BotnnucilJoumal ofthr Lznnean Sone& (1997), 124: 1-120. \Vlth 33 figures
The geographical relationships of British and
Irish vascular plants
CHRISTOPHER D. PRESTON AND MARK 0. HILL
Institute of irevestrial Ecolog, Monks Wood, Abbots Ripton, Huntingdon, Cambs. PEI 7 213
Recezved 3 4 1996; accepted f o r publication October 1.996
Classifications of British and Irish vascular plants into floristic elements are reviewed. Only
H.C. Watson and J.R. Matthews have attempted to devise a more or less comprehensive
classification, based on the British range of the species (Watson) or the European distribution
(Matthews). A new classification of 1481 native species is presented, based on their range in
the Northern Hemisphere. Species are classified by their occurrence in one or more major
biomes (Arctic, Boreal, Temperate, Southern) and their longitudinal distribution (Oceanic,
Suboceanic, European, Eurosiberian, Eurasian, Circumpolar). The distribution of species in
the floristic elements is illustrated by coincidence maps for the British Isles and Europe. The
British and Irish flora is dominated by Boreo-tempcrate, Temperate and Southern-temperate
species, with the Temperate species being the most numerous. Species with continental
distributions (Le. species which are rarer than expccted in western Europe) are listed; most
of these are in the Boreo-temperate and Temperate elements. The floristic elements are
discussed in relation to the life-form spectra, habitat preferences and altitudinal limits of the
component species, and analysed in terms of Ellenberg indicator values for temperature and
continentality. The new classification is compared with that of Matthews. An additional 48
species which are endemic to the British Isles are listed. The scope for extending this method
of classification to other organisms and for adapting it for use outside the British Isles is
discussed.
0 1997 Thc Linnean Society of London
ADDITIONAL KEY WORDS:-Atlantic zone - biodiversity - biogeography - biome
chorology continentality endemic - floristic elements - indicator values - - temperate.
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CONTENTS
Introduction . . . . . . . . . . . . . . . . . . . . . .
Historical review . . . . . . . . . . . . . . . . . . . .
The distribution of plants within Britain . . . . . . . . . .
The distribution of plants within Ireland . . . . . . . . . .
The wider distribution of British arid Irish plants . . . . . . .
Conclusions . . . . . . . . . . . . . . . . . . . .
A new classification of British and Irish vascular plants into floristic elements
Taxonomic scope . . . . . . . . . . . . . . . . .
Geographical data . . . . . . . . . . . . . . . . .
Choice of method . . . . . . . . . . . . . . . . .
Outline of the new classification . . . . . . . . . . . . .
0024-4074/97/050001+ 120 $25.00/0/ht960084
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Floristic elements in Britain and Ireland . . . . . . .
The Arctic-montane elements (1 3-1 6) . . . . . .
The Boreo-arctic Montane elements (2 1-26) . , , .
The Wide-boreal elements ( 3 6 3 6 ) . . . . . . .
The Boreal-montane elements (4 1-46) . . . . . .
The Boreo-temperate elements (51 --56) . . . . .
The Wide-temperate elements (61-66) . . . . . .
The Temperate elements (7 1-76) . . . . . . .
The Southern-temperate elements (81-86) . . . .
The Mediterranean elements (91-93) . . . . . .
Species with a continental distribution . . . . . .
Ecological characteristics of the floristic elements . . . .
Life-form spectra . . . . . . . . . . . . . .
Habitat preferences . . . . . . . . . . . .
Altitudinal limits . . . . . . . . . . . . . .
Comparison with other classifications . . . . . . . .
Ellenbcrg’s indicator values . . . . . . . . .
Matthews’ elements . . . . . . . . . . . .
Young’s classification of Arctic species . . . . . .
Dupont’s classification of Atlantic species . . . . .
Numerical analyses . . . . . . . . . . . .
Classification of other species . . . . . . . . . .
Endemic species . . . . . . . . . . . . . .
Species confined to the Channel Islands . . . . .
Discussion . . . . . . . . . . . . . . . . .
Methodology . . . . . . . . . . . . . . .
Geographical relationships of the British and Irish flora
Further applications of the classification . . . . .
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Acknowledgements
References . . . . . . . . . . . . . . . . .
Appendix 1. List of species in each floristic element . . .
Appendix 2. Alphabetical list of species. . . . . . . .
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107
INTRODUCTION
The present geographical distributions of plants are the result of climate, habitat
availability and dispersal history. Other factors such as competition, diseases and
pests are vitally important to plant survival, but, insofar as they affect distributions,
they normally reflect differences in habitat and climate. It is generally accepted that,
at the largest scales, the limits of distribution are determined mainly by climate and
dispersal history (Cain, 1944; Birks, 1987). Habitat is less important, because suitable
habitats are normally available within the broad envelope of a species’ range.
Within Europe, there are three main climatic factors that determine plant
distributions, namely summer warmth, winter cold and summer drought (Hintikka,
1963; Pigott, 1989; Prentice et al., 1992; Pigott & Pigott, 1993). The distribution of
most species is correlated with one or more of these climatic variables (Iversen,
1944; Dahl, 1951; Conolly & Dahl, 1970; Willis, 1985) and the effects of climate
have been demonstrated experimentally in some cases (e.g. Pigott, 1968; Woodward
& Pigott, 1975; Woodward, 1975). Pleistocene history has also had an enormous
influence, resulting in a much smaller, essentially Arctic, flora above the climatic
treeline in northern Europe than that found in the Alps or in mountains to the
south (Ozenda, 1994; Korner, 1995; Walker, 1995). In addition, geographical
isolation of populations has resulted in the evolution of vicarious species such as
Salix lapponum (widespread in northern Europe) and S. heluetica (mainly in the Alps)
kLORISTIC ELEhI62 15 I N R R I T V h AND IREIAKD
3
and the European and American beeches (Fagus glvatica and E grandzjilia; cf-.
Huntley, Bartlein & Prentice, 1989). These have similar climatic tolerances but nonoverlapping geographical ranges.
The purpose of classifying plant distrihutions is to establish categories that reflect
similarities of range (McLaughlin, 1994). Few, if any, plant species have geographical
distributions which are identical at all scales. For phytogeographical analysis it is
necessary to generalize by grouping species together into jloristic dements. A floristic
element contains a group of species which have similar geoqaphical distributions
at the present day in a specified area. Floristic elenients are “convenient approximations or reference points that describe easily detectable clusters within
the distributional continuum, and should not be regarded as representing real
discontinuities” (Birks, 1973). The definition of a floristic element adopted hcre is a
strictly geographical one. Many authors have used this (or analogous terms) to
describe species which not only have similar distributions at the present time but
are also assumed to have a common distributional history, migrating together in
response to changing environmental conditions (Cain, 1947).
Given these facts, a natural and gencmlizahle approach to the categorization o f
plant distributions is to specify them in two parts, the first corresponding to climatic
tolerance and the second reflecting dispersal history, i.e. defining the part of the
area with potentially suitable climate t h a t is actually occupied.
Floristic elements can be defined in rclation to any specified area, hut the most
useful categories for general purposes arc those based on large areas. Knowlcdgc of
the wider geographical distribution of specks may pro\4dc insights into their
distribution and ecology which cannot he obtained from investigations which are
limited to a particular study area. This is true for study areas which are as small as
the Welsh county of Radnorshire (M’oods, 1993) or the Isles of Scilly in south-wcst
England (Lousley, 1971) or as large as ‘I’urkcy (Davis, 1965-85) or the Middle East
(Zohary, 1973). The wider geographical range of species is particularly r t h m t
when considering the classification of thc British and Irish plants, which form part
of the vast Holarctic floristic kingdom which covers Europe, North Africa, Asia
outwith the tropics and most of North iimerica (Takhtajan, 1986). The Holarctic
Kingdom may be divided into subkingdoms and regions, and the whole of the
British Isles falls into the largest of all floristic regions. the Circumhoreal, which
includes Europe (expert for the Meditrrrancan region), Siberia, Kamchatka, most
of Alaska and a large part of Canada.
Matthews (1937, 1955) classified the flora of the British Isles into ‘geograyliical
elements’ which were based primarily on the distribution of the species in Europe.
Matthews’ classification is still widely usrd, sometimes with modifications, lo describe
the composition of the flora of particular areas (e.g. Wynne, 1993; Trueman, hlorton
& Wainwright, 1995) and of defined plant communities (Rodwell, 1991-95), or to
analyse the distribution of plants within tlie British Isles (e.g. Pcrring, 1996).HOWC\W,
knowledge of the taxonomic relationships and tlie wider distribution of the flora of
Britain and Ireland is much greater now than it was when Matthews wrote, and
recent authors have drawn attention t o the need to revise his work (Perriyg, 1985;
Graham, 1988). In this paper we revirw the previous studies of the geographical
relationships of the British and Irish flora, including the classification proposed hy
Matthews (1937, 1955) and the modifications suggested by later authors. \#Ye then
outline a new classification of the vascular plants of the British Isles into floristic
elements.
C. D. PRESTON AND M. 0. HILL
4
TABLE
1. The ‘types of distribution’ of British plants defined by H.C. Watson (1835, 1847-59, 1868)
on the basis of the distribution of the species within Britain. The later accounts (Watson, 1847-59,
1868-70) are followed where the treatments differ
Twe
Distribution
British
Highland
Scottish
Intermediate
English
Germanic
Atlantic
Local or
Doubtful
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Examples of species included
Widespread in both England and
Scotland
Mountains, especially in Scotland
Confined to Scotland, or more
prevalent there than elsewhere
N. England and S. Scotland
Confined to England, or more
prevalent there than elsewhere
S.E or E. England
S.W. or W. Britain
Few localities, not conforming to
above elements
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Percentage of total
flora in element*
Alnus glutinosa, Lotus cornirulatuc
3 7 ‘In
Gentiana niualic, Saliv hebacea
Mertensia maritima, Trientalis eumpaea
8%
6%
Primula farinosa, Saxijraga liirculus
Ceterach oficinarum, limut communis
3%
29’/0
Puhatilla uulgariJ, Silent conica
Sedum anglicum, Sibthorpia twopapa
h a h a nizoides, Eriocaulou aquatirum
9%
59 0
3 ?6
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* Percentages calculated from Watson’s (1847-59,
4 509) lists; the species listed as intermediate between two of
the above elements are included in the element which Ll’atson considered they most closely resembled.
HISTORICAL REVIEW
‘The distribution ofplants within Britain
Hewett Cottrell Watson (1 8 0 6 1 8 8 1) devoted the major part of his botanical
career to the careful and critical compilation, presentation and analysis of the
distribution of plants in Britain (Egerton, 1979; Allen, 1986); his work did not cover
Ireland. He first presented a detailed classification of the distribution of species
within Britain in his book Remarks on the geographical distribution ofBritish plants (Watson,
1835), subsequently modifying it slightly in Cybele Britunnica (Watson, 1847-59).
Watson recognized eight ‘types of distribution’ in his modified classification (Table
1). These types were floristic elements in the narrow sense of the term; he did not
regard them as linked by a common migrational history and he specifically stated
that the Germanic type, for example, “is not applied with reference to any supposed
origin from Germany, but simply as indicating the tendency of the species to a
distribution connected with those provinces of England which are bounded by the
German or North Sea eastward” (Watson, 1847-59, 1: 50). In discussing the types,
Watson (1847-59, 1: 54) stated that “no decided lines of separation can be drawn
between them. They may be said to pass gradually into each other; because the
distribution of some species is of such an intermediate character as to render the
choice of type to express it either dubious or optional”. Nevertheless, he was able
to assign nearly two-thirds of the British flora to his primary types; the remaining
species were listed as intermediate between two of these types (Watson, 1847-59).
Watson’s ‘types’ became the standard floristic elements used by 19th century
British botanists (e.g. Trimen & Dyer, 1869; Briggs, 1880; Baker, 1885), despite the
fact that his system took no account of the distribution of British species in Europe.
Moss (1914) commented that “it would be easy to criticize adversely this scheme of
classification, and it is curious that the scheme met with such widespread approval”.
This approval presumably reflected the fact that Watson’s classification was the only
one available for practical use, as it was the only one which assigned all the British
FLORISTIC ELEMENTS IN BRITAIN AND IREIAND
5
TABLE
2. The ‘types of distribution’ of Irish plants defined by R.L. Praeger (1902) on the basis of the
distribution of the species within Ireland
~
,r)ve
Distribution
Examples of species included
General
( I ) throughout Ireland
(2) widely scattered, but with no d r a r pattern
(3) common maritime plants
Central
XIarginal
Ultonian*
hlumonian*
Lagenian*
Connacian*
Central plain
Confined to margin, absent from crntral plain
Northern
Southern
Eastern
\Yestern
BtIlis peremis
LStar/y bdonica
Obrlilearia ofirinali,\
Andromeda polfoha, Stdlnna palrr~tnr
Hvpencurn eloda. Lohrltn doltinannu
Cicitln r’imJn, Saarlfrnga oppo iztlfilin
Blarkstonza perjolintn, Nnguii ula gaiidlflorn
Srilla wmn, Tnjolium glom~rntum
Adiantum rapillu-ornm\, Dabomn inntnhnin
* These names are derived from the province$ if Ircland (lilstcr, Xlunster,
Lcinster and C:otitiaiight1
species to a specific category. Its exclusively British basis was not a significant
disadvantage to those who simply wanted to place the flora of small areas (such as
counties) in the context of Britain as a whole. Moss (1914) followed his critical
comments by a ‘modern classification’ which simply stated that the British flora
could be placed in three ‘main groups’ (an Arctic-Alpine group, a Western group
and an Eastern group); this did not provide a viable alternative to Watson’s scheme.
Watson presented data on the European and wider range of British species in
three of his books (Watson, 1832, 183.5, 1868-70). In the Remarks (1835) he divided
Europe into Polar, Arctic, Boreal, Temperate and Mediterranean zones, and listed
the range of each species in these terms (e.g. Bor.-Med., Tem-Med.). Similar data
were provided separately for the distributional range of British species in North
America. Watson also indicated the longitudinal range of the species by listing the
occurrence in nine belts, ranging from western Europe through Asia to eastern
North America. These data could have formed the basis of a classification of the
British flora into elements based on their wider distribution. Howejrer, Watson never
made use of them in this way, even though he was well aware that “in limiting our
attention to the island of Britain exclusively, some of the species must be referred
to types of distribution, under which they would not be placed if our views took in
a larger geographical space” (Watson, 1847-59, I : 55; cf. Burkhardt & Smith, 1991 :
53).
The distribution of plants within Ireland
H.C. Watson’s detailed records of the distribution of plants in Great Britain were
complemented in Ireland by the work of Colgan & Scully ( 1 898) and Praeger (1 90 1).
Colgan & Scully (1 898) used Watson’s floristic elements, based on the distribution
of species in Britain, to compare the flora of England and Wales with that of Ireland.
They demonstrated that 98% of the members of the most widespread element in
England and Wales, the British type, also occurred in Ireland, whereas only 12%
of the species in the most easterly element, the Germanic type, did so.
Praeger (1902) also discussed, and mapped, the distribution in Ireland of the
species in Watson’s types. In addition, he classified species on the basis of their
distribution in the 40 vice-counties of Ircland, recognizing seven elements (Table
C . D. PKESI’ON AND M. 0. HILL
6
TABLE
3. The ‘floras’ defined by E. Forbes (1846b) on the basis of the distribution of the species in
the British Isles and Europe
Flora
Distribution in Britain and Europe
Examples of species included*
West Irish,
or Asturian
S.W. or W.Ireland; nearest European sites in
N. Spain
Dabon ia r.ontnbnra, Enro markaiana
Devon, or
Norman
S.W. England. S.E. Ireland, Channel Islands;
S. Europe
Ial11~hispiduu“,Rubia peregrinu
Kentish,
Chalk of S.E. England; opposite coast of France
h u l a lury<ar, P&ma
Alpine, or
Scandinavian
hlounrains, especially in Scotland; Scandinavia
Centinria n m h , Saxijoga ceinuo
Germanic, or
Central European
Widespread in Britain or with variously reytricted
distributions; western and ccntral Europe
Brlh permiis, dldampjtum cnsiotum
orhiculan
or north French
~
* There are numerous
~
~
~~~
errors in the examples citrd by Forbcs (l846h): spccies which conform to his definitions
have been selrcted here.
2). Five of these, groups of widespread, northern, southern, western and eastern
species, were analogous to Watson’s types. The remaining elements, one group of
species occurring in the calcareous central plain of Ireland and the other found
around the predominantly non-calcareous and often mountainous edge of the island,
had no analogues amongst Watson’s types in Britain.
nze wider distribution of
British and Irish plants
Forbes’ ‘bnlliant memozr’, 1846
Edward Forbes (1815-1854) was a polymath who made major contributions to
the study of the British and European flora and fauna in his short career (Preece &
Killeen, 1995). His major biogeographical work was initially delivered as a lecture
to the British Association in Cambridge in 1845 (Forbes, 1845a, b, c, 1846a), and
later elaborated as a major paper (Forbes, 1846b).
Forbes was writing at a time when the widespread glaciation of northern Europe
had only recently been appreciated: the major statement of the glacial theory,
Agassiz’s Etuder sur les glaczers, was published in 1840 (Rudwick, 1969). Forbes
attempted to account for the origin of the flora and fauna of the British Isles in the
light of these newly discovered geological changes. He assumed that all species had
a single centrr of origin (and had not, for example, been created in more than one
area) and that they had reached their current distributional range by migration o\rer
land from that centre. A disjunct distribution was the result of the fragmentation of
a previously contiguous range. With certain limited exceptions, Forbes did not
accept the possibility of long-distance dispersal.
In order to account for the colonization of the British Isles, Forbes (1846b) divided
its vascular plants into “five well-marked floras” (Table 3). The floras were defined
on the basis of the distribution of the species in the British Isles and Europe, and
some examples of species belonging to them were provided. The “great mass of the
flora and fauna ofthe British Islt-s” was placed in the “Germanic or central European
type”, and Forbes held that these migrated from Central Europe after the glacial
FLORISlIC ELEME‘C 15 IN UKI IAIN AND IRCL-WI)
7
period but before the British Isles became isolated from mainland Europe; some
species reached Britain but failed to reach Ireland before it was cut off by the sea.
The “Alpine flora, or Scandinavian type” was widespread at the time of the glacial
period but subsequently retreated northwards or on to mountain summits. The
“west Irish flora, or Asturian type” was considered to have reached these islands
before the glacial period by means of a former land mass extending from the west
of Ireland to the north of Spain, and westwards to incorporate the hlacaronesian
islands. Most of the members of this element had been exterminated from the British
Isles during the glacial period; only a few of the hardiest species had survived in
S.W. Ireland. Forbes was less certain about the history of the “Kentish flora, or the
north French type” and the “Devon flora, or Norman type”; he suggested that these
species had also arrived in pre-glacial times but accepted that they might have
arrived at the same period as the Germanic flora.
Forbes’ paper forms a strilung contrast with the work of Watson in the lack of
detailed distributional data and the sweeping hypothcses he proposed to account
for the origin of the British flora (and fauna). His work was severely criticized by
the notoriously cantankerous Watson (1 847-59, 1: 47 l), who considered that “it
absolutely teems with errors in its botany-inconclusive arguments, inconsequent
logic, inept illustrations, and the guesswork of the imagination put forth ostensibly
as the ascertained facts of science”. Nevertheless, it proved to be a stimulating
contribution to plant geography and contemporaries such as Charles Lyell, Charles
Darwin and J.D. Hooker were able to assent to at least some of his conclusions (cf.
Burkhardt & Smith, 1987, 1989). Forhes’ lasting achielrement lay in his explanation
of the distribution of Arctic-Alpine plants, which is now supported by a mass of
palaeontological evidence (Godwin, 1975). Darwin had independently arri\.ed at the
same explanation before Forbes, and nwer ceased to regret that he had neglectcd
to publish the hypothesis himself (Barlow, 1958: 124--5).He cited Forbes’ explanation
at some length in the Origin ofspecies (Darwin, 1859). Even Forbes’ “Atlantis theory”
of the origin of the species confined in the British Isles to S.W. Ireland received
support from many biologists (although none from Darwin). Holvever, in this case
the geological evidence has not providcd any support for his theory, and the history
of these species in our islands has not yet been completely resolved.
A renexial o f interest, 1911-1 6
Interest in the wider distribution of the British and Irish flora revived in the early
20th century, when floristic elements wcre again discussed in the context of the perglacial survival and post-glacial immigration of the flora. In 191 I Clement Reid
introduced a discussion at a meeting of the British Association attended by the
members of the International Phytogeographical Excursion, who were then visiting
Britain (Reid, 191 1; British Association for the Advancement of Science, 19 12).
Reid pointed out that fossil evidence acquired since Forbes’(1846b) memoir provided
“brilliant proof’ of his explanation of the distribution of the Arctic-Alpinr specirs.
However, Reid contended that during the glacial period “any survival of our
flowering plants, except in the case of a few arctic and alpine species, was quite
impossible” as the pre-glacial flora would have been “swept away almost as completely
and effectually as the celebrated volcanic eruption wiped out the plants of Krakatoa”.
He contended that many species (including the species in the Atlantic element,
which were separated from continental populations by the sea, and species of
8
C. D. PRESTON AND M. 0.
HILL
calcareous and aquatic habitats, which were surrounded by dissimilar terrain) had
reached Britain by “chance introductions of seeds during thousands of years” which
were “now mainly due to birds” although before Britain became isolated “herds of
migrating bison, deer, and horse have played their part”.
Reid’s views were challenged, in whole or part, in the discussion following his
paper, and subsequently by Drude ( 1912), Scharf€ (1 9 12) and Stapf (19 14, 1917).
Stapf (1914) emphasized that it was essential to treat the British and Irish flora “as
a section of the flora of Western Europe”, as in Forbes’ (184613) “brilliant memoir”,
rather than as a “detached unit”. He provided a detailed analysis of the “southern
element” in which he included 150-1 60 species of southern distribution which were
absent from Central Europe. He demonstrated that the Atlantic species could be
separated from the Mediterranean-Atlantic element. The Atlantic species predominated over the Mediterranean in Ireland, whereas in southern England the
Mediterranean species tended to be more frequent. The species of wet habitats
tended to have an Atlantic distribution whereas those of light, well-drained soil had
Mediterranean affinities. Stapf (19 14) accepted that the species in the southern
element had not persisted through the glacial period in our area, but rejected the
view that they had arrived “singly and independently” by chance dispersal.
Studies stimulated by the ‘age and area’ hypothesZr, 1923-26
J.R. Matthews (1923, 1924, 1926) examined the relationship of the flora of the
British Isles to that of the European mainland by plotting the distribution of species
in the vice-counties of Britain and Ireland and in the neighbouring countries of
Europe. He showed, for example, that a group of 266 ‘English species’ which were
confined (within the British Isles) to England and Wales were concentrated in southeast England; in mainland Europe, their greatest concentration was in France. He
interpreted these results in terms of the ‘age and area’ hypothesis of J.C. Willis
(1922). Willis suggested that in an area where there were no major barriers to
migration, the largest areas were occupied by the oldest species. Matthews (1923)
concluded that the English species were a recent flora which had migrated from
France; particular concentrations revealed the migration routes they had followed.
The flaws in Willis’ hypothesis were soon detected: Wilmott (1930), for example,
pointed out that “species may have small areas at birth, but they have equally small
areas at death” and that the concentration of species observed by Matthews (1923)
in south-east England “could equally be due to climatic and geologic limiting
factors”. Salisbury (1932) made the same point, accepting that the theory that the
English species had migrated into south-east England was “not in itself improbable”
but pointing out that their presence “is adequately accounted for by a peculiarly
favourable combination of soil and climate”. However, Matthews’ interest in the
broader distribution of the British and Irish flora had been aroused, and it was to
lead to more significant publications.
Floristic elements dg5ned tg~E.J. Salisbuy & J.R. Matthews, 1932-55
In his presidential address to the Norfolk & Nonvich Naturalists’ Society, E.J.
Salisbury followed the example of Moss (1 9 14) and Stapf (19 17) in criticizing H.C.
Watson’s types of plant distribution, which “from their purely insular bias, are liable
to obscure the true distributional affinities” (Salisbury, 1932). Unlike earlier critics,
Salisbury went on to classify the flora of the British Isles into nine components,
9
FLORISTIC ELEMENTS IN BRITAIN AND IRELAND
TABLE
4. The components and elements of the flora of the British Isles recognised by Salisbury ( 1 932),
with their equivalents in Matthews’ (1937, 1955) classification. For the definition of Matthews’ elements,
ser Table 5
Component
(a) Alpine
(b) Northern
(c) Southern
(d) Oceanic
Element
(1) Arctic
(2) Northern
(3) Continental-Northern
(4) Southern
(5) Mediterranean
(6) Continental-Southern
(7) Western
(8) Southern Oceanic
~
(e) Continental
(f) Western Central’
(9) Endemic
(h) Rccent Immigrants
(i) Generally distributed
(9) Steppe
(10) Continental
(1 I ) Northern Continental
hlatthews (1937, 1955)
Alpine
Arctic-Suharctic and Arctic-Alpine
(:otitincntal-Northern and Northcrn-Montane
Continental Northern
Continental Southern
hlediterranean
Continental Southern
Oceanic West European
Oceanic Southern
Oceanic Northern
Chntinental
Continental
Continental Northern
Endemic
European, Eurasian and lorthern
misphere (or Wide).
* Described by Salisbury as an ill-defined group of a frw species with distributions intermediate hetween oceanic
and continental, e.g. Apium inundatum. Cirsium tubemum.
“based upon the areas in which they characteristically occur on the Continent”.
Some of these components were subdivided to gwe a total of 16 geographical
elements (Table 4).Although these elements were devised for Britain and Ireland,
Salisbury considered only the East Anglian representatives in his paper. He discussed
the elements present in East Anglia in detail, with the sole exception of the “species
of general distribution”, which he dismissed with the comments that they are “too
numerous to cite individually” and “have little significance for our present purpose
of comparative study of geographical distribution”. Salisbury was concerned with
the historical aspects of plant geography, but he was even more interested in the
ecological factors limiting the distribution of the members of the floristic elements
which he recognized.
The floristic elements devised by Salisbury (1932) were modified and extended
to cover all the flowering plants in Britain and Ireland by Matthews, initially in his
presidential address to the British Ecological Society (Matthews, 1937, 1955).
Matthews (1937) identified 15 elements and provided a detailed treatment of 1 1 of
them, listing the component members and mapping their distribution in vice-counties
(Table 5). The 11 elements treated in detail included 45% of the British flora,
including the “species which show some noteworthy feature in their geographical
connexions”. In 1955 he separated the North American element from the Oceanic
Northern element and listed the members of this. He did not list the remaining
55% of species which were members of the small endemic element or the large
European, Eurasian and Northern Hemisphere (or Wide) elements. Unlike Salisbury
(1 932), Matthews did not cover pteridophytes. Matthews’ elements are considered
later in this paper in relation to the elements we recognize.
Matthews (1955: 121) considered that since the flora of the British Isles “is
essentially immigrant and apparently derived from different parts of Europe it should
be possible to analyse its several components and indicate the probable paths of
C. D. PRESTON AND hl. 0. HILL
10
TABLE
5. The geographical elements in the flora of the British Isles defined by Matthews (1937, 1955)
primarily on the basis of the European distribution of the species. Matthews (1955) is followed where
the two treatments differ
Element
Distribution
Wide
Eurasian
European
N. hemisphere
Europe and Asia
Europe
Mediterranean
Mediterranean region, extending
north through LV. France
S. and W. Europe, including
Mediterrancari region
W. Europe
Oceanic Southern
Oceanic Wect
European
Continental
Southern
Continental
Continental
Northern
NorthernMontane
Oceanic Northern
North American
Arctic-Subarctic
Arctic-Alpine
Alpine
Endemic
Examples of species included
Percentage of total
flora in element
t Carm ro rtmta, Potarnn@on
natan i
tcvmnadenta conopsea, Pruntu pndus
t Cratagw hi’zgula, Potenhllfl
neumannzana
Aibutus unedo, Neotziiua mnculnta
13%
3 1O h
9 ‘In
Ilex aqutJblium, Ruhta perepna
5%
Erica ciliaris, Genista anglica
6%
S. and Central Europe
Hippocrepi.i mmnsa, Orc his simta
8 Yo
Central Europe, extending into Asia
Central and h-.Europe; montane in
south*
N. Europe and rnouiitains of C . or
S. Europe*
N.W” Europe
Main centre of distribution in
America
Northern Europe, especially
Scandinavia
Arctic regions, and on mountains
further south*
Mountains of Europe. absent from
N. Europe and arctic regions
Endemic
Carpinus betulu, T4mnicn Jpkatn
Carex limoJa, Andromeda poliJolia
6 ‘10
6%
Linnaea boreali5, .Vzcphnr puntiin
2%
Lubelia dnrtmanna, h&i’ra galr
Erioraulnn aquafirum, .Fpiranthes
rnmanzofiana
Diapemia lapponica, Sazfiuga
rioularit,
D y a s octopetala, Samfiaga
oppositifolia
Centiana orma, J4iiiuartia .iedoides
#Fumaria accidentalis, Limontum
3%
2Yo
0.5%
2oh
5%
0.5%
1n/‘
rerunium
* hlany species circumpolar
or circumhoreal.
t Matthews (1937, 1955) did not list the species in these elements; we have selected these examples.
#Matthews (1937, 1955) provided no examples; these are taken from Wilniott (1930), which he cites.
invasion”. In the following year Godwin’s The histoy ofthe Britishjora was published,
with its significant subtitle A factual basis for jhytogeography (Godwin, 1956). Godwin
(1975) later explained that he wished to provide facts to replace “the historical
speculation, up to that time prevalent, based on nothing more than comparison of
present-day distributional ranges”. Godwin’s book effectively ended the tradition,
begun by Forbes (1846b), of defining floristic elements in order to obtain an insight
into the history of the flora of the British Isles.
Revisions of Matthews’ elements, 1973-95
Few British phytogeographers attempted to modify Matthews’ elements in the
first 30 years after their initial publication in 1937. Good (1947) rearranged the
elements into seven major groups: Wide, Eurasian, European, Southern, Northern,
Continental and Arctic-Alpine. Some authors who accepted the Matthews’ elements
incorporated many species which had not been classified explicitly by Matthews
himself. Hyde & Wade (1957) and Ellis (1983), for example, allocated almost all the
plants native to Wales to Matthews’ elements.
FLORISTIC ELERiE57 S IN BRIT AIK AND IK!L\ND
II
In a phytogeographical analysis of the flora of the Isle of Skye, Birks (1973) departed
from Matthews’ scheme in two significant ways. He replaced the Mediterranean and
the three Oceanic elements by a new classification which recognized Atlantic and
Sub-Atlantic elements, the former subdivided into Southern, Northern, Widespread
and Mediterranean Atlantic elements and the latter into Southern, Northern
and Widespread Sub-Atlantic. This classification owed much to Ratcliffe’s (1968)
phytogeographical analysis of Atlantic Ixyophytes in the British Isles. Birks (1973)
also amalgamated Matthews’ European, Eurasian and Northern Hemisphere (or
Wide) elements into a single Widespread element. These categories were used 13)Goode (1974) in an analysis of the Shctland flora, and aspects of Birks’ revision
were adopted by Jermy & Crabbe (1 978) and Graham (1988). Graham’s analysis
of the flora of Co. Durham is notable as he allocated many additional species,
including numerous introduced taxa, t o hlatthews’ elements.
Perring (1 985) amalgamated Matthew’ European and Eurasian elements. on the
grounds that most species which are w-idcsprcad in Europe also extcnd into western
Asia as far as the Urals or the Caucasus. However, unlike Birks (1973),he retained
the distinction between the Eurasian and the \Vide element. Perring (1 985) also
revised the composition of the elements, adding or reallocating 168 species which
occurred in his study area, the Shropshire region. Perring’s modifications werr
followed by Woods (1 993) and Truenian et a/. (1995).
Atlantic, Subatlantic, Mediterranean-f~rlarrnticand Arctic-Alpine species in Ireland
were listed by Webb (1983), who adopted narrow definitions of thcse terms.
JVumerical anabses
Birks & Deacon (1 97 3) used techniques of numerical analysis to imrestigate whether
the phytogeographical gradients within the present flora of the British Isles were
also present in the Late-Devensian, mid-I+landrian and late-Flandrian floras. Recent
records and data from Quaternary fbssils were listed for 12 geographical regions,
and the percentage occurrence of each of the floristic elements recognized by Birks
(1 973) calculated. Non-metric multidiinensional scaling of coefficients of dissimilarity
demonstrated strong differentiation within the present flora of the British Isles, Ivith
the montane and northern elements prcdominating in the north and west and thc
continental and southern elements in the south and east. A similar analysis of fossil
records revealed only weak dflerentiatioii between regions in the Late-Devensian,
when the northern and continental clcmcnts were fairly uniformly distributed. In
the mid-Flandrian there was more evidence for a north-south gradient, and by the
late-Flandrian the phytogeographical gradient was as marked as it is in the present
flora.
Birks (1976) analysed the distribution of European pteridophytes by minimumvariance cluster analysis and principal co-ordinates analysis, using summarized data
derived from Atlas Florae Europaeae (Jalas & Suominen, 1972). He recognized 21
floristic elements, of which 10 are reprcseiited in the British Isles. Myklestad & Birks
(1993) also used summarized data from Atlas Florae Europaeae (Jalas & Suomincn,
1976) in a TWINSPAN analysis of thr distribution of European M i x species. They
identified nine floristic elements, eight of‘which were represented in thc British Isles.
