps and photographic illustrations. The
distribution maps are excellent. There is one for every species; they are all the same size, large and
very clear; black dots stand out sharply from a background which indicates the contours of the
land. It is instructive to compare the different distributions of species which occur in both our
countries. For example, Rubus dasyphyllus is limited to one comparatively small area in the north
of Westphalia, though evidently common there, and R. nemoralis, though scattered throughout
Westphalia, is rare. There are only twelve recent records. On the other hand R. sprengelii is
abundant throughout the province, especially in the hillier areas.
Most of the illustrations are photographs of herbarium specimens and as good as such photos
probably can be. They show the shape, dentition and disposition of the leaflets, but the stem
clothing and armature and the floral characters are usually lost. The leaves of R. mucronulatus and
R. adspersus are well illustrated. Sometimes close-up photographs of parts of the living plant are to
be preferred. For example, the stem of R. radula, the rachis of R. infestus and the panicle of R.
sprengelii are well illustrated in this way. It is always a big advantage to know the plant before you
come to the photograph! Then the finer points of distinction become visible. Compare, for
example, the illustrations of R. nessensis and R. scissus.
E. S. EDEES
Unlucky plants. Compiled by R. Vickery. Pp. 92. The Folklore Society, London. 1985. Price £4.95
(ISBN 0-903515-10-S).
This short book is concerned with plants which are believed to bring misfortune if picked or taken
indoors. Roy Vickery has compiled this collection from information supplied in a survey carried
out by the Folklore Society. Their survey elicited 524 items, which related to 90 species of plant.
The result takes you right back to your childhood, triggering recollection of submerged beliefs —
though in many cases they are not far below the surface. I know many people who today will not
have lilac or may blossom in the house. There are several instances of transferred epithet, as for
instance is the case with lilies and snowdrops and white flowers in general which, associated with
funerals, have become the bringers of death. Sometimes it is the dangerous character of a place
342 BOOK REVIEWS
itself that seems to have attached itself to the reputation of the plants which grow there. Bog
asphodel, called by Linnaeus ossifragum (bone-breaker) and dreaded by fell farmers because it
‘softened sheep’s bones’, grows in acidic bogs, perilous for sheep. Similarly, it is possible that
children are told that the seductive blooms of the yellow water lily bring ill luck, to protect them
from the dangers of drowning in the waters where it grows. Mistletoe is not one of the plants under
consideration, but I should like to know whether the reluctance to take mistletoe inside a church is
anywhere explicit. For the most part, Mr Vickery steers clear of interpretation; but where he does
make forays, they are always enlightening: the relationship between primroses and poultry-raising
for instance. This interesting collection of folk belief seems like the tip of the iceberg. I hope it will
provoke further study in this area, eliciting the why as well as the what.
F. GREENOAK
Flora of the Isle of Man. D. E. Allen with the assistance of L. S. Garrad and M. Devereau. Pp.
xi+250, with topographical map and six figures. The Manx Museum and National Trust, Douglas,
Isle of Man. 1984. Price £13.75 (ISBN 0—901106—23-2).
This Flora is a traditional local Flora with no dot maps, and by current standards it is a modest
volume. However, it is a book full of scholarship and reflects the author’s study of the Isle of Man
flora over 35 years.
The book is divided into six main chapters that consider the physical setting (nine pages), history
and composition of the flora (eight pages), habitats (ten pages), the districts (two pages),
discoverers of the flora (14 pages) and systematic list (178 pages). In addition, there are appendices
on species and subspecies which might occur in Man and references to work on the lower plants,
together with an extensive bibliography, acknowledgments and an index to plant names only.
In considering the introductory chapters the author gives competent and adequate accounts for
the physical setting, and the history and composition of the flora. The chapter on the habitats of
the Island gives the reader a good overall impression of what to expect on a visit to the Island, but I
think more descriptive detail with lists of significant species for selected sites would have improved
this section. The chapter on the discoverers of the flora is especially good. It takes the reader from
the earliest recorders of the flora in the 16th Century through the problems and triumphs of the
19th and early 20th Centuries, to the present day in a most readable and informative manner.
The systematic list forms the main part of the Flora and avoids what, to the reviewer, is the
annoying habit of only including recent records. This is a compilation of all records, and a
distinction is made between pre- and post-1950 records; but it is not clear how thoroughly the
Island’s flora has been surveyed in recent years or to what extent the accounts are based on casual
as opposed to systematic and quantifiable recording techniques.
Of interest to the reviewer is the citation of herbarium specimens. This is of immense value to
future students and allows the present author’s work, particularly in respect of critical groups, to be
re-evaluated at some future date.
The records are organized according to six districts based on natural eco-geographical
subdivisions, a scheme which in the context of the Isle of Man makes sense. Nevertheless, I would
have liked to see a few selected dot maps which could have been used to advantage to show the
distribution of certain species or change in status of others.
The author stresses the importance of assigning each species a status, and native, colonist,
denizen or hortal categories are used. This is an area of considerable difficulty for botanists, but it
is important to record change, to point out what is an introduction and to show how it might have
arrived. It is the story behind the record that is of value, and in this David Allen has succeeded
admirably.
The sequence and scientific names are those of Dandy’s List of British vascular plants (1958) but
recent changes have been incorporated. A feature of the Flora is the infraspecific detail that is
provided, but unfortunately many of the names are not in Dandy nor in many of the more recently
published standard Floras. Of particular note is the section on Rubus, which formed a special
study.
Common names are included and usually follow Pocket guide to wild flowers (McClintock &
BOOK REVIEWS 343
Fitter 1956). In addition, Manx names are given where these have been reliably identified with
particular species.
There are six figures and a topographical map, which are adequate except for Figure 3, where
the detail is lost. There are no reproductions of photographs and paintings except for the striking
colour photograph of henbane on the dust jacket. |
This is a Flora of succinct detail. It gives the results of considerable historical research into
both people and their study of the flora, into the origins and uses of Isle of Man plants, and it
gives much detail on the exact identity of the plants found in the Island. It is a worthy companion
to other local Floras and indispensable for anyone interested in the Isle of Man flora.
E. F. GREENWOOD
A guide to the vegetation of Britain and Europe. O. Polunin & M. Walters. Pp. 238, with 60
plates of line drawings, 110 colour plates and 53 text figures. Oxford University Press, Oxford.
1985. Price £17.50 (ISBN 0—19-217713-3).
This is primarily a descriptive account aimed at the ‘intelligent layman’ of the vegetation of
Europe (excluding Spitsbergen). It is a task for which the much-travelled authors, especially the
late-lamented Oleg Polunin, were well-equipped. The main section of the book, Part II, which
follows brief accounts of ecological concepts, soils, climates and vegetational history, deals with
the subject mainly on a regional and climatic basis. Chapters 6-11, therefore, cover Arctic,
Boreal, Atlantic, Central European, Mediterranean and Pannonic vegetation. However, these
are followed by three dealing with Alpine, Fresh-water and Coastal plant communities. This
approach, a mix of climatic and vegetational types, is the authors’ compromise solution to the
difficult problem of dealing with such a varied subject. In their view the vegetation of the last
three is essentially azonal, occurring in more than one of the preceding climatic zones. The six
preceding chapters cannot, therefore, be considered complete in themselves.
The problems of a regional approach are well illustrated by the treatment of mires. These
receive their main discussion in the Boreal chapter, although arctic mires have already been
mentioned, blanket bogs appear in the Atlantic one and raised bogs and fens in both this and the
Central European and Alpine chapters.
The important maps in the Introduction are regrettably neither numbered nor provided with
clear titles. The title is either hidden in the map with the key or, in one case, missing altogether.
At the beginning of each of the first six chapters in Part II is a map showing the particular region
in question, and these maps are based on the climatic zones shown in the map on p. 9. However,
the picture is confused by the map of “potential vegetation’ on p. 20, which is said on the
following page to show the main regions. This shows the distribution of “poorly drained wet-
lands” (are these the fresh-water wetlands of chapter 13?), and a boreal area which, contrary to
the climatic map and that in the Boreal chapter, includes, very sensibly, north-eastern Scotland
and southern Sweden. There are further discrepancies in relation to the Mediterranean region,
especially in the Balkans, and the Pannonic region isn’t shown at all. These mistakes may be
relatively minor but they are irritating and they do create a general impression of careless
editing.
The longest chapter is, not surprisingly in view of the senior author’s interests, that on
Mediterranean vegetation. After a general introduction to the region and its climate, there are
sections on evergreen oak woods, deciduous and semi-evergreen oakwoods, laurel woods, pine
woods, montane coniferous woods, sweet chestnut woods, horse chestnut and walnut woods, fir
woods, cypress and juniper woods, wet woods, bush communities and grasslands. Within each
section there are maps of the dominant trees and many short lists of species to illustrate regional
variants. For example, under evergreen oaks there are lists of Quercus ilex associates from
southern Spain, Italy, the Adriatic coast of Yugoslavia and Greece, the briefer accounts of Q.
suber, Q. coccifera and Q. macrolepis woods.
The communities are given English names which are not intended to conform to any particular
classification, and the emphasis in the text is on English names, even for non-British species, such
as the Bug (not Bog) Orchid, Star-fruited Fenugreek and Small-headed Blue-eryngo. The many
344 BOOK REVIEWS
lists inevitably result in a certain amount of repetition and their number does lead to a considerable
loss in readability, with the result that this is inevitably more of a reference book.
As already mentioned the authors’ aims are descriptive, but here and there are more informative
sections dealing with the relationship between vegetational types, successional changes and the
effects of man. This is especially true of the Mediterranean chapter. By contrast, the role of man
and his animals in modifying montane vegetation and the tree-line is dismissed in a few lines. It is
curious that the Alpine chapter makes no reference to altitudinal zonation within the alpine zone.
There is confusion in the authors’ use of the subalpine zone, which, in the glossary is said to be
“usually above the tree-line”’ and to consist of “‘dwarf shrubs and stunted trees’’; this is the lower
alpine zone. The Arctic chapter is very unsatisfactory in that the map shows it as including the
Scandinavian mountain chain and half of Iceland, but there is very little in it relating specifically to
these areas and the only reference to Iceland in this Arctic chapter is to birchwoods! As a guide to
Scandinavian or Icelandic montane vegetation this chapter is virtually useless. Why, incidentally,
do the authors use the term ‘arctic-alpine tundra’ in preference to the more familiar ‘fell-field’?
Another irritating feature is the authors’ failure to decide whether the Alpine chapter is Alpine or
Arctic. Having included the Scandinavian mountains in the Arctic, we find the occasional
reference in the Alpine chapter to northern examples such as Scottish tall herb communities and
Scandinavian snow-patch communities “in the Boreal region’’; it is all very confusing.
The decision to adopt a mainly regional approach means that the pinewoods of north-east
Scotland are treated as Atlantic rather than boreal, and the chalk downlands of south-eastern
England are similarly regarded as Atlantic, whereas their flora has a not unimportant central
European and continental element. The authors’ examples could sometimes be improved upon.
There is no mention under Atlantic heaths of the very characteristic grass Agrostis curtisii, nor is
there special mention in this chapter of Digitalis purpurea. Montane calcareous flushes in Scotland
are said to be characterized by Cratoneuron commutatum, Saxifraga aizoides, Festuca rubra and
Cardamine pratensis. This is true, but more widely distributed, much more interesting but
unmentioned are the open flushes with many arctic-montane rarities, such as Juncus alpinus,
Kobresia simpliciuscula and Carex atrofusca, in addition to Juncus triglumis and Thalictrum
alpinum.
There is a third part to the book, a list of national parks and nature reserves arranged
alphabetically by country. It is, in fact, an appendix, a useful one, but not an integral part of the
text. The maps of the individual countries are poor, so reduced as to be well nigh illegible.
This is followed by a selected bibliography, a glossary and indices of Latin and English names of
illustrated species and a final one to the plant communities, which is fine if you are attuned to the
authors’ terminology and are aware of dwarf fan-palm garigue and hedgehog heath alpine
communities.
The book concludes with a fine set of colour plates which are both attractive and informative. I
particularly like those of a palsa mire, of Atlantic oceanic heath (showing Agrostis curtisii, not
Deschampsia setacea) and the Mediterranean woodlands. That of chalk grassland on the Sussex
Downs is almost unrecognizable, and the penultimate plate is of Crithmum maritimum, not
Crambe maritima.
Throughout the chapters are 57 pages of line drawings by Rosemary Wise. These are an
excellent feature, clear and life-like, although generally rather faint. The diagrams and block
diagrams of vegetation types are mostly unsatisfactory, being fuzzy and too stylized. The black &
white plates are similarly rather poor, lacking in contrast and adding little to the text, for example
those of Ulex gallii (Fig. 16), maquis (Fig. 35) and Paliurus (Fig. 36); the goats in Fig. 37 could be
crossing a snow-patch rather than suffering the blistering heat of the Mediterranean hills.
Mis-spellings are very few; minor errors in the text include sub-tropical (p. 20) for sub-
Mediterranean, “the grass Kobresia myosuroides’’, (p. 162), the occurrence of Galium pumilum in
Iceland (p. 27) and the inadvertent inclusion (p. 69) of Atlantic acid grasslands within the
calcareous ones. Epipactis atrorubens is not a characteristic species of machair (p. 190).
The main problem of this book is the sheer scale of the subject and the difficulty of organizing
and condensing the material into a readable, comprehensible whole. Inevitably many communities
get only the most cursory treatment — nine lines for sea-cliffs, for example. The task is virtually
impossible in a book of this size. Parts I and II might occupy 191 pages but there are only 90 of
actual text and half of these are taken up with lists.
BOOK REVIEWS 345
Nevertheless this is a useful and informative book. The authors must be applauded for their
willingness to undertake such a monumental task and for the impressive scale of their compilation,
and the publishers for the attractiveness of the format and the reasonable price.
G. HALLIDAY
Mountain flora of Greece, Volume 1. Edited by A. Strid. Pp. xxx+822, with 50 text figures.
Cambridge University Press, Cambridge. 1986. Price £65.00 (ISBN 0-521—25737-9).
Some years ago my wife and I climbed to the Hellenic Alpine Club’s Gamila hut in the northern
Pindus. We were the first visitors of the year; the bedding was damp, the chimney blocked, so that we
ended the night on the floor under a pall of smoke. But that is another story. . . The flowers around
were plentiful, and I scrambled up the steep wall of Mount Timfi, near the hut, to its near-flat plateau
with long views to Gamila, Smolikas, and the great gash of the Vikos gorge. Among the many good
plants here was Saxifraga oppositifolia.
This seemed to me rather easterly of its Alpine range, so on return home I checked it in Flora
Europaea. It was not recorded for Greece, so I felt I had a small plus. The first thing I did on opening
Mountain flora of Greece was to look up Saxifraga oppositifolia. And there it was, a 1954 record of
the late Dr Goulimis, that indefatigable Greek amateur botanist, confirmed by Professor Strid, from
Timfi alone.
In fact, of some 926 taxa in the first volume of this notable work, about a third were either not
recorded for Greece in Flora Europaea, or were there under different names or with different rank.
30 further taxa in Flora Europaea are omitted as erroneous or synonymous. This seems an
astonishing proportion for a country relatively well explored by such great botanists as Heldreich,
Grisebach, Orphanides and Rechinger among many others, with such as Dr Goulimis, Werner
Greuter and Elli Stamatiadou in recent years.
The activities and publications of these botanists are cited and described in an Introduction which
elaborates also the work of modern Swedish botanists and finally those of Arne Strid (Professor of
Botany at the University of Copenhagen), Per Hartvig and their co-workers, who collected nearly
20,000 specimens which have been the main source for the descriptions in this volume. The criterion
for inclusion, incidentally, is that of plants growing more or less exclusively above 1800 m.
The introduction ends with a brief but excellent outline of geography, topography and geology of
the region, and a check-list of the mountain peaks according to district. These mountains include
those of Crete and the highest Ionian and Aegean islands, even though some of these do not rise to
1800 m. In this section there is an account of the phytogeographical elements making up the Greek
mountain flora, with their significant connections with the Alps and Carpathians and the mountain
floras of Italy and Anatolia.
Fifty specially prepared line illustrations complete the volume, many of these showing several
species or their diagnostic features, like the siliculae of Alyssum species, or the leaves and flower
profiles of violas. There are nine for Umbelliferae!
Needless to say the keys, descriptions and locality citations are exemplary in this first volume,
which encompasses vascular cryptogams, conifers, and angiosperms from Fagaceae to Plumbagi-
naceae, in its 800—-plus pages. It is the first serious critical Greek flora since Hayek’s Prodromus
Florae Peninsulae Balcanicae, completed in 1933.
A. HUXLEY
Index of collectors in the Welsh National Herbarium. S. G. Harrison. Pp. 140. Amgueddfa
Genedlaethol Cymru/National Museum of Wales, Cardiff. 1986. Price £12, or £13.10 by post from
the National Museum of Wales, Cathays Park, Cardiff CF1 3NP. (ISBN 0-7200-0295-8).
The National Museum of Wales was founded nearly three-quarters of a century ago, and the
formation of the Department of Botany herbarium began with the transfer in 1912 of 3500+
mounted plants from the Welsh Museum of Natural History, Arts and Antiquities (originally Cardiff
346 BOOK REVIEWS
Museum and Art Gallery). Since those early days this embryonic collection has now increased to
nearly 265,000 specimens gathered in many parts of the world by hundreds of collectors, although
the more important herbaria are British with a strong bias to the Principality. The more valuable
Welsh herbaria include those of the Rev. Hugh Davies (1739-1821), author of Welsh Botanology
(1813), J. E. Griffith (1843-1933), compiler of The flora of Anglesey & Carnarvon (1895) and C.
T. Vachell (1848-1914) & E. Vachell (1879-1948).
The Index of Collectors in the Welsh National Herbarium covers not only Spermatophyta and
Pteridophyta, but also Bryophyta, Fungi, Lichens, Algae, Myxomycetes and Plant Galls. It lists
alphabetically by surname the collectors whose dried plants are preserved at Cardiff, with details of
forenames, years of birth (and where applicable, of death), in many cases data on the regions or
areas whence the gatherings originated, and the years in which they were made. A selected
bibliography is given, and appendices provide a selection of herbarium abbreviations, a numbered
list of the British and Irish vice-counties, and some examples of collectors’ handwriting. This is an
excellent addition to the rapidly growing series of books cataloguing the plant collections at
University and Museum Botany departments, which began in 1897 with the Druce & Vines
account of the University of Oxford herbaria.
D. H. KENT
Habitat handbook no. 1. Hampstead Heath flora. J. Bellamy et al. Pp. 14+35 pp. of distribution
maps, a list of extant and extinct species and two appendices, with 15 coloured photographs and 31
black & white figures. Department of Recreation and the Arts, Greater London Council. 1986.
Price not stated (ISBN 7168—1653-9).
In this work the flowering plants are recorded on 100-metre squares, 25 of which form larger 500-
metre squares, identifiable by means of a system of letters and numbers, and based on the National
Grid. There is a general introduction, and two appendices contain lists of species in various
habitats. There is evidence that the compilers consulted London’s Natural History (Fitter) but
none that they consulted recent Floras of Middlesex (Kent) or London (Burton). Nevertheless it is
a painstaking piece of work most attractively presented, and it is to be hoped that it will be
followed by subsequent editions in which not only will many additions to the existing maps be
made, but the distribution of trees, shrubs, ferns, rushes, sedges and grasses be added.
Nomenclature appears to be largely accurate though at least two errors have been noted.
K. G. MESSENGER
Flowers and ferns around Huddersfield. M. J. Lucas & J. Middleton. Pp. viiit 115, with 23 pages of
line drawings and two maps. Kirklees Leisure Services, Huddersfield. 1985. Price £2.95 (ISBN 0-
9502568-8-9).
Since no previous attempt has been made to produce a complete list of flowering plants and ferns
of the Huddersfield district, the authors of the present book are to be congratulated on their efforts
to remedy this omission. It is somewhat surprising that in such a well-worked area such a list has
not been produced before.
The authors acknowledge the help of past and present botanists in the use of their records, but
one could wish that additional names had been mentioned, particularly those of the late Dr T. W.
Woodhead of Tolson Memorial Museum, Huddersfield, the Rev. T. A. Jefferies and Dr J.
Grainger, all of whom carried out work in the district, though mainly of an ecological nature.
The map of the area surveyed would perhaps have been of greater value had use been made of
the tetrads, which could have been done without endangering species. The map of the river basins
is of geographical interest.