The relationship of the elements identilird in these studies to those we recognize is
discussed later in this paper.
12
C. D. PRESTON AND M. 0. HILL
Conclusions
There have been only two comprehensive analyses of the distribution of British,
or British and Irish, plants. H.C. Watson provided a classification based on the
distribution of species within Britain which was widely used in the 19th century,
but fell into disfavour because it failed to take into account their wider distribution.
It was eventually replaced by Matthews' classification.
For over a century, the main motivation for the derivation of floristic elements
was the search for some evidence about the history of the British and Irish flora.
The elements tended to be defined on an ad hoc basis without a clear statement of
the principles on which they were based. There has been a general consensus that
widespread species are of little interest phytogeographically, a view which has been
held at least as strongly in recent years as previously.
A NEW CLASSIFICATION OF BRITISH AND IRISH VASCULAR PLANTS
INTO FLORISTIC ELEMENTS
Taxonomic scope
We have allocated to floristic elements all the species which are believed to be
native to the British Isles, or have occurred as natives since 1600 but are now
extinct, with the exceptions detailed below. Taxonomy and nomenclature of British
and Irish plants follow Stace (1991). In deciding on the native status of species we
have usually followed Stace (1991).
In allocating species to floristic elements we have based our classification on the
native range of the species, including all infraspecific taxa. Thus, in classifying Vu/ulpia
ciliata we have taken into account the total native distribution of subsp. ambigua
(which is native to Britain) and subsp. ciliata (which is recorded only as an introduction);
in classifying i'hehpteris palustris we have taken into account the distribution of the
Eurasian var. palustis and the American var. pubescens. We believe that this is
appropriate in view of the large scale of our analysis. We have not classified the
component subspecies separately, as these are often poorly recorded and inconsistently treated in different areas.
Many species which occur in the British Isles are very variable in their wider
range. They are often closely related to plants from elsewhere which different
authorities may recognize as allied species, or as subspecies or varieties, or may
simply include in their circumscription of the taxon present in our area. If these
segregates have vicarious distributions, the category into which the distribution of
the species is classified may depend on the taxonomy followed. Thus 7ientalis europaea
is a boreal species which if interpreted narrowly is confined to Eurasia. However,
if the concept of 7: europaea is expanded to include 7: arctica it becomes a disjunctly
circumpolar species occurring in Eurasia and western N. America, and an even
broader interpretation of the species to include the eastern North American T.
borealis gives it a completely circumboreal distribution. There are numerous other
cases in which the choice of taxonomy affects the longitudinal distribution of the
taxon. There are also cases where the latitudinal distribution is affected by the
taxonomic treatment followed, although they are less frequent. Chlysosplenium alternfolium, for example, is represented in the Arctic by a taxon (C. tetrandmm) which is
FLORISTIC ELEMEN'I'S IN BRITAIN AND IRELAND
13
sometimes treated as subspecies of this species and sometimes as separate species.
In facing these taxonomic problems we have usually adopted a broad species
concept, often following Flora Europaea (Tutin et al., 1968-80, 1993) or the works of
Hulttn (1 958, 1962, 197 1) and Hulttn & Fries (1 986). We have, therefore, included
Trientalis arctica as a subspecies of 7: europaea but treated ir: borealis as a distinct species,
the taxonomy recommended by Hulttn (1971) and H u l t h & Fries (1986). We have
included the Arctic segregate of ChgJsosplenium altern$olium within our definition of
this species, following Hulttn (1971) and Hulten & Fries (1 986) rather than Tutin
et al. (1993) who regard it as specifically distinct. It is noteworthy that in both cases
our broad species concept contrasts with that of Czerepanov (1995), illustrating the
much narrower species concepts which often prevail in eastern Europe.
It is impractical to document in this paper all the taxonomic decisions we have
made in classifying the distribution of individual species. Where the decision makes
a substantial difference to the floristic element in which a taxon is placed, our
taxonomic concept ought to be apparent from the element to which we have
assigned it. The few cases where we have adopted a narrow species concept are
identified in the species' lists (Appendices 1, 2).
Some species which are native to the British Isles have not been included in our
analysis. They fall into the following categories.
(1) The microspecies of Hieracium, Rubus and Taruxncum. The aggregate species
Hieracium murorum, Rubushticosus and Tnruxacum oficinale are allocated to elements.
(2) A few critical taxa for which there are insufficient reliable data on their world
distribution: Cystopteris dickieana, Erophiln glabre.ycens, E. majuscula, Poa humilis, Rorippa
microphylla, Utricularia ochroleuca, U. spgia and Zostera argustZfoliu. We have included
these taxa in the broadly defined species Cjstopterisfiagilis, Erophila uerna, Poa pratensis,
Rorippa nasturtium-aquaticum, Utricularia intermedia and Zostera marina.
(3) Species which are endemic to the British Isles.
(4) Species which are included in Stace (1 99 1) because they occur in the Channel
Islands but not in the British Isles proper.
Our classification is based on the native ranges of species, not the range as
modified by introductions. However, we have indicated species which are now well
naturalized outside their native range, and sometimes provided the appropriate
floristic element for the total current range.
The analysis of the number of species in each floristic element and their distribution
and life-history characteristics is based on the 1481 native species defined above.
There is some doubt about the native status of some of the species included in the
analysis; these doubtfully native plants are indicated by a section symbol (5) in the
species lists (Appendices 1, 2). We have also classified the distribution of the endemic
and Channel Island species but these are excluded from the numerical analyses.
Geographical data
Our major source of information on the geographical range of species is Hultkn
& Fries (1986), backed up by a more detailed treatment of selected species (Hultkn,
1958, 1962, 1971). Hu1ti.n & Fries (1986) provide maps of the distribution in the
northern hemisphere of species which are native to or naturalized in Scandinavia,
and these include 75% of the species which are native to the British Isles. Our task
would have been much longer and more laborious if we had not had access to this
14
C. U. PRESTON AND hl. 0. HILL
invaluable compilation. Distribution maps of additional species which occur in
Central Europe are provided by Meusel, Jager & Weinert (1965),Meusel et al. (1978)
and Meusel & Jager (1992). The European distribution maps provided by the
continuing Atlas Florae Europaeae project (Jalas & Suominen, 1972-94) are most
valuable.
Most of the taxa which are not covered by the above sources are plants which
in Europe are confined to the west and south, and which have not yet been covered
by Atlas Florae Europaeae. Data on the distribution of most western species are provided
by Dupont (1962). Data on the southern species have been taken from Flora Europaea
(Tutin et al., 1968-80, 1993), Flora of T u r k 9 (Davis, 1965-85) and the as yet unfinished
Med-Checklist (Greuter, Bur-det & Long, 198&89), as well as floras of the eastern
Mediterranean countries (e.g. Meikle, 1977-85; Strid, 1986; Strid & Kit Tan, 1991;
Turland, Chilton & Press, 1993).
For data on the distribution of species in the former U.S.S.R. we have consulted
the summaries provided by Czerepanov (1 995). For the distribution and status of
taxa in North America we have consulted the recently published Flora $North America
(Morin, 1993) for pteridophytes, and Gleason & Cronquist (1963) and Fernald (1970)
for flowering plants.
In cases of doubt or difficulty we have also consulted other sources, including
regional floras and accounts of individual species and genera, which are too numerous
to be cited individually. The sources of our information on the distribution of aquatic
plants are given by Preston & Croft (1997).
The compilation of data on the geographical ranges of species is fraught with
difficulty: erroneous records, and records of a species made years ago when taxonomic
concepts were different, tend to be perpetuated from work to work. In classifying
the wider distribution of the vascular plants of the British Isles it is only practical
to rely on secondary sources, and one has to accept the risk that some of the data
they present are erroneous. The definition of the native raiigr of taxa which have
been widely introduced is often difficult and sometimes virtually impossible: we have
followed Hulten & Fries (1!386), but it must be recognized that there is a substantial
element of doubt about the native range of many of the species which are noted as
being widely naturalized.
Choice o f nzethod
We have already noted that the major purpose of defining floristic elements is to
establish categories with similar climatic requirements and which may have at least
some similarities of dispersal history. The problem of definition can be tackled either
by classifying distributions directly or by classifying the climates that they represent.
Data sufficient to specify the macroclimate at any point in the world have recently
become available, e.g. the dataset used by Prentice et al. (1 992) to define theoretical
biomes. Thus, for any species it is in principle possible to convert geographical
ranges to climatic limits, often called envelopes. This technique has, explicitly or
implicitly, been used by plant geographers for many years. However, to be effective,
it requires that altitudinal data are available for records from mountainous districts
(Beerling, Huntley 81 Bailey, 1995). It also requires digitized maps of the distributions
of species.
Digitized world distributions have been used extensively by Lausi & Nimis (199 1)
FLORISTIC ELEhIEh 15 1U BKI I N N AUL) I K E W h L )
li
in the preparation of chorograms (coincidence maps) of the occurrence of sets of
species from the Yukon Territory. Digitized world distributions are not, however,
available for the whole British flora, although European data are now available for
species mapped for the Atlas Florae Europneae project (Jalas & Suominen, 1972-94).
It is thus not feasible at present to classify more than a small proportion of
distributions numerically, let alone to define their climatic requirements accurately.
For these reasons, attempts to define floristic elements by numerical methods
have been few (Birks, 1987; McLaughlin, 1994). Classifications based on distribution
alone, not taking explicit account of climate, have indeed been attempted for selected
species groups, using European distributions (Birks, 1976; hlyklestad & Birks, 1993;
see above), but they are somewhat provisional and have not been based on world
distributions. Attempts to define floristic elements taking account of both spatial
distribution and climate are even fewer (but see Brisse & Grandjouan, 1974).
It is doubtful, in fact, whether a numerical analysis of distributions de now would
be especially useful. Unless it took account of climate as well as of species ranges
(cf. Carey et al., 1995), it would almost certainly pay too much attention to differences
between range size in species whose climatic requirements are similar. If it did take
account of climate, then the resulting climatic zones ought-if they are to be
credible-to reflect the well-known zones of European and world vegetation (Walter,
1979; Walter & Breckle, 1989; Ozenda, 1994). These main vegetation zones can
already be identified from their climate by numerical criteria (Woodward, 1987;
Prentice et al., 1992), and there would he little merit in defining subtly different new
zones. We have therefore used the standard existing vegetation zones directly.
Outline 0sthe new cla.r.r$ication
We have attempted to devise a classification of British and Irish plants which is
applicable to all species, whether they have restricted or widespread distributions.
We have aimed for a general classification which can be adapted for use elsewhere
in western Europe, and which we hope will be easily comprehensible outside the
British Isles. We have applied it to the bryophytes (Hill & Preston, unpublished)
and hope that it is applicable to other groups of plants and perhaps also to animal
groups. We have adopted aspects of thr classification from a number of different
sources, including H u l t h (1950) and Uouchard et al. (1991). In concept our system
has many similarities to the temperature and contineiitality indices (<e'eip.emerte) of
Ellenberg (Ellenberg, 1988; Ellenberg P t al., 199 1 ; Lindacher, 1995). However, the
Ellenberg indices are defined for Central Europe only and take little account of the
distributions of species in the rest of the world. A comparison of our elements with
Ellenberg's indicator values is given below.
Species are classified by two criteria: occurrence in one or more of the four major
terrestrial biomes (Arctic-montane, Boreal-montane, Temperate and Southern) and
eastern distributional limit (Oceanic, Suboceanic, European, Eurosiberian, Eurasian
and Circumpolar). The floristic elements are derived from a combination of these
two criteria and have been named accordingly, e.g. Circumpolar Arctic-montane,
European Boreo-temperate, Oceanic Temperate. In addition the major biome
categories (MBC) and the eastern limit categories (ELC) have been numbered with
a single digt number. The two digit number made up from the biome number
16
C. D. PRESTON AND hf. 0. HILL
followed by the number for the eastern limit forms a short-hand way of referring
to the element.
Our procedure has been to start with the European distribution and to classify
the species on the basis of its European distribution and any wider contiguous range.
We have placed major emphasis on the main range and tried not to give undue
emphasis to small and disjunct populations. Major disjunctions in the world range
are described by qualifiers. Thus, a species which occurs from Europe eastwards
throughout the boreal zone to eastern Asia is described as Eurasian whereas a
species which occurs in Europe and disjunctly in Central and eastern Asia is
European, with the disjunct occurrences indicated by the appropriate qualifiers.
As our classification is of the vascular plants of the British Isles, all species native
to our area occur by definition in western Europe; the western limit of a species is
therefore much less significant than the eastern limit. However, we have used a
qualifier to indicate those species which are rare in western Europe but commoner
further east and are therefore described as continental.
Class$cation by major biome
The major terrestrial biomes of the world are defined by potential vegetation
types which themselves are governed by climate. They are recognized by most
biogeographers (e.g. Walter, 1979; Cox & Moore, 1993; Ozenda, 1994) and are
often subdivided. North of the tropics, they are basically distributed in latitudinal
belts, which are termed zonobiomes (Walter, 1979). This latitudinal zonation is
complicated by the presence of mountains; increasing altitude has a similar (although
not identical) effect to increasing latitude, and Arctic or Boreal orobiomes are
therefore found on mountains in temperate or tropical areas. The zonobiomes tend
to be less apparent in oceanic areas such as the British Isles than in more continental
climates (Tuhkanen, 1987).
The four major biomes used in our classification are defined below; their European
distribution is mapped in Figure 1:
(1) Arctic-montane. Species with their main distribution north of or (on mountains)
above the tree line.
(2) Boreal-montane. Species with their main distribution in the coniferous forest
zone. They may occur in the Boreal zonobiome, and/or in the coniferous forest
zone on mountains to the south.
(3) Tmperate. Species with their main distribution in the cool-temperate, broadleaved deciduous forest zone, often called the nemoral zone by European authors.
These species may occur in cool steppes in continental interiors.
(4)Southern. Species with their main distribution in the warm-temperate zone
south of the broad-leaved deciduous forest zone. In Europe the warm-temperate
zone is characterized by a Mediterranean climate with summer drought. These
species may also occur in warm steppes in continental interiors.
Many species are confined to one of these zones. The zones contain numerous
different habitats and a species of the Boreal zone does not necessarily grow in
coniferous woodland: it may grow, for example, in bogs or fens, on rock outcrops,
as a submerged aquatic or as a weed. Species may be classified as belonging to a
single zone if their main range lies in that zone even though they extend to a limited
extent into the neighbouring zone to the north or to the south, or both. Species are
classified as belonging to two zones if they occur widely in both zones, or narrowly
FLORISTIC ELEMENTS IN BRITAIN AND IRELhNU
17
Figure 1. hlap showing the approximate s o u t h r ~ ~limits
ii
of the Arctic
and Borcal (C) zoiiohionies,
arid the approximate northern limits of the Trmprrate (B) and Southern (D) zonobiomcs, in Europr
arid western Asia. The distribution of land O\YI- I 0 0 0 1x1 is shown by shaded 0.5 x 0.5"grid c-ells.
but to a n equal extent in each (Fig 2) The following categories arc retoqnized for
species occurring in more than one hiome.
(1) Boreo-arctzc Muntunr. Species occur1ing in the Arctic-montane and Borealmontane zones.
(2) Hide-boreal. Species with a distriliiition which is (entercd on thc Borcal mnr
but which also occur widely in the AI ( t i c and Temperate zones
(3) Buwo-tempeiate. Species which occ 111 inor e or less equall) in the Boreal m d
Temperate ~ o i i e sor, if absent from tlic Boreal zonohiome, nscend to the s u l ~ d p i i ~ ~ ~
Lone on mountains.
(4) 1/[email protected] with n disti ilmtion which is centcrcd 011 thc Tcinpcr<itc
zone but which also occur widel) in t l i r Korenl and the Southern (hleditet I ancan)
ZOllC5.
(<5)Soutlzein-temn~ert?te.Species \Lhich a i t' found inore or Irss cclu.ill\. 111 thc Tciiipci 'itc
m d the Southern (l\lediterrnnean) zoiw\.
Some specirs occur in diffrrent ones, 01 coniliinntions o f /ones, i n difkreiit pa1 t s
C. D. PRESI'ON AND M. 0. HILL
18
A
1
2
3
4
5
Figure 2. Diagammatic representation of the distribution of species in the Arctic (A), Boreal (B) and
Temperate (T) zonobiomes. Species 1, 2 and 3 would be classified as Boreal and species 4 and 5 as
Boreo-temperate.
of their range. In classifying these species we have given preference to their
distribution in Europe. Sparganium natans, for example, is a characteristic species of
the Boreal zone in North America (Cook & Nicholls, 1986) but in Europe it is
widespread in the Temperate zone as well; we therefore classify it as Circumpolar
Boreo-temperate. G p h a latzjilia has a Southern-temperate distribution in Europe
and we classify it as Circumpolar Southern-temperate despite its Wide-temperate
range in North America. However, we have taken the North American range into
account when classifying species which are very rare in Europe but much more
widespread in North America (e.g. Potamogeton epihydms, Sisyrinchium bermudiana).
Classtjcation by eastern limit
The categories used to define the eastern limit of taxa are defined below; the
eastern limits of the Oceanic and Suboceanic categories arc mapped in Figure 3.
(1) Oceanic. Species which are restricted to the Atlantic zone. Oceanic species do
not (or only just) extend eastwards in Europe to Germany; they are western in
Scandinavia. They may occur in the Boreal, Temperate or Southern biomes. In the
Temperate biome the Oceanic zone is exemplified by the distribution of Erica cinerea.
We use the term Oceanic to apply to a defined area of western Europe, not to
describe species with coastal affinities.
(2) Suboceanic. Species with a main distribution which extends beyond the Oceanic
zone to the western Mediterranean, western Central Europe or Sweden.
(3) European. Species with a mainly European distribution; they may extend
eastwards to the Caucasus, Pontic Asia and the Middle East but do not occur east
of 60" E. As Perring (1 985) pointed out, relatively few species arc strictly confined
to Europe as defined, for example, by Flora Europaea; he therefore amalgamated
Matthews' European and Eurasian floristic elements whereas we have chosen to
define Europe rather more widely to incorporate those areas in western Asia to
which primarily European species characteristically extend.
(4) Eurosiberian. Species with a main distribution which reaches its easterly limit
between 60" E and 120" E. Species with a southerly distribution qualify for this
category only if they extend east of Iran, Iraq and the Arabian peninsula to reach
their eastern limit in or beyond Afghanistan.
F i p r r 3. 'l'he appi-oxiinate eastrrn limits of
t l i r ( )ccariic. (L4)and
Sulmccanic (B) categorirs.
(5) Euraszan. Species with a distribution which extends across Asia to an eastcrn
limit east of 120" E.
(6) Circ-urrzpolur. Species which are Ihund in Europe, Asia and North A4i11eriCii.
Specks qualify as circumpolar cven if' thcy arc absent from part of one of tlicse
continents (e.g. absent from eastern North Amcrica); in this case they- are descrilied
as disjunctly circumpolar.
2% n~edzterranean-rltlantzcand Aledit~nanrmrriiontanr rlrmenti
Two elements h a w been devised for sprcics which are found 111 the Temperate
zonc in western Europe but arc confined to the Southern mile further east. Thew
elements combine an Oceanic 'Tempcl <itc and a Europeaii Southern distrihuiion.
T h e hlcditerranean-Atlantic element includes species whk h are fairly strictl) conhied
to the Mediterranean zone and the Atlantic fringe of Europe. 'I'he\. &re absent from
Central Europe. Species must occur in I)otli the western m d eastern Meditcrranem
to qualify as Mediterranean-Atlantlc: thc
foLlrld ill the AtInI1tic zollc J,ut on]) 111
the western hfediterranean are classifiecl <i$ Suboceanic Southern-tempeldie. The
20
C. D. PRESTON AND hi. 0. HILL
species in the Submediterranean-Subatlantic element have a broader distribution
than those of the Mediterranean-Atlantic element in both the Atlantic and the
Mediterranean zones, and often extend into the south-western parts of Central
Europe. Some of them are rarer in the true Mediterranean zone than in the
Submediterranean region.
The distinction between the Southern-temperate and the Mediterranean-Atlantic
elements is relatively straightforward for most species: the Southern-temperate species
are not markedly western in the Temperate zone. However, the distinction is difficult
to make for coastal species as the distribution of coastal habitats in Europe is
essentially Mediterranean-Atlantic: the only exceptions are the Arctic, Baltic and
Black Sea coasts. Species which extend into the Baltic or have outlying inland
populations in Central Europe are classified as Wide-temperate or Southerntemperate; those which are strictly confined to the Atlantic, Mediterranean, North
and Black Sea coasts are classified as Mediterranean-Atlantic.
The species in the Mediterrancan-montane element are found in mountains in
the Mediterranean zone, but in habitats which are too warm to he described as
Arctic or Boreal. They also differ from the Arctic-montane and Boreal-montane
species in occurring at low altitudes in the Temperate zone.
List gjlooristic elements
The floristic elements recognized in this paper are listed below. The first digit of
the number denotes the maior biome category (MBC), the second dipt the eastern
limit category (ELC).
13
14
15
16
European Arctic-montane
Eurosiberian Arctic-montane
Eurasian Arctic-montane
Circumpolar Arctic-montane
61
63
64
65
66
Oceanic Wide-temperate
European Wide-temperate
Eurosiberian Wide-temperate
Eurasian Wide-temperate
Circumpolar Wide-temperate
71
72
73
74
75
76
Oceanic Temperate
Suboceanic Temperate
European Temperate
Eurosiberian Temperate
Eurasian Temperate
Circumpolar lemperate
81
82
83
84
85
86
Oceanic Southern-temperate
Suboceanic Southern-temperate
European Southern-temperate
Eurosiberian Southern-temperate
Eurasian Southern-temperate
Circumpolar Southern-temperate
2 1 Oceanic Boreo-arctic Montane
23 European Boreo-arctic Montane
24 Eurosiberian Boreo-arctic Montane
26 Circumpolar Boreo-arctic Montane
34 Eurosiberian Wide-boreal
35 Eurasian Wide-boreal
36 Circumpolar Wide-boreal
*
41
42
43
44
45
46
Oceanic Boreal-montane
Suboceanic Boreal-montane
European Boreal-montane
Eurosiberian Boreal-montane
Eurasian Boreal-montane
Circumpolar Boreal-montane
51
52
53
54
Oceanic Boreo-tcmprrate
Suboceanic Boreo-temperate
European Boreo-temperate
Eurosiberian Boreo-tempcrate
9 1 Mediterranean-Atlantic
92 Submediterranean-Subatlantic
93 Meditcrranean-montane
FLORISTIC ELEhlhh 15 1U BRL 1AIN AND IRLLXNU
55 Eurasian Boreo-temperate
56 Circumpolar Boreo-temperate
Qualijiers
The following annotations are used to qualify the floristic elements:
+
(after the numerical code for the clcnient) Disjunctly circumpolar. This may be
used to qualify species classified as Oceanic, Suboceanic or European which are
also found in eastern Asia and North America. It is also used to qualify species with
a circumpolar range with some gaps (e.g. species absent from eastern North America).
(after the numerical code for the element) Continental, i.e. notably rarer in the
oceanic zone of Europe than one would expect from its habitat requirements.
-
Am1 Also occurring in North Am(-rica. This qualifier is applied to Oceanic,
Suboceanic, European and Eurosiberian taxa; it is applied to Eurasian distributions
only if the taxon is very rare in North America; otherwise the distrihution would
be circumpolar. This qualifier may be replaced by four others which provide further
details of the distribution in North America:
Am2 Occurring in Greenland but not in continental North America.
Am3 With a very restricted range in mainland North America; more widespread
in Eurasia.
Am4 Occurring in North America but only in the west.
Am5 With a very restricted range in Europe; more widespead in North America.
As1 Also occurring in Central Asia.
As2 Also occurring in eastern Asia.
Bol Arctic or Boreal species (other than exclusively coastal taxa) which are absrnt
from mountains which rise from the Temperate zones of Europe and western Asia
(they may occur in the mountains of (kntral Asia or North America).
Bo2 Arctic or Boreal species whic.11 occur on mountains which rise from the
Temperate zone of Eurasia but are alxent from the northern Arctic or Boreal zones.
Col Species which have a coastal distribution throughout their range
Co2 Species with a primarily coastal distribution in the British Isles.
Djt Species with markedly disjunct distributions. This qualifier is replaced by
+
for disjunctly circumpolar distributions.
Sst Species interpreted taxonomicall)- in a narrow sense (.s~nsu.rfr.icto).
Tax Distributional data for this species iiicotnplctc or unreliahlc because of taxonomic
difficulties.
Tso Also occurring in the tropics (including tropical mountains) or the Southern
Hemisphere or both.
Wid Widely naturalized outside tlic native rangc. 'l'hc total rangc. native and
22
C:. I).I’KESTON AND hl 0.
HILL
naturalized, is sometimes classified in brackets after this symbol, i.e. [76] indicates
that the species now has a Circumpolar Temperate distribution.
Allocation o f species to Joristic elements
Species have been assigned to the appropriate biome primarily by visual inspection
of distribution maps or, where these are not available, from other distributional
data. Although a species is classified by the biome(s) in which it occurs rather than
by distribution per se, the appropriate biome(s) can usually be deduced from a
distribution map. We have also made use of information on the habitat and altitudinal
range of the species in those regions (such as the mountainous countries around the
Mediterranean) where several biomes may occur in close proximity. The eastern
limit, a purely geographical classification,has been established from the same sources.
Grouping species into floristic elements by eye rather than by computational
methods has been criticized by Jardine (1972) on the grounds that there may be
tendencies to discover patterns where none is present, to group distributions round
hypothetical ‘types’ and to bias the classification because of preconceptions about
the factors determining plant distributions. We have tried to avoid these pitfalls by
defining the categories before embarking on the classification, then by reviewing
the distribution maps of species in each element in turn to ensure that they all
conform to these definitions. However, as Watson (1 847-59, I : 54) emphasized,
there are no discontinuities between elements, which “pass gradually into each
other”. Many decisions have had to be made about borderline cases, and it follows
that the exact position of a single species in the classification should not be interpreted
too rigidly, as there may be an arLguablecase for placing it in one of the adjacent
elements.
We have also had to accept small-scale distribution maps at face value, although
the production of such maps itself involves numerous subjective decisions, and “the
elegant loops and curves which we see in so many maps” must, as Webb (1965)
suggests, “conceal ambiguities and ignorance”. We ourselves have had to decide
whether to treat fringe occurrences as part of the main, classified, distribution of a
species or deal with them as disjunctions which are identified by an appropriate
annotation. There are also a few species with anomalous distributions which do not
fit comfortably into the classification we have devised (e.g. Allium schoenoprasum,
Buniuni bulbocastanum, Minuartia uerna, Tiphroseris palustri,r).
Coincidence maps
The distribution of the specics in the floristic elements that we have defined is
illustrated by coincidence maps showing the proportion of each element in grid
squares.
The maps of Britain and Ireland are based on the coniputer database managed
by the Biological Records Centre at ITE Monks Wood. This holds most of the
records collected for the Atlas ofthe Britishjorrr (Perring & Walters, 1962), updated
by the results of more recent surveys (e.g. Rich & Woodruf’f, 1990; Stewart, Pearman
& Preston, 1994; Preston & Croft, 1997). The proportion of the species in each
element is shown in 10-kn2 grid squares. No symbol is plotted in squares with less
than 10% of the species in the relevant element; the smallest symbols denote squares
with 10-19% of the species and successively larger symbols are used for successive
10% intervals, except that the largest symbol is used for squares with 80-100%.
FLORISTIC ELMFA I \ IN BKI r
m AND
23
IRELQD
TABLE
6. The number of British and Irish native species in each floristic elenient
Eaatcrri limit caregor)
Slajor hiomc category
I
Ocea
2
Suho
1 Arctic-montanr
2 Boreo-arctic hlontane
3 \Vide-boreal
4 Boreal-montane
.5 Roreo-tempcrate
6 /Vide-trmprrate
7 Tcmperatc
8 Southrrn-temperate
9 h'lcditerranean
Total
1
29
6
I0
2
I
1
27
8
X
-111
:3
2R
297
I06
1
48
25
69
I59
5+
47
I12
1
0
67
II
I20
81
3
-
1
3
38
5I1
64
1.!
3
38
17
6
526
41
25
I7
26
I3
~
297
107
250
T h r eastern limit categories are as follows: 1 O t c a , Occnnic; 2 Subo. Suhoceanic: 3 Euro. E u q x - n n ; 1 E d ) ,
Eurosiberian; 5 Easi, Eurasian; 6 Circ, Circumpolar. Fur the hlcditerranean elements, 91 drnotes S\lcditci-ranean,\tlantic; 92, Suhmcditerranean-Subatlantic.; 93, ~Irtlirci-r;inean-montanr.
TABLE
7. The percentage of species in rach tloristic rlcmrnt. For key to castern limit catrqories. see
'1'at)lC 6
Major biomc category
I Arctic-niontanr
2 Borro-arctic Montane
3 LVide-hoi-ral
4 Borcal-inontanc
5 Boreo-trmperatc
6 \Vide-temperatr
7 'I'empcrate
8 S(iiithern-tcmprrare
9 Rlcditcrraneaii
Total
The maps have been plotted using thc UMAI' program written by Dr iZJ. Morton.
The European maps are based on a computer database with rccords fiom =i/lus
Florue Europaeae, volumes 1-9 (Jalas & Suomiiicn, 1972-9 1). Distributional data for
29 1 of the 1481 species classified by 115 are iiicluded in this datdhdw, and the maps
are based on this subset. Symbols of in( t ca4ng yize indicate the prewicc of 1 1 23'Y",
26-40"/0, 41-55%, 55-70% and over 70% of the species in each element i n 50-km
grid squares. Grid squares in eastern Europc tend to be under-recorded (Holuh,
1981; Webb, 1988) and this must br Iwrne i n mind in interpretiiig the E U I O ~ C ~ I ~
coincidence maps.
The species allocated to each floristic element are listcd in ,lppcndix 1. 1 he
number of species allocated to each cl(micnt is g i \ m in Table 6: for thesc figurcs
expressed as proportions, see I'able 7 The Europcan di5trihutioii of a snmplc of
24
C. D. PRESTON AND M. 0.HILL
species in the six largest major biome categories and in four eastern limit categories
is shown in Figures 4-13 (see pp. 25-29). The different elements, grouped by major
biome category, are described in the following accounts.
The Arctic-montane elements (13-1 6)
The Arctic-montane elements consist of a relatively well-differentiated group of
79 species which are confined to the Arctic zonobiome or to the corresponding
orobiomes to the south. There are no Arctic species with Oceanic or Suboceanic
distributions (although three montane endemics have been classified as Oceanic
Arctic-montane and are listed in the Endemics section below).
In the British Isles the Arctic-montane species are concentrated in the mountains
of Scotland, particularly in the Breadalbane range in the Central Highlands, the
Cairngorms and the mountains around Lochnagar (Fig. 14) (Figs 14-33 are grouped
at the end of the paper on pp. 65-85). They are found in a range of habitats at
high altitude, including grassland, exposed montane heaths, cliffs, screes, rocky
detritus, flushes, springs and streamsides; some species are calcicole and others
calcifuge. The European map demonstrates major concentrations in the Arctic, the
Alps and the Pyrenees (Fig. 4).
Most of our Arctic-montane species are found both in the Arctic zonobiome and
in the mountains to the south. We have distinguished by qualifiers those which are
confined (at least in Europe and western Asia) to one or other of these areas. The
species found in both regions show every gradation from those which are frequent
in the mountains, e.g. Saxzjaga oppositfolia, to those which are rare or even, as in
the case ofJuncus biglumis and Saxzjaga niualis, confined to single localities in Central
Europe (Tutin et al., 1968-80; Webb & Gornall, 1989). Koenigia islandica, which is
now confined to the Arctic zonobiome, is known from fossil evidence to have
occurred in Central Europe in the past (Hultkn & Fries, 1986).
The high proportion of Arctic-montane species in the Circumpolar element (52‘10)
reflects the lack of regonal differentiation in the flora of high latitudes. Hooker
(186 1) identified “one general arctic flora” and recent estimates suggest that 60%
of Arctic species have a circumpolar distribution (Walker, 1995). However, some of
the species classified as Circumpolar Arctic-montane are absent from part of the
circumpolar area, notably Dyas ortopetala which is absent from eastern North America
(Hultkn, 197 1).
The European and Eurosiberian species also show tendencies towards a wider
distribution. Six of the 35 species in thcse groups have a disjunctly circumpolar
distribution, being found in Europc, North America and as isolated populations in
Central or eastern Asia. A further 19 spccics are amphi-atlantic, being found in
mainland North America or Grecnland, and one species is recorded from Europe
and Central Asia. The only species which are confined to Europe and its environs
are the Arctic plants Arenaria noruegicn, rlrtemisia noruegica, Salh(ix arbuscula and S. myrsinites
and the montane taxa Coehlearin pyrenaica, Minuartia recurun and lid. setlodes; some of
these are members of taxonomic complexes with a widcr distribution. The presence
of the distinctive hlznuartia sedoitles is notable as so few of the species of the mountains
of Central Europe are found in the British Isles but not in the Arctic.
FLORISTIC ELEMENTS IN BRITAIN AND IRELAND
25
FiLgure4. The distribution in Europe of species in the Arctic-montane major hiome category-. The
map is based on a sample of 25 species which occur in the British Isles arid are mapped by Jalas &
Suominen (1972-91). Symbols of increasing sire indicate that 11-25, 26-40, 41 -55, 56-70 and
71-100'% of the species are found in the 50-km squares of the U T h l grid in which they are plotted.