Certain plant species not likely to be indigenous are said to be ‘‘native’’; these, especially trees,
include Pinus sylvestris, Taxus baccata, Carpinus betulus, Sorbus aria agg., Clematis vitalba and
Swida sanguinea. Species such as Medicago falcata, Foeniculum vulgare and Echium vulgare can
BOOK REVIEWS 347
hardly be more than casuals in the area. Critical groups are generally dealt with in the widest sense,
the brambles for example being all under Rubus fruticosus agg.
The Huddersfield canals have been intensively studied and are probably some of the best known
botanically in the country. Along with many native species grow several of what the authors call
“goldfish aliens’, introduced aquatic species now well established, including Egeria densa,
Lagarosiphon major and Elodea nuttallii. Alisma lanceolatum is stated “not to be recorded again as
far as we know, till Angus/Kincardineshire’’. It does, however occur in both canal and river Calder
not very far away. Is Eleocharis acicularis a rare plant?; it also grows in some quantity in the
Calder.
The 23 pages of drawings by Derek Toms, though pleasing in themselves, seem to answer no
useful purpose. The book is not intended to be a guide to identification, and in any case only a brief
selection of species is illustrated. Possibly a few more detailed drawings of critical species would
have been of more use; those of Egeria, Elodea and Lagarosiphon show some useful distinguishing
features.
A few species present in the area are not included in the list. Perhaps at some later date a more
complete and enlarged edition will be forthcoming.
F. MURGATROYD
Concise dictionary of biology. Edited by E. Martin. Pp. 256, with numerous line drawings. Oxford
University Press, Oxford. 1985. Price £7.95 (ISBN 0-19-866144-4).
This dictionary is derived from the Concise science dictionary published by O.U.P. in 1984. It
includes all the entries from that work relating to biology and biochemistry, together with those
entries from geology necessary for an understanding of palaeontology, and a few entries relating to
physics and chemistry which are required for an understanding of those aspects of biology. In
addition there is a small selection of words used in medicine and palaeoanthropology.
The dictionary includes an extensive system of cross references, which means that each entry can
be placed within a broader context: an initial enquiry may then lead to constructive browsing within
its c. 3000 definitions.
In a field in which there are already many competitors this hardback may seem an attractive
bargain. My only reservation is whether the blotting-paper on which it appears to have been
printed will stand up to the frequent use for which worthwhile dictionaries must surely be
prepared.
F. H. PERRING
Guide to mountain and moorland. B. Brookes. Pp. 128, with numerous colour and monochrome
photographs and line drawings. The British Naturalists’ Association Guides. The Crowood Press,
Marlborough, Wiltshire. 1985. Price £7.95 (ISBN 0-946284-76-8).
This little book, like its fellows in the series of guides to British habitats by the B.N.A., looks
attractive, handles nicely, and is reasonably priced. Like the rest, the present volume seeks to
attract by an abundance of illustrations — they occupy exactly half the book. Brian Brookes
combines expertise in his subject with a clear and lively style. Six chapters cover Mountains,
Moorlands, Moorland Wildlife, Mires, Forests, and Conservation. (Why was so much space
devoted to ‘Forests’? The accounts of the development of moor from forest and the re-
instatement of forest on moor are interesting, but surely description of the wildlife of those forests
is beyond the scope of this book, especially when a companion volume covers ‘“‘Woodlands’’.)
Predictably, in only sixty pages of script, the author has had to cover some topics in a somewhat
superficial fashion, and the strength of the book lies in the success with which he has compressed so
much into a small compass. In fact, for my money I would have preferred more Brookes and less
pictures. Whilst some plates are excellent and evocative, others are too small, too ‘muddy’, or
nothing to do with the subject. (A monochrome otter has a rather lame caption “. . . which
348 BOOK REVIEWS
ee |
sometimes inhabits upland sites...”!). Featured flowering plants include many common
species, but botanists will be attracted by a tiny flowering Tufted Saxifrage (Saxifraga cespitosa)
and frustrated by a single straggling stem, with bright pink flowers, of the Blue Sowthistle
(Cicerbita alpina). Colour-balance is variable. Corrie Kander appears in perfect muted grey and
brown; Racomitrium lanuginosum comes out a lurid green. In a book so dominated by
illustrations, their lack of consistency is a major handicap.
The ‘interested layman’, and his (older) children, will, I think, enjoy the book, and the
author’s warnings on the fragility of upland ecosystems are both timely and appropriate:
conservation will only be achieved when those who use the uplands have some empathy with the
natural processes which create and sustain them.
F. J. ROBERTS
Infraspecific classification of wild and cultivated plants. Edited by B. T. Styles. Pp. xiv+435.
Clarendon Press, Oxford. 1986. Price £47.50 (ISBN 0—19-857701-X).
The conference of which these are the published proceedings was held in September 1984 in
Oxford by the Systematics Association in association with four organizations concerned with
cultivated plants: the Royal Horticultural Society (RHS), the International Union of Forest
Research Organizations (IUFRO), the International Board for Plant Genetic Resources
(IBPGR) and the International Union for the Protection of New Varieties of Plants (UPON). It
provided a forum for the exchange of information and viewpoints on plant variation at the
infraspecific level.
Such a bridge-building exercise is greatly to be welcomed because, as Professor Hawkes and
Professor Heywood agreed (in their introduction and summary, respectively), polarization of
views between botanists and cultivators must be avoided. A considerable divergence of views is
revealed as to how much the classifications used by the two groups of workers can or should
connect, ranging from those of Lewis, who outlines a universal scheme of incorporating wild and
cultivated plants and adding extra ranks (subspecioid, race, major group, cultivariant), to those
of Turnbull and Griffins, who doubted whether foresters should use the Jnternational Code of
Nomenclature for Cultivated Plants (ICNCP) at all.
Several speakers show that variation in some crop plants (e. g. Brassica rapa, Solanum
aethiopicum) can be classified using cultivar groups; in others (e.g. annual Bromus, Lens, Beta)
such groups can be treated as subspecies and integrated with the botanical classification, at the
same time illuminating the variation induced by cultivation. In chile peppers, groundnuts and
faba beans Barbara Pickersgill distinguishes three stages of such variation: (i) changes due to
initial domestication, (ii) subsequent partition of variation in cultigens through geographical
isolation, and (iii) ecological adaptation or human selection.
Continuous variation is another perennial taxonomic problem. Honor Prentice concludes that
a single general-purpose classification cannot include information about complex infraspecific
variation, and that supplementary maps and diagrams may be necessary to convey such
information. On the other hand, Chris Brickell mentions the problem of the introduction to
cultivation of plant from different parts of a continuously variable wild population. Such
introductions often remain distinct in cultivation but not invariable; hence they cannot be
classified as cultivars. In such cases he strongly favours the use of the botanical rank forma; I
support him in this (cf. my treatment of Hypericum olympicum in The Plantsman, 2: 193-200,
1980).
In reading this book, I was impressed by the efforts that are being made to bridge the wild/
cultivated gap in plant taxonomy, despite the sometimes very different problems that have to be
faced in each field of study. It is therefore to be recommended to Watsonia readers not only as an
invaluable source book for taxonomic data, techniques and opinions, but also (and primarily in
my view) as a demonstration that taxonomy within the garden wall, the field margin and the
forest fence is very much alive and should not be ignored by the field botanist.
N. K. B. RoBsSon
BOOK REVIEWS 349
The flora and vegetation of Britain. Origin and changes — the facts and their interpretation. Edited by
J. L. Harley & D. H. Lewis. Pp. 1-128, with numerous text-figures and two frontispieces.
Academic Press, London. 1985. Price £5.00 (ISBN 0—12-325570-8).
This Festschrift celebrates the eightieth birthday of Professor Arthur Roy Clapham, being the
proceedings of a symposium held at the University of Sheffield on 19th May 1984. The volume
under review is actually a reprint of part of Volume 98 Part 1 of the New Phytologist (1984), of
which Clapham was an editor for 30 years (1932-1961). To the original publication have been
added in this volume a very brief index (not covering plant names), a frontispiece photograph of
Professor Clapham (unfortunately undated but I believe c. 1970), and a photograph of the
symposium participants (the two un-named faces are both B.S.B.I. members — no. 11 is Dr D. A.
Ratcliffe and no. 16 is Mr T. C. G. Rich).
The main part of the volume consists of seven excellent review articles on ecology (2),
vegetational history (3) and cytogenetic (1) and distributional (1) aspects of plant taxonomy.
Coverage of mainstream taxonomy is singularly lacking but, as C. D. Pigott points out, Clapham
was not primarily a taxonomist, so the bias towards ecology is fully justified.
Probably of greatest interest to B.S.B.I. members are the papers by S. M. Walters on the
relation between the British and European floras, by T. T. Elkington on cytogenetic variation in
the British flora, by C. D. Pigott on the ecology and conservation of the British flora, and by D. A.
Ratcliffe on human influence on British vegetation since Medieval times. All of these, as well as
the other three articles, make very interesting reading equally suitable for the bed and for the
laboratory. My personal preferences are Ratcliffe’s and Elkington’s contributions, as these contain
more ‘meat’ than some of the others and represent valuable assessments of important topics by
leading active researchers. These two articles in particular are strongly recommended; I think no-
one actively interested in the British flora will fail to gain new ideas or attitudes from their study.
C. A. STACE
Olands och Gotlands Véxtvarld en ekologisk och Kulturhistorisk Flora. U. Ekstam & R. Jacobson.
Pp. 336, with 232 colour photographs, 15 maps and 6 text figures. Bokforlaget Natur och Kultur,
Stockholm. 1984. Price 195 S.Kr. (ISBN 91-27-01319-7).
Gotland and Oland are two low-lying botanically interesting large limestone Swedish islands in the
Baltic between 56°N and 58°N. Oland lies close to mainland Sweden, with which it is now
connected by the longest bridge in Europe, but Gotland is more isolated. On account of their
geology and maritime climate, together they possess a flora distinct from that of mainland Sweden,
which is granitic, by sheltering a number of species unknown on the mainland, notably Globularia
vulgaris, Helianthemum oelandicum, Adonis vernalis and various orchids. The two islands have
floristically much in common, with approximately the same number of species on each, but some
occur on the one which are not found on the other or are more abundant on the one than on the
other. Their botanical exploration began in 1741, when Linnaeus visited them both with a party of
five young gentlemen who came at their own expense, as his assistants in making and recording
observations. He published in 1745 a detailed interesting account of their travels, Olandska och
Gothlandska Resa, of which an annotated English translation by Asberg and Stearn has been
published in Biological Journal of the Linnean Society, 5: 1-220 (1973). 2
Linnaeus wrote in his account for 1st June 1741 that “As soon as we touched the shore of Oland
we realized this was a land which was altogether different from the rest of the Swedish provinces.
Thus we decided to record everything we would see on this island more meticulously’. Gotland
they surveyed with the same eager curiosity and diligence, and not only as regards natural history.
Linnaeus even noted that the speech of the Gotlanders “‘was somewhat different from ordinary
Swedish and was somewhat like Norwegian in accent”; its phonology, according to Gudmund
Schitte’s erudite linguistic work, Our Forefathers (1933), gives it ‘‘a claim to be regarded as a
separate branch co-ordinate with Swedish and Danish’. In keeping with this evidence of isolation,
Pulsatilla vulgaris subsp. gotlandica from the mainland is an endemic pasque-flower.
Linnaeus found on the two islands a number of species which he had not seen wild elsewhere,
350 BOOK REVIEWS
and he based his diagnoses in the Species Plantarum on Gotland or Oland material by references
back to his Olandska och Gothlandska Resa. Thus Gotland is the restricted type-locality for
Artemisia rupestris, Cladium mariscus, Geranium lucidum, Lactuca quercina, Melica ciliata, Pyrola
minor, Schoenus nigricans, Scutellaria hastifolia and Tofieldia calyculata. Oland is the restricted
type-locality for Atriplex hastata, Carex arenaria, Globularia vulgaris, Helianthemum oelandicum,
Hornungia petraea, Oxytropis campestris and Tetragonolobus maritimus. In view of this Linnaean
association, but above all for the intrinsic interest of their plants, the two islands have received
much attention both floristically and ecologically from Swedish botanists, culminating for Oland in
Sterner’s Flora der Insel Oland (1938). The literature relating to them is very extensive. Oland och
Gotlands Vaxtvarld, profusely illustrated in colour from photographs by Marrus Mattson and Tord
Porsne as well as by the two authors, is a welcome and innovative addition which can be thoroughly
recommended as a guide to the plant world of these unique islands.
The work is not however, a conventional flora with all the species in one taxonomic sequence
family by family. Instead they are grouped according to habitats, as the title partly implies. The
first part deals primarily with general ecology, including human influence, for the islands have
been inhabited for a long period. Here the different vegetation types are described and illustrated.
Then follow concise accounts of the individual species, all illustrated with habit photographs, with
notes on habitat, distribution and characteristic features. The little squares accompanying these
neatly summarize occurrence: O (=Oland), G (=Gotland), S (=sillsynt; rare, uncommon), M
(=mattlig; moderately common), a (allman; common) as explained on p. 10, according to the
number of places where found. The species are placed under 19 well defined habitats, which should
be a convenience in fieldwork. Since this book would be invaluable for non-Swedish visitors to
these islands now so accessible, the publishers might do well to issue with it a leaflet translating the
page-headings, e.g. Ang och betesmark=meadow and pastureland; Vagkant=roadsides; Trask
och vattendrag=marsh and watercourses, etc. Visitors from the British Isles will find here many
plants with which they are familiar, e.g. Potentilla fruticosa in Oland growing on limestone where
water stands during the winter, as it does in the west of Ireland. More exciting are such specialities
as Globularia vulgaris and Helianthemum oelandicum, relicts of a warmer period with their nearest
representatives in southern Europe. Twenty-nine species of Orchidaceae, including Dactylorhiza
sambucina with red and yellow forms often growing together and known as ‘Adam och Eva’, occur
on the islands; although often abundant, they are rightly protected by law, as are Adonis vernalis,
Anemone sylvestris, Hepatica nobilis, Pulsatilla vulgaris, P. patens and some other species. There
appears to be no reference to Stora Karls6 and Lilla Karlso off Gotland, the Charles Islands, where
Linnaeus discovered Artemisia rupestris and Lactuca quercina. For anyone wishing to repeat
Linnaeus’s travels or simply to have a pleasant botanical holiday on Oland or Gotland, this book
will be an excellent companion, the wealth of illustrations compensating for any linguistic
difficulty.
W. T. STEARN
Cladistic biogeography. C. J. Humphries & L. R. Parenti. Oxford Monographs on Biogeography,
No. 2. Pp. xii+98, with 67 text figures. Clarendon Press, Oxford. 1986. Price £19.50 (ISBN 0-19-
854576-2).
Undoubtedly, Hennigian cladistics are an important tool of the modern systematist; likewise,
cladistic biogeography, with its biological area-cladograms, is a useful approach to answering
certain questions, particularly at the world scale, which is only one of many levels at which the
biogeographer works. However, the strident and bombastic efforts of protagonists in these fields to
make this approach the only acceptable one in biogeography are a nonsense and I, for one, refuse
to be bounced into a paradigm shift. I find the methods inductive, mechanical, unrelated to
process, retrospective (and thus dubiously ‘historical’), lacking in any true sense of geography
beyond that of latitude and longitude, and jargon-ridden in the extreme. Nevertheless, this brief
book is an important contribution to biogeography, not the least because one can actually
understand it, dense though the discussion is, and also because we are not treated to the same ad
hominem abuse and rhetoric which has so far characterized much of the writing on the topic. It is,
BOOK REVIEWS ‘sk
in my opinion, precisely the type of controversial contribution which should grace the Oxford
Monographs on Biogeography.
Of course, the argument remains largely unchanged and, as normal, progresses through
contrived opposites (dispersalist/vicariance biogeography — I give allegiance to neither), through
assertion (e.g. “geological hypotheses do not test biogeographical patterns”) and through the
divorce of form or pattern from process (i.e. plants and animals don’t actually need to live
anywhere, but express their areas on isomorphic surfaces). The book presents a potted history of
spatial biogeography, which naturally reaches the right cladistic conclusions, gives an excellent
survey of methods employed by cladistic biogeographers, and then gets slightly obsessive about a
lost ‘Pacifica’. Particularly annoying is the in-built assumption that the convergence of pattern has
eluded past biogeographers. What has certainly eluded the cladistic biogeographers is cartography;
the maps are, quite frankly, appalling, being usually non-equal-area and lacking even scales. The
authors also believe that the hypotheses of cladistic biogeography are ‘testable’ in the true scientific
sense. They are, of course, testable only if you enter the philosophical circle of cladistic thought
and then all is well, although when converted you must be especially careful not to fall into the trap
of telling a ‘story’ or you will be roundly chastized, as is the fate of poor Rosen on p. 30.
Nevertheless, this volume is the best introduction so far to the ‘vicariance approach’ in
biogeography, although at c. 20 pence per printed page members of the Society might not find the
tough reading worth either the effort or the cost.
P. SToTT
Supplement to the Flora of Gloucestershire. S. C. Holland, H. M. Caddick & D. S. Dudley-Smith.
Pp. xx+196, including 8 colour plates and 20 black & white drawings. Grenfell Publications,
Bristol. 1986. Price £11.50 (ISBN 0—948715-—00-6).
The facsimile reprint in 1974 of The flora of Gloucestershire by H. J. Riddelsdell, G. W. Hedley
and W. R. Price, published in 1948, helped to stimulate interest in the rich flora of the county and
made increasingly clear the desirability of an account of the changes in plant distribution and status
over nearly four decades. During that time, as elsewhere, construction of roads, abandonment of
railways, quarrying, adoption of modern agricultural practices and especially drainage have
affected the vegetation of many habitats and resulted in the loss of nearly one hundred species, a
loss matched almost equally by gains in the flora. Since 1950, extensive fieldwork, the more recent
based on a 2X2 km grid, has been undertaken by the Gloucestershire Naturalists’ Society. In this
Supplement Mrs Holland has acted as an energetic Executive Editor, with a Committee of Miss
Caddick and Mrs Dudley-Smith and also, up to 1982, Miss K. E. Ludbrook.
Regrettably, except for Rubus and Charophytes, the Supplement does not cover the same area as
the 1948 Flora but relates to the Administrative County of Gloucestershire established in 1974.
The decision to restrict the recording to the new county boundary means a lack of comparability
with the 1948 Flora as only part of Botanical District 5 is now covered, most of this falling in Avon.
In terms of Watsonian vice-counties, this means that a floristically-rich part of v.c. 34 is excluded.
Most of the book constitutes detailed records complementing the entries of the earlier Flora,
especially as regards occurrences in previously under-recorded areas. Perhaps inevitably the
Supplement has to be read in conjunction with the 1948 Flora to obtain a picture of the overall
distribution of a species in the county. For Anacamptis pyramidalis, for example, only two districts
are mentioned in the Supplement and the reader may wonder to what extent this orchid occurs
elsewhere. Some information given would have desirably been further updated; the colony of
Monotropa hypopitys on Cleeve Hill is reported as about three hundred plants in 1964, but there is
no information of its performance since.
Of particular interest are the details of changes in abundance and about species of restricted
occurrence outside Gloucestershire. Distribution maps are given for some 16 taxa, including
Herminium monorchis of diminished distribution and Spiranthes spiralis of increased abundance.
The value of B.S.B.I. initiatives in surveys of particular species is reflected here in the maps and
entries for Populus nigra and Sorbus torminalis among others. The contrast in distribution of
Gagea lutea, Saxifraga granulata and Thesium humifusum is evident from the ‘dot’ maps. Although
352 BOOK REVIEWS
only a few maps are given, numbers of grid squares are routinely included for new localities, so that
future mapping by computer can be undertaken.
A very attractive feature of this book is the high quality colour plates. Particularly notable
among them are Cuscuta europaea on nettle (a survey of hosts is included but one queries whether
haustorial contact was made with a ‘host’ such as Prunus spinosa) and a prolifically flowering
colony of Utricularia vulgaris. Some of the black & white drawings usefully show taxa restricted to
and of particular note in Gloucestershire and for which illustrations are not readily available.
Among these are Corrigiola telephiifolia, new to the British alien flora but now feared lost, and a
form of Stellaria nemorum intermediate between subsp. nemorum and subsp. glochidisperma.
Some of the other plants, e.g. Anagallis tenella, are illustrated much better in well-known sources.