26
C. D. PRESTON AND M. 0. HILL
Figure 6. The distribution in Europe of species in the Boreo-temperate major biome category. The
map is based on a sample of 53 species which occur in the British Isles and arc mapped by Jalas &
Suominen (1972-91). For explanation of symbols, scc caption to Figure 4.
Ficgure 7. The distribution in Europe of species in thc ‘I’cmpcrate major biome catcaory. The map is
based on a sample of 1 13 sprcies which occur in the British Isles and are mapped by Jalas & Suominen
(1972-91). For explanation of symbols, see caption to FiLpre4.
27
Figure 8. 'Thc distribution in Europe of spccio 1 1 1 thc Southrrn-rrnipcratc major I k m r categor);. ' h e
map is based on a sample of 72 species which ( I ( ('111- in thc British Isles and arc mapped h> J a l a h &
Suomiiieii (1972-91). For cxphnation of- symbtrls, set' caption to Fi,gire 4.
Figure 9, T h e distribution in Europe of specir\ iii tlir ~lrditrrI.aiieaii-.~tlaiiitic
floristic. e h i v n t . 'llic
map is bascd on a samplc of 16 spcrics whicli ( I ( c u r i i i thv British Isles and arc niappccl IJY,Jala\ 8r
Suominrn (1972- 91j. For rxplaiiation of symlx)la. sce caption t o Figure 4.
28
C. D. PRESTON AND M. 0 . HILL
Figure 10. The distribution in Europe of species in the Oceanic eastern limit category. The map is
based on a sample of 21 species which occur in the British Isles and are mapped byJalas & Suominen
(1972-91). For explanation of symbols, see caption to FiLpre4.
Fi<gure 11. The distribution in Europe of species in thc Suboceanic rastern limit category. The map
is based on a sample of 23 species which occur in the British Isles and are mapped by Jalas &
Suominen (1972 91). For explanation of symbols, ser caption lo Figure 4.
FLORISTIC: ELERlEN 1h IN BRITAIN AND IRLIAND
Figure 12. T h e distribution in Europe of spr.cir.s in the European eastern liinit catcgoi);. 'l'hr rnap is
based on a sample of I 1 1 species which OCCLII' in the British Islcs and arc mapped by Jalas tk Suominrn
(1972-91). For explanation of symbols, see captiorl to Figure 4.
a
30
(:. 1). PRESI'ON 4 N D R I . 0. HILL
T h e Boreo-arctic Montane elements (21-26)
There are 38 species in the Boreo-arctic Montane elements. No species are
classified as Suboceanic or Eurasian Boreo-arctic Montane. The European species
are more widespread in Scandinavia than the corresponding species in the European
Arctic element, and the Circumpolar species are more widespread in the Boreal
zone as a whole.
The greatest concentrations of Boreo-arctic Montane species in the British Isles
are found in the Scottish mountains which support most Arctic-montane taxa.
However, the members of the Boreo-arctic elements are more widespread than the
purely Arctic species in both Britain and Ireland (Fig. 15). The Boreo-arctic species
also have a greater altitudinal and habitat range than that of the Arctic-montane
species: in addition to species of open, montane habitats they include plants of sandy
or rocky seashores, lowland as well as upland heaths and swamps and bogs.
The proportion of Boreo-arctic species in the Circumpolar element (66%) is even
greater than the equivalent proportion for Arctic species (52%). In addition, the
tendency of the European and Eurosiberian Arctic species to have a disjunctly
circumpolar or amphi-atlantic distribution is even more marked amongst the Boreoarctic species. The only species which is confined to the European area (including
adjacent parts of south-west Asia) is Euphrasin salisburgensis, although Juncus balticus
would also be placed in this category if we had classified this species in the narrow
sense rather than as a member of the circumpolar J . arcticus complex.
'There are four Boreo-arctic species which have a predominantly or exclusively
coastal distribution: Lathyrusjuponicus, Lymus arenarius, Ligustrum scoticum and Mertensia
maritimu. These are all classified as European Boreo-arctic and are absent from the
northern coast of Central Asia but they are found in east Asia (sometimes as distinct
subspecies)and across the Bering Sea into North America, where they are widespread.
Almost all the European, Eurosiberian and Circumpolar Boreo-arctic species
which are found in non-coastal habitats are found in both the Arctic region and
the mountains to the south. Carex aquatilis and Cornus suecica are the only exceptions,
as they are confined to the Arctic and Boreal zonobiomes. Some species show some
taxonomic differentiation with latitude and altitude: Eccinium uliginosum and I? vitisidaea are represented in the Arctic by small-leaved subspecies, for example, and a
hermaphrodite subspecies of Empetrum nigrum extends further north and to higher
altitudes than the monoecious plant (Tutin et al., 1968-80). Euphrasza sahburgenszs
has arctic and alpine ecotypes (Yeo, 1978) and in Britain coastal and montane
populations of Equisetum uariegatum differ morphologically (Stewart et al., 1994).
A single species, Euphrasia ostenfeldii, is classified as Oceanic Boreo-arctic. It is
endemic to the northern Atlantic area (Britain, Faeroes and Iceland) and is one of
several Arctic or Boreal species in the genus, some of which are endemic to the
British Isles (see below).
The morphologically and cytologically variable species Allium schoenoprasum has
been assigned to the Boreo-arctic major biome category, but there are several
anomalies in its distribution. Although it has a primarily boreo-arctic and montane
distribution, it penetrates into lowland regions along river valleys in mainland
Europe. It also occurs in some coastal habitats and in sparsely vegetated rocky
habitats which are wet in winter but bone-dry in summer (Hulten &r Fries, 1986;
Stewart et al., 1994). Unlike the other Arctic and Boreo-arctic species, it has become
widely naturalized outside its native range.
FLORIS ric ELESIEN I 4 IN BRITMY AND IRELAND
31
'The Wde-boreal elements (34-36)
The Wide-boreal species are found in the Arctic, Boreal and Temperate zones.
This is a small group of 19 species, which are frequent throughout the British Isles,
especially around the coast and in northern England and Scotland (Fig. 16). They
are ecologically heterogeneous, and some (e.g. Equzsetum aruense, Fe..rturn ruhra) may
occur in a wide range of habitats.
The Wide-boreal species can be divided into two major groups. Five species,Armeria
maritima, Cochlearia oficinalis, Honckenya puflloides, Plantago maritima and Tipleurospermum
maritimum, are predominantly coastal in western Europe, although Cochlearia ?ficinalis
and Plantago maritima in particular also occur in inland and montane habitats. These
five species extend in Europe from Spitsbergen or Arctic Russia southwards at least
as far as north Spain. Most of the remaining species are very widespread in a range
of inland habitats throughout the Arctic, Boreal and Temperate zones. They tend
to be morphologically variable and taxonomically heterogeneous: examples include
Cardaminepratensis, Deschampsia cespitosa, Eriophorum angusttfolium, Festucn rubra and Luzula
mu/t$ora. The two Eflhroseris species arc- rather exceptional: both the T. intepfilius
complex and T palustris tend to occur in the Arctic and the Temperate zones but
are rare or absent in the intervening Boreal. The arctic variants of 7 paluJtris often
differ morphologically in their dwarf habit, strongly pubescent inflorescences and
large capitula, but they are not clearly differentiated from the plants of the Temperate
zone (Hultttn, 197 1).
The single Eurosiberian Wide-boreal specics, Plantago maritima, is also widespread
in North America and just crosses the Bering Sea into eastern Asia. The single
Eurasian species, Ranunculus acris, is \videly naturalized in North America and when
both native and non-native occurrences are taken into account its distrihution is
now completely circumpolar. The remaining 17 species are in the Circumpolar
Wide-boreal element.
'The Boreal-niontnne elements (41 46)
? >
I here are 103 species in the Bo~-cal-niontaneelements. In Britain thr Boreal
spc-cies are widespread in Wales, northern England and Scotland. The Boreal and
Boreo-arctic species occupy virtually thc. same area, but the Boreal species are rather
evenly spread with a less marked concentration in the Central Highlands and the
Cairngorms (Fig. 1 7). In Ireland the Borcal sprries are infrequent, and concentrated
t. In western and Central Europe thc- main
in the mountains of the north and
concentrations of species are in Scandinavia and the Alps (Fig. 5).
'The Boreal-montane species are e v c n more ecologically diverse than thosc of thr
Roreo-arctic elements. They include Z'inu.s ~~~1oe.stii.s
and some characteristic herhs of
coniferous forests, but also species of tall herh communities, bogs, moors and hcaths.
screes and rock outcrops, <grassland,swamps, and aquatic?coastal and weed) habitats.
The Oceanic and Suboceanic Borcal-inonlane elements contain both European
endemics (e.g. Saxzj?qa hypnoides, S o ~ h u \ rupicoln) and amphi-atlantic species. 'I'he
amphi-atlantic plants include Erioraulon gunti ti cum and LYpirantlie.\ roninn,:ojinnn, sprcirs
which are much more widesprcad in North America than they are in Europc, and
have traditionally been included in t h v 'North American element' in the British and
Irish flora.
32
C. D. PRESTON AND M. 0.HILL
Although most of the species in the European Boreal-montane element are
confined to Europe in the broad sense (18 species), there are species which are
amphi-atlantic (6), are found in Europe and E. Asia (1) or are disjunctly circumpolar
(2). Only two of the Eurosiberian species are recorded from North America, Isoetes
lacustris and the disjunctly circumpolar Carex buxbaumii, although two others are now
naturalized in Greenland (Cirsium heterophyllum, Vaccinium myrtillus).
Most of the species in the European Boreal-montane element are found in
mainland Europe both in the Boreal zonobiome and in the mountains to the south;
only two species are confined to the zonobiome (Euphrasia scottica and Lamium
confertum). A slightly larger group, comprising Lhyopteris remota, Meum athamanticum,
Oxytropis halleri, Ribes alpinum, irhlaspi caerulescens and Viola lutea, is found in the
mountains of Central Europe but not in the Boreal zonobiome; some of these species
extend north to southern Scandinavia and two, Meum athamanticum and Thlmpi
caerulescens, are naturalized further north. Virtually all the Eurosiberian, Eurasian
and Circumpolar species are found in both the zonobiome and the mountains of
Central Europe. The only exceptions are Rubus arcticus, which is absent from the
mountains, Potentilla3uticosa, which has a remarkably disjunct distribution in Europe,
extending north only to southern Scandinavia, but which occurs in the Arctic in
east Asia and North America, and Primulafarinosa, a montane rather than boreal
species.
A few species listed in the Boreal-montane elements have rather anomalous
distributions. Some (e.g. Coeloglossum viride, Trichophorum cespitosum) show a tendency
to extend into the Temperate zone in oceanic western Europe. Bbsmus rufus,
classified as European Boreal-montane, has an amphi-atlantic and primarily coastal
distribution but also occurs in saline habitats in Central Asia. Minuartia uerna is a
very variable species which has a disjunct Eurasian distribution, primarily in the
mountains but reaching the Arctic in northern Russia; it also occurs in isolated
lowland populations, especially in areas rich in heavy metals. Matthews’ (1955)
classification of the species as Arctic-Alpine was rejected by Webb (1983) and the
Boreal-montane element appears to be the least inappropriate category in which to
place it.
The Boreo-temperate elements (51-56)
The Boreo-temperate elements make up the third largest group in the British and
Irish flora; only the Temperate and Southern-temperate elements contain more
species. Many ofthe 233 Boreo-temperate species are widespread in both the Boreal
and the Temperate zones. However, other species have a more restricted distribution,
occurring in a rather narrow band along the southern part of the Boreal and the
northern part of the Temperate zones. Examples of these restricted species include
Pohyala amarella, Rhynchosporajma, Sagina nodosa, Stratiotes aloides and Kciu sylvatica. At
least one species, Pkyteuma orbiculare, is taxonomically heterogeneous: f? orbiculare sensu
strict0 has a Boreal-montane distribution in Central Europe whereas the segrcgate
P tenerum, although regarded as conspecific, occurs at low altitudes in western
Europe. Boreo-temperate species are frequent throughout the British Isles (Fig. 18),
and occur in a correspondingly wide range of habitats. In Europc they are frequent
from central Scandinavia to the Alps (Fig. 6).
The Oceanic Boreo-temperate elemcnt is similar to the Oceanic Boreal in
FLORISTIC ELEhfCN 1 5 IK BR11 h I U . W U IRLIAND
7s
containing both European endemics (c.g. Alleronopsis cambrics) and amphi-atlantic
species, the latter including two plants which are much more widespread in
North America than in Europe (Limor~llaaustralis, Potamogeton epihydrus). Most of the
Suboceanic Boreo-temperate species ha17e amphi-atlantic distributions, reaching
Greenland (Callitriche hamulata) or maillland North America.
The European Boreo-temperate specitas in the British Isles show a tendency to
be more frequent in the north and west of Britain and Ireland than in the English
and Irish Midlands (Fig. 19). Coincidence maps (unpublished) for the Eurosiberian,
Eurasian or Circumpolar species show no discernible trends within either Britain
or Ireland. The Eurosiberian and Eurasian elements, in particular, contain sonie of
the most frequent species in the British and Irish flora (e.g. drlzillea rnill~filium,
Cerastium fontanurn, Ranunculus repem, Tr$dium rqhis, Urtira dioica). l h e rare species in
these elements often (but by no means always) have a ‘continental’ distribution in
Europe, being rare throughout western Europe (e.g. Carex elongata, Luzula pallidula,
Rumex aquaticus).
hiany species in the European and Eurosiberian elements occur naturally in
North America. Others are now thoroughly naturalized there. Indred, there is some
doubt whether some of the very widespread and rather w c d y specics are native or
not: Fernald (1970), for example, treated Saginn ,brorurnbpns as native whereas Crow
(1978) suggests that it was introduced soon after European settlement. The Eurasian
species are by definition absent or veiy rare as natives in North America. Some
(e.g. Oxalis aretosella, Potaniogeton compre.\ ~11,s)arc replaced in North America by closely
allied species and many others have bcconic naturalized there and ha\.e attained a
circumpolar distribution artificially.
Fewer of the naturally circumpolar spvcies arc quite so widespread and ecologically
tolerant as the Eurosiberian and Eurasian herbs listed above. hlany are plants of
aquatic habitats: 42% of the Circumpolar Boreo-temperate species may grow as
hclophytes or hydrophytes compared to 12?6 of species in the Eurasian and
Eurosiberian Boreo-temperate elements. l’he other species are ecologically diverse.
‘lhe 34 species which occur exteni\ cly in the Boreal, Temperate and Southern
zones are placed in the Wide-tempcrntr elements. None of them is cla5sified
a5
Suboceanic \.Vide-temperate. The spec ics arc frequent throughout the British Isles,
and many 10-km squares in Britain and Ircland contain at lrast 80% of the nienilxm
of the group (Fig. 20).
T h r single Oceanic W‘ide-temperate species, Szs_vnnthium hpirnudianti, i, I estricted
as an apparently native species in Europc to western Ireland. This 5pecics, if hroadly
defined, extends in North America from Greenland and Alaska to Florida. Texas,
California and perhaps Mexico (hlosquin, 1970), hence the classification as Widetemperate. The taxonomic relationship of thc plants in Ireland to those in North
America is unclear; some authors ha\r suggrsted that the Irish plant is both nati\e
and endemic, bhereas others regard i t as m introduction (Stacr, 199 1).
‘The three European Wide-temperntc species are the M idrspread aquatic Ranunrztli~r
peltatur and two coastal species, Cakzle nzaiitznin and Pohgonum o y p p m m i t n . The latter
extend from Arctic Russia south to thc shores of the hlediterrancan and the R l x k
Seas: if’we recognized an element for sprcies occurririg in all four miiobionies, c hey
34
C . D. PRESTON AND M. 0. HILL
would be placed in it. Both species are variable and are represented by a number
of geographical subspecies in Europe (Tutin et al., 1993)
Almost all the species in the Eurosiberian and Eurasian elements are common
weeds, although Atriplex patula and A. prostrata occur naturally in saline habitats and
become increasingly restricted to coastal sites in the Boreal zone. Virtually all the
weedy species have become naturalized in North America and in other areas where
they do not occur as nativrs; many have attained a circumpolar distribution. Hu1ti.n
(1971) describes Chenopodium album as probably “the most common introduced taxon
of all around the globe” and Plantago major became known as “Englishman’s Foot”
because of its propensity to accompany European settlers (Josselyn, 1672: 86;
Berkeley & Berkeley, 1965: 24.) The main exceptions to these generalizations are
the Eurosiberian aquatic Schoenoplectus lacustris and the Eurasian Eleocharis palustris,
which are replaced by similar species in North America.
Some 43% of the Circumpolar species are helophytes or hydrophytes, compared
with 13% of the Eurosiberian and Eurasian species (figures almost the same as those
in the corresponding Boreo-temperate elements). The remaining species include
weeds, plants of coastal habitats and herbs and grasses with broad ecological
tolerances.
irhe Emperate elements (71-76)
The Temperate elements include 557 species, over 35% of the flora of Britain
and Ireland. These species are found throughout the British Isles, although they are
most frequent in England and very sparsely represented in the Highlands and Islands
of Scotland (Fig. 21). The concentration of species in temperate Europe is well
shown in Figure 7.
irhe Oceanic and Suboceanic Temperate elements (71, 72)
The 48 Oceanic Temperate species are widespread in the British Isles, being
most frequent in western, and particularly south-western, Britain and Ireland, and
least frequent in the English and Irish Midlands (Fig. 22). Nevertheless, the presence
of appreciable numbers of‘ Oceanic Temperate plants in eastern England, even in
relatively ‘continental’ areas such as Breckland, emphasizes the fact that the whole
of Britain lies within the oceanic zone.
There is considerable variation in the distribution of the Oceanic Temperate
species in the British Isles. One group of species is strictly coastal; examples include
Cochlearia anglica, Limonium binenlosum, $pergularia rupicola and Tr$dium occzdentale. These
are characteristic species of the Atlantic coast of Europe, but many of them are
found on both western and eastern coasts of Britain and Ireland. Species which are
more or less restricted to western areas include plants which grow in humid and
often shaded microclimates (e.g. D y p t e r i s aemula, Sibthorpia europaea, Trichomanes
speciosum) or in more open sites (e.g. Pinguicula lusitanica, Sedum anglicum, Ulex gallit).
The latter group merges with species which are widespread in Britain and Ireland,
but rarer in the English and Irish Midlands than elsewhere (e.g. Carex binervir, Erica
cinerea, Myosotis secunda). There are also species which are widespread throughout
Britain and Ireland (e.g. Conopodium mqjus, Hyacinthoides non-scrzpta, Ulex europaeus) and
even plants with an easterly bias (e.g. Festuca arenaria, Genista anglica, Oenanthejuviatilis).
The widespread and eastern plants provide striking and oft-cited examples of the
mismatches which can occur between the distribution of species within the British
Isles and at the wider European scale.
hlany members of the Oceanic Tcmperate element are confined to Europe or
occur elsewhere only in hfacaronesia or northwest Africa. Very few occur naturally
in North America. However, there arc some species with disjunct populations outside
the Atlantic zone of Europe. Hymenop/~yllumtunbrieense is very rare in North America
and also occurs in temperate and tropical regions elsewhere (hlorin, 19!13), and
Szbthorpza europaea is found in western Europc and in the mountains of Grcccc, Crete
and tropical Africa (Hedberg, 1955). This suggests that at least some members of
this element have relict distributions, a conclusion also reached by \.Yebb (1 983) and
Rumsey P I a/. (1991) because of the fragmented distribution of many species within
the Oceanic zone. In the case of Trzrhonmnrs speriosum, the tiny garnetopliytes 1ia1.e
a wider distribution than the sporophytcs, and perhaps persist at sites from which
the sporophytes have disappeared follocz.ing dcforestation (Rumsey et nl., 199I ) .
IJlex europaeus is exceptional amongst the Oceanic Temperate species as it has
spread from its native range eastwards into Europe, and now has a Suhoceanic
Temperate distribution. It has also Ixxmiiic naturalized in western North America
and in the southern hemisphere (Hu1ti.n 8r Fries, 1986).
The 28 species in the relatively small Suboceanic Temperate elcrncnt are more
evenly distributcd in Britain and Ireland than the Oceanic Temperate species, ivith
no appreciable concentration along t tit, western seaboard and with a less iiiarked
reduction in the English and Irish Alidlaiids (Fig. 23). Thcrr is only one coastal
species in the Suhoceanic element (Skipltirlzum mnritzmunz), hut otherlvise thc specks
hare as wide an ecological range as those in the Oceanic element.
hlaiiy of the Suboceanic 'Temperatc species are endemic to Europe. Two species
extend westwards to Greenland and three arc found in mainland North Amrrica.
n~European %2pemte
element (73)
The 297 European Temperate spec ies form the largest of the leinperate elements,
and much the largest single element i n the flora of the Britisli Isles. 7'hcrc is
considerable Lariation in the distribution of mdi\idual 5pccics: sonic. extend into the
Boreal zone and thus approach the Eiu opcaii Boreo-temperatc range- (c.g G+ttia
Jruztnns) whereas others are absent froin the northerii part of thc Temper 'ttc ~ o n c
arid protide d link with the corresponding Southei n-temperate element (e.g. Eu,bhoibia
amqiSdalozda). The species also differ in tlicir longitudinal limits, Mitli sonic ha\ iiiq a
'c ontinental' distribution in Europc (c.g lIzatit/iuc gratzano/~olitanuc)cvhcrcas others ; I r e
frequent west to the Atlantic coast of Irela~id(c.g. Rnnuntzilu~J7nn7miila).In the British
Isles the European Temperate species ~ r almost
r
ubiquitous, beinq niost frequcmt
in England and rather sparsely distributcd in the Highlands and I\lnnds of Scotland
(Fig. 23).
This element encompasses specirs \I ith ninny difhent habitat requit cmcnts
Howe\~er,it is notabk that most of thr trees and shrubs arc Europenn Tempcrntc
species: the trees include ilcer ramn/ieJtw, arpzniiy hululus, F o p y'ltahrn. Fjarznuc ~ n ( ~ 1 ~ 1 0 1 ,
Qurirur spp., Sorbus toimmnlzc, Tnnus hnccata nnd C'lmu~ spp. and the shrubs (:0tnii$
snnguznen, Cbllyluc azlrllann, Crutaeguc spp Enonynus twoparus, Liguctium r~u(qnr~.
Airnu\
spznosa, almost all the Rosa spp. and lihiirntim lnntann. hlany \\oodland herbs (r.9
Galium odorntum, Lnmzastrum gulrobdolon. 2 \ l ~iirzali,
~ ( p r r r m i ~ )nre also inemhers of this
eleinent.
36
C. D. PRESTON AND h4. 0. HILL
Some species in the European Temperate element are endemic to Europe sensu
strict0 but many extend eastwards to the Caucasus and the mountains south of the
Caspian Sea. Only a small proportion (6%) occur naturally in North America,
although many more (at least 25%) are recorded there as naturalized aliens. Some
15% of the species are recorded at disjunct localities in central or eastern Asia,
although information on the status of the species in these localities is difficult to
obtain and this figure may include some species which are present there only as
introductions. Some species (4% of the total in the element) are found in both North
America and central or eastern Asia, and thus have a disjunctly circumpolar
distribution.
irhe Eurosiberian irmperate element (74)
There are 120 species in this element. In the British Isles they have a similar
distribution to those in the European Temperate element, although the coincidence
map (Fig. 25) shows a more marked gradient between the south-east of England,
where the greatest concentration is found, and the north-west of Scotland.
The Eurosiberian element contains both trees (e.g. Alnus glutinosa, ?ilia cordata),
shrubs (e.g. Frangula alnus, Rhamnus cathartica), woodland herbs (e.g. Anemone nemorosa,
Neottia nidus-avis) and one species described by Rackham (1980) as circumboscal
(Melampyrum cristatum). However, both woody species and plants which (at least in
Britain) are primarily woodland herbs are less well represented in this element than
in the European Temperate group. Most of the Eurosiberian species are plants of
more open habitats, including still or flowing waters (e.g. Hydrocharis morsus-ranae,
Oenanthe aquatica, Potamogeton lucens) and swamps and fens (e.g. Carex acut$mnis,
Ranunculus lingua, Sium lalfolium). The species of drier habitats, including open
grassland, include some which remain frequent in the oceanic zone of Europe (e.g.
Artemisia vulgaris, Carex capvphyllea, Senecio jacobaea) and others which are markedly
continental in their distribution (e.g. Artemisia campestris, Hypochaeris maculata, Phleum
phleoides). The Eurosiberian element also includes species which are coastal in their
distribution in Britain and Ireland, although found in inland habitats further east
(Althaea o$cinalis, Atriplex pedunculata), and weeds (e.g. Fumaria vaillantii, Sinupis aruensis,
Eola aruensis).
Only 6% of the Eurosiberian species are native in North America, although
almost half the remainder have been recorded there as naturalized aliens. Some
species which are believed to be restricted as natives to the Eurosiberian area are
now naturalized in the remainder of the temperate northern hemisphere (e.g. Sonchus
arumsis, Tnj%lium bratense, Tripleuraspemurn inodorum).
Although the Eurosiberian species extend by definition from Europe to central
Asia, some are represented in the oceanic or European zone by distinct taxa. An
extreme example is Asparagus oficinalis, found in western Europe as the coastal subsp.
prostratus which has an Oceanic Temperate distribution; the cultivated subsp. oficinalis
is believed to be native to the Eurosiberian area although its native distribution is
obscure. A less extreme example is provided by Anemone nemorosa, which is represented
by subsp. nemorosa in Europe and western Asia and by subsp. altaica further east
(Hulttn & Fries, 1986).
irhe Eurasian and Circumpolar '(emperate elements (75, 76)
There are 38 Eurasian and 26 Circumpolar Temperate species. The coincidence
maps for these species in Britain and Ireland (unpublished) are very similar to the
map for the Eurosiberian species (Fig. 25).
The trends shown in passing from the European to the Eurosiberian species are
continued into the Eurasian element. 'rhe only shrub is Rosa pimpinellfoliu and the
only trees are two species (Salix tn'undra, S. uiminalis) which are doubtfully native in
our area; Meikle (1984) considers that S. viininalis is probably native to Russia but
has been carried westwards by basket-makers. The Eurasian element contains a
number of markedly continental spccies (Carex humilis, Inulu sulicina, Kolu rupestris).
Although no species are native to North America, over half of them have become
established there as introductions.
The Circumpolar element is the smallest of the Temperate elements, a markcd
contrast to the elements in the more northerly major bioine categories where it is
usually the largest. Some members of this element are only disjunctly circumpolar.
The ferns Asplenium ruta-murun'u, Ophioglossum uulgatum and TnehpteriJ pulustris are
absent from western North America whereas Alismu graminwn and Astragulus d a r k s
are rare or absent in eastern North America. Several Circumpolar species arc
represented in America by taxa which are distinct at varietal or subspecific level,
and are sometimes treated as separate species by some authorities, e.g. ;ThP&tPris
pulustris var. pubescens and Mburnum opulus subsp. tn'loburn.
??ie Southern-temperate elements (81-86)
These elements contain 296 species, more than any other group except for the
Temperate elements. The species are widespread in the British Isles but: not
unexpectedly, have a more southerly hias in Britain, being most frequent in southeast England, rather coastal in Walrs and north-west England and v e r y sparsely
distributed in upland Scotland (Fig. 26). These trends are apparent in western
Europe from Iceland and Scandina\ia south to the Pyrenees (Fig. 8); the patterns
elsewhere are less clear, perhaps because of under-recording in southern and eastern
Europe, but the European maps demonstrate that the Southern-temperate species
are more frequent than the Temperate species in the hlediterranean region.
No species have been classified in purely Southern elements, so the Southerntemperate and Mediterranean-Atlantic elements contain the British and Irish specics
with the most southerly ranges. Specics with very few temperate localities, such as
the Oceanic Southern-temperate Fumarau reuteri and the hlediterranean-Atlantic
Neotineu maculutu, could arguably bc assigned to separate Oceanic Southern and
European Southern elements, but thc presence of these plants in some temperate
stations, the obvious links with the Southern-temperate and Mediterranean-Atlantic
elements and the very small size of a n y such Southern groups all providc reasons
for adopting the system used here.
The 25 Oceanic Southern-temperate species have distributions with a more
southerly bias than those of the Oceanic Temperate species, but the distinction is
not always easy to make, especially for species with disjunct distributions. 'The
Oceanic Temperate species are usually confined to the northern part of the Iberian
peninsula, whereas the Oceanic Southern-temperate species extend further south.
Most of the Oceanic Southern-temperate species reach their northern world limit
in our area, the most marked exceptions being Anugallis tenellu, which extends north
to the Faeroes, and Eleogzton Juitans, which reaches southern Sweden.
Although there are difficulties in classifying individual taxa, the distinctly southern
and western distribution of the Oceanic Southern-temperate species in both Britain
38
C. D. PRESTON AND hI. 0 . HILL
and Ireland (Fig. 27) contrasts with that of the much more widespread Oceanic
Temperate species. Many of the Oceanic Southern-temperate species are plants of
‘Atlantic’ habitats such as heathland and damp grassland; the others are found in
a range of other habitats including coastal and inland rock outcrops, calcareous
grassland and ruderal sites.
Some of the Oceanic Southern-temperate species have markedly disjunct distributions in Europe. The ericaceous shrubs Daboerza cantabrzca and Emu engena are
absent from Britain, and their Irish localities are at least 900 and 1 100 km respectively
from the nearest populations in mainland Europe (Webb, 1983). There are similar
disjunctions in the Oceanic Temperate species Erica mackazana (1 100 km), Pznguzculu
grandy70ra (950 km), Saxzjiaga hzrsuta (950 km) and S. sjxzthularzs (900 km). Explanations
for these remarkable distributions have oftcn been proposed. Many authors, from
Forbes (1846b) to Webb (1983), have favoured the theory that species such as
Daboecza cantubnca and Enca mackazana are relics which survived the last glacial period
in or near their present sites. This theory cannot explain the distribution of the
frost-sensitive Enca engena: Webb (1983) was at a loss to account for this disjunct
distribution, but Foss & Doyle (1988) have suggested that it might have been
introduced to Ireland in the 15th century by traders or pilgrims travelling between
the Iberian peninsula and Ireland.
Many Oceanic Southern-temperate species are endemic to western Europe, or
to western Europe and nearby regions of north-west Africa and Rilacaronesia.
Ranunculus tnpartitus has recently been dircovered in the Aegean island of hlykonos
(Dahlgren, 199l), which suggests linkr with the Mediterranean-Atlantic element,
and Anagallzs tenella has isolated localities in Greece and Crete. The Atlantic
distribution of Eleogztonjuitans is mere11 a part of a widespread range in Africa, Asia
and Australia.
The 54 Suboceanic Soutliern-temperate species vary from those which are rather
generally distributed in the Temperate and southern zones of western Europe (e.g.
Azra praecox, Cirszum tuberosuni, Scutellaria minor) to species which occur along the
Atlantic seaboard and also occur in the western Mediterranean, often reaching Italy
and sometimes with outliers in Greece (e.g. Asplenzum marznum, Oenanthe larhenalti,
Parapholzr rtrgom). This last group clearly approaches the distribution of the Mediterranean-Atlantic species. In Britain the Suboceanic Southern-temperate species
are widespread in England and Walcj, but les5 frequent in Scotland and Ireland
(Fig. 28).
The European Southern-temperate element includes 106 species. Some of these
have northern limits in southern England or at similar latitudes elsewhere in Europe
(e.g. LactuLa raltgna, Lzthospermum purpurocaeruleum, Eolu kztazbelzanu), but many others
extend northwards to southern Scandinavia. Some species are synanthropic weeds
which have spread northwards to central Scandinavia (e.g. Fumana ofirinah, P a p a w
rhoeas, Raphunus raphanzstrum). The European Southern-temperate species are widespread in Britain arid Ireland, and the coincidcnce map (unpublished) is similar to
that of the Southern-temperate elements as a whole (Fig. 26). Most are species of
dry, open habitats, including rocky open ground, coastal and weedy communities,
or are plants of marshes and open water. The element includes some very common
species which arc frequent throughout the British Isles (e.g. Holrus lunutw, Hypochaeris
radzcuta, Juncur efusus).
Only a few members of the European Southern-tempelate elements are native
to North America: most of thcse aquatics or wetland plants such ar Equzretum telmateza
FLORISTIC: CLEhIEN I 5 Ih BRI I 4IN AND IREL\ND
3C)
(confined to western North America, where it is subspecifically distinct), Lrtnna gibba
and Ludwigia palustris. However, many species have become naturalized in North
America or throughout the circumpolar Southern-temperate zones.
The 8 1 species in the Eurosiberian Southern-temperate element have a similar
distribution in the British Isles to the European Southern-temperate element, and
thus to the Southern-temperate elements as a whole. However, the coincidence map
for the Eurosiberian species (unpublished) shows a more marked reduction in
numbers in upland areas compared with that for the European species, similar to
the difference between the European and Eurosiberian Temperate species noted
earlier. The Eurosiberian species grow in a similar range of habitats to the European.
As with the European element, many species have become naturalized outside their
native range: at least 50% are well established in North America and some 25%
are naturalized throughout the northern hemisphere.
The Eurasian Southern-temperate ( 1 7 species) and Circumpolar Southern-temperate elements ( I 3 species) are small. Like the European and Eurosiberian species,
many Eurasian species have become naturalized outside their native range and our
representatives include some which are only doubtfully native in our area. hlost of
the Eurasian species are weeds or wetland plants, and the Circumpolar list is
dominated by wetland species. Many of the latter are found not only in the northern
hemisphere but in the tropics or southern hemisphere as well.