Attempts have commendably been made to explain changes in distribution, but the reasons
advanced for these do not always seem probable. The decline of Parnassia palustris at Brassey
Nature Reserve, which is linked with a large increase of Juncus subnodulosus, appears more likely
to be associated with eutrophication than with climatic change as suggested. Pinguicula vulgaris,
reported as “‘not seen in the 1970s and may have been lost” at Puckham Marsh was known to be at
this site in 1975 (Dr B. D. Wheeler, pers. comm.).
Many records refer to aliens, and a list of well over a hundred casuals new to Gloucestershire is
usefully included. Care has been taken over nomenclature but one wonders about the value of
giving an English name such as ““A Dodder”’ for Cuscuta campestris!
Despite some defects, this Supplement has much to commend it. The standard of production is
high, typographical errors are few and there is much here for the plant lover and those interested in
changes in plant distribution.
A. J. WILLIS
Watsonia, 16, 353-360 (1987) 353
Obituaries
RONALD ARCHIE BONIFACE
(1913—1985)
Ron Boniface, a B.S.B.I. member since 1950, died in West Middlesex Hospital on Christmas Eve,
1985, from a brain tumour. Ron was born on 9th July 1913, in Eastbourne, Sussex, where he lived
with his parents until World War II. In the thirties, besides being a keen racing cyclist, he became
interested in plants as a hobby, and soon became an expert on the flora of the Eastbourne area. He
contributed a number of interesting records to Wolley-Dod’s (1937) Flora of Sussex. After a period
on war work in Staffordshire, where he rediscovered the interest of Chartley Moss, he and his
parents moved to Chiswick in West London, where he worked in building and as a motor engineer.
He spent his spare time botanizing, his main interest lying in re-discovering plants that other
botanists had given up as lost, and in exploring areas that were currently unknown botanically. He
refound Pinguicula vulgaris for Hampshire in this way.
I first met Ron in 1947, and became most impressed with his keenness and remarkably critical
knowledge of the British flora. Without formal training, he became in my opinion one of our most
knowledgeable field botanists. About this time, Ron became interested in bryophytes too, and did
excellent work on them in south-eastern England: he published a paper on the ‘“‘Hepatics of the
London Area” in the London Naturalist in 1956.
His contributions to botanical knowledge, however, were not to be measured by the few papers
he actually published. His excellent herbarium has gone to the National Museum of Wales.
For many years, often accompanied by Ted Wallace, he was my regular companion on field
trips. We had many long excursions to such places as the Scottish Highlands, Teesdale, East Anglia
and other places. He was a good field companion, modest and even-tempered, and had a quiet,
gentle sense of humour. When the weather was bad or the going strenuous a cup or two of really
strong tea would always revive him, preferably the sort in which the spoon would almost stand up
on its own!
Ron married Iris Bagshaw in 1959, and they had one daughter, Janet. I am indebted to his step-
daughter, Mrs Carol Watson, for some information on his earlier life.
Those of us who knew him miss his gentle field companionship and his extraordinary knowledge
of British plants and their habitats.
F. ROSE
RONALD MELVILLE
(1903—1985)
Ronald Melville, I.S.0., B.Sc., Ph.D., M.P.S., Ph.C., F.L.S. died at the age of 82 on 6th August,
1985. Though an occasion for sadness, his death marked the end of a long, happy and fulfilling life,
a distinguished career and a period of association with the Royal Botanic Gardens, Kew, spanning
50 years.
Ronald Melville was born in Bristol on 12th March 1903, the second of seven children, and first
son of Edgar and Florence Melville. Only six months after his birth, the family moved to Cardiff
and it was there that he spent his childhood. He originally intended to follow a career in medicine
but, as his father died when he was 13, he left school after matriculation in 1919 to train in his
uncle’s pharmacy in Cardiff. In 1924 he became a member of the Pharmaceutical Society and, at
the age of 22, he gained the Society’s highest diploma, a qualification which now ranks as a degree.
While practising pharmacy in retail shops and at the London Hospital, Whitechapel, he studied
for an honours degree in Botany at Birkbeck College and graduated in 1931. In 1930 he became a
354 OBITUARIES
research assistant under Professor V. H. Blackman of Imperial College, and gained a scholarship to
study for a Ph.D. at the Research Institute at Cheshunt. During this time he was lecturing in the
evening at Birkbeck College and also engaged in research on aquatic Phycomycetes under Dame
Helen Gwynne- Vaughan and Dr B. Barnes. He was awarded his doctorate in 1934 for a thesis on the
growth relationships of seedling tomato plants and joined the staff at Kew in November that year.
At the Royal Botanic Gardens he worked in the Museums with William Dallimore and then John
Hutchinson, with whom he was later to collaborate in writing a book, The story of plants and their
uses to Man, published in 1948. During his years in the Museums he became responsible for the
identification of woody plants in the Arboretum at Kew and the conifers in the National Pinetum at
Bedgebury. In 1936 he made an intensive study of the British elms and became an acknowledged
authority. Problems in the elm complex and in intergrading Ulmus hybrids led, in turn, to studies in
leaf morphology and ontogeny and the interaction between leaf-shape patterns in hybrids.
Through the 1939-1945 War he represented the Royal Botanic Gardens on several committees
including the Vegetable Drugs Committee of the Ministry of Health. In this capacity, when the
country’s supply of vitamin C from citrus fruits was threatened, he recommended the use of rose hips
as an alternative. With Dr Magnus Pyke working on their chemical assay, he then undertook a
detailed survey of the British wild roses and examined the problems of classification in this group of
pseudosexual unbalanced polyploids. Having first proposed the use of rose hips in the preparation of
the now familiar vitamin-C-rich syrup, he became deeply involved in directing the development of its
production and served on the Scientific Committee of the Rose Hip Products Association. His was
indeed an extremely important contribution to the dietary welfare of the Nation.
After 16 years in the Museums at Kew he transferred to the Herbarium in 1950 to take charge of
the Australasian Section, a responsibility he retained until his retirement. His expertise was greatly
stimulated by a year spent in Australia from 1952 and 1953, when he was based in Melbourne Botanic
Garden under the auspices of the Maud Gibson Trust and collected more than 20,000 specimens
from many parts of that Continent, especially Victoria. Later, in 1961, he spent six months in New
Zealand as a D.S.I.R. Senior Research Fellow at the Herbarium of the Botany Division, Lincoln,
near Christchurch. While in Fiji on the return journey in 1962, he and his wife collected a rare
Glochidion (Euphorbiaceae) which Kenneth Airy-Shaw in 1971 named G. melvilliorum; and, in
1978, an Australian Acacia was named A. melvillei by Dr Pedley of the Queensland Herbarium.
As a taxonomist, Ronald Melville sustained his early interest in hardy trees and shrubs, continuing
his specialization in Ulmus and Rosa and becoming the B.S.B.I. Referee for both genera. He will be
remembered with gratitude by the many members to whom he explained and demonstrated the
intricacies of rose and elm classification on the field meetings that he led; and until shortly before his
death he was always ready to name rose or elm specimens for members, no matter how large the
parcel.
From 1956 he was engaged in researching floral evolution. A study of floral vascular systems by
clearing techniques, begun in 1959, led to the formulation of the Gonophyll Theory to explain the
nature of the flower and its evolution, and in developing his ideas he took into account the
reproductive structures in extinct plants. Considering the reproductive organs of late Palaeozoic
Glossopteridales (sensu lato), he came to regard the alleged bisexual structure of Glossopteris as
occupying an important, key position in his theories, and believed that the occurrence of primitive
structures in many extant Angiosperms supported the various intermediate evolutionary stages of
both the male and female organs that he postulated in his ‘New theory of the angiosperm flower’.
Running parallel with his work on floral evolution, he studied the evidence for continental drift,
which led him to a reconstruction of the Mesozoic Gondwana continent and an explanation for the
migration routes of early Angiosperms from the southern to the northern hemisphere. Some of his
preliminary studies of the phylogeny of several genera and the impact of continental drift on the
distribution of species were discussed at the Gondwana symposium in Montevideo in 1967.
He retired as a Senior Principal Scientific Officer in March 1968, with more than 100 published
papers to his credit. Nevertheless, he continued to be actively involved as a botanist, writing many
more papers and working regularly at Kew until he suffered from declining health in 1985.
As a consequence of his work as a member of the Survival Service Commission for the
International Union for Conservation of Nature and Natural Resources, he was responsible after
retirement for compiling the first ever Red Data Book of threatened plant species, published in 1970,
and he continued working for the I.U.C.N. until 1973. During his retirement, among his many other
OBITUARIES 355
botanical interests, he turned his attention to studies of leaf venation, which, by 1970, had led to
the discovery of Angiosperms with leaves closely resembling members of the fossil Glossopteridae.
He related leaf, petal and sepal venation patterns to this fossil group and eventually formulated his
Remoration Theory of evolutionary retrogression. Linking his findings with his theories on
phytogeography and palaeomigration of plants, especially in the Pacific, and correlating evidence
provided by floral anatomy, ontogeny, leaf structure and continental drift, he never ceased in his
attempt to find a comprehensive solution to the problems of Angiosperm origin and evolution.
In 1935, he had married Elsie Sharpington, herself a botany graduate of Westfield College,
University of London. They had two children, Fenella and John, and four grand-children and
enjoyed a happy and tranquil family life. Later, his daughter also became a botanist and, in 1969,
they together published a paper on fenestration in the leaves of Monstera.
From 1936 he served on various pharmaceutical committees and assisted in the production of a
new edition of the British Pharmacopoeia (1949).
As a result of his work and interest in British trees and shrubs he had become a member of the
Royal English Forestry Society in 1935, and from 1939 had served on committees of the
Systematics Association, becoming its Botanical Secretary in 1946. In that year he was also elected
to the Council of the Botanical Exchange Club (soon to become the Botanical Society of the
British Isles), and in later years he was a member of the Richmond (Surrey) Scientific Society,
serving as its President from 1975 to 1977 and continuing afterwards as a very active committee
member. He was elected a fellow of the Linnean Society in 1938, and became a member of its
Council from 1942 to 1946. In the New Year’s Honours List of 1968, the year of his retirement, Dr
Melville was made a Companion of the Imperial Service Order in recognition of his considerable
and various contributions to science, including his key role in rose hip syrup development and his
work on pollen analysis of honey.
Ronald Melville’s important war-time proposal to use rose hips as a source of vitamin C, and his
subsequent involvement in the development of rose hip syrup production will never be forgotten,
but perhaps he will be best remembered among botanists for his theories of angiosperm evolution.
There is no doubt that, borne of a very active and enquiring mind, his controversial theories and
ideas on this and other topics have always stimulated discussion, and will continue to be subjects
for keen debate. He was exceptional in his depth of knowledge in a wide field of botanical
disciplines, and all who knew him during his many years at Kew will remember him with great
affection.
ALEXANDER STUART WATT
(1892—1985)
Alexander Stuart Watt, F.R.S., Ph.D. was born near Turriff in Aberdeenshire on 21st June 1892
and died at Cambridge on 2nd March 1985 at the age of 92. Of farming stock, he graduated from
the University of Aberdeen in 1913 and came to the Botany School at Cambridge to work under
the pioneer ecologist Sir Arthur Tansley. After two years military service he returned briefly to
Cambridge, but did most of the work for his Ph.D., which he obtained in 1924, while he was a
lecturer at Aberdeen. In 1929 he was appointed Gurney Lecturer in Forestry at Cambridge, but
when that department finished in 1933 he returned to the Botany School, where he was a lecturer
until his retirement in 1959. He was President of the British Ecological Society in 1947, was elected
to a Fellowship of the Royal Society in 1957 and received the Gold Medal of the Linnean Society in
£975.
Watt’s profound knowledge of vegetation was based on meticulously recorded observations and
experiments made in the field. His first work on the regeneration of oakwoods was followed by
studies of the beech-woods of southern England and the heaths of the Cairngorms. But his greatest
love was his study, continued long after retirement, of the grasslands and heaths of the East
Anglian Breckland, on some of which he made continuous recordings for over 50 years. His
knowledge of Bracken, Pteridium aquilinum, was encyclopedic, but the work that is probably most
356 OBITUARIES
interesting to B.S.B.I. members was his research on the rare species characteristic of Breckland. He
was sceptical of the Society and its activities pre-war, but joined in 1952, because he understood
“they were now going to do some proper botany”’. He gave me a lesson in this when he made me dig a
great hole in almost solid chalk so that he could demonstrate the massive tuberous stock of Bryonia
cretica subsp. dioica. He quite rightly said that that would ensure I would not forget the fact.
Alec (or, as he was known to his contemporaries at Cambridge, ‘Sandy’) Watt was a truly modest
and kindly man, but a wise councillor to all who asked his opinion. With his wife Annie he kept open
house to many generations of students who remembered them with great affection.
P. D. SELL
MARY McCALLUM WEBSTER Oi,
(1906—1985)
Scottish botany lost an outstanding enthusiast when Mary McCallum Webster died on 7th November
1985 at the age of 78. She was born in Sussex on 31st December 1906 of Scottish parents. Her father,
who was in the Army, was the son by her first marriage of Mrs M. L. Wedgwood, a friend of George
Claridge Druce. She was educated at home until the age of 15 by a record number of governesses, 19
in 7 years. Finally, she achieved her ambition to go to boarding school — ‘‘to play games and wear a
gym tunic’’, first at West Heath and then Ham Common. After her schooling she went to Brussels to
‘finish’. She had always been interested in flowers, and botany was her best subject at school.
Although she did various domestic jobs, played a lot of tennis and qualified for Wimbledon, and also
played hockey for the district, she still found time to botanize a great deal. When war came, Mary
joined the A.T.S. and was attached to the 10th Batt. Gordon Highlanders. She was sent to train at
the first school of cookery at Aldershot from which she was posted to Orkney and later Shetland,
where she was cook-sergeant at the time of the Battle of Narvik. The next move was to the
Bournemouth Officers’ Cadet Training Unit. She ended up as a Staff Captain in Field Marshall
Montgomery’s H.Q. in Germany in a branch disbanding the German Army.
After the war, she worked partly in hotels and partly free lance. While employed by friends in a
hotel in Kinlochbervie, a party came to stay which included Humphrey Jennings and the famous
ornithologist the iate James Fisher. Stephen Potter pretended to be the botanist of the group. He
used to spread out the daily finds under the kitchen table while she was busy concocting something
special for dinner, and there would be a cry “Cookie, what is this?””. And down she would get on all
fours, naming the plants. From then on she decided that work was for the winter, and the summers
were to be spent botanizing. When the B.S.B.I. mapping scheme was in progress for the Atlas of the
British flora, having no car, she walked a 100 miles a week for several summers, recording the species
in 266 grid squares in the north of Scotland.
In 1958 she visited her brother in Natal and then went on to a seven-month safari in Northern
Rhodesia and Tanganyika, where she collected 5000 specimens for Kew. One of them is the holotype
of a new species, Digitaria melanotricha W. D. Clayton, collected on Nsange Mountain, Ufipa, and
another from Lake Chila, Abercorn District, the holotype of Eragrostis mariae Launert. The next
three winters were spent at Kew helping to identify her African collections and doing some general
curating. This was followed by four winters in the Cambridge herbarium. After a short time she
decided that the technicians there were not working quite hard enough; with a mixture of firmness
and cajolery she dragooned them into a formidable force, and the usually sedate herbarium was
transformed into a hive of activity. It was in the happy days when extra labour was obtained by the
University at an hourly rate, and she would often be found already at work at 8.30 a.m. and could still
be found there at 10 p.m. When Mary came to fill in the cards for the Hieracium maps for the Critical
Atlas, Cyril West came to sort out the specimens and advise on particular records. Both started early,
both went without lunch and neither stopped very early for supper. The productivity was frightening.
The first of the various societies she joined was the Wild Flower Society in 1915. She was a faithful
member all her life, and became the Secretary for the Scottish branch in 1968. So many beginners can
thank her for her botanical help. She did not suffer fools gladly, but if one was willing to learn, Mary
couldn’t be more patient or instructive. Many ‘Woofs’ know their grasses from her saying: ““Holcus
mollis with hairy knees or Poa nemoralis with black knees”. Her comments to those who ought to
OBITUARIES 357
know better were very forthright. When P.D.S. altered the description of Ranunculus ficaria
subsp. bulbifer in her Flora from larger flowers to smaller flowers, she left it. But when later he
found that she indeed had large-flowered plants with bulbils in her area and confessed, he got the
most tremendous ticking off. She joined the Botanical Society of Edinburgh in 1954, the Inverness
Botany group in 1966 and was a member of the Moray Field Club. She was a Fellow of the Linnean
Society from 1960 to 1974.
She became a member of the B.S.B.I. in 1936, serving on the Council from 1960-1964 and the
Meetings Committee from 1960-1976. She was County Recorder for vice-counties 94 (Banff), 95
(Elgin) and a portion of 96 (Easterness). On a committee her remarks were very often very much
to the point, and her frequent presence on field meetings usually meant that some very obscure
plants would be recorded in the area for the first time.
Her first publication was a contribution with Peter Marler to the Flora of West Sutherland. In
1975 she produced a Check list of the flora of the Culbin State Forest, a place dear to her heart,
where she took so many people to see Goodyera, Corallorhiza, Orthilia and her favourite Moneses
uniflora.
Her most outstanding contribution to Scottish botany, however, was her Flora of Moray, Nairn
and East Inverness, published in 1978. Mary was highly individualistic and this is clearly shown in
her Flora. Using her life savings and helped by the financial support of two friends, she produced in
her own words “a Flora how I want it done’’. It differs from all other county Floras in listing
specimens and where they are to be found for nearly all the taxa. It contains a great many notes,
keys and helpful drawings about critical taxa provided by her many friends, and the twenty plates
of coloured photographs add a nice finishing touch. The maps with their dot distributions, which
form such a dreary part of most modern county Floras, were banished to the end of the book.
Although a dedicated ‘square-basher’ Mary was well aware that most botanists wanted to know
where a plant grew, and a dot in a 10 km square did not help much. In size the book was also
different from many of its large two- or even three-column contemporaries, fits easily into the hand
and is graced with a delightful dust jacket. Her faith in the lay-out of the book was justified for, as
she said, “It sold like hot cakes”. The first 800 copies paid for all her expenses, and with the
remaining money she was able to make two visits to Australia. The copyright of the Flora and the
remaining 70-80 copies have been left to the B.S.B.I.
Latterly she was working on a check-list for Banffshire, and in 1983 led a large field meeting
based on Tomintoul where over 15 N.C.R.s were recorded and 300 pink cards made out. That was
the last of the many successful meetings she led for both the B.S.B.I. and the W.F.S. to Arran,
Skye, Cairngorm and many other Scottish localities. She instilled into those who went into the field
with her that the best way to help conservation was to mind where one put one’s feet.
A visit to her cottage at Dyke made one aware of her varied interests: the pile of driftwood at the
gate collected for flower arrangement, her garden full of unusual flowers where even the weeds
were rich and rare, the porch full of recording cards and papers, dried flowers hanging on any
available hook for her dried flower arrangements, and her press with specimens for the herbarium
or her pressed flower pictures. All these she did with success, winning prizes for her flower
arrangements and firsts for her dried flower pictures exhibited at the Royal Highland Show.
The only time Botany took second place was during Wimbledon fortnight when she was glued to
the television, or during the tennis tournaments in north-east Scotland, particularly at Nairn and
Elgin, where she competed until three years ago. A tennis cup in her cottage is a reminder of how
good a tennis player she was. She was given it to keep after she won the Ladies’ Open Singles at the
North of Scotland tournament four years running during the 1950s.
Having seen virtually every wild plant growing naturally in the British Isles, she became greatly
interested in shoddy plants, and she loved the excitement of finding an unknown species on the
shingle at Galashiels or in the shoddy-manured fields in the south. She found the knowledge gained
from looking at these aliens was a help in identifying the grasses she saw on her two trips to
Western Australia. These two trips made a sort of grand finale to a life in which the love of flowers
played such an important part and to which the glory of the Australian flora brought such great
pleasure in her last years.
Her British specimens are divided between the Department of Botany, University of Aberdeen
(ABD), the Botany School, University of Cambridge (CGE) and the Royal Botanic Garden,
Edinburgh (E). Cambridge is particularly rich in critical taxa of a wide range of genera, especially
358 OBITUARIES
Hieracium or “‘hawkers”’ as she called them. Many of her colour slides are also preserved in the
Cambridge herbarium.