‘Three Mediterranean elements are recognized: hlediterranean-Atlantic, Submediterranean-Subatlantic and Mediterranean-montane.
The Mediterranean-Atlantic element is one of the most distinctive of the floristic
elements in Britain and Ireland and the ‘Mediterranean’ or Mediterranean-Atlantic
species, like the ‘Arctic-Alpines’, have been recognized for many years. The 69
species in this element are widespread in the hlediterranean region and extend
northwards along the Atlantic coast (Fig. 9). They are absent from central Europe.
Many reach their northern world limit in our area, although others extend further
north along the Atlantic coast to southern Scandinavia. In Britain the species are
concentrated in the south and west; in Ireland, where they are less frequent, they
are concentrated on the south and east coasts (Fig. 29).
The Mediterranean-Atlantic species include some plants which are frequent ori
undisturbed, sandy Mediterranean heaches and find a more precarious niche in
similar habitats on the stormy Atlantic coasts (e.g. Euphorbiapeplis, Polygonurn triaritiniurn,
Otanthus maritzmus). Other coastal spccies, including plants of sea-cliffs and salt
marshes, are more widespread in Britain and Ireland (e.g. iltripler portulaioides,
Crithmum maritimum, Limoniurn vulp-are). Many of the Mediterranean-Atlantic species,
however, occur inland in the Mediterranean region, but tend to become restricted
to the vicinity of the coast towards the northern edge of their range (e.g. Ophioylossurn
lusitanicum, Pobcarpon tetraphyllum, Rornulta iolumnae). Some of the more widespread
species have a south-western distribution in the British Isles (e.g. Po&mhn canzbricum,
Rubia peregrina, Umbilicus rupestris)and two species, Asplenium onopteris and Arbutus u~zedo,
are confined to Ireland. Arbutus unedu is known from pollen analysis to he a native
species which was formerly more abundant and probably more widespread (Mitchell,
1993). Other Mediterranean-Atlantic species have a southern or south-eastern
40
C. D. PRESTON AND hi. 0. HILL
distribution in Britain, and are rare or absent from Ireland. These include a
few species of seasonally flooded habitats or permanent water (Callitriche tmncata,
Damasonium alisma).
The Mediterranean-Atla.ntic element includes a high proportion of therophytes,
many of them winter annuals. The native populations of some of these species,
including Gastridium ventricosum, Tifolium incarnatum and vU&a ciliata, show marked
fluctuations in numbers in response to fluctuating climatic conditions at the northern
edge of their range (Martin & Frost, 1980; Lovatt, 1981; Stewart et al., 1994).
Most of the Mediterranean-Atlantic species have a distribution centred on the
Mediterranean basin. However, there are a few which are more widespread, and
are found elsewhere in the northern or southern hemispheres (e.g. Adiantum capillusveneris, Juncus acutus, Polygonum maritimum). Other Mediterranean-Atlantic species have
also become naturalized outside their native range and a few are only doubtfully
native to the British Isles. These include species which may be ancient relics of
cultivation (Allium ampelopasum and possibly Erodium moschatum) and Matthiola incana,
which is almost certainly a naturalized garden plant (Rich, 1991). The native
distribution of Z'om'lisnodosa, as interpreted by Tutin et al. (1 968-80), is MediterraneanAtlantic but the species has become naturalized in Central Europe and now has an
approximately Submediterranean-Subatlantic distribution.
The 47 species in the Submediterranean-Subatlantic element ha\re a broader
distribution in the Mediterranean and Temperate zones than the strictly Mediterranean-Atlantic species. They characteristically have a northern limit which
extends diagonally from northwest to south-east Europe, often talung in the southwestern part of central Europe. Some of these species (e.g. Daphne laureola, Luzula
forstem) are much rarer in the true Mediterranean region than in the Submediterranean
areas to the north. Some have populations extending eastwards into central Asia.
The Submediterranean-Subatlantic species are much more widespread than the
Mediterranean-Atlantic species in the British Isles (Fig. 30), where they have a
markedly southern distribution in Britain and a somewhat less markedly southern
distribution in Ireland.
There is a very small group of six species which have montane distributions in
southern Europe and the Alps, where they are characteristically found in dry, rocky
habitats, but which descend to lowland habitats towards the northern edge of their
range. Helianthemum canum, for example, occurs in calcareous montane grassland in
southern Europe and north Africa and at disjunct localities at lower altitudes further
north (Proctor, 1956). We have been able to accommodate these only by placing
them in a category of their own, the Mediterranean-montane element. The flora of
the mountains of southern Europe is very rich and contains many European
endemics; like the species ofthe Alpine orobiome of central Europe, very few species
extend north to the British Isles. All six Mediterranean-montane species are rare in
Britain, where they have been well-studied (Proctor, 1956; Pring, 1961; Gilbert,
1970; Kay & Harrison, 1970; Wilson, Whittington & Humphries, 1995), but only
Helianthemum canum is found in Ireland. All tend to grow in open, well-drained and
usually base-rich rocky habitats.
Species with a continental distribution
Species which are surprisingly rare in the Oceanic zone of Europe are described
as 'continental'. We have not placed these in a single floristic rlemcnt but regard
FLORISTIC EIX1\IEhTS IN BKI1:IIN AND IREIANI)
41
'continentality' as a trait which might !in theory) be shown by species bvhich belong
to any of the floristic elements except thosc which require the species to be fi-equent
in the Oceanic zone (i.e. the Oceanic, Suboceanic and Mediterranean-Atlantic
elements). The definition is essentially one which depends on relative frequency
rather than presence or absence (an absolutely continental species would not occur
in Britain or Ireland). There is, therefore, a limit to the extent to which continental
species can be identified from small-scale distribution maps, and it might be possible
to refine the definition using statistical criteria once Lgridmaps are available for the
plants of western Europe.
Most of the 96 continental species bclong to the Boreo-temperate and Temperate
elements; few are Boreal-montane or Southern-temperate. The concept is not
especially valuable for Arctic-montane slxcies as there is riot much habitat for them
in the Oceanic zone anyway, but A/o,twcrrru.\ borealis (absent from Scandinavia and
known only in Europe from Spitsbergcn, Arctic Russia and the Urals) and Oytropib
campe.rtri.r (which has a predominantly castern distribution in Scandinavia) are
continental Arctic-montane species, and L-lllium.schoPnoprasum and (f'areu chordorrhiza
may perhaps be described as continental Rorco-arctic species.
The British and Irish distribution of the continental and the oceanic species in
all floristic elements is compared in I;igure 31. The scattered distribution of the
continental taxa, with the only marked concentration of plants in thr Brcckland
and adjacent chalklands of East Anglia, forms a striking contrast to the widespread
distribution of the oceanic plants. The habitat preferences of the continental species
are discussed below.
The distribution of Raunkiaer life-fi)rms in each of the major bionic categories
is detailed in Tables 8 and 9. Life-forms foi the species are taken from thr Ecological
Flora Database (cf. Fitter & Peat, 1 Wl), although we 1ial.e seprnted bulbous
geophytes and nanophanerophytes from other geophytes and phnnerophytc.s, nnd
made minor additions and corrections. Species which mny exist as t i t o life-forms
have been scored as 0.5 under each catcqory, a i d the figures in Table 8 nrc rounded
to whole numbers.
Most species in the British and Irish flora arc hemicryptophytes. althouqli thcy
are outnumbered by therophytes in thc more southerly elements (Jt'ide-temperate,
Southern-temperate and Mediterraneaa). Thc chamaephytes make up a significnnt
proportion of the more northerly rlrments. The rel&~cly high proportion of
helophytes and hyclrophytes in the Boi cnl-niontnne and Boreo-temperate elements,
and of hydrophytes in the Wide-tempcmtc elcrnents, is notew ortliy. Most of the
larger phanerophytes, not surprisingly, ar e Temperate species. The bulbous geoph) tes
are concentrated in the coldest and warmrst lioines. Other geophytes arc M.idesprcnd
and there is a particularly high pcrccntage in the Boreal-montane elements, ti\ othirds of which are orchids.
C. D. PRESTON h Y D hf. 0. HILL
42
TABLE
8. Number of species in each Raunkiaer life-form for the major biome categories (MBC)
MBC
Life-form
cham
1 Arctic-montane
2 Boreo-arctic Montane
3 Wide-boreal
4 Boreal-montane
5 Boreo-temperate
6 Wide-temperate
7 Temperate
8 Southern-temperate
9 Mediterranean
Total
epip
gh
3 2 7
2
14
17
1
1
-
-
-
-
1
0
I
9
7
-
5
1
23
135
helo
-
4
3
1
14
24
1
1
15
22
1
19
14
13
116
-
33
20
go
2
33
11
1
93
hemi
3
5
hydr
12
20
7
28
20
3
91
10
211
97
3 1
592
mnip
3
-
5
-
-
1 9 1
4
1
34
Ill
np
5
II
-
1
8
~~
~
25
7
34
3
2
48
-1
60
ther
Total
4
2
79
38
19
103
233
34
557
296
122
1481
-
8
20
14
104
117
53
322
The life-forms are as follows: cham, chamaephyte; epip, epiphyte; gb, bulbous geophyte; go, other geophyte:
helo, helophyte; hemi, hemicry-ptophyte; hydr, hydrophyte: np, nanophanerophyte; mmp, micro- and megaphaiierophyte; \her, therophyte.
TABLE
9. Percentage of species in each Raunkiaer life-form for the major bionic categories (MBC).
For key to life-forms, see Table 8
MBC
Life-form
cham
epip
1 Arctic-montane
11
2 Boreo-arctic Montane
18
11
-
3 Wide-boreal
4 Boreal-montane
5 Boreo-temperate
6 Wide-temperate
7 Temperate
8 Southern-temperate
9 Mediterranean
Total
11
7
7
8
-
I
2
-
-1
9
<1
2
-
5
8
5
-
~
i l
helo
-
3
-
1
6
I
3
~
-
go
gb
1-1
9
3
9
5
11
8
5
5
I4
10
6
6
4
1
6
hemi
4
4
hydr
-
0
1
7 4 5
32
12
48
9
’ 3 0 1 9
- 1 3
5
3 2
7
2 6
2
4 0
6
lip
mmp
ther
Total
-
5
5
I00
100
100
4
3
-
5
4
I
4
5
2
3
-
4
1
6
2
3
-
8
9
41
19
1 0
19
22
100
I00
I00
I00
100
10u
100
Habitat preferences
Ellenberg et al. (1991) assign the species of Central Europe to eight broad habitat
categories, excluding only some species with very broad habitat tolerances. We have
used this classification to illustrate the habitat preferences of species classified by
major biome category (Tables 10, 11) and eastern limit category (Tables 12, 13),
and for the continental species in all elements (Table 14). The habitat is that
occupied by the species in Central Europe; it is not always appropriate to the British
Isles but this does not matter as we are considering the habitats in relation to the
broader range of the species. A more serious drawback is the high proportion of
Oceanic and Mediterranean species which are absent from Central Europe and
therefore from Ellenberg’s lists. We have attempted to classify these species in those
groups where fewer than half of our species are classified by Ellenberg et al. (1991),
and figures including our classifications are presented separately (in italics) in the
tables.
FLORISTIC: E:I,EI\IEN I X I N HKI IAIN AND IRELAXI)
13
TABLE
10. Numbers of species in each major Iiiome category (hIBC) according to Elleiihcrg habitat
ratcgories
hIBC
1
1 Arctic-montane
12
2 Boreo-arctic- llontane
3 \Vide-boreal
12
2
33
59
6
72
?
I
I
4 Boreal-montane
5 Boreo-temperate
6 \Vidc-tcniperatc
7 Tcmpcratc
8 Southern-trinpPrate
9 \lediterrancan
Y Jludzfmnneari
Total
2
1
7
228
3
4
Yi
1
I
5
7
2
2
I5
2
3
I
I
2
(5
7
I1
210
5
I8
58
1
I3
1
22
12
3
I
13
(i
3
6
20
'3
117
44
27
8
I
4
2(iO
(is
37
8
-
3
I
li
10
11'3
1 IJ2
18
*
8
-
Iti
8
26
17i
-
9
T.~BLE
11. Proportions of species in each major l)ioiiie catrgoi? (1LIKCl) according to Ellrnlicrg 1ial)itat
catrgorirs, ignoring thosr species not classified I)) 13Irnl)rrg rf a/. (19!>1). For kr)- t o 1ial)itat catrgories.
see Tablc 10. Thc line in italics includrs yircics assigncd to thr hahitat catcgorirs by its
hlRC
The high proportion of aquatic and \\.ctland spccies in the Boreo-arctic and Borcalmontane elements is demonstrated in Taldc 1 1, as is the significant representation of
coastal species in the Wide-boreal and \Vide-tcmpcratc elements. Tht. concentrations
of plants of alpine habitats in the Arctic-montane elements, coniferous \voodland in
the Roreo-arctic arid Boreal-montane &merits and broadleaved \.\.oodland in the
Temperate elements are not surprising-. 'The largest habitat category in the N'icletemperate, Southern-temperate and hlctliterranean major Iiiome categories is that
of disturbed places, but this appears to IIC a particularly heterogeneous category
which includes not only weedy species (\\-liiclidominate the \Vide-temperate clcmentsj
but also plants which (in Britain, at least) are typical of shallow, droughted soils ( ~ . g .
Juncu.~rapitatus, Omithogalum angustfoliuni) and many species of seasonally-Noodcd sites
C. D. PRESTON AND M. 0. HILL
44
TABLE
12. Eastern limit category (ELC) in relation to Ellenberg habitats. For key to habitat categories,
see Table 10. The line in italics includes species assigned to these categories by us
Ellenberg habitat category
ELC
1 Oceanic
I Oceanic
2 Suboceanic
3 European
4 Eurosiberian
5 Eurasian
6 Circumpolar
Total
1
8
19
18
54
41
16
91
228
2
3
4
5
6
10
6
28
24
1
6
8
23
4
4
23
63
13
27
26
146
66
2-k
30
305
1
4
30
7
17
7
3
14
58
101
93
25
20
269
7
3
5
21
-
2
16
17
8
3
47
*
8
117
31
36
66
35
18
36
308
100
3
4
30
8
1
18
15
177
26
Total
159
159
142
526
297
107
250
1481
TABLE13. Eastern limit category (ELC) in relation to Ellenberg habitats, presented as proportions.
Species not assigned to a habitat by Ellenberg P t a/. (199 1) are disregarded. For key to habitat categories,
see Table 10. The line in italics includes species assigned to the habitat categories by us
Ellenberg habitat category
ELC
n=
1
2
3
4
5
1 Oceanic
42
128
I06
460
262
89
214
1 I73
19
15
24
30
6
4
3
15
22
23
22
35
28
9
23
2
5
7
5
2
4
II
5
31
21
25
32
25
27
I Oceanic
2 Suboceanic
3 European
4 Eurosiberian
5 Eurasian
6 Circumpolar
Total
17
II
I6
18
43
19
4
7
5
14
6
7
Total
7
100
100
LOO
I00
2
-
.Y
-
4
2
3
6
9
I
2
20
22
II
1
9
8
7
15
I
4
26
8
2
1on
100
100
1 no
TABLE
14. T h e habitats of species with a continental distribution, in categories defined by Ellenberg
et al. (1991). T h e percentage of continental species in each habitat category is compared to the
percentage for all species, ignoring species not assigned to a category. For key to habitat categories,
see Table 10
Ellenberg habitat caregoq
Number of continental species
Continental species ( O h )
ALL species ("/a)
1
2
3
4
16
1 9
0
0
5
8
9
23
1
1
5
19
5
38
4
5
26
6
7
8
*
Total
3
4
4
4
5
2
14
17
12
96
100
15
100
(e.g. Limosella aquatica, Mentha pulegium). Species in the heath, meadow and pasture
category show rather few trends.
The wide longitudinal range of species of alpine habitats, conifer woodland and
aquatic and wetland sites is apparent in Tables 12 and 13, as is the restriction of
most of the plants of broadleaved woodland to the European area. The species with
a continental distribution are predominantly plants in the heath, meadow and
pasture category; plants of disturbed places are under-represented and there are no
15
FLORISTIC CLEMEN r4 I N BRITAIN AND IRELAND
TMLE15. Altitudinal upper limit in relation to major biome category (hlBC). Valucs are nurnbers of
species in each MBC, grouped in 300 m uppcr-altitudinal bands, or for which the maximum altitudc
was not known (*). Mean maximum altitudes \\\%recalculated for all species for which a numerical
maximum value was aailable; the numbers of such species in) are lower than the sum of the first five
rows, because many species were known to he rrstricted to the lowlands (1300 in) but lacked a prrcise
maximum altitude
Maximum altitudc
0-299 m
300-599 m
600-899 rn
900--I I99 m
1200- I343 m
*
Total
hlean (in)
SD [ni)
SE (m)
I
2
1
3
12
36
26
1
79
1066
'214
21
77
n
3
6
2
6
18
4
2
5
1
6
7
8
9
178
82
23
3
-
10.5
I1
'Total
3
2
I!)
33
9
I!)
I4
230
203
I
7
6
27
6.1
5.5
35
1
61
4
10
2
30
2
I6
233
1
31
10
t
584
400
I91
I80
i3
73
3.1
5.57
296
I22
1181
560
338
63
21)
464
250
13
38.5
371
233
236
570
178
3-1-0
IR
I66
26
II
Io:?8
-
38
19
913
308
54
32
961
3.56
89
16
27
:i
8
IO'i
714
32:
'i.3
Hi
68.5
1315
22
200
1
48
TABLE
16. Lowest altitudes of species in relation to major bioine categories. Values arc numbers of
species in each category; the altitude class
*
is used for specics lacking lo\vest-altitude data
5lajiir. Iionir category
Loivrst altitude
1
2
3
t
0-99 111
100-299 rll
300-5119 m
600-899 m
900-1 199 rn
17
1.5
15
45
6
20
17
I
6
5
-
?
1
-
'I'otal
18
79
11
38
*
3
2
1.54
I
6
7
30
-
315
-
188
-
$1
60
7R
4
233
31
-
21 1
557
-
108
296
Total
867
23
27
-
t9
l0:i
1
-
4
19
8
62
122
18
I
j4.i
1481
coastal continental species. The ecology of the continental species was discussed by
Walters (1953), who pointed out that they tended to be plants of open habitats and
base-rich soils.
Altitudinal limit3
It is to be expected that the major biomes in which species are found should be
reflected in their altitudinal range in Britain and Ireland. We compiled altitudinal
data from a variety of sources, notably Wilson (1 956), the Ecologzcal Flora Database
(Fitter & Peat, 1994), Stewart et al. ( I 9Y4), and, for aquatic plants, Preston & Croft
(1997). In our database, upper limits are available for about 95% of species (Table
15), subject to the proviso that for many species known to be restricted to low
(<300m) altitudes, no precise value is available. Lower limits are less well known,
with values for only 63% of species (Table 16).
46
C. D. PRESTON .4ND hf. 0. HILL
TABLE
17. Arctic and Boreal species (major h o m e categories 1-4) that are confined to low (<300 m)
altitudes
Species
Elemriit
(a) Coastal specks
Atnplex longpa
Atnplex praecox
Rlysniu rufus
Carex ruantima
Corm recto
Ei~phrsiafuulaerisi.,
Honrkmya pep1uide.s
L.ut/ywja,bnoicuJ
&qynius orenanus
Ligwtimm sroticiini
.\fp/tpnsia mantima
Trijleuro sprnnwrn mantimum
(b) Inland specics
41liuni schoeiiuprasum
Cnre.r busbaiimii
13
13
13
16
11
11
36
21
23
2'3
23
16
Specics
Element
26
43
15
16
16
46
16
16
13
42
13
16
4 '3
41
36
44
26
Upper altitudinal limits show a clear trend from the mainly lowland Mediterranean
(MBC 9) species to the Arctic-montane (MBC 1) species, whose mean upper limit
exceeds l000m (Fig. 32). A slightly unexpected result is that whereas the Boreoarctic Montane category (MBC 2) has its average limit about halfway between those
for the Arctic-montane (MBC 1) and Boreal-montane (MRC 4) categories, the Wideboreal category (MBC 3) has on average a higher average maximum altitude than
the strictly Boreal-montane category (MBC 4). The difference is less than one
standard error and is not statistically significant.
Many of the arctic and boreal species that are confined to low altitudes are either
coastal or are plants of wet places (Table 17). Three lowland species (Epipogium
upbllum, Moneses ungora, Pobgonutum aerticillutum) are Boreal woodland plants and one
(Polygonum boreule) is a Boreal weed. The one remaining species, Allium sckoenoprusum,
is partly coastal, and has a southwestern British distribution (Stewart ut al., 1994)
that is totally anomalous fbr a Boreo-arctic Montane species. It has a Continental
distribution in Europe, contrasting sharply with the few other Continental species
in MBC 1 and MBC 2, which arc montane in Britain. Perhaps its ecological
requirement to be bonc-diy in summer and wet in winter is hard to achieve in the
wet climates of northern Britain.
Two Temperate species, Ericu rinprea and Gulium saxutile, reach altitudes above
1200 m (Table 18), and thc Submediterranean-Subatlantic Geranium lucidurn reaches
760 m. All three species occur in western Norway, but their occurrence at relatively
high altitudes in Britain would not be expected from their world distribution. The
presence of Erica cinerea in this group is based on a single record, made by an
inexpert botanist, of very stunted plants growing at 1220 m 011 bare rocky ground
in the Grampians (Stewart, 1891). This record must be regarded as very doubtful
as the next highest occurrence is at only 790m, on Purple Mountain, Co. Kerry,
and even at this altitudc the species is very stunted (Scully, 1916).
Lower altitudinal limits for the Borro-temperate, Temperate, Southern-temperate
and Mediterranean species (MBC 5-9) are almost all below 100 m (Table 16). The
FLOKIS I IC: ELEXIENT 5 Ih BKI 1 1IV .\NI) IKtIY\ND
TABLE
18. Species belonging to 'Tetnperatc and Southern eleincrits (major
biornc categories 7-9) that ac1iin.c an unusually high altitridiiial limit
Species
Element
hlaxirnum altitude
En(n rinerea
71
1220
Galiuni saxatile
Curdamine hmuta
L o k r rornrculatus
Rnnunru1u.c oniiopl&\
Gwanium luridurn
H y p r i ( u m andro.\armum
72
1310
84
I I60
915
85
82
92
92
(ni)
1005
760
635
only exceptions are Lychnzs uzscanu and AZltconopsir cambrzq and the latter descends
to lower altitudes as an escape from cultivation. The highest lower limit is that of
Saxzjzga rzzduns, a Circumpolar Arctic-montane species not found below 9 15 ni.
The boreal Cicerbzta abznu has not been rccorded below 530 m, and it may be extinct
at its lowest sites, but it may have been driven relatively recently to this high altitude
by intense grazing pressure (Marren, Payrie & Randall, 1986).
In summary, while there is a marked tendency for Arctic-montane species to
occur at high altitudes and for Temperate, Southern-temperate and hlediterranean
species to avoid them, Boreal-montanc and Borco-temperate species occur at a wide
range of altitudes and are not, as group, lirnitcd altitudinally in the British Isles.
Ellenbergi indicator tlaluw
The floristic element assigned to each species by us has been compared with the
indicator values (zeigerwerte) for temperature (T) and continentality (K) ascribed to
that species by Ellenberg el al. (1 99 1). 'I'he temperature values (Zmptrutur:ahlen)
range from T = 1 (cold-indicator, only in high mountains, i.e. in the alpine and
nival zones) through T = 5 (temperate-submontane)to T = 8 (submediterranean)and
T = 9 (extremely warm indicator). Likewise the continentality values (Kuntinentalitatzahlen) range from K = 1 (euoceanic, scarcely reaching Central Europe)
through K = 5 (intermediate, from weakly suboceanic to weakly subcontinental) to
K = 9 (eucontinental, absent from westcrn Central Europe and rare in the east). No
British species is rated K = 9, but two species, Lpidium latfolium and Ruppia marititnu,
are rated K = 8 . Neither of these is especially continental and these K values are
probably erroneous.
Contingency tables (Tables 19, 20) relating Ellenberg's and our categories show
a broad level of agreement between his T values and our major biome categories
(MBC); agreement between his K values and our eastern limit categories (ELC) is
less good.
About 17% of species in the British Hora have not been given Ellenberg indicator
values, mainly because they do not occur in Central Europe. Most of the species
without Ellenberg values, amounting to 1 1 O/O of the British flora, are Oceanic or
Mediterranean-Atlantic (ELC 1; Table 20). A smaller category, 3% of the British
C. D. PRESTON AND hi. 0. HILL
48
TABLE
19. Numbers of species in Ellenberg temperature (T) categories in relation to the major biome
categories (MBC) defined in this paper. Values in italic correspond to the species with unexpected T
values, listed in Table 21. The Ellenberg category x signifies wide temperature amplitude; * siLpifies
species not included in the Ellenberg enumeration; mean values Tor T are taken over rows 1-9
hlajor hiome catcgory (hIBC)
1
Ellenberg T
1
2
3
4
5
6
7
8
9
6
3
7
8
I
I
21
15
55
I
24
1
140
16
59
17
29
9
3
2
62
122
I
Mean 'Y
SD
SE
n (rows 1-9)
2.2
0.9
0.12
52
9
2
4
Total
*
4
11
24
13
I
26
79
S
3
5
I0
1
19
I0
38
3.7
0.29
24
98
22
103
233
10
4.3
5.3
0.9
0.9
0.12
0.08
63
125
5.8
0.7
0.2.5
8
1.4
17
13
2
34
13
42
296
77
36
557
5.7
0.5
0.10
19
5.8
0.7
0.03
414
6.3
0.8
0.05
241
6.9
1.0
0.14
58
Total
12
30
30
78
227
483
1 k7
28
3
216
21 1
1481
5.6
1 .3
0.01
1033
TABLE
20. Numbers of species in Ellenberg contincntality (K) categories in relation to the eastern limit
categories (ELC) used here. Values in italic correspond to the species with unexpected K values, listed
in Table 22. ? signifies species for which T (cf. Table 19) is given but whose K value is missing; x
signifies wide continentality amplitude; * signifies species not included in the Ellcnberg enumeration;
mean values for K are taken over rows 1 8.
Eastern limit categor) (ELC)
Ellenhrrg K
3
4
5
8
7
27
5
1
I32
179
I00
52
3
4
4
6
2
29
11-2
36
6
.i26
297
2..3
0.9
0.09
I07
3.2
I .0
0.05
470
1
2
26
5
6
I I
70
I8
5
1
I
1
-
-
X
3
2
*
111
Total
I59
Mean K
SD
SE
n (rows I - 8 )
I .8
I .3
0.20
41
14
I 0I
37
73
15
17
I
3
36
4.2
1.3
0.08
252
21
3
15
-
3
23
1
I07
4.5
1.7
0.19
80
6
Total
I
'38
4
.
5i
10
47
6
30
I
6
64
26
226
386
I59
I96
30
67
2
23
141
21 I
250
1481
4.6
I .6
0.13
I54
3.6
I .5
0.04
1104
flora, consists of Arctic and Boreal species which are scarce or abscnt in Central
Europe. The remaining 3% is made u p of Suboceanic species (2%) and a miscellaneous category (1 ' i n ) .
The mean T values for each major Iiiotnc category show a general increase from
Arctic-montane (hfBC 1, mcan T = 2.2) through Boreal-montane (RIBC 4, mean
T = 4.3) and Temperate (MBC 7, mcaii T = 5.8) to Mediterranean (1LfRC 9, incaii
T = 6.9). Modal values for these categorim arc T = 2, T = 4,T = 6 and T = 7. T h e
Ellenberg wide-amplitude category ‘I‘ = x is particularly well represented foi- the
categories hIBC 5 (Boreo-temperate) and MBC 3 (Wide-boreal). Because ‘r= s is
an unspecified wide-amplitude categor), it has to t x treated as a missing \ d u e when
calculating means.
Iblean values of T for the composite categories, Boreo-arctic hlontanc (hIBC 2).
Wide-boreal (MBC 3), Boreo-temperate (h,IBC 5) and Southern-temperate (RIKC:
8), arc not significantly different from the averages of thc incaiis for the major
biomes that make them up. On the other hand, the mean ‘I for hlBC 3 is 5.8.
which is the same at that for the Tempcratc clcments with MBC 7. Givcn that the
\.Vide-boreal cleincnts include a high jirojiortion of coastal species, \vhicli are $so
fucto not montane, the occurrence of a tclativcly high mean T is understandable.
The mean values for K in the eastc*t-nlimit catcgorirs show a similar h i t less
clear progression from Oceanic plus R Icditcrraneaii-Atlatitic ( E K 1 iiiraii K =
1.8), through to Circumpolar ( E I C 6, incan K=4.6; ‘Table ‘LO). l h c less clear
progression reflects the fact that in cotitincntal Europe the kvw-cast gradient in
floristic composition is wcakcr than the nortli-south o m . .4nadditional factor is that
the definitions of K values for K 2 6 sp,cify that the frryucncy dccwases from c
to west in Central Europc. Given t h a t the castern range limits for Eurosibcrian
(ELC 4), Eurasian (ELC 5) and Circrinipolar (ELC 6) lic a loyg way t o thr rast of
Central Europe, it would be surprising if’ diih-enccs Ix.twren thcir mean K \ . a l u c ~
were large. O n the basis of E1lenl)crg:’s dcfinitions, one ~vould expect a closc
correspondence bcttwccn K = 7 and o i i r Coiitinental catcgor).. N o such closc cot.respondence exists; indeed the proportion of’ sjxcics rated by us ;is (:ontincntal is
about 1”h for K = 3 , 1 7 % for K = 4 a i i t l K = 5, and 25% for I.;= (iand I( = 7.
.
C. D. PRESTON AND hf. 0. HILL
50
TABLE
21. Species with an Ellenberg T value which differs markedly from the mode for the major
biorne category (MBC)
Ellenberg T value larger than expected from hlBC
h4BC
T
Species
1
1
4
4
1
1
2
2
2
5
4
4
Carex norvegim
Cochlearia pyrenaica
ilrlinuartia stnrta
Sedum msea
Carex aquatiliJ
Lathyrus japonicus
Lymus arenariu.t
Pilostlla peleteriana
Cmtoptlyllum submersum
6
6
6
8
8
7
Ellenberg 1‘value smaller than cxpected from MBC
hlBC
T
Species
Carrx capil1ciri.i
Silme uii$ora
Alchemilla glabra
Phptuunia orbirulare
A’arcztrus pseudonarrissu~
Dyjopterir subniontnna
2
1
5
5
2
3
5
8
3
4
2
9
TABLE
22. Species with an Ellenberg K value which differs markedly from the mode for the eastern
limit category (ELC)
Ellenberg K value larger than expected from E1.C
ELC
K
Species
1
4
1
4
5
6
5
Beta vuigans
FeJtuca lemanii
Frstuca arenana
Glanczum flauum
Fzlago pyramidata
Koelena uallesiana
Rzbes rubrum
Hzppophae rhamnoidei
Pulmonana obsura
CLuercuJ robur
Malva neglerta
Aftnuartin r m n o
Ribes Jpicatuni
Silene otita
L.epzdium lat@bunz
1
1
2
2
2
3
3
3
3
3
3
3
4
5
7
6
6
6
7
7
7
7
8
Ellenberg K valuc sinnller than expected from ELC
ELC
K
Species
4
4
4
4
4
1
1
4
5
i
2
2
2
2
2
2
2
2
2
2
Ah.sma lanceolatum
Apiuiii gracrolrns
Arabis glabra
C a m otrnbae
Duc trlorhiza marulata
1.toelr.s lnrurtizs
1ioi$)i.s srtacea
Liiuiiirn alpzna
Arnbir petraea
5
2
5
i
5
5
2
2
2
2
6
1
6
6
2
2
6
6
2
2
Qmnadenia ronopJw
Herarlrum sphondylium
Po!ystzclium aculeahnz
17t,irularia australir
< o i t ~ i ( i nultii
Lqiinzarhia tlys$ora
Annma maritima
JUTU
u.\ baltzru.!
Tricliophurum respilo w n
<ostura manna
Cflrer elfltn
lowlands in western Europe. The two remaining mismatches are Sedum rosea, which
we consider to be truly Arctic-montane (not compatible with T=4) and Carex
capillaris which we treat as Boreo-arctic Montane (not compatible with T = 1).
The mismatches for continentality (Table 22) are not so simply explained. Only
four of the mismatched species, A h m a lanceolatum, Beta vulgaris, Festuca lemanii and
Pulmonaria obscura, are indicated as having doubtful K values by Ellenberg et al.
(199 1). Taxonomic differences or nomenclatural confusion can probably explain the
discrepancies for Ameria maritima, Carex otrubac, Festuca arenaria, Filago pyramidata,
Heracleum sphondylium, Juncus balticus, Trichophorum cespitosum and Utricularia australis.
Doubts about native range can explain the discrepancy for Ribes rubrum.
Of the plants for which K appears iincxpcctcdly large, tlie discrepancy for K i h
spzcatum and SzlenP otiter is more apparent thnn real; both s p e c k are gcnuincl)
Continental. O n the other hand, the vnlues K = 6, 8, 7 and G for Glouczuni /Iaz~um,
Lepidium lat!filium, Malva neglrcta and Qietru5 robur arc simply wrong. The remaining
three species, H$pophae rharnnozdP5, KoeGtiri r ~ ~ l l c ~ i and
n n a L41muaitza te(utTa, ha\ e dirjunct.
partly montanc distributions, and arc r‘ithrr c-lifficult to classifj A. zallcmzna could
fall into our Mcditcrranean-montanc cl(wicnt, except that it is alw found at lo\\
altitudes in the Mediterranean region.
Scveral plants for which K appears uncxpectcdl~sinall ha\ c dislunc tions or xc
v e q rare in Central Europe although thcy cxtcnd furthcr cast T h u s .Irahzi p e t t r i m
is disjuncti\.e, Isopter lacurtrzs and I6ron1i o a11)zna extend eastwards mainly i n northern
(not Central) Europe, and Isolepis wtaccw and Po!y,tzchum aculeatum cxtcnd \\ell t o the
cast but as rare plants. This Icavc$ a icsiduc of secwi spccics, Aiabic globin, (,arm
data, Dacglorhzza marulata, Gjmnadrnza r onopwa, Ipmachza thy?szfZoia, ZooJteici niatzna and
Z. noltzz, for which the Ellenherg K \ ‘due< appear to be too lo\\. l‘he value K = 1
for 1,. ttyszjorn is presumably due t o ‘I copying crroi; in Ellcnbcrg (19711) it is
accorded K = 7.
111 conclusion, Ellenberg’s <egcrwprtf 111c
x tde n valuable external check. In try iiiq
to rcconcile our elements with them, n t ’ werv led to re-eudminc the distribution of
scvcral spccics and made a fcw change\. Most rrmaining discrepancies hctw ccn the
,7rzgmwrte, hlRC and ELC can bc atti il)utrd ritlier to taxonomic difeieticrs or t o
Ellenberg’s Central European focus. ( Inly lor t\\o species do wc disagrer 11itli the
value of T and only for twelve species nit11 the \ alue of K
Matthews (1995) allocatcd 599 of tltr- 148 1 spccics classilied Iiy LIS t o floristic
elements; the remaining 882 species (GO‘’?, of the total) were allocated to onc‘ of the
groups hc defined but did not list ( l $ X e , F,urasian, European and Endemic) o r wcrc‘
not included in his treatment. The fcrns and f c w i allies were amongst thc cscludrd
species. The relationship of the floristic, rlerncnts defiiicd hy Alatthru~s( 1 955) t o
thosr recognized here is shown in ’l’alilt, 23. \/Ye have excluded sliwics classified I)?,
Matthews but now rcgardrd as aliens ur rcduccd to infraspecific tam. hIatthclvs’
elements are discussed individually in tlic fbllowing paragraphs.
hlattliews defined tlie Mcditcrrancaii clement as spccics with their cliiet’ centre
of distribution in the Mediterranean rcgion, and most of‘the spccics that he inclutlcs
in it are classified by us as r\leditcrratic,nn-Atlantic. The only cxceptions arc Filqo
pjrajrizduta (Suhmediterraiieati-Suhatlanric), Frankenia 1nezIi.r (Suboc-canic South(mitemperate, as it is restricted to tlie Lvcstcrii hlcditerranean). 1,znmriuni hrllid~diun~
(Eurosibe rian Southern-t cmper atc) a ntl Z’~~~~.so.~t,~rmiiit7
i.omubien.i P ( European Tc i n perate). Limoniuni bellzdzjdiunz has a l\lcditcrraiieati-Atlalitic distriliution but also
occurs inland on saline soils in eastcrn Ilurope, liencc its classification as Soiltlicrntemperate. I? cornuhierisp is one of tliv inorc southerly members of the Europcaii
Temperate element. It rcachcs its nortlicmi limit in soutliern England, Hun#nry and
south-central Russia (Tutin et nl., 1 YOU 80) and tlieref’orc lacks the c-haractrristic
N\Y-SE northern limit of the neditrri.ntican-i\tlantic species, and i n ‘I’urke), it is
an upland species \vliich occurs in Coy/ii,s scrub and on forest triargiiis ac ;tltitridcs
bet1\7een 500 and 1500 metres (Davis. 1 !
C . D. PRESTON AND M. 0. HILL
52
TABLE
23. Occurrence of the species in floristic elements defined by Matthews (1955) and in the
elements recognized in this study. The Matthews' elements of species with a continental distribution
are also given. The elements defined in this paper are listed in numerical form; for the names of the
elements, see the text. Matthews' elements are numbered as follows: 1, Mediterranean; 2, Oceanic
Southern; 3, Oceanic West European; 4, Oceanic Northern; 5, Continental Southern; 6, Continental;
7, Continental Northern; 8, Northern Montane; 9, North American; 10, Arctic; 1 1, Arctic-Alpine; 12,
Alpine; *, not classified by Matthews
Element according to Matthews
Element
1
2
3
4
5
6
7
8
9
10
12
II
*
Total
C h t i n e n t a l spccics
-
-
-
3 2 6 1 7
6
-
1
1
42
96
FLORISIIC ELEhlENl‘s IN BRITAIN AND IREIAND
53
The Oceanic Southern species occur “chiefly in southern Europe, including the
Mediterranean region and in western Europe” and are “essentially a south-west
European group” (Matthews 1955: 123). The Oceanic Southern species fall into a
range of our Southern-temperate and Mediterranean elements, from MediterraneanAtlantic (e.g. Damasonium alisrna) and Submediterranean-Subatlantic (Medicago arubzca)
to Suboceanic Southern-temperate (C,urduus tenuzjorus) and European Southerntemperate (Lotus angustissimus). The conspicuous exception is Pobgonum oxypermurn.
Matthews (1955) misclassified P raii as Oceanic Southern-it extends from northwest France to Arctic Russia (Tutin Pi a/., 1993)-and this species is now included
in the more broadly defined l? oxyApemzum which has a Wide-temperate coastal
distribution.
Matthews’ Oceanic West European element includes species with a strongly
western distribution in Europe. Almost all of these fall into our Oceanic or
Suboceanic categories. Most of the exceptions, which have European, Eurosiberian,
Mediterranean-Atlantic or even Circumpolar distributions, are plants which are
now more broadly defined taxonomically than they were when Matthews wrote, or
have since been discovered outside the oceanic zone.
The relatively small Oceanic Northern element was created by Matthews for
species which were chiefly characteristic of north-west Europe. They are spread
thinly in our Boreo-arctic, Wide-boreal, Boreal-montane, Boreo-temperate and even
Temperate elements. Four of the six l’emperate species (iltriplex laciniata, Cochlearia
anglica, C. danica, Seriphidium maritimum) extend from Portugal, Spain or south-west
France to sourthern Scandinavia and are not markedly northern in their distribution.
Matthews’ Continental Southern element contains plants which are found in
central and southern Europe, thinning out northwards. One would expect them to
belong to our Temperate and Southern-temperate elements and most of them do,
although Matthews’ element also contailis plants which we regard as MeditcrraneanAtlantic (e.g. Glauciumjauum, Pobcarpon teti-aphyllum)and numerous SubmediterraneanSubatlantic species (e.g. Bvonia dioica, Xuscu.r aculeatus).
Matthews’ Continental species are “characteristic of central Europe, thinning out
westwards, but frequently extending east through Russia into Asia”. Most are classified in our European and Eurosiberian Boreo-temperate, Temperate or Southcrntemperate elements; a few are Eurasian Temperate. The exceptions are the Circumpolar Wide-boreal Zphroseris integefiilia and the Suboceanic Southern-temperate
Bunium bulbocastanum and Moenchia erecta. B. bulbocastanum is an anomalous species in
having a Suboceanic distribution in Europe but, as Dupont (1962) pointed out,
being rare or absent in large expanses of the oceanic zone.
It is surprising that only 26 of thc 81 species in Matthews’ Continental element
are annotated as having a continental distribution in our classification. Some of thc
species we do not annotate are borderline cases which could be regarded as
continental by a less stringent criterion (e.g. Astragalus g(~q~phyllos,
L>$sacu.s pilom.~,
Scleranthus perennis). However, many arc frequent in the Oceanic zone of Europe and
some of these show little or no tendency towards an eastern distribution even in
Britain (e.g. Anagallis minima, Genista tinc-toriu, Quercus petraea,
robur). The Continental
element even includes species with a rather western distribution in Europe (e.g.
Tee.dalza nudicaulis). It is difficult to know why Matthews placed these species in the
Continental rather than the European rlement (species with a wide distribution in
Europe), and the composition of his Continental clement appears to be unsatisfactory.
The Continental Northern element is described by Matthews (195.5)as essentially
a
54
C . D. PRESTON .\ND k l . 0. HILI.
TABLE
24. Percentage of spccies in thc floristic elements which occur as natives in North . h c r i c a .
Figures are provided for only the Oceanic (Occa),Suboceanic (Subo),Europcan (Euro) and Eurosiberian
(Esib) eastern limit categories; species in the Eurasian category are by dcfinition rare or absent from
N. America and all Circumpolar species occur there. FiLguresfor elrnieiits \vhich have fewer than 10
species arc placed in brackets
h h j o r home category
Eastern limit c ategoq
Otca
Arctic-montane
Borco-arctic Montane
M’ide-horcal
Boreal-montanr
Boreo-temperate
Wide-temperate
Tcmpcrate
Southern-temperate
Mediterranean
Sub0
Euro
Esib
Total
71
83
100)
39
22
13
7
3
3
boreal; it comprises species which are found in central and northern Europe,
decreasing or becoming montane in the south and including many species which
have a more or less circumboreal distribution. Not surprisingly, most species fall
within our Boreal-montane and Boreo-temperatc elements, with smaller numbers
in the Boreo-arctic and Wide-boreal elements. There are, however, also 17 species
which we classify as Temperate. One of these (Qrhnzs uzscarza) cxteiids into the Boreal
zone and several are characteristically found in the northern parts of the Temperate
zone (e.g. AstragaluJ danzrus, Crassula aguatzca) but some have typical Temperate
distributions (e.g. Eleochan multzcaulzs, Hottonza palustrzs) and there are even two species
with a relatively southern distribution in the Temperate area (Hypunrum hzrsutum,
Srrophularza umbrora). The Continental Northern clement also includes one plant that
we classify as Eurosiberian Southcrn-temperate ( Z u c n u m seordtum).
Matthews’ Northern Montane clement is closely related to the Continental
Northern: it includes species of northern (rather than central and northern) Europe
which reappear in the mountains further south. One would expect that most of
these species would fall into our Boreal-montane elements and that proves to be
the case, with smaller nunibers of Boreo-arctic and Boreo-temperate species. iYe
classify Eola rupestrzs as Temperate although it is one of the more northerly of the
Temperate species; Walters (1953) pointed out the affinity of this species to those
in Matthews’ Continental element. ‘ l h c other non-boreal species in the Northern
Montane element is Potmtzlla rupstnr, which we place in the Mediterranean-montanc
element although it is the most northerly of the species in this small group.
The smallest of Matthew’ elements is the North American element. in which he
included six species with their main centre of distribution in North America, and
which are very restricted in Europe. One of these, JuncuJ tenuy is now recognized
as an introduction in Europe. Hyfieneum canadensf, excluded from this paper as an
alien in deference to Stace (1 99 1) arid Kent (1 992), provides another example if
Webb & Halliday’s (1973) arguments for regarding it as a native species are accepted.
‘The unusual feature of the distribution of the North American species is the
extreme asymmetry of their distribution (Heslop-Harrison, 1953). hlany apecies in
the Arctic and Boreal elements which have Oceanic, Suboceanic, European or
Eurosiberian distributions are also found in North America (Table 24). Most of
FLORISTIC ELEhIEN 1-5 Ih BKI I h I N AND IREIAND
55
these are widespread in both continents whereas the species of the North American
element are widespread in N. America but restricted to the Oceanic fringe of Europe.
Unlike Matthews, we have not created a special element for the predominantly North
American species. We classify the species in Matthews’ N. American element as
Oceanic Boreal-montane (Enocaulon aquaticum, Spiranthes romanzofiana), Oceanic Boreotemperate (Limosrlla austrulis, Potamogrton epilydrus) and Oceanic Wide-temperate
(Siyrinchium bemudiuna). The northern affinities of this group are not surprising in
view of the northern affinities of the entire amphi-atlantic assemblage. Whether the
distribution of some or all of these sprcies requires an explanation which differs
radically from explanations of other aniphi-atlantic distributions is a debatable point.
There are also species which are more frequent in Europe than in North America,
or present in Europe and Greenland but absent from mainland North America, but
it is particularly difficult to establish whether many of these are native to North
America.
The species of the Arctic zono- and orobiomes are divided by Matthews into three
elements, the Arctic-Subarctic, the Arctic-Alpine and the Alpine. These elements, as
represented in the British Isles, are closely related. Some species qualify as ArcticAlpine rather than Arctic solely because of the continued existence of one or two relict
montane populations, suggesting that the chances of post-glacial sunival may play a
significant part in determining which of these two distributions a species currently has.
The Arctic-Alpine and Alpine species are also linked by species such as Llqvdia .rerotinn,
which have montane distributions in Europe but are found in the Arctic elsewhere.
We have therefore included all these in a single group of elements, the Arctic-montane?
although the strictly Arctic and the strictly montane species are marked by appropriate
qualifiers. Most of the species included by Matthews in these three elements Fall into
our Arctic-montane or Boreo-arctic Montane elements, although there are a few which
we classify as Boreal-montane (e.g.ArctoJtaphv1o.cuva-ursi, Carex recta, Gcerbita alpina, Kubus
chamaemorus). Matthews’ Alpine element is more heterogeneous than the others and
also contains species which we regard as Oceanic Temperate (Saxfiaga hirsuta, S. spathularis) and Mediterranean-montane (Arab&.rcabra, Draba aizoides).
The above analysis suggests that hlatthews’ system and ours are in closest
agreement for the more extreme (and perhaps, therefore, more distinctive) floristic
elements in the British and Irish flora: thc species with markedly northern, western
and southern distributions. Matthews’ treatment of the less distinctive elements
appears to us to be unsatisfactory. His Continental Northern and Continental
Southern elements include species with predominantly central and northern, and
central and southern, distributions. However, the Continental clement does not
include, as one might expect, all species with neither a northern nor a southern bias
but only those which are ‘continental’ in the sense that they decrease in frequency
in western Europe. This restriction was perhaps applied to avoid the inclusion of
many wide-ranging temperate specics which he regarded as too widespread to be
of interest. In practice, he appears to have applied the criterion rather erratically.
Most phytogeo<qaphers have accepted the concept and have cited the truly ‘continental’ species when discussing this element: IYalters ( 1953) commented that
“perhaps the most-quoted example is Ifvonica .$icata”.
Young:s clu.,.+/ic.ution of!f‘Arctic.s,bccie.\
The Arctic was divided latitudinnlly b j Young (1971) into four zones from the
Low Arctic (zone 4) to the northernmost islands in the Arctic Ocem (zone 1). Young
C . 1). PRESTON AND X i . 0 . HILL
56
TABLE
25. Comparison of northern limit of Arctic species, as defined by Young (1 97 I), with occurrence
in major biome categories. Zone 1 is the most northerly of the arctic zones
N limit
hlalor bionic category
Arctic-montane
Zone
Zone
Zone
Zone
1
2
3
4
Borcw-arctic hfontanc
Widc-boreal
1
2
5
2
8
4
4
7
2
13
2
Boieal-montanr
Borco-temperatr
~
~
1
~
I
7
5
J
TABLE
26. The distribution of Euatlantic and Subadantic species, as defined
by Dupoiit (196‘4, in floristic elements. ’The floristic elements arc given in
numerical form; for the names of the elements, w e the text
F l o r i k clrmriit
41
Euatlantic
Suhatlantic
Total
1
1
2
51
1
I
2
71
1
7
12
2Y
72
81
3
I:!
I5
82
2
5
7
93
26
2
5
7
Total
1
1
37
63
used these zones to classify the flora of Saint Lawrencc Island, Alaska, according to
the northern limit attained by the specics. ‘The relationship of this classification by
northern limit to our classification by total range is shown in Table 25 for the 67
species which are common to Young’? list and ours. There is a clear relationship
between the two classifications, with most of the species which reach only Arctic
zone 4 being classified by us in the Borcal-montane or Roreo-temperate elements
whereas all the species which reach zone 2 are placed by us in one of the categories
containing Arctic plants. hlost of the species rcaching zone 1 fall into our Arcticmontane elements.
In LuJIore utluntipe europienne, Dupont (1962) presented a dctailed analysis of spccies
and subspecics confined to thc Atlantic zoiic of Europe. As defined by Dupont, thc
Atlantic zone is broadly equivalent to our Oceanic Temperate arca: it excludes the
Boreal and Southern regions. Within the Atlantic zonc Dupont recognized t\vo
categories. Euatlantic taxa are ~irtuallyconfiiied to the Atlantic zone, and sometimes
have a limitcd distribution within it, wliereas Suhatlantic taxa arc found throughout
the Atlantic zone but extend txyond it t o a greater or lesser dcgrer, although Lvitli
increasiiig distalice fi-om the zone they Ixcomc. progressi\-clj- rarer. Dupont (1 962)
also discussed in detail man)’ specics which he cxclLided fi-om thesr catcgorics fbr
onc rcason or anothcr. He cxcluded. for csample, spccirs \rliich arc restricted to
the Atlantic zone in Europc, h u t also havc a liroadcr range elseIrhcrc.
In Table 26, Dupont’s classification is comparcd to ours for the 63 Euatlantic
and Suhatlantic species wliicli arc prcwnt in the British Islcs. \Ye ha\^ c.xcluded
specics which Dupont regarded as pos\ibly Euatlantic and prohahl) Subatlantic,
but which he excluded from the dcfiniti\c lists because of doubts about their
taxonomy or distribution. We havc also ex( luded subspecies, including species 1% IIK h
have been reduced to infraspecific ranh sincc I h p o n t (1 962) wrote.
Most of the Euarlantic species are incnibers of our Oceanic Temperate elernelit
The only species which we regard as Occ,niiic Box ral-montane is D a r ~ ~ ~ l o i l i i ~ a p t ~ r ~ u r c . l l a .
described by Dupoiit as northern Euatlnntic, and the only Ocemic Boreo-temper ate
species is Meconopsu cambim, dcscribcd I)) Dupont as Euntlaiitic montane. ’The
Occnnic Southern-temperate species ‘irv C,’c.n/nrrizum scillozdc.s, Dabwcin cnnfabiira m d
Erzzca ~a~qans.
It is surprising that four sprcics ~iicludeda4 Euatlantic 11) Uupont <ire
classified as Suboceanic in our s) stem: tlir>scare the Suboceanic Soutlierii-tempei ,ite
Euphorbza IEyberna and Ranunculus omzo~ihyllu\m d the Suboceanic Temperate S ~ ~ ~ f i / i ~ d i u n ~
marzfimum and I’izcza orobus. In the case of Southern-tcmperate spec ics, the d i w cpanc\
arises because we include western l\/leditcrraiiran records \ v h ~ c h\t ere disregarded
by Dupont because he had a narronri tn.ionomic concept of the species (E//F//oihn
hybc./na) or because he treated records ‘1s duhious which ha\ r sul)sequcntl) lieen
conhrmrd (Ranunculus onmp/yllur). I3ifli.i t~iicc5i n taxonomic trcntnicnt or additioiial
geographical info1Ination probably exp1,iin the different classifications of h i / ) h i d / u m
maiztzmum: Dupont does not cite the lo( dities in Estonia nnd crntral Gci rmin
Finally, TEza orobu, is a borderline c n w of ‘I species which Dupont descrihd as
Euatlantic despite the fact that it hay \itc’s in German) and Snitmlnnd.
1he Subatlantic specics fall aliiiost cyua11) into our Oceanic Teiiipcr ntc ‘ind
Oceanic Southern-temperate element\, although a minorit) ni e clas5ihrd < i s Suhoceanic. apart from thcse, the oril)
eptions are the Oceanic Horeal-niont,inc*
Saxfrugn Iypnozdej, the only species wliic h is dcscribed b\ Dupont (1062) a\ nortlic i n
Sulintlantic, the Oceanic Boreo-teinpc 1,itc Loitherzum 0 5 5$ntqim, \\ hich he dcscril~c~s
as Subatlaitic \\ith montane a i d n o r t h u l i tcmdcncicc, nnd A 1 r / h j u o h / > . \ \ l i i O i
Dupont describes as Subatlantic niontnnc~<ind\z c classifj as n~cditerrnncnti-iiiOiit,~iic.
Tliirr) -fivr specie\ which fall into O U I O( v‘inic Boreo-tcmpci ntc. l’ciiipri ate nnd
Southern-temperatc clements ai c not i i i ( ludcd ,unongst the atlantit f l o r n a< definrd
by Dupont (1 962). Twelt e of these ‘ii c. ( I i t i c nl o r rccenh dcsci Il)cd t r i u \ ~ h i c hLirii
not listed by Dupont (Euphram spp lf(\/uttr spp., @u\ coidcrt(z, Scilrcoiiim s p p nnd
7r2filrum otczdoi/a/ei and n further eiqlit c plnnts h i c h lie list\
pos\ililJ or
prolAdy AAtlantic.Of iiiorc interest < i i ( the y>c‘cics dcxrilied as pscudontlantit 111
Dupont: these are Atlantic in Europr, I N I I licit P an aniph~~itlniitit01 $2 id(*r ranqc.
(At1$lei Iuriniatci, Eleogiton jluzimz 5. Hz nzc uo/)// 11ii/?i zc ils on 1 1 . Atb/hor/iitr europm N , 7i /to/ru/itt>\
,,he( /o\uin, H ~ I I ~ z P
/iciibrgc.n\r
~ ~ o ~i\ ~c zv I~idetl
J I ~from
~ ~this
I ITI~oup <ifi t h a s outl\ iiiq
locditics outside the Atlniitic m n c ) l,/m\dlo uu\/iol/A m d 131/ofamo~gu/oirr p / g dill\ ‘11 c‘
not listed bp Dupont but rtoiild p r c ~ i i i i i ~ i l1,h111 i n t o 1 1 1 i \ psciidoatl,iiiti( cdtcgoi 1
i sliccics are excluded on [ l i t grounds thnt die\ ha\ c
\zidc‘i disti i t x i t i o n >
Suh,itlCrntit-suliai ctic itlic, 1
h co-(cnipvi‘it(, A/c(zitdlzri i i / u ) z / / m / i ) .hi\c
isoLitcd populatioii~iii thp hlcditei 1 ‘ u i ( ~ i I icyion 01 thr. l\llddlc l,,ist (h\ r spc ( I C \.
dlthougli thcw 1 ccords of H ) / j e i i ( r m u d / / 111 < l l l d l ’ l l ( I / ? / d / / ~ / I Z i / ) O / / / \ r i l l ’ c ’ 1 1 0 1 1 1 O U \ )
oi <irrtttl\tei 11 but not iltlnntit (S~dzo)i/(I
,\lost of ~ L I ISuboccnnic 1empci ntc ~ i i i dS o ~ i ( l i c r i i - t r r i ~ I ) ~spct
~ r ‘ ~iv\
t ( ~< I I c i i c ) t c11
cla\sihed n s Sulxitlnntic In Dupont ( 1 0 0 2 ) 111 i n m \ c dsc‘s thk 1s I x ~ ~ t tlir~
i q
l i ~ (t
11idcl distnt~utionthat Iic- I S prcpii (.(I to <I( c vpt ‘ l I 1 c x T c m p ~(itc
I ’ 1 ) ~I( 4 IM\
too
niCiii\ outh irig loc&ies
outside tlic ,\tlrintic ioiic and n i m \ 01 t h r Sourlic’ii i trmpci n t c species eutcnd l i l t 0 the \ \
I1 Al(’dltc‘lIdlledll ‘ \ ~ l t l l O l \ Ilh(. lhlpl)111
.
L~~
(
(3
( X \ t ( \ l
C. D. PRESTON APJD M. 0. HILL
58
TABLE
27. Distribution of Sulix species in the floristic elements defined by Myklestad
& Birks (1993) and those defined in this paper. MyMestad & Birks’ elements are
denoted by letters, as follows: A, Suhx pleugnos element; D, Southern Widespread
element; E, Widespread element; F, Northern Widespread element; G, Arctic-Alpine
element; H, Northern element; I, Northern Boreal element. Our elements are given
in numerical form; for their names, see the text
Myklestad & Birks’ element
Floristic
element
A
D
E
F
G
H
I
13
16
24
26
44
53
54
55
74
75
81
84
(1962), and Webb (1983),who draw up lists of single elements but do not attempt
to classify all the species in an area, tend to adopt stricter criteria than those such
as ourselves who have to classify all species into one group or another.
Numerical anabses
The TWINSPAN analysis of the distribution of European Salix species by
Myklestad & Birks (1993) is compared with our classification of the same taxa in
Table 27. Two species classified by Myklestad & Birks, S. atrocinerea and S. cinerea,
are treated as subspecies of S. cinerea by Stace (1991). For this comparison we have
replaced our classification of the species (as Eurosiberian Boreo-temperate) by
separate classifications of subsp. cinerea as Eurosiberian Boreo-temperate and subsp.
olezjilia (S. atrocinerea) as Oceanic Southern-temperate. Myklestad & Birks also classify
the Irish endemic S. hibernica but as this is now regarded as a synonym of the
widespread S. phyZic$olia we have excluded it from the comparison.
There is a very close correspondence between the seven floristic elements in
which Myklestad & Birks place our S a l k species and the eleven elements that we
recognize. We classify, for example, the five taxa in element A as Temperate or
Southern-temperate and the six species in elements G, H and I (A4rctic-Alpine,
Arctic and Scandinavian) as Arctic-montane or Boreo-arctic Montane.
The floristic elements in which Birks (1976) places our pteridophytes are more
complex than those recognized for Salzx as the pteridophytes are a larger group and
they show strong distributional gradients from west to east as well as from north to
south. A comparison of Birks’ (1976) numerical classification of pteridophytes and
TABLE
28. Distrihution of ptcridophytes in thr flol-istic rleiiients defined by Hirks f 1 !I761 a ~ thow
~ d
dcfincd in this paper. Birks’ elements arc reprcxwtctl l)y Irttrrs. Oui. elements arc’ givw in ~iiimcrical
form; for theii iiaiiic~src the text
ours is giken in Table 28 This exclutlcs C;y\tof),l,terz\ dzcXeuno, nhich \ . ~ eha\ e h e m
unable to classify Tlicrc i, a clow coi I cyondence betwcen man\, of liis g r o u p nnd
ours: Birks’ group A contains four Ocraiiic species, for example, and groups hf and
I% contain 6 hlediterranean-Atlantic spccies and onc fiom each of tht. Sulimediterraneari-Subatlantic, hleditert diic,rrn-t~ioiitaiic and Sulmceanic Souther nternperdtc elements. Othcr elements ar c inorr di\ erse, but 110 more \o t l i m onc
would expect from BirLs’ description\ of them iZ few species appcnr to b c . out 01
place: tlie Boreo-arctic hlontarie Hupn ;rtr \ d q o in group P, which coiitciiiisthc most
widespread species. the ‘Temperate D ~q / )J/ o T z \ cii\tntn in group 7’ (species of nor thcrn
Europe and the Nps) arid the Boreal-iiioiitaiic D!yopterz\ oretlda i n gi oiip L (\pet ies
widcsprcad south to the northern hleditci 1 nnean) Inspec tion of the zlt/cr\ I + J ~ ~ P
Euwpneae maps s u g c s t s explanations lor these apparent aiioinnlics. Hufie,;ia \ P / ( L ~ o
has scattered recor ds in lowland Eut ope (many mappcd as probnhlc extinction$)
which appear frcqucnt at the 50-km s c ~ l eand ebcn niorc frequent nt the larycr
scale of Birks’ analysis. Dyofiten\ c~zstntci15 one of the more northerh ol the Tenipei(rtc
species, and D. oretrdeJ is is coricentratcd 111 tlie Borenl ~ o i i chut theic ~ r scattcrrd
c
records from elsemhere and neither thr tii‘ip 1101 any other data pro\ ide\ ‘r conkinc iiig
picture of the distribution of this p o o ~ l )icvxded taxon
60
C. D. PRESTON AND hl.0. HILL
CXASSIFICATION OF OTHER SPECIES
Endemic species
The 48 species endemic to the British Isles are listed in Appendix 1. The list
excludes Fumaria purpurea, which is found in the Britain, Ireland and the Channel
Islands. The species have been allocated to the most appropriate floristic element:
three montane species are classified as Oceanic Arctic-montane (Athyrium $exile,
Cochlearia micacea and Euphrasia cambrica) and the remainder divided between the
Oceanic Boreal-montane (1 1 species) and Oceanic Temperate (34 species) elements.
Sagina boydii cannot be classified as it is known only in cultivation, although it is
thought to have been collected somewhere in Scotland.
This list of endemics differs substantially from the list of species endemic to the
British Isles given by Walters (1978), which was based on Flora Europaea. Six of the
plants listed as endemics in 1978 do not appear on our list. Coincya monensis, described
by Walters as a unique example of British endemic which was known to pre-Linnaean
authors, is now regarded as only subspecifically distinct from a more widespread
European plant, although the long history of the British plant means that the epithet
monensis is retained for the species (Leadlay & Heywood, 1990).Other endemic species
cited by Walters (1 978)which have now been reduced to subspecific rank or to synonyms
of more widespread taxa are Ep$actis dunensis (Stewart et al., 1994)) S a h hibernica
(Meikle, 1984) and Saxzjaga hartii (Webb, 1987),whilst Fumariapurpurea was discovered
in Guernsey in 1970 (McClintock, 1975) and Bromus pseudosecalinus is treated by Stace
(1 99 1) as an introduction of unknown origin. There are numerous taxa on our list
which were not included by Walters, including the recently described species Cotoneaster
cambricus (Fryer & Hylmo, 1994),Epipactisyoungiana (Richards & Porter, 1982))Limonium
britannicum, L. dodartzjimne, L. loganicum and L. pamum (Ingrouille & Stace, 1986))and
several taxa which have long been recognized but are currently regarded as species:
Athyrium $exile, Cerastium nkrescens, Cochlearia micacea, Limonium procerum and Ulmus plotii.
We include Spartina anglica, which is introduced from the original British stock into
other European countries, Euphrasia anglica, E. cambrica and E. heslop-harriionii, species
which were excluded by Walters for various reasons, the extinct Centaurium latzjilium,
which is doubtfully distinct from C. elythraea, and the enigmatic Sagina boydii. We also
list the endemic Sorbus species, a genus which Walters (1978) did not consider.
Any assessment of the number of plants endemic to the British Isles involves numerous subjective decisions, both about the rank of individual taxa and, more importantly,
about the genera to be included. Our list is dominated by the narrowly defined species
of Euphrasia and the apomictic species of Limonium and Sorbus. Some authors adopt a
broader species concept in Euphrasia (e.g. Silverside, 1991). Alternatively, one could
adopt an even narrower species concept in the Limonium binenlosum ,group, and there
are unpublished doctoral theses which subdivide the British Ranunculus auricomus and
Ulmus aggregates into microspecies: any of these courses would increase the number
of endemics considerably. The apomictic species of Hieracium and Zraxacum and the
facultatively apomictic Rubus species are excluded from consideration on account of
the large and critical nature of these genera, which contain hundreds of endemics.
Species conjned to the Channel Islands
There are twelve species which are included as native members of the flora of
the British Isles solely on the basis of their occurrence in the Channel Islands. These
FLORISTIC ELEMEN I S I N BRITAIN AND IRELASD
61
can be classified into floristic elements using the same criteria as the plants of Britain
and Ireland. The elements to which they belong are the Oceanic Temperate
(Festuca arrnorkana, Lirnoniurn norrnannicum), European Temperate (Schoenoplectuspungens),
Eurosiberian Temperate (Orchis lax$ora), Oceanic Southern-temperate (Limonium
auriculae-ursgolium), Suboceanic Southern-temperate (Arrneria arenaria, Exaculurn pusillum)
and Mediterranean-Atlantic (Anogrammcl leptophylla, Linaria pelisseriana, Milium uernale,
Myosotis sicula, Ranunculus paludosus). The list clearly indicates that most of these
species have southern affinities.
Schoenoplectus pungens is included in the above list on the assumption that it is native
in Jersey but not in Lancashire; it is now extinct in both sites but it has been reintroduced to localities in Lancashire. I,znaria pelisseriana is doubtfully native, even in
the Channel Islands.
In this paper we have attempted to devise a system of classification based on
general principles, and then to apply it to individual species. These two aspects of
the work need to be discussed separately.
The use of the generally accepted major biomes provides a latitudinal classification
which should be comprehensible to botanists throughout the northern hemisphere.
The longitudinal gradients in plant distribution are much less strongly marked,
especially in continental interiors, and these categories are therefore more subjective.
Our choice of classes is inevitably influenced by the fact that we are classifying the
flora of an area which lies at the western edge of the Eurasian land mass. The
combination of the latitudinal and the longitudinal categories to give floristic elements
provides a non-hierarchical ordination. In discussing the division of areas into
floristic regions, Webb (1965)has pointed out that a single region is normally related
to several surrounding regions, but this can be distorted by hierarchical classifications
which suggest that some of the relationships are more significant than others.
Similar considerations apply to floristic elements. Although our classification is not
hierarchical, the different elements can be grouped together in different ways for
special purposes. In the discussion of thc different floristic elements in this paper we
usually grouped them by the major biome category, but they can also he aggregated
by eastern limit category.