Mary McCallum Webster was a remarkable person, who at both work and play never gave less
than a hundred per cent, nor would she tolerate slackness in others. Her forthright views often
brought disagreements, but her many friends will best remember her for her robust good humour
and lasting loyalty. Her life was dedicated to her love of the Scottish flora and is best expressed by
the words of S. M. Walters in his foreword to her Flora ““One whose heart was always, and
remains, ‘in the Highlands’. There are really few better places for one to be.”
O. M. STEWART
P. D. SELL
CYRIL WEST
(1887—1986)
Cyril West, O.B.E., M.A., D.Sc., F.L.S. was born at Forest Hill, near Sydenham, Kent on 16th
December 1887 and died in hospital at Linton near Maidstone, Kent on 25th March 1986 at the age
of 98. His father, John Stapylton West, was a tea dealer in the City who also lived to a great age, of
96. Cyril West’s school education was at St Olave’s in London, after which he went to the Imperial
College of Science from 1907-1914 and 1916-1919. He took honours in Botany and received his
B.Sc. in 1911 and his D.Sc. in 1918. S
Soon after the outbreak of the First World War he was commissioned in the Royal Artillery; but
an accident with a horse seriously injured his knee, and after a long stay in hospital a medical board
recommended his transfer to the Special Reserve of Officers, whereupon he returned to the
Imperial College. In old age the damage to his knee brought about trouble with his ankle, which
finally prevented him walking into the mountains after hawkweeds.
At Imperial College, West began research under V. H. Blackman on the dormancy of seeds,
then moved to Cambridge to continue it with Franklin Kidd, both of them working under
Blackman’s more distinguished brother, F. F. Blackman. At St John’s College they became
associated with George Briggs (1893-1985), and the three of them collaborated in studies of plant
growth which resulted in two famous papers in Annals of Applied Biology in 1920, discussed by G.
C. Evans in chapter 13 of his The Quantitative Analysis of Plant Growth (1972). The work on seed
dormancy led to their study of the effects of carbon dioxide on plant metabolism, and thus to the
controlled atmosphere storage of fruit, first in the laboratory, then in tests at Chivers’ farm at
Histon; and their results finally came into commercial use in 1928. This project led to the
establishment of a Food Investigation Organization with a main laboratory in Cambridge known as
the Low Temperature Research Station. Kidd and West joined its staff; but soon afterwards
money became available to investigate the transport of fruit from abroad and a laboratory was built
at Ditton, near Maidstone in Kent. West was Superintendent of this laboratory from 1931 until
1949, when he returned to Cambridge and remained there until his retirement in 1951. Throughout
this period Kidd and West published a long series of joint papers, and in 1963 they were presented
with Kamerlingh Onnes Medals at Leiden for their work on cold storage. A full report on this was
published in Mededelingen van de Nederlandse Vereniging voor Koeltechniek in 1963. This is the
only time, as far as I am aware, that West went outside the British Isles.
While working at Cambridge between 1919 and 1922 West demonstrated to students in the
Botany School, and Sir Harry Godwin in his autobiography Cambridge and Clare (1985) says how
impressed he was by West’s conviction of the supreme value to biological students of examining
fresh material at every opportunity. He took his B.A. in 1921 and his M.A. in 1926.
Cyril West’s interest in the British flora was quite different from his interest in plant physiology.
It gave him intense pleasure to look at a plant just for its beauty; he would make long journeys to
see a plant for the first time, and much of his spare time was spent looking at them. His love of
plants is perhaps best illustrated by an incident on what was his last visit to a mountain locality.
Archie Kenneth had managed to get him up the chair-lift at Allt nan Guibhas in Glen Coe. On
returning from a short walk Archie found Cyril sitting on a rock stroking the hairs of Hieracium
holosericeum, one of the most beautiful of all hawkweeds.
OBITUARIES 359
In his very long life he had travelled the length and breadth of the British Isles and he probably
saw more British species of plants living than any other botanist has ever done. I do not think he
had any ordinary species left to see; he had seen nearly all the species of Hieracium, W. C. R.
Watson had shown him over 300 of the Rubi he recognized, and I had checked off most of the
Fumariae, Sorbi and Euphrasiae and some of the Taraxaca for him. Towards the end of his life he
was shown many aliens, and I told him about many varieties. As a young man he was very fond of
rock climbing in North Wales, which enabled him to see such plants as Lioydia at close range. He
only collected plants other than Hieracium which he thought might be useful to the Cambridge
herbarium or if I especially asked him to get something.
Cyril West’s great contribution to systematic botany was his work on the genus Hieracium. It was
a group he had always been interested in and which he had discussed with F. J. Hanbury and H. W.
Pugsley. Only when he was about to retire and Pugsley’s Prodromus had just been published did he
begin the study of the genus seriously. In 1950 I had just returned from National Service and was
considering what critical genus to take up. I finally decided on Hieracium and was examining the
specimens in the Cambridge herbarium when in walked Cyril West with the same object in mind.
We joined forces, little knowing that our partnership was to last for over thirty years.
In the early days when he was regularly in Cambridge working at the Low Temperature Station
we spent an enormous amount of time putting the Cambridge specimens in order and getting to
know the species recognized by Pugsley. Later, throughout the winter, when he was mostly in Kent
he would come up for four days a month. It was then I discovered his remarkable staying power.
Our days were usually from 9 a.m. to 9 p.m., but it was I who was drooping at the end of the day
and completely exhausted after four days, but he, even when at the age of 90, went off to catch his
train as though nothing had happened. And he never had anything to eat between breakfast and
supper and then not very much. When not at Cambridge he would spend much of his time in the
room he still had in the Ditton Laboratory 35 years after he had retired, poring over his specimens,
or visiting the British Museum (Nat. Hist.) to work out a whole series of problems to be discussed
on his next visit to Cambridge. As I had other official work to do, much of the donkey work was
left to him. Many local flora writers and recorders owe it to him for passing on to them information
we had worked out. Much of the most meticulous proof reading was also carried out by him. The
52 pages of Hieracium in Flora Europaea were letter by letter, figure by figure, twice checked back
to the original manuscript over a period of two very long days. This gave him a severe headache,
but when I offered him medication he would only take half an aspirin. Although his main interest
was the British species of Hieracium he readily gave up the time to write the more important
accounts for Flora Europaea and Flora of Turkey.
During the summer months he spent much time searching for hawkweeds in the field. In the
1950s his companion was Major J. W. Cardew, who rode a Sunbeam twin, West sitting in the
sidecar with all the equipment. When necessary the sidecar was taken off and they went up remote
mountain tracks on the motor bike. In later years he regularly visited Archie Kenneth in Kintyre.
Archie would go into the mountains to collect and he would examine and press the specimens, a
task which often took half the night. In the early days he was loth to collect specimens at all
because of his strong feelings about conservation, but he eventually realized it was necessary and
his later gatherings are all first class. His grid references must be considered with care. He would
never take a map into the field and often put vague localities on his specimens. When I pressed him
to include references he did so, but only afterwards from memory. Those specimens collected by
Archie Kenneth usually have very precise grid references. His work on Hieracia in relation to other
work on the genus and lists of published papers is set out in a paper I have written, now awaiting
publication in Watsonia.
He originally joined the old Botanical Society and Exchange Club in 1914, resigned in protest at
the way in which Druce collected such a large number of specimens, and joined again after Druce’s
death. He was made an Honorary Member in 1973. He was elected a Fellow of the Linnean Society
in 1913 and an Honorary Fellow in 1978. He was a very prominent conservationist, joining the
Kent Naturalists’ Trust in 1961 and sitting on its council from 9 March 1961. He was a founder
member of the Kent Field Club, of which he was made President when over the age of 90.
West always wore a dark grey suit, stiff white collar, grey tie, white silk scarf, lightweight black
boots, and a black trilby hat. You were just as likely to meet him so dressed in the middle of the
Cairngorms as in the centre of London. If it was wet he might tuck his trousers in his socks and put
360 OBITUARIES
on a mackintosh and sou’wester. If he got wet through he would still appear for dinner dressed
exactly the same, though changed. He referred to his good suits and his bad suits, ee they all
looked exactly the same to me.
He had a great love of cricket and was privileged to see the two great innings by Frank Woolley
against the Australians at Lords in 1921. I could not resist mentioning it occasionally, for the
memory of it never failed to bring tears of joy to his eyes. Woolley played for his beloved Kent,
and in his eyes neither Don Bradman or Jack Hobbs could touch him for elegance and grace.
Cyril West was one of the kindest men I ever met. The technicians in the Cambridge herbarium
would do anything for him. They packed his parcels, supplied him with all his needs and brought
him his tea. And in return they all got boxes of chocolates for Christmas. We celebrated his 90th
birthday in the Cambridge herbarium with a great iced cake which was one of the few items of food
of which he was very fond. I only once heard him speak of a botanist with any ill-will and that was
George Claridge Druce. When I suggested we named one of the new Shetland hawkweeds after
him he flatly refused, saying that compared with E. S. Marshall and H. W. Pugsley he was very
second rate and his F.R.S. was a scandal.
He will be much missed by his many friends.
P. D. SELL
Watsonia, 16, 361-363 (1987) 361
Report
ANNUAL GENERAL MEETING, 10TH MAY 1986
The Annual General Meeting of the Society was held in the rooms of the Linnean Society of
London, at 12 noon, with 83 members present. Mr D. E. Allen M.A., F.L.S., President, took the
chair.
Apologies for absence were read, and the Minutes of the last Annual General Meeting, as
published in Watsonia, 16: 109-110 (1986), were approved unanimously and signed by the
President.
REPORT OF COUNCIL
The Report for the calendar year 1985 was accepted nem. con., and some highlights for the Society
during the year were reviewed: a record attendance at the Annual Exhibition Meeting in London;
a successful conference on Critical Groups in the flora of the British Isles held at the Merseyside
County Museums; and the setting up of the Conservation Association of Botanical Societies. It was
noted that the B.S.B.I. Conservation Committee would be retained. Plans for a new mapping
survey, proposed for 1987, had progressed well during the year.
' TREASURER’S REPORT AND ACCOUNTS
The Report was accepted nem. con. Comment on the steady but slow increase in membership was
discussed, and the possibility of a more vigorous recruitment policy suggested to increase income
for ambitious publications. Experience had shown however that many of the new members joining
in large numbers following publicity stayed with the Society for about one year only. The general
feeling was not to change the Society to attract more members.
ELECTION OF OFFICERS
From the chair, Mr Allen proposed the re-election of Mrs M. Briggs M.B.E., F.P.S., F.L.S., as
Honorary Secretary General and Mr M. Walpole F.L.A., F.L.S., as Honorary Treasurer, and
these were carried with applause. Mr Allen, thanking all the Society’s Officers, took the
opportunity to acknowledge the Secretaries of the Permanent Working Committees as the
workhorses behind the scenes, upon whom the Society depended for the success of its varied
activities. Mrs J. Robertson, retiring Honorary Meetings Secretary, was warmly thanked for her
contribution to the Society’s programmes, and Mrs A. Lee was welcomed as her successor, with
reference to her help in past years, in particular her contribution to the 1985 Annual Exhibition
Meeting. Sincere thanks to the Editors of the Society’s publications were coupled with a particular
note of acknowledgment to Mr E. D. Wiggins, retiring Editor of B.S.B.I. News, for his
achievement in producing 27 numbers of this publication which, under his editorship, has become
such a popular and valued feature of the life of the Society.
ELECTION OF COUNCIL MEMBERS
In accordance with Rule 10, nominations had been received for Lady R. FitzGerald, Dr P. A. Gay
and Mrs S. Oldfield. Their election was proposed by Mr F. H. Brightman, seconded by Dr C. P.
Petch, and carried unanimously.
362 REPORT
ELECTION OF HONORARY MEMBERS
Council had nominated Professor D. H. Valentine and Mr E. D. Wiggins, and the President was
delighted to propose their nomination from the chair. This was carried with warm applause.
RE-ELECTION OF HONORARY AUDITORS
The re-election of Grant-Thornton (formerly Thornton-Baker) as Honorary Auditors was proposed
by Mr D. J. McCosh, seconded by Mr D. H. Kent, and carried unanimously with the Society’s
gratitude.
. ANY OTHER BUSINESS
Mr Walpole, commenting on this significant occasion of the Society’s Sesquicentenary, thanked the
President for conducting the Proceedings of the day, and for his Presidential Address, which had
given opportunity for the Society to take stock at the Sesquicentenary, and also for writing his
excellent book The Botanists — the history of the Botanical Society of the British Isles. The Society was
deeply grateful to him for all the hard work and research for this.
There being no further business, the meeting closed at 12.50.
M. Briccs
AFTERNOON SESSION
During the afternoon following the A.G.M., short discussion papers were presented by the
Chairman and Secretary of each of the four Permanent Working Committees. 30 minutes was
allotted to each committee, in which a summary was presented of current activities, followed by a
discussion with comment and suggestions invited from the floor.
Dr N. K. B. Robson, Mrs A. Lee and R. Smith (Field Meetings Secretary) presented the activities
of the Meetings Committee. They put forward the proposal of a symposium-type meeting, with
papers, on the day following the Annual Exhibition Meeting. It was noted that field meetings had
increased from 27 in 1982 to 35 in 1986, but the number was still possibly insufficient for the demand.
The Society’s publications were reviewed by M. Walpole and A. O. Chater (Publications
Committee), with emphasis on the journals, notably the contents of Watsonia vis-a-vis B.S.B.1.
News. The possibility of publishing more on intraspecific variants and less on aliens in B.S. B.I. News
was discussed.
For the Records Committee, D. E. Allen introduced D. A. Wells, who outlined plans for the new
mapping scheme scheduled for 1987-88. There was discussion of this, and on the many demands on
vice-county recorders — which should be monitored to prevent overloading recorders with work.
Mr Allen remained in the chair for the presentation of the work of the Conservation Committee,
and Miss L. Farrell outlined the more important issues. A number of threatened species were under
consideration for the Quinquennial Review of plants scheduled in the Wildlife and Countryside Act
of 1981. Some useful information on these was provided by members present, and it was agreed that
a note in B.S.B.I. News requesting specific information would help with the problem plants, as each
member’s knowledge could contribute and be of value.
CONVERSAZIONE
In the evening, after the formal business of the day had been completed, 55 members and guests
attended a Conversazione in the Library at the Linnean Society’s rooms. We were pleased to
welcome representatives from the following Societies: the Linnean Society of London, British
Bryological Society, British Lichen Society, British Mycological Society, British Pteridological
REPORT 363
Society, Wild Flower Society, and the Royal Society for Nature Conservation. Mr Allen welcomed
five other presidents and other representatives of these Societies, and thanked the Linnean Society
for the use of their premises and Miss E. Young for her outstanding flower arrangements. The
President of the Linnean Society, Professor W. G. Chaloner F.R.S., served the party with wine
and light refreshments.
M. BriGGs
EXCURSION HELD IN CONJUNCTION WITH ANNUAL GENERAL MEETING
SHEFFIELD PARK AND BLUEBELL RAILWAY. 11TH MAY 1986
A small but enthusiastic party of some 40 B.S.B.I. members and guests gathered at 11.00 hrs at
Sheffield Park, near Haywards Heath, in Sussex. The party was given a guided tour of the
extensive woodland gardens, which support a varied ‘wild’ flora, notably good quantities of
Ophioglossum vulgatum, which is being encouraged by the gardens’ staff. The fine collection of
trees and shrubs was admired, as were various aliens such as Claytonia sibirica, nicely in flower.
After a picnic lunch, the group assembled at the nearby terminus of the Bluebell Railway,
thence embarking on the superb steam train. As the train made its leisurely progress across the
green, spring countryside, we were able to enjoy a varied display of wayside and woodland flowers.
The afternoon was a celebration of the most traditional aspects of our subject, with the show of
spring flowers seen in the context of a lively ‘charabanc’ type of outing in good company and, for
many of us, an excellent cream tea served in the traditional dining-car. At the other terminus there
was an opportunity to stretch legs and to examine the weeds in the station flower-beds. Segregates
of Veronica hederifolia provoked lively discussion. Alas, the Bluebells were late after a cold spring
and were far from their best.
Several of the party had come in period costume, and at the end of the ride, back at Sheffield
Park, the assorted clergymen, ladies and gentlemen, and a ruffian with a cudgel, posed for
photographs. Chris Preston’s clergyman role was a great success, but nearly led to his downfall a
few minutes later when the car in which he was travelling stopped to ask the way from a real
gentleman of the cloth! As usual, a smal! band of diehards set out afterwards to visit nearby
botanical sites: they were rewarded by numerous tiny plants of Montia fontana subsp.
chondrosperma on a damp corner of a village green, but in general the late season was against
them. It was a most enjoyable day!
J. R. AKEROYD
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INSTRUCTIONS TO CONTRIBUTORS
Papers and Short Notes concerning the systematics and distribution of British and European
vascular plants as well as topics of a more general character are invited.
Manuscripts must be submitted in duplicate, typewritten on one side of the paper only, with wide
margins and double-spaced throughout. They should follow recent issues of Watsonia in all matters
of format, including abstracts, headings, tables, keys, figures, references and appendices. Note
particularly use of capitals and italics. Only underline where italics are required.
Tables, appendices and captions to figures should be typed on separate sheets and attached at the
end of the manuscript. Names of periodicals in the references should be abbreviated as in the World
list of scientific periodicals, and herbaria as in Kent & Allen’s British and Irish herbaria. Line
drawings should be in Indian ink on good quality white card, blue-lined graph paper or tracing paper.
They should be drawn at least twice the final size and they will normally occupy the full width of the
page. Lettering should be done in Letraset or by high-quality stencilling, though graph axes and
other more extensive labelling are best done in pencil and left to the printer. Photographs can be
accepted only in exceptional cases.
Contributors are strongly advised to consult the editors before submission in any cases of doubt. 25
offprints are given free to authors of papers and Short 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 articles.
Papers and Short Notes should be sent to Dr R. J. Gornall, Botany Department, The University,
Leicester, LE] 7RH. Books for review should be sent to Dr N. K. B. Robson, Botany Department,
British Museum (Natural History), Cromwell Road, London, SW7 5BD. Plant records should be sent
to the appropriate vice-county recorders. Reports of field meetings should be sent to Dr B. S. Rushton,
Biology Department, The University of Ulster, Coleraine, Co. Londonderry, N. Ireland, BT52 1SA.
B.S.B.1. Publications
1. BRITISH FLOWERING PLANTS AND MODERN SYSTEMATIC METHODS
Ed. A. J. Wilmott, 1948. 104 pages, 18 plates. Wrappers. £4.25
2. THE STUDY OF THE DISTRIBUTION OF BRITISH PLANTS
Ed. J. E. Lousley, 1951. 128 pages, illustrations and maps. £4.25
a SPECIES STUDIES IN THE BRITISH FLORA
Ed. J. E. Lousley, 1955. 189 pages, 2 plates and 23 text figs. £5.25
ye EROGRESS IN THE STUDY OF THE BRITISH FLORA
Ed. J. E. Lousley, 1957, 128 pages, 4 plates and 9 text figs. £5.25
8. THE CONSERVATION OF THE BRITISH FLORA
Ed. E. Milne-Redhead, 1963. 90 pages. £5.25
11. FLORA OF A CHANGING BRITAIN (Reprint, 1973)
Ed. F. H. Perring, 1970. 158 pages, 21 text figs. Paperback. £3.50
ISZ-sPEANTS WILD AND ‘CULTIVATED
Ed. P. S. Green, 1973. 232 pages, 8 plates and 24 text figs. £3.20
14. THE OAK: ITS HISTORY AND NATURAL HISTORY
Eds. M. G. Morris and F. H. Perring, 1974. 376 pages and 8 plates. £8.25
15. EUROPEAN FLORISTIC AND TAXONOMIC STUDIES
Ed. S. M. Walters, with the assistance of C. J. King, 1975. 144 pages and 4 plates. £3.80
iG) THE. POLLINATION OF FLOWERS BY INSECTS
Ed. A. J. Richards, 1978. 213 pages and 31 plates. £40.00
17. THE BIOLOGICAL ASPECTS OF RARE PLANT CONSERVATION
Ed. H. Synge, 1981. 586 pages and numerous text figs. £42.00
is /ELANT LORE STUDIES
Ed. R. Vickery, 1984. 260 pages. Proceedings of a joint conference of BSBI and Folklore Society in
1983. £7550
iS ARCHAEOEOGY AND THE FLORA OF THE BRITISH ISLES
Ed. Martin Jones, 1986. Proceedings of a joint conference held in 1984.
SPECIAL OFFER: 13 and 15 may be bought together for £3.50
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