I he classification of individual species into elements is sometimes straightforward,
but there are many borderline cases and other examples where the choice ofelement
has been hampered by a lack of geographical data. (The availability of data is to
some extent correlated with major bioine, as there is a dearth of synthesized, fineresolution data for many species in thc Southern-temperate and Neditcrranean
elements). The classification is therefore likely to be more useful in assessing the
phytogeographical affinities of assemhlages of species than in providing information
about individual taxa. However, there is no reason why the classification of individual
species proposed here should not be testcd by the analysis, by traditional or numerical
methods, of other datasets which m a y hc a\&lable IIOW, or which may become
availablc in the fiiture.
r
,
62
Geographiral relationships
the British and Irish ,flora
The main results of our analysis can be summarized rather simpl). The flora of
Britain and Ireland is essentially temperate. It is the Temperate major biome
category which dominates the flora of England, Wales, lowland Scotland and Ireland
(Fig. 33). The essentially temperate nature of our flora is perhaps concealed by
classifications such as that of Matthews (1937, 1955)which concentrate on selected,
small floristic elements. The Boreo-temperate elements replace the Temperate as
the largest single major biome category in the Highlands and Islands of Scotland
and in scattered 10-km squares in Ireland, particularly in the north and west. The
Southern-temperate elements form the largest major biorne category in some coastal
10-km squares in England, Wnles and Ireland, and the Mediterranean elements
form the largest group in one square in the Isles of Scilly. The preponderance of
these southern elements in coastal squares with only a small land area may be an
artefact in at least some cases, as these fra'gmentary squares are not always well
recorded and there may be a tendency for rare or scarce species to be overrepresented. However, we have excluded from Figure 33 any 1 0-km square in which
fewer than 50 species are recorded.
The majority of the species in the Boreo-temperdte, Temperate and Southerntemperate categories are restricted as natives to Europe or Eurasia. However, many
of these species have become naturalized in areas of the New World where the
climate is temperate and therefore similar to that of Europe. The colonization of these
'neo-Europes' (which include North America, southern South America, Australia and
New Zealand) by the Temperate and Southern flora and fauna of Europe is discussed
by Crosby (1986).
The relatively few species which are absent from the temperate zone fall into the
Arctic-montane, Boreo-arctic Montane and Boreal-montane elements. The majority
of these species have more or less circumpolar distributions, but have not spread as
aliens outside their native range. These elements appear to have been relatively
immobile in historical times, although not, of course, when conridered against the
longer timescale of the Holocene.
Further applications
of the class$cation
The classification presented liere has been developed for application to the nati1.e
vascular plants and the bryophytes of Britain and Ireland: the classification of
bryophytes will be presentcd elsewhere. Application of a similar clas5ification of the
two groups is essential if their phytogeography ir to be compared. Although
introduced species are not covered in this paper, they might also be classified
according to the Tame criteria as long as they are sufficientl) widcsprrad to indicate
their potential distribution. The South African Carpohrotus edulzs, for example, has a
Mediterranean-Atlantic distribution in Europe whereas the Asian Fallo@a japonzi a
has a Temperate range. Species which are native in mainland Europe but not in
the British Isles can be classified according to their total range: Solezrolin solezrohz,
endemic as a native to thc western Mediterranean area, now ha5 a Suboceanic
Southern-temperate distribution, whereas the naturalized populations of .dyer pseudoplatanus in Britain are mercly
extension of the native Boreo-trmpcrate range of
FLORISTIC ELEhIEN I S 1% RRI'141N AND IRELAND
61
that species in Central Europe. There is also scope for applying the classification to
animal groups as well as to plants.
We anticipate that this classification of British and Irish plants could be applied
with little or no modification to the flora of other areas in north and north-west
Europe. A more formal separation of Arctic and Alpine taxa might be desirable in
regions where the Alpine element is morr numerous and more distinct. A difficulty
in extending the classification to countries further east is its inability to deal with
species which reach their western limit in central or eastern Europe. The species in
the Pontic or Pontic-Sarmatic, Irano-Turanian and East Mediterranean elements
discussed, for example, by Harder et a/. (1965), fjtefureac (1965), Davis (1971),
Wendelbo (197 l), Zohary (1 973) and Carlstrom (1 987), would fall into this category.
In order to accommodate these species, it is necessary to introduce the western limit
as a third criterion used to define the floristic elements. The classification, extended
if necessary to accommodate the more eastern species, might also be applied to
the southern countries of Europe. The lowland Mediterranean species could be
accommodated in a series of Southern elements, but the species which are found
above the tree-line on the mountains of the Mediterranean repon in vegetation
which cannot be described as arctic do not fit comfortably into our classification.
The Mediterranean-montane species, sometimes referred to as Oro-hlediterranean,
are a significant feature of the flora of the Mediterranean region, and present "acute
problems" to the phytogeographer (Davis, 1965-85). In adapting our classification,
one might consider creating a separate RIediterranean-montane biome of equivalent
status to the other major biomes, rather than place the species in a single. special
category. There is some overlap between the species which have easterly western
limits in Europe and those in the Mediterraneati-montane category, as many IranoTuranian species which extend westwards to the Mediterranean regions of Turkey
and to the Aegean islands are found thew at high altitudes in the mountains (Davis,
1965-85; Runemark, 197 1). The Meditcrranean-montane climate miniics the harsh
winters and hot dry summers characteristic of the Anatolian plateau.
We have not attempted to solve these detailed problems because they are scarcely
relevant to the elements of the British flora. A more thorough solution must depend
on better taxonomic and geographical data, and take account of climate as well
as of geographical distribution. In our view, climate is pre-eminent; major
biomes should be the basis of any classification into floristic elements. If the OroMediterranean climate resembles that of steppes to the east, then the species that
are present there are best classified as steppic. Inconveniently, this means that in
mountainous areas some account must be taken of the altitude at which species
occur, not just of their general area of occurrence. In an extended system, some of
our own elements would have to be altrred. For example, a few British and Irish
species such as Astragalus danicus are mainly steppic and ought perhaps to be assigned
to specifically steppic elements with westcrii Europe as a mere outpost.
Superimposed on a climatic definition of elements must be a sprcificatioti of the
floristic kingdoms, provinces and regions where species occur. For many species
distributions, the floristic regions of Takhtajan ( 1986) provide about the right l e \ ~ l
of resolution.
Finally, the use of the species as the unique level at which elemcnts are defincd
can be inadequate, especially for neo-mdcmics. Neo-endemics should hc vicwed as
members of species groups in their br-oaclcr context. Plant geographers regularly do
this, and often assign genera and sections of genera its ~vellas spccirs to floristic
61
C . D. PRISTON AND hl. 0. HILL
elements. This would be a valuable exercise for the British and Irish flora, hut it is
bcyond the scope of the work presented herc.
We acknowledge the financial support provided by the NERC through its TIGER
(Terrestrial Initiative in Global Environment Research) pro,gramme, award number
T91/197. We are also grateful to Dr B.K. Wyatt for encouraging us to embark on
this project, to T. Lahti for supplying us with digitized data from Atlas Floiae Europaeae
and to D.B. Roy and S.M. Wright for plotting the coincidence map5 for the British
Isles and Europe respectively. Other information or assistance has been provided
by Mrs V. J. Appleby, Ms M.D. March, J.O. Mountf'ord, T.H. Sparks and Professor
C.A. Stace. We are grateful to Professor B. Huntley, A.O. Chater, Dr G. Halliday,
Dr F.H. Perring, Dr S.M. Walters and Professor A.J. Jl'illis for comments on a
draft of' this paper.
F1,ORISTIC ELEMENTS IN BRITAIN AND IREI,4ND
I
*
16-23
24-31
32-39
40-47
48-55
56-63
Figure 14. The distribution in Britain and Irrland of species in the Arctic-montane mapi- biomr
The smallest symbols denote 10-kni squarrs in Lvhich 10-1 9% of the sprrirs are recorded:
successively larger symbols are used for successi\.r 1 O?’U intrivals, excrpt that thr largeht symliol is used
for squares with 80-100°h of the specks.
category.
66
C. D. PRESTON AND hf. 0. HILL
8-11
12-15
16-19
20-22
23-26
27-30
Figure 15. The distribution in Britain and Ireland of species in the Boreo-arctic Montane major biome
category. For explanation of symbols, see caption to Figure 14.
67
*
2-3
4-5
6-7
8-9
10-11
0 12-13
14-15
16-19
Pigitrc 16. 'I'hc distribution in Britain and Irclnntl i)f spccics in thc llYc-t~orcalmajor Iiiomc i-atcgor)
For cxplanatioii of symbols, scc caption t o F'igi1i.t. I + .
68
C. D. PRESTON AND M. 0 . HILL
11-20
21-31
32-41
42-52
53-62
63-72
73-83
Figure 17. The distribution in Britain and Ireland of species in the Boreal-montane major biome
category. For explanation of symbols, see caption to Figure 14.
69
FLORISTIC ELEMENTS IN BRITAIN AND IRELAND
7
*
24-46
* 47-69
0
70-93
94-116
117-139
140-163
164-186
187-233
Figure 18. The distribution in Britain and Ireland of species in the Boreo-temperate major biome
category. For explanation of symbols, see caption to Figure 14.
70
C. D. PRESTON i\ND hf. 0. HILL
10-14
15-18
19-23
24-28
29-32
0 33-37
38-48
Figure 19. The distribution in Britain and Ireland of species in the European Boreo-temperate floristic
element. For explanation of symbols, sec caption to Figurr 14.
il
F1,ORISTIC F,I.EhlENT\ IN BRIT.2IN AND IREIANII
t
021-23
24-21
28-34
Figure 20. The distribution in Britain and Ircland of species in the Wide-temperate major Iiome
category. For explanation of symbols, see caption to Figure It.
72
C. D. PRESTON AND M. 0. HILL
112-167
168-222
223-278
279-334
0 335-389
390-445
Figure 2 I . The distribution in Britain and Ireland of species in the Temperate major hionie category.
For explanation of symbols, see caption to Figure 14.
73
FLORISTIC ELEMENTS IN BRITAIN AND IRELAND
*
5-9
10-14
15-19
20-24
25-28
29-33
34-38
39-48
.... I.:.:..:.
++++
..........
r l i ; J&
. i i i:
............
* -
I.
Figure 22. The distribution in Britain and Ireland of species in thr Oceanic ‘Temperate floristic
element. For explanation of symbols, see caption to Fi,pre 14.
74
C. D. PRESTOK AND M. 0. HILL
9-1 1
12-14
15-16
17-19
20-22
Figure 23. T h e distribution in Britain and Ireland of species in the Suboceanic Temperate floristic
element. For explanation of symbols, see caption to Figure 14.
60-89
90-118
119-148
149-178
179-207
208-237
g f l..
..... ...
*. * *,
. . ..
Figure 24. Thc distribution in Britain and 1rchntl of' spwies in the European 'Trrniiu"tr floris;lic
rlrment. For cxplanation of s)mhols, see caption ( ( I b'ixurr 14.
76
C. D. PRESTON AND M. 0. HILL
. 13-24
I
25-36
37-48
49-60
61-72
a 73-84
a 85-96
97-120
Figure 25. The distribution in Britain and Ireland of species in the Eurosiberian Temperate floristic
element. For explanation of symbols, see caption to Figure 14.
77
FLORISTIC ELEMENI'S IN RRI'IAIN AND IRELAND
q
90-1 19
120-149
150-178
179-208
Figure 26. The distribution in Britain and Ireland of species in the Southern-temperate major biome
category. For explanation of symbols, see caption to Figure 14.
78
C . D. PRESI’ON AND hl. 0. HILL
0
8-10
11-13
14-15
16-18
19-20
21-25
Figure 27. The distribution in Britain and Ireland of species in the Oceanic Southrrn-temperate
floristic element. For explanatioii of symbols. scc caption to FiLqrc 14.
79
FLORISTIC ELEMCN 1 5 IN BRITAIN AND IREI-\ND
11-16
17-21
8 22-27
28-32
0 33-37
38-43
*
/w. ....
-d L.
I
.
........
..
-
..
.......
g...................
i:: :I: :::..
..................
Figure 28. The distribution in Britain and Ireland of species in the Suboceanic Southern-temperate
floristic element. For explanation of symbols. TTC ( nption to FiLgure 14.
80
C . D. PRESTON AND M. 0. HILL
Q
*
14-20
21-27
28-34
35-40
41-47
48-54
Figure 29. The distribution in Britain and Irrland of species in the Mediterranean-Atlantic floristic
element. For explanation of symbols, see caption to Figure 14.
10-14
15-18
19-23
24-28
29-32
33-31
0 38-47
P
82
C. D. PRESTON AND M. 0. HILL
A
d
19-27
28-36
37-45
46-54
55-63
64-72
Fi,yre 3 1. The distribution in Britain and Ireland of species in the Oceanic castern limit category (A)
compared with that of species with a continental distribution (B). For rxplanation of symbols, see
caption to Figure 14.
B
- 10-18
*
w
19-27
28-37
38-46
47-55
56-65
66-74
75-96
84
C. D. PRESTON AND hl. 0. HILL
I
-
1400
n
E 1200
1
1000
800
600
400
200
0
I
I
I
2
*
I :i;4
I
3
*
*
*
*
1
Y
I
I
1
I
I
4
5
6
7
8
I
9
Major biome category
Figure 32. The maximum recorded altitude for species in each major biome categor). The horizontal
line represents the median value and the rectangle the quartiles. The total range is represented by the
vertical line and by asterisks; the latter identify species with maximum altitudes which lie outside 1.5
times the interquartile range.
FLORISTIC EI,EXIENT\ IN BKI 1AI?; AN13 IKELZNI)
85
Temperate
o Southern-temperate
Figure 33. The doininant major bionic catcpoi-y OIRC:) in the 1 0 - h i squares in Britain imd I r e h i d .
T h e major biome category with the largest riumlier of species in each squarr is shoitn Ily sinall circles
(Temperate: hZBC 7), large circles (Boreo-t~inpcratr,RIBC 5), sliadetl square\ (Southerri-tenrper~~~~,
hlBC 8) and black squares (Mediterranean, hlBC: 9). Squares from ivhich fewer than 5 0 species are
recorded are excluded.
86
C I) P R E S I O N AND 1 1 0 HlLL
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-
92
C. D. PRESTOIV AND M. 0. HILL
APPENDIX 1: LIST OF SPECIES IN EACH FLORISTIC ELEMENT
K$J to symbols
9 Doubtfully native
7 Endemic to British Isles
Present in Ireland, but not in Great
t
+
??
Am1
Am2
Am3
Am4
Am5
As 1
As2
Bo 1
Bo2
c o1
co2
Djt
Sst
Tax
Tso
Wid
Britain
Channel Islands, not Great Britain or Ireland
Disjunctly circumpolar, but belonging to the indicated element in Eurasia
Continental in Europe
Not assigned to an element by us
Also in N. America
Occurring in Greenland but not continental N. America
With very restricted range in N. America, more widespread in Europe
Occurring in N. America but only in the west
With very restricted range in Europe, more widespread in N. America
Also occurring in C. Asia
Also occurring in E. Asia
Arctic or boreal species (other than exclusivelycoastal taxa) absent from temperate-zone mountains
of western Eurasia
Montane species absent from boreal and arctic in western Eurasia
Coastal throughout range
Mainly coastal in British Isles
Distribution disjunct
Sensu strict0
Taxonomically difficult
Occurring in tropics or Southern Hemisphere or both
Widely naturalized (may be followed by present distribution in square brackets)
European Arctic-montane
Alchemilla alpina
t Arenaria ciliata
Arenaria nomegica
Artemisia nomegica
Bartsia alpina
Carex microglochin
Cerastium alpinum
Cerastium arcticum
Cerastium cerastoides
Cochlearia pyrenaica
Draba nomegica
Epilobium alsinfolium
Gentiana nivalis
Gentiana uema
Gnaphalium norvegicum
Gnaphalium supinum
Luzula urcuata
Luzula spicata
t Minuartia recurua
Minuartia sedoides
Phyllodoce caerulea
Poa Jexuosa
Salix arbuscula
Salix herbacea
Sali~xmyrsinites
Saxij?aga aizoides
Saxzjaga stellaris
Am2
Am2
Bol
Bol
Am1
Am1 As1 Tso
Am1
Am1 Bol
Am1 As1
Bo2
13 Am1 Bol
13 Am2
13 Am1
13 As1
13 Am1 As1
13 Am1 As1
13-t Am4 As2 Bol
13-t Am1 As1
13 Bo2
13 Bo2
13+ Am1 As1 As2
13 Am1
13 Bol Sst
13 Am1
13 Bol Sst
13 Am1
13 Am1
13
13
13
13
13
13+
13
13
13
13
Silene acaulis
konica fmticans
13-t Am1 As2
13 Am2
Eurosibm'an Arctic-montane
Arabis alpina
Erigemn boreah
Euphratia figida
Juncus tnjdus
Pznguicula alpina
Eronica alpina
14
14
14+ Am1 As1
Eurasian Arctic-montane
Arabis petraea
Oxytropis campestris
Saussurea alpina
15
1515
Circumpolar Arcticmontane
Alopecurus borealzs
Arctostaphylos alpinus
Astragalu alpinus
Athyrium distentlfolium
C a w atrojiusca
Carex bigplowii
Carex lachenalii
Carex maritima
Carex norvegzca
14
14
14
Am1 Sst Tax
Bol
Am1
Am1
16- Bol
16
16
16+
16
16
16
16
IF
Tso
Co2 Tso
I'LORISTIC CLERIEN I 5 I N BKI r
Carex ranjma
Caier rupestns
Gain saxatilzs
Dzcipensia lapponzca
DFhasznstrum alpznurn
Dnas octopetala
Eprlobium anagallzd~nhimi
Junc us biglumzs
J u n ~ u scartaneus
j%nrur tngluinis
Kobresia simplzczumila
koenigia zslandzra
Llqdia serotzna
Lmeleuria prorumbens
dlliniiartza rubrlla
atlznuaitza stncta
Lb@nsntis alpestm
Oyria dzgyna
Poa alpinn
Sagina nivali,
Sagina saginoide~
Sahw lanata
Salzx wtziulata
Sarfiaga cernua
Sa xifiaga cerpztosa
J a u ~ a g anzilalzs
Saxlfrnpzl oppoJztijblza
Sarfraga nuulans
M u m rorea
Sibbaldia pmrunibens
Thalietrum alpinurn
Qieldza pu~zlla
16
16
Bol
Calamagrostis strictci
(.'mu aquatilis
Careu atrata
( ~ I F Xcapillaris
( h e y chordorrhiza
( . ' h x dioira
C a w isaginata
Enipetrum iiigruni
EquisPtum aariegatuni
Eiiophoruin uaginatutn
Ftstuca i:iiizpara
16 Bol
16 Bol
16 +
I6 f Am4
+
16
16
16
16
16
16
16
+
+
16
16
I6
16f
16
16 f
16
16
16
16
16
16
16
m .LYD I R E L ~ N D
~~i 'rso
Am4 Bo2
Huperzia selago
\junrus ba1ticu.i
I p p o d i u m annotinurn
Pemcnria rim$ara
Phleum alpinum
Poa glauca
.Yali.r phq'licij2ia
,S'ar$raga h i r c u h
Ihrciiiium ul&inosuni
I hri.iniurn ZtitiJ-idaeci
I l b d i a alpiria
1lbodsia iluensis
Bol
Bol
801
Y3
26
26
26
26
Tso
Bo1
26 26
26
26
26
26
26
26
26
26
26
26
26
26
26
Tax
Tso
Tro
Tso
26
26
2b
26
Bol
Eiiiosibunan It?& boieat
Phiitago tnaritima
I6
16 Bol
16
16
16f
+
Euimnn I lid?-boreal
Rannnculus acrit
33
\$'id [36]
16
Oceanic Boreo arrtii Montane
Euphra\za ostenfpldzi
21
Bol
European Boreo-aretzc '21ontane
Ah hemzlla glomerulans
23
(.ornu\ suecica
23f
Draba incana
23
7 Euphraua salzsbuTenszs 23
L.ath>inisjaponirus
23+
Lg rnus arenanu
23
Ligusticum rcoticum
23f
Qrhnz r alpina
23
,Ilertensa mantima
23f
Sudurn uiliosum
27
Am1
Am1 ,4s2 B o l
h i 1
Am1
,452
C:oI
+ Am1 As2 C:o?
Euiotzberztin Boreo-arctic Alontane
Potmtzlla crantziz
24
S t i h lapponum
2-1.
Am1 As2 C ~ J I
< h l
Am1 As2 C ~ J I
Am3 Sst
Am1
Circ uinpolar Roreo-arctic .Ifontone
.Illrum \clioenopmrum
2h - \Yid
Brtula nana
26
36
36
36
36
36 +
36
36
36
36
36
\$'id
Col
\Vid
'rso
c:01
94
h@osotiJ stolontjra
Saxfiaga hypnoides
5)iranthes romanzoJana
C. 1).PRESTON AVD hl. 0. HILL
41
Bo2
41
41
42
42
42
42
42
European Boreal-montane
Ajuga pyramidalis
Alchemilla jilicaulis
Alchemilla u:ichurae
Atr$lex longipes
Atriplex praecox
Bhsmus rufu.
Carex ornithopoda
Cirerbita alpina
Cpptogramma crispa
Darglorhiza lapponica
Dlyokteris remota
EleoclzariJ austriacn
Euphrasia Jcottica
Lamium confertum
Lobelia dortmanna
Melainpyrum Jylvaticum
i W u m athamanticun?
O.u_ytropishalleri
Pohgonatum uerticillatum
Potamogeton rutilus
Aeudorchir albida
Ribes alpinum
Schoenusferrugineus
Spaiganium angustfolium
Thlaspi raerulescens
fiollius europaeus
Kola lutea
43
43
43
43
43
43+
43
43
43
43
43
43
43
43
43
43
43
43
43
43
43
43
43
43
43
43
43
Sst
Bo2
44
Tax
EuraJzan Boreal-montane
h&ogzum aphyllum
Mznuartza uerna
Pznus Jylvestm
Tax
Am1 Bol
Bol
Am1
Am2
Col
h 1 2 c:ol
Am1 .4sl co2
As2 Sst
Bo2 Tax
Bol
Am2 Bol
Lhl
€302
Bo2
Am3
Bo2
+ Am1 As2
Bo2
44k Am1 As2 Tso
44
44
44
44
44
Am1
Sst
44
44
45
45
45
45
Bo2 Djt
45
WlCl
.hj
Suboceanzc Boreal-montane
b o p t e n s oreades
Galeum stemen
Pobgonum boreale
Saxfiaga rosacea
Sorbics rupzcola
Eurosibenan Boreal-montanr
Calama<grostispurpurea
Carex buxbaumii
Cirsium heteroptyllum
Geranium sq’lvaticum
Isoetes lacustris
Qrola media
Ror$pa dandira
Salix rnyrhtfalia
Kzccinium myrtillus
Pnniula f a i v m a
Rumer long:filzuj
Djt
Cin urnpolar Borrnl montaizr
ilrtaea spirata
46
Andiomeda polzfoha
46
&tostaphqhc utla-urjz
46
Asplenzum tnchornanelramosum
46
Collitrzche hermaphrodztzta 46
Cora curta
46 Tso
Catex Instocarpa
46
Carer lzmoha
46
Carey magellanzta
46 Tso
Cf’areupaurgora
46
Circafa nbma
46
Coeloglasmm u m l e
46
Corallorhiza trlfida
46 CJ Jtopteizr nzontano
46
Dzplicuiastncm iomblanntum 46 Ilro rera longfolza
46
Dpoptens expanba
46+ Tax
Eyui return PratenJP
46
Goog)>eturepem
46
Hammarba paludo!a
46
Hzmrhloe odoratn
46 Iroeks echmorpora
46+ T a x
Juntus alpznoartzrulatu,
46
Jun( us filfomzs
46
Lnmaea borealzs
46
Liltera roidata
46
Lyszniarhia ttystjlora
46
hfoneses ungora
46
.4ala~j e
46
Niiphar pumzla
46
Oithzlra serunda
46
Pinguzcula LuLqariJ
46
PohJtzchum IonrhztiJ
46
I’otamogetnn alpznia
46
Potamogeton f i l f o n n ~
46
Potatnogeton praelongia
46
Potmtzlla fnitirosa
46+ Bo2
19ioIa mznor
46
Ranunrulub rfptatiF
46
Rubies arctirus
46 Bol
Riibic~chamaemoru
46
Sctieiirhzena palustnJ
46 Selagnella selagnozder
46
Subularza ayuatzta
46
Zzihophorum alpznum
46 Trzihophoncm cespitoruni
16 Tao
Tnentalzs europaea
+6+ Am4
C‘trirulana zntermedia
46
k i rinzum miiio~aipi~in
16
Eztznaicm o ~ x ) . i o i c o ~
16
+
+
+
+
+
+
+
+
Oceanic. Boreo-tmpi-ate
Euphrasia arrtica
il
FLORISTIC ELEhlEST Is IN HRITAID; AND IRELISI)
Euphraia conjusa
Ffimenophyllum utilsonii
Limosella austrulis
hleronopsis canibrica
.iVam-thecium ossfragum
Potninogeton rpifydrus
Puc,cinellia maritima
51
Suboceanic Boreo-temperate
Atnplex y l a b n u d a
Calhtmhe hnmulatn
,2$rica gale
.\fiiiopbllum ultern$oruni
Ophioglossuni azorirum
PzloJella peletenana
Rhqtu hoJpora f u x a
Mene un@oia
52
52
52
52
52
32
52
52
Eu'urt@vm Borro-tempelate
All liemilla acutzloba
Alchfmzllu glabra
illchemzlla glaucrsren c
4 Icherni/la grarz Izs
ilkliemilla montzcola
Alrhernzlla rub( renata
dlo/ircut uc genzculatus
rintlpllis culneraria
Calluna iulqaris
C a m dzgztata
Chrer echinata
Cme~java
C m e x panicea
aquatzc a
Corn allaiia mayizs
Crfpis paludora
Drcthampsia pexuo ra
Eleoiharz~quinquefluiu
Gtophorum latfoliuni
Euphrasia rostkoriana
Gnlqb r i c tetiahit
Galium molluqo
Gentiandla rampe5triI
H~,hpopIiarrhamnoide $
FJjpenium maculatum
JuiiruJ bulbomr
LPontodon autumnah c
~ i o p o d i r l l niniindata
.\Jontzn jontana
VntduI ctncta
€ ' e h ulari, palurtris
Phyteuma orbi( ulaie
Pilo wlla jagellarts
Pob ynla amarella
PolJpodiuni r~rtlgnrr
Rnnuntulus auricotnti I
Rhtnanthui niinnt
Rzha Jpiititiim
CntnbroJa
51
51
51
51
51
51
Bol
Sst
Am5 Tso
Bo2 "id
Am5
Am3 Col
.4ml
Am2
Ainl As2
.4ml
co2'Tax
41nl
Co2
53 53
53 53 53 53 53
53
53
5353
53
53
53
53
53
53
33
53
53
53
53
53
53
53
+
53
53
53
53+
53
53
33
53 53 53
53
53
53 -
\\Y
Wid
\%'id
As1 \%'id
As2
:\m1 Sst
Am1 As1
As1
Am1 As 1 T S ~ J
As2 Wid
Am1 4 s 2 Tso
Am1 As1 A d
Tax
b'id
As1 \Vid
i1
\%'id
Am 1 As?
Am1 As1 As2
,451 \Vid
h
Bo2
Sst
LYid
Tax
9i
96
5 Kibes n i p i m
Rurne.r aretosa
Sagina nodosa
Sagina procumbens
Sa@ttaria sa<gittfolia
Salix cinerea
Salix pentandra
Sa1i.i repenc
Scutellaria galericiilatn
Stellaria graminea
Stratiotes aloidees
Thalirtrum ,flaoum
Trfolium medium
fizjoliiim repens
liissilngo fagara
lhtica dioica
Lfironica chamaedy
Vpronica scutellatn
?+cia sepium
?$cia .lylrmtira
Kola canina
C D. PRESTON AVII hf. 0. HILL
54
54
54
54
51
“Id
\Vld 1561 S,t
Am1
\!’Id
S\t
54
54
54
lVld
54
54
5454
54
Am1
!\’id
\Yld
54
54
54
\Vld
LYid 1561
Wld
Wid
W’id
Am1 Ltid
LVld
54
54
Am2
54
54
54
[Jtnculana austrah5
klenana o&( inali 5
Tiria rtacca
55
55
55
Tso
Wid
Wid [56]
56 +
56 +
56
56
56
Tso
56
56
56
56
56
Tso
56
Tso
+
56 Tso Wid
.
6
i
56-
‘rso
56
5(3
56
Euiaxan Boreo-temperate
Achillea rnill$olium
Achzllea ptavnzrn
Antennana dzozba
Anthmcus splziestnc
Calamagrostis epzgqw
Cicuta zliroJa
Festuta ouina
Fil@endula ulmaria
Galeopszc blfida
Galium uliginosum
(lalzum velum
Geranium pra/rn,e
Glrrtioma hedrratea
Gnaptialium uligznoum
Qmnadenia conoprra
Hrracleum >phon&zurn
Lamium album
Lmana riu(gnrzs
Luzula pallidula
Alaianthemum bzfoliutri
,Clelim nutanes
Oxalis aretosella
Peiszt ana birtorta
Platanthela bifilza
Populus tremula
Potamogeton comprpssir 7
Prunus padus
Ranunculu, rtptn,
Rubus jaxatilic
Saliv raprea
Solidago ozl;gaurra
Sol bus aurupana
Stellaria palustnr
Tanaceturn oulgarp
Thulictrnm minus
55
55
\$’id [56]
\Vid
J.1
rr
55
55
55 55 \Vid
55 \Vid
_-
31)
55
5.5
55
55
55
55
\;l’id [56]
Wid
\.$’id
\Vid [56]
Wid [56]
55
\$‘id
55 LVid
55 P\W [56]
55- Am3 \Vid
55 -
5.555
55
55
55
55
55
55
55
Am4 M’id 1.561
Sst
Wid [56]
i h 1 2 Sst
55
55
55
55
55
55
\.\’id
\Vid [56]
56
5(i
56
56
56
56 56
56
.56
56
56
56
56
56
Co2
+
‘Tso
Bol
5(5
5 6 t Bo2
56 ‘rso
56 \fid
56
56 +
56+
56
56
56
56
56
56
56
56
56
\\‘id
\$I%
+
Tax
56+
56s Tso
56 \Vid
56
56+ \$:id
56 +
FLURISTIC EI.ERIENI'S IN BR17':IIX AND I R E W l ' D
Rorirpa palustris
Rubus idaeus
Rumrx aquaticus
Satguisorba o8cinalis
Spaiganium emersum
Spaiganium natans
Starhys fia1ustri.r
Triglochin maritimum
Triglochin palustre
/Ttririilaria minor
Thnira scrpq'lllfolia
56
56
56 56+
56
56
56
56
56
56
56
Taiaiarum ofirinale a g
zodrra marina
\.\'id
97
66
66
Tuo
(,o1
Am4 \Vid
Am1 Col
C02
Tso
Tso
Wid
c:0 1
C h1
Ot Panic It ide-ternfirrate
t Sis) nnchium bermudzana
Am5
61
Tso
Eumpmn 1 lfdetpn2/letate
C'akil~maritima
Po!>gonunz oy p m i u t n
Ranunculus peltatus
63
63
63
Col
Am1 Col
I>jt
c:o2
c:o 1
COl
T'.AX.
Euincibmarr Ilfde-tmipate
Anttioxanthum odoratum
.4trz/,lex patula
dtriplex prortrata
C@~ l l abuiJa-/jn rtons
EL]t i zgza repens
Fallopia rourolruluc
Poa annun
Poa trmalzs
Rumm acptocella
Srhonioplectus lac 11rit is
kSjrrgula m e n u \
Cirrurrpofar IIidp-tpmperate
Agio rtz 5 rtolontfPm
Alicina plantago ayiratrra
Juiit us bujonzus
Juni us go-ardzr
Phra<pziPsaurtralir
Pan pratensiJ
Polygonum aoiculair
Pofamogeton prriinatus
Ptunrlla aulganc
Ruppza czrrhosa
64
64
64
64
64
64
64
64
64
64
64
.4s2 Wid [661
65
65
65
65
65
\$'id [66]
\$'id
LVid
LVid [66]
Wid [66]
Wid [66]
Wid [66]
\$'id [66]
Wid [66]
Syt
Wid [66]
Wid 1661
Ll'id Tax
\$'id [66]
66
66
66
66
66
66
66
66
\Yid
Tso Wid
Tso FVid
C02
Wid Sst
\Yid
66
( h l Tax
Col T a x
Col Tax
(h2
1-90
71
71
(lo1
ChI
\Vid [72]
71
71
Tso
7'50
Wid
C02 Tso
+ Am4
Tax
Ruppia marziima
(:ol
7I
71
71
71
l'so
66
66
rim 1 Tso
71
71
71
71
71
71
71
71
71
71
71
71
7I
71
71
71
71
'202 Tso
Subactmzc Empeiate
iipiuni
inundaium
72
72
72
T\o
98
C. D. PRESTON AND M. 0. HILL
Centaurea nigra
72
Chlysosplenium oppo~itfolium72
72
Drosera intermedia
Eleocharis multicaulis
72
Erica tetralix
72
Festuca jilformis
72
Galeopsis segetum
72
Galium saxatile
72
Gbceria declinata
72
Helleborus viridis
72
Hypericum pulchrum
72
Juncus sguamsus
72
Linaria repens
72
Littorella ungora
72
Luronium natans
72
Lysimachia nemorum
72
Ornithopus pequsillus
72
Pilularia globullfera
72
Pobgala serpyllfolia
72
Pobpodium inteljectum
72
Pvtamogeton pobgon@lius 72
Pvtentilla sterilis
72
9 Ribes rubrum
72
Seriphidiurn marztimum
72
Kcia ombus
72
European Temperate
‘4cer campestre
Aconitum napellus
Aethusa ynapium
Agrimonia procera
Agrostis vinealis
Ajuga reptans
Alehernilla xanthochlora
Alliana petiolata
Allium oleraceum
Allium scomdopasum
Allium ursinum
Allium vineale
Anagallis minima
Anthriscus caucalis
Aphanes amensis
Aphanes inexspectata
Apium repens
Aquilepa vulgaris
Amoseris minima
Arrhenatherum elatius
Arum maculatum
Asperula cynanchica
Asplenium adiantum-nigrum
Asplenium septentrionale
Aster linosyris
Astragalus &yphyllos
Atmpa belladonna
Bellti perennis
Berberis vulgaris
Berula erecta
Blechnum spicant
Wid
Am 1
Am2
Wid
Wid
Am2
Am3
Am3
Wid
Col Sst
73
73
73
73
73 Am1 Tax
73
73
73 As1 Wid
73 Wid
73 - Wid
73
73 N’id
73 Am1 Tso Wid
73
73 Wid
73 Wid
73 Tax
73 Wid Sst
73
73 Wid
73
73
73 Am3 As 1 Tso
73 Am4 As1 Djt
73- G o 2
7 3 As 1
73
73 Wid
73 Wid
73 Am1 As1
73 Am4 As2
+-
Bbsmus cvmpressuJ
Brachypodium glvaticum
Brassica nigro
Briza media
Bromopsis benekenii
Bmmopsis erecta
Bmmopsis ramosa
Bromus rommutatus
Bromus racemasus
Lallitriche plagcarpa
Callitriche stagnalis
Campanula latfolia
Campanula patula
Campanula trachelium
Cardamint amara
Cardamine bulbgera
CardamineJiexuosa
7373
73
73
7373
73
73
73
73
73
73
7373
73
73 73
As1
As1 As2 Wid
Wid
As1
Wid
Wid
Wid
Tax
Wid Tax
As1
Wid
As1
Am3 As1 As2
Tso
Carex arenaria
73 c02
Carex davalliana
73- Sst
Carex hirta
73 Wid
Carex hostiana
73 Am3
Carex montana
73- As1 As2
Carex paniculata
73
Carex pilullfern
73
Carex remota
73 As1 As2
Carex spicata
73 WidTax
Carpinus betulus
73
Centaurium littorale
73 c 0 2
Cephalanthera damasonium 73
Cephalanthera lonszfolia
73 As1
Cephalanthera rubra
73
Cerastium dlffsum
73 Tax
Cerastium pumilum
73
Ctrastium semiderandrum
73 Wid
Chaenorhinum minus
73 Wid
Chaerophyllum temulum
73
Circaea lutetiana
73+ Am1 As1 As2
Cirsium acaule
73
Cirsium eriophorum
73
Clematis vitalba
73 Wid
Clinopodium acinvs
73 As1
Clinopodiurn ascendens
73
Clinopodium menthfolium 73
Colchicum autumnale
73
Cbrnus sanguinea
73
Corylus auellana
73
Crambe maritima
73 Col
Crataegus laevigata
73
Crataegus monogyna
73 Wid
Crepis biennis
73
Citpis capillaris
73 Wid
Crepis mollis
73 Cynoglossum gmanicum
73 Cynosurus cristatus
73 Wid
cytisus scoparius
73 Wid
Dacglvrhiza majalis
73 Tax
Danthvnia decumbens
73
7 3 M'id
7 37 3 Wid
73
73
7 3 As1 Wid
73
73
73
7 3 + Am1 As2
7 3 As1 As2
73
73
7 3 Tax
7 3 Tax
73
7 3 \$'id
7 3 As1
73
7 3 - \Vid
73
73
73
73
73
7 3 As1
73
Filngo minima
73
73
Fro xinus e n PIJior
Fritillann nzeleagri c
73
7 3 - As1 As2
Gatyen lutea
Gulropszs ungustifolza
73
&diiirn aparinr
7 3 \$'id [7G]
Galiicrn odoraturn
7 3 As1 As2
Gnlzurn punzzlrttn
73
Gellitfa pdosa
73
Genitta tinctonn
73
Gentinnella gemianzra
7 3Gentinnella ufiqino3n
73
Geiniiiuin coliitnbinum
7 3 \Yid
9 Geinnium pj ienaiiiiin
7 3 LVid
Gernnnirn robrrtznniini
7 3 i\sl As2 \Yicl
Geinniurn Jarguinrutn
73
7 3 \Vid
G!) trna notata
7 3 As1
Giornlandia dm ra
73
Helinntliemutn numtnulnrium 7 3
Helictotrirhon pi-ntme
73
Helidoti-dionpubesrrns
7 3 As 1 \$'id
H$i,bocrepis c.omosa
73
Ho1cu.r mollis
73
Horde!yrnuJ cumpaezc.r
73
Hordeurn Jrcalinum
73
Hornungia p e t m a
73
Hottonin palustri.5
73
H ~ ~ i r i i c u rliumfu.wm
n
73
Hyprriczrrn montcinurn
73
Hyperirurn tetrnptemni
73
Dianthus armena
Dianthus gratzanopolztanuJ
Dtplotaxi r tenufolia
Dqracus fullonurn
Dtptncus pzlorus
Dinba rnuralz
DToptens a@nzr
Dg~opterz~
dilatata
Elatine hexandra
Eleorliaris paniiila
Epilobiurn montanurn
Efiilobzurn obscuritrn
Epzlobzurn pamtflorurn
Epipat ti^ leptochiln
Ep$m tis pl;yllantlier
Epipactu puipuiata
EuorcirnuJ curopaeur
Eupatorzurn (annabinurn
Euphorbia anggdafoideJ
Euphorbia (YParisczaJ
Euphotbin I zlloia
Euplirasza rnirranttin
Eupliraia nernorosn
Faguc yloatzra
FeJtuc n alti rema
Fertura gzgnutea
Filngo IuteJienJ
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
Am3
\$'id
\Vid
:\sl \Yid
As I
\$'id
Am 1 As I As2
\\.id
-,
13-
73
73
73
73
73
73
73
73
73
73
-,
1373
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
7:373
73
73
73
73
73
73
73
73
\Vid
Am3
\\id
\\id
l\id
\\id
\\id
\\id
A52
. t n l \\id
SSl
A\l
-
.\\I
A d 3 .Ad
\\'id
100
Poa compressa
Pobgala riulgaris
Pobgonatum multy7orum
Pobgonum ruriuagum
Potamogeton acuttjilius
Potentilla anglica
Potentilla neumanniana
Pllmula elatior
Pnmula vulgaaris
Prunus avium
Prunus spinosa
Pulmonaria obscura
Pulsatilla vulgaris
Quercus petraea
Quercus robur
Radiola linoides
Ranunculus aquatilis
Ranunculus Jammula
Ranunculus Juitans
Ranunculus peuicillatus
Ranunculus sardous
Ribes uva-crispa
Rorippa gilvestris
Rosa agrehs
Rosa amensis
Rosa caesia
Rosa caninn
Rosa micrantha
Rosa obtustjilia
Rosa rubkinosa
Rosa sherardii
Rosa .r$losa
Rosa tomentosa
Rumex lydrolapathum
Rumex obtusfolius
Rumex palustris
Rumex sanguineus
Sagina subulata
Salvia pratensis
Sambucus niga
Sanicula eumpaea
5 Saponaria o$lcinalis
Saxtjaga granulata
Scabiosa columbaria
Scleranthus annuus
Sclerauthus perennis
Scorzouera humilis
Sedum acre
Selinum cannfolia
Senecio aquaticus
Senecio syluaticus
Senecio uiscosus
Serratula tinetoria
Silene noctZfEora
Silene otites
Sorbus aria
Sorbus torminalis
C. D. PRESION A I D hl. 0. HILL
73
73
73
73
73
73
73
7373
73
73
7373 73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
73
7373
73
Wid
Spirauthes aestzvalu
Sta(lys alpzna
As1 As2
Tax
Am3
As1
Wid
Wid
Am4Asl As2
Djt Tso
Wid
Wid
Wid
Wid
Wid
Wid
Wid
As1 Sst
Wid
Wid
Wid
Wid
Sst
Statlys germanica
Starlys o$lcznabs
Stellai ia neglecta
Stellana ulzgznosa
73
73
73
73
73
73
As1
As1 As2
Am3 As1 As2
Tso Wid
As1 Wid
$mphytum ojicinale
@mphyturn tuberosum
E x u s barcata
Efsdalia nudzcaulz,
Zucnum botys
7hymus pulegzozdes
7ilza pla$J$yllos
Tnfolzum dubium
Trfolzurn orhroleucon
Tnsetum Jauescen r
Lrlmus glabra
L’lmus minor
8 lilmus procera
Lhlenana diozca
Vnleizanella dentata
1 hlmanella locusta
Vnleriandla rzinosa
Vrbascum brhnztzs
L’eronica agrestu
L’eronzca montana
konica tnphylloJ
lFbumum lantana
Lkza hirsuta
Hiia lathyroides
Hi za tetra ,perma
Kola odorata
Kola rezchmbarhzana
Kola nuinzana
Kola trzcolor
k u m album
73
73
73 Sst
73
73 73 Tax
73
73 Wid
73
73 Wid
73
73 Wid
73 Tax
73 4 m l As1
73 As1
73 Wid
73
73
73 \Yid
73
73 73
73 Wid [76]
73
73 Wid [76]
73 Wid
73
73
73 Wid
73 As1 As2
Euroszbenan Zmperate
A n u s glutzuosa
Althaea o@cinalu
Anrhusa arvenizs
Anemone nemoroJa
Arabzdopszs thalzana
Arabis glabra
Arctium lappa
Aitemzsia absznthiuni
Artemzsza campestns
Artemzsza uulgans
Asparagus o#cznahs
Atnplex pedunculata
Bubarea uulganr
Braclypodzum pznnaturn
Butomus umbellatus
Carduus cnspus
Carduus nutans
Carex acutfomis
Carex cayophyllea
74
74 Co2 Wid
74 Wid
74
74 Wid [76]
74 Wid [76]
74 Wid [76] Tax
74 Wid
74 74 Wid
74 Wid
74 Co2 Djt
74 Wid
74
74 FYid
74 Wid [76]
74 Wid
74
74
102
Impatiens noli-tangere
t h u l a salicina
Qsimachia uukaris
Lythrum salicaria
Myosotis sylvatica
Myriophyllurn spicatnm
J%jmphoidespeltata
Odontites uernus
Persicaria maculosa
Persicaria minor
Picris hieraciaides
Pluntago media
Pobgonatum odoratum
Po!vstichiirn aculeatum
Ranunculus rircinatus
Ronppa amphibia
Rosa pirnpinellfolia
9 Salix triandra
3 Salix viminalis
Schoenoplectus triquekr
Sedum telephiurn
Seseli libanotis
Thlaspi amense
Zrilis japonica
Kola rupestris
C. D. PRESTON AND M. 0. HILL
75
75 75
75
75
75
75
75
75
75
75
75
75
75
75
75
75
75
75
75
75
75 75
75
75 -
Am4
Wid
Wid [76]
Wid
Wid [76] Sst
Wid
M‘id
Wid [76]
Wid
Sst
Wid
Djt Wid
Wid
Tso
LVid
Wid [76]
Wid
Circumpolar Tmperate
Ali.rrna gramineum
Aspleniiim ruta-miiraria
Astragalus danicus
Atriplex littoralis
Bidens cernua
Cabstegia sepiuni
3 Chenopodium glaucurn
5 Chenopodium hybridum
Clinopodiurn vulgare
Crassula aquatica
D yopteris Jilix-mas
E1eochari.r uniglumis
Galium spurium
Ghceiia maxima
Kbeleria marrantha
h n a trisulca
Monotropa tLyfiopi&s
12.lyrioplyllum oerticillatum
Ophioglossum vulgatum
Persicaria lydrop$er
Pteridium aquilinum
Rumex maritimus
S p a p n i u m erectum
The&terij palustris
konica catenata
Eburnum ofiu1u.r
76$ Sst
76
7 b $ Am4
76+ Co2
76
76 Tso
76+ Am4 Tso \Yid
76
76+ Tso
76
76
76 T5o
76- Tso Wid
76 M’id
76
76 Tso
76
76
76
76 Wid
76 Tso
76 Tso
76+ Am4 Tso
76+ Tso
76f Tax
76
Oceanic Southemtemperate
Agrostzs curtzsii
81
+
+
Carurn verticallaturn
Centuunum srzllozdes
t Daboecza cantabnca
EleogztonJlzcztans
Enca cilzam
7 Eirca engena
Ei-zca vagans
Euphorbza portlandzca
Furnana muralis
fiumana reuten
Heniana czlzolata
HjFencunz linanZfohum
Hvpencum undulatum
LPpzdznm heterophyllurn
Lobelza urenr
Po!lgala calcarpa
Puicznellza rupestns
Ranunculus tnpartztus
Salzrornza nztenr
Scrophulana scorodonin
Sedum f o i rtenanuni
Ulex minor
Whhlenbeigza hedeiacea
Suboceanzc Southern temprrate
Azia praeror
Aloperurus bulbosur
Asplenium mannuin
Baldrllzn ianunt uloides
Rrauira oleracea
Bunium biilbotaJtanum
Callztrzciie obtubangula
Cardiius tenulfEoniJ
Carex pun1 tata
Chamaemelum nobile
Ciiszum tuber0 rum
Coinrqa nionenszs
Corngzola litoralir
Dzqitalzs puvpurea
Eiodzum lebelzz
Erodiurn mantzmurn
Euphorbza hjiberna
Frankenza laeuzs
Helleborus foetzdus
Hydroc o p l ~ilulgan,
I h m arnara
Ilex aqufolzuin
Illreebnim vertzczllritiim
Iri \ fortzdzrsima
Jiincur foliorus
khelerza vallesiana
Lartzita wora
hontodon saxatili,
Lonicein penc!ymenurn
Lotus subbZfl0rus
.2hbora minima
,2loenrhia ere( ta
81
81
81
81
81
Djt
Djt Tso
81
81
81
81
81
Djt
81
81
81
81
81
81
81
81
81
81
81
81
81
Col
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
\$‘id
Co2
Col Tax
Wid
Col Tso
Tax
Co2
\4’id
Djt
\Vid
\st
LVid
C02
FLORISTIC ELEhlEN I \ IT BRITAIN .ZND IRELqND
Ornanthe irocata
Oenanthe lachenalii
Ornithopus pinnatus
Orobanche rapum -genistae
O5munda regalzs
Parapholis stngosa
Petrorhagia nanteuihi
Petroielznum segetum
Puccinellia famculata
Ranunculus hederaceus
Ranunculus omtophi,lluJ
Ranunrulus pan$orus
Salicoinia ramosusima
Scrophulana auriculata
Scutellana minor
t Szniethis planlfolia
Spartina mantima
Staclys arvensls
Eucnum Jromdonza
Tnfolium ornithopodioides
Llrbascum virgntum
Euiopean Southern temperate
h a cayophyllea
Ajuga chamaepngJ
Allium sphaerocephalon
A lopecums my05u roides
Ammophila arenana
Aiiacampti r &mmidalis
ilnisantha stenlzs
Anthemis amensw
Anthmzs cotula
Ballota ngra
Bromui hordeareus
Bupleuruni tenuiJJimum
[ arex distanc
Carex extensa
C arex fracca
Carer pendula
Centauriitm eythram
9 Cprastzum braclybetalum
Ceiastium glomeratum
Chpsanthemum segetum
Coronopus squaniatus
Coynephorus ianesreni
Cjperus longus
Ebtngza athenca
Ehtngza juncea
Equisetum telmateia
Eyngium campestre
Eyngium mantimuni
Euphorbza exigua
5 Euphorbia lathyns
Euphorbia peplu~
Euphorbia pla<yphyllos
Euphorbia sermlata
Fzlago uulgans
Fumana denszjlora
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
82
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
Am1 As2 Tso
c oI
Tax
Am1 C02 Tso
Col Tax
COl
Wid
Wid
Wid
Col Wid
Wid
Wid
Wid
Wid
c102
c02
Co 1
Wid
Wid
Wid [86]
Wid
Wid
Go2
Co2
co 1
Am4
Co 1
Wid
Wid
As 1
Fumaria oficina1i.r
Furnoria pawflora
Ckgiaa bohemira
3 Galanthus nioalis
Gmmium diJ.wtum
Geruiiium molle
Hedem heli,x
Uolrus latiatm
Hypochaeris glabra
Hyporhaeris radirata
Ii-iJ pseudaconis
3uncus ambigitus
,Juiicui capitalus
3uncus effusuJ
Junrus marifirrus
3unc.u~JubnodulosuJ
kirk,ria elatine
kIi-kxia spuria
Lactuca saligna
Lgozisia hybrida
LPmna gibba
Lvurojum aestivum
Lithospeimurn
~urpuroraendeum
Lolium perenne
Lotus angurti.rsiinus
Lotus glaber
Ludwigia palustris
M a h i i a r i a rerutita
,\lentha p u l ~ g ' u m
i\!)losotis ramosissima
Oeiianthe j i bfolk1
Orchis simia
O m i t h o g a h angu tfolium
Orobanche artmiisiarrampe.stric
Orobanche miuor
Papaver axemotit
Papaver rhoen3
Auredanum o@cinal
Phleum arenarium
Phleum bertolonii
3 Pirrir erhioides
Potamogeton coloratui
Ranunculus baudotzi
Ranunculus bulbosus
Ranunculus ficaiia
Ranunculus ophioglossijoh i
Raphanus raphanistmni
Rueda lutea
RiibusjuticoJus a g .
Rumex pulcher
Sagitia apetala
.Yagina maritima
4 Sambucus ebulus
.%x$ragu tridac9lite.i
tYenecio iiulgaris
I0:I
83
8:3
83 83
83
83
8:i
83
83
83
83
83
83
83+
83
83
83
83
83
83
83
83
83
83
83
83
83
8:3
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
83
\\'id
IVid
\Vid
\\'id
\Vid
LYid
Wid
Wid
Am1 Tax
\Vid
Am1 r t l ..\a2
M'id
C02 \Vid
lt'id
Am1 .\sI
LVid
\Yid
Xm1 \Vid
\Vid (861
\Vid
LVid
\Vid
Wid
co2
LVid
Wid
co2
Wid
FVid
\Vid
LVid
Col
LYid [86]
'Tso
104
4
Sherardia arvensk
Skymbrium ojicinale
Sonchus asper
Sonchus ateraceus
Svrbus domestica
Spetgularia rubra
Spiranthes spiialis
Zucrium chamaedy
Tnfolium striatum
Trinia glauia
Eluianella carinata
Pironica arvensis
Veronica hederifolia
Vuonica praecox
Kcia sativa
Kola kitaibeliana
C. D. PRESTON AND M. 0. HILL
83
83
83
83
83
83
83
83 83
83
83
83
83
83 83
83
Eurosiberian Southern-temperate
Agrimonia eupatoria
84
Alisma lanceolatum
84
Anagallis arvensis
84
Apera interrupta
84
Apium gravevlms
84
Apium nod@um
84
Arenaria serpyllfolia
84
BvlboschoenuJ maritimus
84
Cardamine hirsuta
84
84
Carex divulsa
84
Carex muricata
84
Carex vtrubae
Centaurium pulchellum
84
84
Clienopodium murale
Chenopodium vulvaria
84
84
Cichvrium inbbus
84
Cladium mariscus
Cvnium maculatum
84
84
Cvnvolvulus amensis
84
Crepisfoetida
84
Cuscuta epittymum
84
Cjpems$scus
84
Dacglis glomerata
84
Daucus carvta
84
Emdium ricutarium
84
Ervphila verna sensu lato
84
Festuca arundinacea
84
4 Galium tricvrnutum
84
Geranium rotundfvlium
84
Gnaphalium luteoalbum
84
Holvsteum umbellatum
84
Hordeum murinum
84
Hyoscyamus niger
84
Hq’pericum perfbraturn
84
Juncus articulatus
84
Juncus injexus
84
Lactuca serrivla
84
Lamium amplexicaule
84
Lepidium latiiolium
84
Limvnium bellidtjivliurn
Wid
Wid [8G]
Wid [86]
Wid [86]
,4m3 Wid
Wid
As1 Wid [8G]
Wid
Wid
Wid
Wid [8G]
C02 Wid
Wid Tax
Wid
C02 Tso Wid
Wid [86]
Tax
Tax
Wid
Wid
Wid [86]
h
l Tso
Wid
M‘id [8G]
Wid
Wid [86]
Wid [86]
Wid [86]
Wid
Wid
Wid
Wid
Wid [86]
Wid [8G]
Wid
iZm3 Wid [86]
Wid
Wid
Wid [86]
C02 Wid
Co2
Lithospemium urvense
Lythrum tyssopfolia
Malva sylvestris
Marmbium vulgare
Mediiago minima
Medicagv sativa
Misopates orontiuni
4 Murcari neglectum
Ononis spinosa
Pafauer dubium
Plantago coronapus
Plantago lanceolata
Pva bulbosa
Pvtamogetvn trirhvides
Potentilla reptans
Pulicaria dysenterica
Ranunculus amensic
Reseda lutevla
Ror$pa nasturtiumaquaticum
Rzcmex conglomeratuJ
Rumex crispus
Salk alba
Salsolu kali
Sangui.cvrba minor
Scandix pecten-uenerir
Srhoenus nkricans
Scirjmides holosrhvenus
Silene conica
S i h e latfolia
Spergulaia media
Stellaria pallida
’Tiucrium scordium
Thlaspi perfohaturn
4 lorilis amensis
Trfolium arvense
Tnyolium campe.rtre
Tnfvlium jagzfPruni
lirtiia wens
Veronica polita
Iklpia myuros
Wolfia arrhiza
Eurasian Southern-temperate
A<pstisgzgantea
Bupleurum falcatum
Epzlobium hzrsutum
4 Equzcetum ramom cimum
Euphvrbia helioscvpza
Hq’dnlla vertzczllata
Lotus cvrnzculatus
Organum vulgar?
Pvtamvgeton cnspus
Schoenoplectus
ta bmaemvntani
Silrnr oulganr
Solanum dulramara
Solanum nigrum
84
84
84
84
84
8484
84
84
84
84
84
84
84
84
84
84
84
Wid
Wid
Wid [86]
Wid
Wid
Wid [86]
Wid
84
84
84
84
84
84
84
84
84
84
84
84
84
84
84
84
84
84
84
84
84
84
84
LYid [86]
Wid [86]
Wid [86]
Wid
Tso Wid
M’id
\\’id
Am1
85
85
85
85
85
8585
85
85
Wid I861 Tax
85
85
85
85
(102 Tso
Wid [86]
Wid [86]
Wid
Wid
Co2 Wid
Wid [86]
Wid
Tso
Wid
h’id
Wid
Wid
Wid [86]
C02 Wid
Wid
Wid
Tso
Wid
Wid
Wid
Wid [86]
M’id
Tso
Wid
Tso
Wid [86]
Tso Wid
Wid
Wid
Tso Wid [86]
FLORISIIC ELESIENTS IN RRl rAIN AND I R E L U D
5
Suaeda maritima
85
Ihbena ofieinalzs
85
k o n i r a anacgallis-aquatica 85
Zoortera noltiz
85
Circunipolar Southowtemperate
86+
Aspleiiium tnchomanes
86
Ceratophqllum demeoum
86
I m n a minor
.Vqas manna
86Persic aiia lapathlfolia
86
Poa angusttfolia
86
Potaniocgeton nodosiu
86
Potanioqeton pu rilluJ
86
Snmolus ziakrandt
86
86
S p e p l a n a marina
Spzrodela pohrhiza
86
q p h n lattfolia
86
Zannic hellia palustris
86
,Zlediteiranean-Atlnntit
AceiaJ anthropophorum
Adiantum capillus-oenenJ
91
91
4 Allznm ampeloprasum
91
7 Arbutus unedo
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
91
drum italzcum
Asplemum obouatum
t Asplrriium onoptens
Atnplrx portulacoides
Beta rulgnnc
Buplrurum baldenJe
Calhtriche brutza
Callitliehe truncata
Cngstgza soldanella
Catapodium marinum
Centaunum tenu$onim
Cnthinum mantimum
Damnsonium &ma
Echium plantagineum
8 Erodzum moschatum
Euphorbia paraliaJ
Euphorbia pepliJ
Fnmana bastardii
Galzum constrictum
Gastiidturn irentrzcomn
Grraiiinm purpureum
Gladiolus zl&ur
Glauc ium jlaiium
Hordenm marmuni
h u l a rnthmoides
Isoetrs hirtrix
Irolepir remua
Juncur acutu,
J u n ~ i tpygmaeus
~
I h a t r i a arborea
Imatera rrptica
Co2 Tso Wid
Wid
\Yid
Col
Tso
Tso Wid
TSO
Wid
Tax
?'so
Tso Tax
Tso
Co2 Tso Wid
Tso
Tso
Tso
Am1 As1 As2
Co2 Tso
Col Sst
Co2
Tax
,4ml Wid
Col
Col
PVid
b'id
Col
Col
Col Wid
As1 C h 2 Wid
Co2
Ainl Co2 Tso
Am4 Co2 Tso
Limoiiium zzilgare
Linum bienne
9 XI(itthiola incana
,21atthiola sinuata
"Z~otinramaculata
Ornanthe pzmpinelloidrr
Oiioni, ieclmata
Ophioglorsum lutztanzc um
OtanthuJ mantimiis
Pnrapholzr zni unw
PorenturPllia L ~ rco
I ra
Poa inJlima
Pull tailon tetraphyllum
Polygonum mantimum
Po!,podium cambi i( uni
Polppogon mon$dienri \
Rcimiilea cohmnae
Kubin prregiina
crlr in I rrbmara
nirotomza perennis
killo nutumnalis
\pr?gulaiia boc conri
h e d a ucra
f i n l i t nodosa
l i i f o l i u i i i bocconei
-lifnhuni glomeintum
7llfolium iniamatuni
Injnliitm squamo\um
I i i / o l i u m suffocatiini
firbrrnna guttata
1 inbilititr iupestnJ
15cui bzthynica
lhl$nl r1lzntfl
I L@ia fascn ulata
Jubniediterranran-.\ubaihntzl
Bloc hrtonin pefolrata
Bg onia dzoicn
Bnw( <emprnirm
( arm depauperato
( nrrx
diuita
(.atupodium ngidum
Ceterach oficinanim
C 11 eiidia filtfornziJ
( Iinoifiodiurn i alamintha
( raJtrila tillaea
Dcipliiie laureola
Epilobium Iancrolatum
Filago pyramidata
Fumaiia capreolata
Gnfiuni paruiense
Geinnium lucidiini
Himontogloiium hirc mum
I l p ,n nm andm saemum
9 I& iu r nphaca
I,uzgil(i f o r s t m
'Zlrdiiago ainbica
.llrdztago po(ymorp1ia
105
91
91
91
91
91
Col
L$ld
91
91
91
91
91
91
As1 S i t
Col
\$'id
91
91
91
91
91
91
91
C o l Tso
LVld
Co2
91
C)
1
91
91
91
91
91
91
91
Co2 - I s 0
As1 \Vid [92]
\Lid
')I
91
91
co2
91
91
91
91
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
c02
M'id
.I51
Ch2
A\I
Azl
LVid
X r l \Vld
Wid
106
Mentha suaveolens
Mercurialis annua
Minuartia hybrida
Ophys apfira
Ophys>c$ora
Ophrys sphegodes
Omithogalum pyrenaicum
Orobanche hederae
Papauer hybridum
Parietaria judaica
Polystichum set+rum
Ruscus aculeatus
4 Sedum album
Silene gallica
Sison amomum
Tamus communis
Triiolium micranthum
Tri$diurn scabrum
Tn9lium strictum
Tnyolium subtmaneum
Verbascum puluerulentum
Ecia lutea
Ecia parugora
Vulpia bromoides
Vulpia unilateralis
Mediterranean-montane
Arabis scabra
Draba aizoides
Lhyopteris submontana
Helianthemum apenninum
Helianthemum canum
Potentilla rupestris
C. D. PRESTON AND h4. 0 . HILL
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
92
93
93
93
93
93
93
B Gentianella anglica
As 1
Wid
Wid
Wid
Wid
Wid
Wid
As 1
Sst
Wid
f Limonium britannicum
f Limonium dodartforme
7 Limonium loganicum
f Limonium paradoxum
f Limonium paruum
f Limonium procerum
7 Limonium recunluni
f Limonium transwallianum
f Primula scotica
7 Senecio cambrensis
f Sorbus anglica
f Sorbus arranenszs
f Sorbus bristoliensis
f Sorbus deuoniensuf Sorbus eminens
f Sorbus hibemica
7 Sorbus lancahensis
7 Sorbus leptophylla
f Sorbus lpana
7 Sorbus minima
7 Sorbus povigentformis
f Sorbus pseudofennica
f Sorbus subcuneata
7 Sorbus uexans
7 Sorbus wilmottiana
f Spartina anglica
f Ulmus plotii
71
71
71
71
71
71
71
71
71
41
71
71
41
71
71
71
71
71
71
71
71
71
41
71
71
71
71
71
41
11
71
41
71
41
11
71
71
71
71
11
41
41
41
71
41
41
71
71
Co 1
Sst
z
Sst
Co 1
Col
Col
Co 1
Col Wid
Species restricted to the Channel Islands
Anogramma leptophylla
91 Am5 As1 Tso
Anneria arenaria
82
$ Evaculum pusillum
82
$ Festuca armoricana
71 Col
Limonium auriculaeursfolium
81
'+ Limonium nonnannicum 7 1 Col
Linaria pelisseriana
91
1Miliurn uernale
91
AGosotis sicula
91
$ Orchis laxzjora
84 $ Ranunculus paludosus
91
$ Schoenoplectus pungens
73 A m 5 Go2
z
Endemic
f Alchailla minima
f Athyrium Jexile
f Bromus intemptus
7 Calamagmstis scotira
f Centaurium latfolium
f Cerastium nigrescens
7 Cochlearia micacea
f Coincya wrightii
f Cotoneaster cambricus
f Epz$actis youngiana
f Euphrasia anglica
f Euphrasia cambrica
f Euphrasia campbelliae
7 Euphrasia heslop-havisonii
f Euphrasia marshallii
f Euphrasia pseudokemeri
f Euphrasia riuularis
f Euphrasia rotundfolia
f Euphrasia uigursii
f Fumaria occidentalis
Col
Col
Col
Col
Col
Col
Col
Col
Species not assiped to elements
Cystopteris dickieana
??
Erophila glabrescens
??
Erophila majuscula
??
3 Oenothtra fallax
??
Poa humilis
??
Ron$pa micmphylla
??
Utricularia ochroleuca
??
Utricularia s&ia
??
f Sagina boydii
??
zoostera angustfolia
??
FLORISTIC: ELE>IENT\ IN IIRI 1,UN &%I)
IRELWD
107
APPENDIX 2 : ~ ~ I , P H ~ \ ~ ~ ~IJST
~ I ' IOF
~ : SPEC:IES
~\I,
Species arc listed alphabetically in this Appcndix. w i i h appropriate qualifiers. For tlir key to symbols,
sec Appendix 1.
Arer rampestre
A r e r a ~anthropophorum
Aihillm mzllpfolium
Arhillen ptarmztn
Aroniturn napellu
Actaea spzcata
Adiantum capillus-oeneiic
Adoua moschatelhna
Aetliu\a cjnapiiim
14gnmonza rupatona
"ipmonza proceia
Agrosti, ranina
Agroctu rapillaru
iigrostii curiicii
Agno \ti r giguntea
.4groctis \tolonzfpra
A4gro,tic.rlznealzr
. h a tayoptpllea
Aiia prarcox
Apga rhnmaepig\
.$uga pyiamidalu
Ali~gareptanc
Alrhrmilla acutiloba
Alcfwnilla akma
14khemillajlzr auha
Ali/irniilln glabm
d I<lirmilla glaures renJ
dlclienizlla glomeiulan r
Alrlietnilla giacilis
7 dlchrmilla minima
~llchrmillaniorihiola
A11 heniilla rubrrenatn
Alrhvmilla u icliuiae
illr/rrmilla uanfhorhloia
Ali\mn ginmineurn
Aksnin Iaricrolatum
AhJnin plantago aquatica
illlzmm p~trolatn
S Allium ampelo/~ramrti
Rlliiitn olerar rum
A l l i u m ~tioenoprariim
=Illium J( oiodoprasiim
a411iiitn sphaenu ephalon
.Alliuni iminum
.4lliritri tineair
Alnua glutinoaa
Altipr urn \ aequali \
73
91
35
55
73
46
91
56
73
53
83
\$'id [56]
\lid
+
i\ml As1 A52
Co2 Tso
84
\$'id
46
71
if
83
7 'I
81
74
81
73
56
54
81
54
74
83
83
+
85
66
73
83
82
83
4'3
ll'ld
91
\.$'id [86] Tax
b'id
Am1 Tax
\.%id
\$'id
b4
M'id [8F]
AlT1i-rso\vld
\\id
\$'id
;\m5 .\\I
TfO
\$id
\\id
A\? \I
id [6b]
77
5.5
\lid
\.Lid
An2
Am1
43
53 \$'d
53 23 ,\nil
5741
53 53 43
7'3
7F*
84
66
73
91
73
83
37
84
it
73
53 -
IS
55
83
\\'id
\Vid
b'id [56]
c.02 \%Id
Col \\Id
84
Cm.2 \$'id
72
lw
84
71
\+id ' l a \
7't~
\.\id 5st
l l i d [76]
41111 S',I T a x
\\Id [ i h ]
73
71
I4
Am2
74
S\t
jb
15
Tso \$'id
4 s l \$'id
93
91
74
-,/ 1
\\id [7G] T a x
\\Id [7h]
IVid
16
26- \Vld
4b
I3
1'3
Am2
84
\\Id
7 3 - \Vid
83
73
73 LVld
74
56
16- €301
82
3h
73
73
74
€301
c 02
\\id
\\id
108
C. D. PRESTON AND hl. 0 . HILL
Artemisia campestris
Artemisia nurvegua
Artemisia vulgaris
Arum italicum
Arum maculatum
Asparagus oficinalis
Asperula cynanchica
Asplenium adiantum-nigrum
Asplenium marinum
Asplenium obovatum
7 Asplenium onopteris
Asplenium ruta-muraria
Asplenium septentriunale
Asplenium trichumanes
Asplmium trichumanesramosum
Aster linogris
Aster tnlpulium
Astragalus alpinus
Astragalus danicus
Astragalus g&cyphyllos
Athyrium distent@ium
Athyrium jilixjiemina
AthyriumJlexile
Atriiplex glabriuscula
Atriplex laciniata
Atriplex littoralis
Atriplex lon&pes
Atriplex patula
Atnplex pedunculata
Atriplex purtularuides
Atriplex praecux
Atn$lex prostrata
Atrupa bdladunna
Baldellia ranunculuides
Ballota nigra
5 Barbarea stricta
Barbarea vuLgaris
Bartsia alpina
Bellis perennis
Berbmk vulgaris
Berula erecta
Beta vulgaris
Betula nana
Betula pendula
Betula pubescens
Bidens cernua
Bidens tnpartita
Blackrtunia pdoliata
Blechnum spicant
B&mus compressus
Bbsmus rufus
Bolbuschuenus maritimus
Botyhium lunaria
Brachypudium pinnatum
Brachypudium glvaticum
Brassica niera
Brassica ulerarea
Y
74 13 Bol
74 Wid
91
73
74 Wid
73
73 Am3 As1 Tso
82 Col Tso
91
91
76 -t
73 Am4 As1 Djt
86+ Tso
+
+
46
7375
16
76+
73
16+
56
11
52
71
76+
43
64
74
91
43
64
73
82
83
54 74
13
73
73
73
91
26
54
54
76
75
92
73
7343+
84
56
74
73
73
82
C02
Co2
Am4
As1
Tso
Am1
Am1 Col
C02
Col
Wid
Co2 Djt
Col Sst
Am2 Col
Wid
Wid
Am1
Wid
Wid
Am1 As1
Co2
Wid
Am4 As2
As1
Am1 As1 C02
Co2 Tso Wid
Tso
4 s l As2 Wid
Wid
Briza media
Brumupsis benekeriii
Bmmupsi,r erecta
Bmmopsis ramosa
Brumus cummutatus
Brumus hurdeacms
7 Brumus intermpius
Brumus racmusus
Bryunia dioica
Bunium bulbucastanum
Bupleurum baldense
Bupleuruni falcatum
Bupleurum tenuissimuni
Butumus umbellatus
Bunus sempervirens
Cable maritima
Calamagmstis canesrrns
Calamagrustis epigejus
Calamagrustis purpurea
7 Calamagrostis scutira
Calamagmstis stricta
Callitriche brutia
Callitriche hamulata
Callitriche hennaphruditira
Callitriche ubtusarigula
Callitrictiepla9carpa
Callitriche stagnalis
Callitriche truncata
Calluna uulgaris
Caltha palustri3
Carystegia sepium
Calystegia soldanella
Campanula glumerata
Campanula lattjiulia
Campanula patula
Campanula rutund$olia
Campanula trachelium
Capsella bursa-pastoris
Cardamine amara
Cardamine bulbtjira
CardamineJlexuusa
Cardamint hirsuta
Cardamine impatiens
Cardamine praterisis
Carduus crispus
Carduus nutans
Carduus tenu$orus
Carex acuta
Carex acutfunnis
Carex apprupinquata
Carex aquatilis
Carex arenaria
Carex atrata
Carex atmfsca
Carex b@owii
Carex binenis
Carex buxbaumii
Carex capillaris
73
73- As1
73 Wid
73
73 Wid
83 Wid
71
73 Wid
92
82
91
85
83 Co2
74 Wid
92 Wid
63 Col
54 55
44 Tax
41
26 Tso
91 Tax
52 Am2
46
82 Tax
73 Tax
73 Wid Tax
91
53 As1 Wid
36
76 Tso
91 Am4 Wid
75 73 As1
73- Wid
56
73 As1
64 Wid [66]
73
73 73 AmJAsIAs2Tso
84 Wid [86]
75 Wid
36
74 Wid [76]
74 Wid
82
54
74
54 26 Bol
73 c 0 2
26
16
16
71
4 4 4 Am1 As2 Tso
26
FLORISTIC ELEMEN1 b I\ BRIT.\IN AND IRELAND
Carex cayophyllea
Carex chordorrhiza
Caiw curta
Carer da valliana
Carex depauperuta
Carex diandru
Curex digitata
Carex dioiia
Carex distans
Cuizr disticha
Carer dizisa
Caw diz'ulsa
Cur-ex echinata
Cartx elata
Carer donguta
C a w Pricetorum
Carer extensa
Carer ,filformis
O a i z .fracca
~
C a m $mu
Carer liirta
Carer hostiana
Carer humilis
Carer la-henalii
Carex laeuigata
Calm lasiocarpa
Calm limosa
Corm magellamica
Cura niaritimu
C a m microglochin
C u m montana
Carex muric.atu
Carex ngru
C a m noruegca
Carex ornithopoda
C a m otiubae
Carer ovalis
Curex pallesiens
Carex paning
Carp\ paniculutu
Carex paurg7oru
Cawu pendula
Carex pilultfpru
Carm pseudog$trus
Care.? pu1icari.r
Care.? punctata
Care"xranJora
Carex recta
C a m rpmotu
C a w liparia
Care.r rostrata
Corm ruprit.;
Carex saxatilis
Carer .rpicuta
Carex sl~gosa
Carrx .yluatira
C a w trinervis
7-126 46 Tso
73- Sst
92 As1
56 Tso
53- As2
26
83 Co2
74
92 Go2
84
53 Am1 Sst
75
54 54- Sst
83 Col
74 83 Wid
53 Am1 As1
73 Wid
73 Am3
75 16 Tso
71
46
46
46 Tro
16 Co2 Tso
13+ Am1 As1 Tso
73- As1 As2
84 Tax
54 Am3
16
43
84 Tax
54 Wid
54 Am1
53 As1
73
46
83
73
74 Am1 As2
72
82 Co2
16 Bol
41 Am1 Bol Col
Tax
73 As1 As2
74 SSl
56
16
16 Bol
73 \Vid Tax
72
75
71 Col
uaginata
ilesiraria
ciridula
mlpina
Carlina vulgarir
Carpinus betu l u ~
Carurn verticillatum
Catabrosa aquatica
Chtapodium marinuin
Catupodium rigidum
C'mlnurpa nigra
LZntaurea scabiosu
(kntnnn'um elythraea
7 (Pntaui-ium latijiliuni
Ceritaurium littoink
C;entaurium pulchellum
Centaurium scilloida
CZntauriuni tenuiflorun
C'ephaluntliera dainusonium
("Pphalanthera longijdiu
C;eplialaanthtra iubra
Cerastium alpinuni
Cmctium arcticuni
( k u t i u i n arveme
4 Cerastiurn brarl;ypetalum
Ceractium cera.rtoide,c
Cemtiuni d$iu.wn
Cerastium fontanurn
Cure.t
Carex
C:clre.r
Carer
I09
26
56
56
74 - 'rdX
74
73
81
53 Am1 As1 'T\o
+
+
92
72
74
83
71
73
84
81
91
73
73
73
4
4
4
LVid
CVid
\\ id
co 1
Co2
Dlt
.As 1
13
13
56
83
13
73
54
83
41
C'eiastium pumiluni
Cera.,tiuni Jemidemndruni
CZrutocapnns rlauiculata
Ceratophyllum denirrmni
Ohatophyllum submerxm
(2terac.h o$icicinaruni
C;hamorhinum minu3
(z7iarroph,~llumteniuluni
~hmuemelumnobile
Cliamerion angusttjolium
Clielidonium niajus
C%enopodium album
(,%mopodium chenopodioides
Chenopodium jicgoliuni
Chopodium glaurum
Ohmopodium hybridurn
(:henopodium ninrab
Chenopodium polyspemum
(.%enopodium rubrum
(:/Ieuopodiumurbirum
Chiopodium uulrnria
Chn'mnthemuni segetum
Chgm'phium altem#oliirni
C%yosplmium
opposit~oliuin
ieridiaj!foomris
rerbita alpina
rhorium inpbus
C'O 1
91
73
73
71
86
71
92
73
73
82
56
_/ n_
65
74
74
76
76
84
74
71
74
84
83
36
Am 1
Am 1 Bo 1
7 so \Vld
A n 1 A51
I'ax
LVid [56]
k\'id [86]
bst
Lt'id
'rso
As 1
LVid
LVid
\\'id [66]
b'id
+ An4 Tso M id
Lt Id
\Vid
h 4 \Vid [76]
LVld
LZ'ld
72
92
43
84
LVid [86]
110
Cicuta virosa
Circaea alpina
Circaea lutetiana
Cirsium acaule
Cirsium amme
Cirsium dissecturn
Cirsiurn eriophorum
Cirsium heterophyllum
Cirsiurn palustre
Cirsium tuberosurn
Cirsium vuhgare
Cladium rnariscus
Clematis vitalba
Clinopodium acinos
Clinopodiurn ascendens
Clinopodiurn calamintha
Clinopodiurn menthfoliurn
Clinopodium vulgare
Cochlearia anglica
Cochlearia danica
7 Cochlearia rnicacea
Cochlearia oficinalih
Cochlearia pyrmaica
Coeloglossurn uiride
Coinga rnonensis
7 Coincya wrightii
Colchicum autumnale
Conium maculaturn
Conopodiurn rnajus
Conuallaria majalls
Convoluulus amensis
Corallortiiza trifida
Cornus sanguinea
Cornus suecica
Coronopus squamatus
Compzola litoralis
Corylus avellana
Corynephorus canescens
7 Cotoneaster cambricus
Ci-arnbernaritima
Crassula aquatica
Cr-nssula tillaea
Crataegus laevigata
Crataegus rnonogna
Crepls biennis
Crepis capillaric
Crepisfoptida
Crepk mollis
Crepis paludosa
Crefis praemorsa
Crithmum maritirnurn
Cmciata laeuipes
Cryptogramma crispa
Cusruta epithymum
Cuscuta europaea
Cjnoglossum genanicum
@mglossum ojicinale
(ijnosuruh cristatus
C. D. PRESTON AND M. 0. HILL
55 46
73+
73
75
71
73
44
54
82
74
84
73
73
73
92
73
76+
71
71
Am1 As1 As2
Wid [76]
Wid
Wid
h i 1 Tso
Wid
As1
Tso
Col
11
36
13
46
82
71
73
84
71
53
84
46 73
23+
83
82
73
83
71
73
76 +_
92
73
73
73
73
84
73 53
74 91
74
43
84
74
73 74
73
802
Wid
As2 Wid
Wid [86]
Am1 As2 Bol
Wid
Co2
Sst
Col
Mld
Wid
Col
As2 Sst
Wid
Wid
Wid
Wid
Cyperus fUSCILJ
Cyperus longus
Cypripedium calceohs
Cystopterk dickieana
Cystopteris fragilis
Qstopteris montana
Qtisus scoparius
t Daboecia canfabrica
Daclylis glomerata
Dartylorhiza jiurhsii
Dactylorhiza incarnata
Dacplorhiza lapponica
Dacplortiiza maculata
Darplorhiza rnajdis
Dacplorhiza praetenlssa
Darplorhiza purpurella
Dacglorhiza traunJteineii
Damasonium alirrna
Danthonia decurnbens
Daphne laureola
Daphne rnqereurn
Daucus rarota
Desciiampsia cesflitasa
Descliarnpsiafrexuosa
Dwhampsia setarea
Dianthus anneria
Dianthus deltoidej
DianthuJ gratianopolitanus
Diapensia lapponica
Dgitalis puqurea
Diphasiastrum alpinum
Dibhasiastrum rombianatum
Diplotaxis tenufolia
Dipsacus Jitllonum
Dip.racus pilosus
Draba aizoides
Drahn inrana
Draba rnuralis
Draba noniegica
Drosrra intennedia
DroJera longfolia
DroJera rotundlifolia
Dlq'as octopetala
Dryopteri.5 aernuln
Dryopterir sfinis
DryopterU carthusma
DryopteriA cristata
DTyopterU dilatata
Dryopteri, expanso
Dyopteris,fili.u-ma.,
Dlyopteric oreades
D?yopteri.s rpmota
Dlq'opteriJ submontana
Ediium planta<+witrn
Echiurn uulgan
Elatine he.randra
Elatine Itydrop$er
ElPocharis nciculari2.c
84
83
56 ??
36
46
73
81
84
74
54
43
54
73
71
4I
54
Tso
Wid
Wid [86]
Tax
Tax
Bol
Tax
91
73
92
54
84
36
53
71
73
5473 16
82
16 +
46 73
73
73
93
23
73
13
72
46
56
16+
71
73
54
7473
Wid
Wid [86]
Wid
Am1 .4s2 Tso
Tso
Wid
Wid
Bol
Wid
Wid
Am1
As1 Wid
Am1 Bol
Am1
Tso
Am4
h l l
lzml
4G+ Tax
76
42
43
93
91
74
73
54 56
Tax
Bo2 Tax
Sst
Wid
M'id
FLORISTIC ELEMEN I S I”u BRITAIN AND IRELAhD
Eleot hans austriaca
Eleot h a m multicaulis
Eleorhans palustris
Eleoclians paroula
Eleorhans quinquepora
Eleocharis uniglumis
Eleogiton Juitans
Ebmus caninus
Ebtri&a atherira
Ebtrigia juncea
Ehtr@a repens
Empetrum nigrum
Epilobium alsinEfoiiuni
Epilobium anagallidijolium
Epilobium hir.rutum
Epilobium lanceolatum
Epilobium montanum
Epilobium obscurum
Epilobium palustre
Epilobium pam$orum
Epilobium roseum
Epilobium tetragonun
Epipactis atrorubens
Epipactis hellebori,ie
Epipacti.7 leptochila
Epipactis pa1ustri.r
Epipactis phyllanthes
Epipactis purpurata
7 Epipactisgoungiana
Epipogium aphyllum
Equisetum ammse
Equisetum ,fluviatile
Equisetum lyemale
Equisetum palustre
Equlsetum pratense
Equisetum ramosissimum
Equisetum gbiaticum
Equketum telmateia
Equisetum uariegatum
Erira riliaris
Erica cinerea
Erica engPna
Erica mackaiana
Erica teti-a1i.x
Erica uagans
Engeron acer
Erigeron borealis
Eriocaufon aquaticum
Eriophorum angusttjblium
Enophorum gracile
Eriophorum lat$lium
Eriophorum uagnatum
Erodiuni cicutarium
Erodium lebelii
Erodium maritimum
3 Erodium moschatum
Erophila glabrescens
Erophila mapcula
43
72
65
73+
53
76
81
54
83
83
64
26
13
16-t
85
92
73
73
56
73
74
74
54
75
73
74
73
73
71
45
36
56
56
56
46
85
56
83
26
81
71
81
71
72
81
56
14
41
36
56 53
26
84
82
82
91
??
??
Am1 As2
Am1 As1 As2
Tso
Djt Tso
Co2
Col
Wid [66]
An12
Wid
As1 As2
Wid
Tax
Tax
Tso
Am4
Djt
Djt
Bo 1
Am5
Wid [86]
Djt
Gid
Eropliila uema senm lato
E y 7gium ranipestre
E y g i u m mantimuni
Euogmus europaeuj
Eupatorium canuabinum
Euphorbia amygdaloides
Euphorbia QpanJsias
Eubhorbia exgua
Eupliorbia hehost opia
Euphorbia h~~berna
4 Eubhoibia lathyiis
Euphorbaa paralias
Euphorbia peplu
Euphoibia peplui
Euphorbia plagphylloJ
6uphorhia portlaiidu a
Eiqhorbza sermlata
Euphorbia azllosa
7 Euphrarza anglita
Euphrasia arctzia
7 Euphrasia cambnca
7 Euphiasia iampbelliae
Euphrasia confusa
Euphrasza foulaenc-ir
Euphrasza jiypda
7 Euphrasia heslop liamsonii
7 Euphiasia marshah
Eicphiasia mzciantlia
Euphratia nemoiosa
Euphrasia ostenjeldii
7 Euphratza pseudokernen
7 Euphrasia nvulans
Eupliiasia rostkooiana
7 Euphiasia rotund~jolza
Euphrasia saiisburgencis
Euphrasia ~iottita
Euplirasza tetrayuetra
7 6uphiasia i!igzcriii
hmrulum pusillum
haps gluatxa
Fallopia convoliulus
Fallopia dumetoium
Feituca altissima
Festura arenana
Feitutn annontana
Festura arundinacea
F e h r a jilEfonni~
Feituia gigantea
FPitur a huonii
Fertuca lemanii
FfJtuca longzfoiia
Fertuca onna
Fktura piatensij
Fettuta rubra
FeJ tuca awlpara
Filago luteicens
Fzlago minima
Filago pyamidata
84
83
83
73
73
73
7383
85
82
83
91
91
83
83
81
83
73
71
51
Col
\.\‘id
As1
\\‘id
M’id [86]
Wid
Col
Col
\Vid
11
41
51
41
I4
41
41
73
73
21
71
41
53
41
23
Col
Bol
Bol Col
Am1
Col
Col
43
Bol
Co2
71
71
82
73
64
75
73
71
71
84
72
73
71
71
71
55
54
36
26
73
73
92
Bol
c:o1
Wid [66]
Col
Col
\.\’id
As1
Col
Tax
LVid
M’id [56]
\.$‘id
’Tax
As1
112
Fi'lagq vulpris
Fil;?endula ulmaria
Filipendula uulgariA
Fragaria aesca
Frangula alnus
Ewnkenia laevis
Fraxinus excelsior
Fritillaria meleagris
Fumaria batardii
Fumaria capreolata
Fumaria densijlora
Fumaria muralis
7 Fumaria occidentalis
Fumaria oflcinalis
Fumaria pam$ora
Fumaria purpurea
Fumaria rpziteri
Fumaiia vaillantii
Gagea bohmica
Gagea lutea
4 Galanthus nivalis
Galeopsis angustfolia
Galeopsis b$da
Galeopsis segetum
Galeopsis speciosa
Galeopsis tetrahit
Galium aparine
Galium boreale
Galium constrictum
Galium mollugo
Galium odoratum
Galium palustre
Galium parisiense
Galium pumilum
Galium saxatile
Galium spurium
Galium stemeri
4 Galium tricornutum
Galium uliginosnm
Galium verum
Gastridium ventricosum
Genista anglica
Genista pilosa
Genista tinctoria
Gentiana nivalis
Gentiana pneumonanthe
Gentiana vema
Gentianella amarella
7 Gentianella anglica
Gentianella campestris
Gmtianella ciliata
Gentianella germanica
Gentianella uliginosa
Geranium columbinum
Geranium dissectum
Geranium lucidum
Geranium molle
Geranium prateme
C . I). PRESTON AND hf. 0. HILL
83
55
74
74f
74
82
73
73
91
92
83
81
71
83
83
71
81
74
83 7383
73
55
72
54
53
73
56
As1
Wid
Am1 Wid
Wid
Wid
\Vid
As1 As2
Wid
Wid [56]
Wid
Tax
Wid [76]
91
53
73
54
92
73
72
7642
84
55
55
91
71
73
73
13
74
13
56
71
53
74 73 73
73
83
92
83
55
Wid
As1 As2
Ainl
Tso Wid
Wid
Am1
4sl
Wid
h'id
Wid
Wid
Geranium pulpureum
Geranzuni purzlluni
3 Geranium pyrenaicum
Geianzum robertiarrunz
Geianzum rotundifalum
Geianzum sanguzneum
Geranium gCatirum
Geum rzink
G e m urbanum
Gladiolus i l h n r u ~
GlnuciumJlncwn
Glaur marzizma
Glerhoma hedrrac ea
Gbrena derlinata
Gbrerza Jluztans
Glyrena maxima
Giyrerza notata
Gnaphalium luteonlbum
Gnaphalzum nontegrum
Gnaphalzum supinum
Gnaphalzum sqlvaticum
Gnaphahum uIginoJum
Goobers repen r
Groenlandia densa
CGmnadenia conopJca
G>mnocarpzum dryopteris
@mnvraipium robertinnum
Hammarba paludosa
Hedera helm
Helianthemum apmninurn
Hebanthemum ranum
Helianthemum
nummulanum
Helxtohrhon pmtenv
Helictotnchon pubercens
Hellebom foetidur
Helleborus vcndis
Herarleum sphondylzum
Heninium monorrhis
Hemiana (iliolata
Hemiana &bra
Hwacium muiorum qg
Hierochloe odoratn
Himantoglossum hiicznuni
H~pporrepis(ornosa
H@pophar rhamnoides
H@puns rlulganc
Holcus lanatus
Holrus mollzs
Holmteum umbellaturn
Hont kenya peplotdeb
Horde!ymur eurojxzeuc
Hordeum mannuin
Hordeum munnum
Hordeurn seralinurn
Homungza petraea
Hottonia palustns
4 HumuluJ hipidus
91
74
73
73
84
73
44
54
74
91
91
56
55
72
73
76
73
84
13
13
54
55
3-6
73
55
56
56
46
83
93
93
Wid
Wid
As1 As2 M'id
Wid
Am1
Col Wid
c02
Wid [56]
Wid
Wid
As1
Wid
Am1 As1
Am1 As1
Am3
Wid [56]
+
+
73
73
73
82
72
55
75 81
74
36
46 92
73
53
56
83
73
84
36+
73
91
84
73
73
73
74 +
4 s l Wid
Wid
Col
As1 Wid
Tso
Wid
Wid
Col
As1 Co2 Wid
Wid [86]
Am 1 As2 Wid
FL.ORIS 1 IC ELEhIEN I5 lr\ H K I 1 \IN rWD IREL,\NI)
Huperzzn dngo
HynrintliozdPs non Irrzpta
Hpdrilla wrtu illata
NjdroihariJ rnorsu: innac
Hj h o c o p f e itulgnii
Hjnienojdyllum tunbngen5e
Hj nimophgllum u ilsoniz
H j oxyamuJ n ger
Hipencum androsnenium
Hjperzcum P I O ~ P J
f$ppnczirn hirtutum
Hj per iciinz hu~nq'iistim
H p n t urn lrnanfolium
H ~ p ~ n r i imarulntuni
m
H$erzciim montnnum
HypPncum perfor atum
H>pericum pukhnim
H)pnrum tetropt~ruin
Hypem urn undtdnium
Hjpocharns glabra
H>pochaPnc nini ulatn
H>poclinrns radimtn
Ibmr amain
1le.r aqufofohuni
Ilk(ebnm wtzrzllatum
Inzpatirns nofz-tnngrrr
Iruiln tonyzne
Inuln rrzthrnoidfs
t Iniila salzcina
In1 fortidztrima
Iris pspudacoru,
Itodes echinospora
Isoptre histnx
IJoPtes Iaruitnr
Isolepi' temua
Isolepis setnrrn
j%n,zone montana
j'utu U I acut2floruJ
j;,mus arutii,
Junrus nlpznoarticiilatu,
Jiinru~nmbiguus
Junrus articulatus
,@ru~ baltirus
JuriiuJ bghimu
Junrus bufonin,
j h us bulborus
JunruJ rapitatus
Juncu, castanrzis
Junrus romprtwic
J i i n r i i ~ ionglompratuc
juniur @lJtiS
Junr us jllformzc
Juni
11s folzosus
3uiicus grrardii
Juncur znJexuJ
&ru\
rnaiitimut
Juncuc p>gnineus
Junrus rqua?ro,iiJ
26
71
8574
82
71
51
84
92
71
74
73
81
53
73
84
72
73
81
83
74 83
82
82
82
75
73
91
75 82
83
46+
91
44
91
74
73
73
91
46
83
84
26
Tso
TSOWld
M'id
Am 1 Tao
Sst
Wid I861
Wid
Wid
LVid
\\'id
Wid
Sst
Am4
<:02
\Vid
Tax
Am1
Am1 Co2 Tso
Wid
1\m3
A m 4 c o 2 l'so
Am1 Tax
Am3 \.$'id [86]
Tso
16
66
53
83
16
74
73
83 +
46
82
66
84
83
91
Tso Wid
Am1
Wid
Wid Tax
Wid
r2mIA~lA.Q\2'1(1
co2
Wid
(202 \\'id
I13
114
f Limonium paradoxum
f Limonium parvum
7 Limonium prncerum
f Limoiiium recumurn
f Limonium transwallianum
Limoaium vulgare
Limosella aquatica
Limosella australis
$ Linaria pelisseriana
Linaria repens
Linaria uukaris
Linnaea borealis
Linuni bienne
Linum catharticum
Linum perenne
Liparis loeselii
Listera cordata
Listera ouata
Lithospennum amense
Lithospennum oficinale
Lithospmum
purpurocamleum
Littorella ungora
Lloydia serotina
Lobelia dortmanna
Lobelia urens
Loiseleuria Procumhens
Lolium perenne
Lonicera pericbmenum
5 Lnnirera xylosteum
Lotus angustissimus
Lotus corniculatus
Lotus glaber
Lotus pedunculatus
Lotus subblflorus
Ludwigia palustris
Luronium natam
Luzula arcuata
Luzula campestriJ
Luzula forsteri
Luzula mulwora
Luzula pallidula
Luzula pilosa
Luzuln spicata
Luzula sylvatica
Lychnis alpina
Lychnis jlos-cuculi
Qchnis viscaria
Qcopodiella inundata
Lyocpodium annotinurn
Lycopodium clauatum
Lq'copu~ruropaeus
Lysimachia nemoiam
Lysimachia nummularia
Lysimachia thyrsEpora
Lysimarhia uulgaris
Lytlirum hysopfolia
L~~thrum
portula
C. D. PRESTON AND R.1. 0.HILL
71
71
71
71
71
91
56
51
91
72
55
46
91
73
56f
74+
46
54
84
74
+
83
72
16+
43
81
16+
83
82
74
83
85
83
73
82
83
72
Co1
Col
Col
Col
Col
Col
4 m 5 Tso
Wid
Wid [56]
Am3
Bo2
Am1 As2
Wid
Wid
Am4 Bo2
Am1
Wid
Wid
Wid
Wid
Wid
Am1 Wid
1 3 f Am4 As2 Bol
73
92
36
5554
13+
73
23
74
74 53
26
56
74
72
73
46
75
84
73
Wid
Am3 Wid
Sst
Am1 As1
Am1
Wid
Am1 As2
Tso
Wid
Wid
Wid
Wid
Qthrum salicaria
dtfaianthemum by5lium
Ma1u.c .glvrstrir
hfalva moschata
Malva neglecta
Malua sylvestris
Marmbium vulgare
izlntricaria i-ecutita
5 Matthiola incana
Matthiola sinuata
Meconopsis cambrica
Medicago arabica
Medicago lupulinn
Medicago minima
Mtdicagn po&morpha
Medicago sativa
Melampyrum arvrnse
Melampyrum cristatum
Melampyrum pratense
Melampyrum sylvaticum
Mdica nutans
Afelica unzjlora
Mdittis melissophyllum
Mentha aquatica
Mentha arvensis
Mentha pulegum
Mentha suaveolens
hi'eryanthes trifDliata
Mercurialis annua
Mercurialis pertmi3
Mertensia maritima
Meum athamanticum
Mibora minima
Milium effu.mm
Milium oernale
iLfinua?-tiahybrida
7 Minuartia recuma
Minuartia rubella
iifinuartia sedoides
hi'inuartia stricta
Minuartia uerna
Afisopates orontium
;Clorhringia trinemia
Mvenchia errcta
hi'olinia caerulea
Alonesex ungvra
Monotropa typopi!y.i
Montia fontana
4 hfuscari neglectum
A+celis muralis
At@uotisalpestri.\
hGosotiJ amensb
hGosotis discolor
,\Gosotis lam
il9osotis ramosissima
itlyosotis scorpioidrs
it!yosotis stcunda
Alyosotis sicula
:
:
75
55 73
73
73
84
84
83
91
91
51
92
74
84
92
8473 74 54
43
55 73
73
73
56
83
92
56
92
73
23+
43
82
56
+
Wid [76]
Wid
Wid
Wid
Wid [86]
Wid
Wid [86]
Wid
Bo2 Wid
Wid
Wid [76]
Wid
Wid [86]
Wid
Wid
Am1 4s2 Col
Bo2
C02
91
92
13
Bo2
Bol
Bo2
16
13
16
45
84
73
Djt
Wid
As2
82
54
46
76
5 3 + Am1 As1 As2
84
73
16
54 Wid
73
56
83
74 lYid
71
91
+
+
C. D. PRESTON AND M. 0. HILL
16
Pilularia globulfia
Pimpinella major
Pimpinella saxfiaga
Pinguicula alpina
Pinguicula grandgora
Pinguicula lusitanica
Pinguicula vulgaris
Pinus ~ylvestris
Plantago coronopus
Plantago lanceolata
Plantago major
Plantago maritima
Plantago media
Platanthern bfolia
Platanthera chlorantha
Poa alpina
Poa angustijdia
Poa annua
Poa bulbosa
Poa compressa
Poa Jlexuosa
Poa glauca
Poa humiliJ
Pan i$ma
Poa nemoralis
Poa pratensis
Poa triclialii
Polemonium caeruleurn
Polycarpon tetraphyllum
Pobgala amarella
Pobgala calcarea
Pobgala serpyllfolia
Pobgala vulgaris
Pobgonatum multgorum
Pobgonatiim odoratum
Pobgonatum verticillatum
Polygonurn arenastrum
Polygonurn aviculare
Pobgonum boreale
Pobgonum maritimum
Pobgonum oxyspmum
Pobgcnum ruriuagum
Pobpodium cambricuni
Pobporlium inteljectum
Polypodium oulgare
Pobpogon monspeliensis
Pobstichum aculeatum
Pobstichum lonchitis
Pobstichum setijirunz
PopuluJ nigra
Populus tremuia
Potamoleton acutfolius
I’otamogeton alpinus
Potamogeton berchtoldii
Potamogeton coloratus
Potamogeton compressus
Potamogeton c.rispus
Potamageton epihydrus
72
73
74
14
71
71
46
45
84
84
65
34+
75
55
73
+
C02 Wid
Wid [86]
Wid [66]
Am1 Tso
Wid
16f
Tax
Wid [66]
Wid
Wid
Am1
86
64
84
73
13
26
??
91
56
66
64
5491
53 81
72
73
73
75
43
65
66
42
91
63
73
91
72
53
91
75
46
92
74
55
73
46
56
83
55
85
51
Wid Sst
Wid [66]
Wid
Am2
As1 As2
Wid Tax
Wid
Am1 Bol
Col Tso
Am1 Col
Tax
Sst
Wid
+
Tax
Sst
Tso Wid [86]
Am5
PotamogetonJilformis
Potamogetonjkesii
Potamogeton gramineu
Potamogeton luceins
Potamogeton natans
Polarnogeton nodosus
Potamogeton obtusfolius
Potamogeton pectinatus
Potamogeton perfoliatus
Potamogeton pobgon folius
Potamogeton praelongus
Potamogeton p u s i h s
Potamogeton rutilus
Potamogeton trichoides
Potentilla anglira
Potentilla anserina
Potentilla aigentea
Potentilla crantzii
Potentilla erecta
Potentilla fruticosa
Potentilla neumanniana
Potentilla palzutm
Potentilla reptans
Potentilla rupestrir
Potentilla sterilis
Primula elatior
Primula farinosa
1Pnmula scotica
Primula vwis
Primula vulgaris
h n e l l a vulgariJ
Prunus avium
Prunus padus
Prunus spinosa
Pseudorchis albida
Pteridium aquilinum
Puccinellia distans
Puccinellia fasciculata
Puccinellia maritima
Puccinellin rupestris
Pulicaria dysenterica
Pulmonaria long@a
Pulmonaria obscura
Pulsatilla vulgaris
e m l a media
orola minor
gYrola rotundtjilia
orus cordata
Quercus petrata
Quercus robur
Radiola linoides
Ranunculus acris
Ranuiiculus aquatilis
46
56
56
74
56
86
56
66
56+
72
46
86
43
84
73
56
74
24
54
46+
73
56
84
93
72
7345
41
74
73
66
73
55
73
43
76
54
82
51
81
84
74
71
73 73 44
46
54
71
73
73
73
35
73
Ranunculus aruenu
Ranunculus auncomu
Ranunculus baudotu
84
53
83
Pulicaria vulgaris
+
+
Tso
Tso
‘rso
Am3
Tso Tax
Tso
Am3
Wid
Wid
Am1
Bo2
Wid
Wid
Am3
As1
Bo2 Djt
Tso Wid
Wid
Wid
Am3
Tso
Co2 Wid
Am1 Go2 Tso
Am3Col
Go2
Am1
Wid [36]
Am4 As1 As2
Djt Tso
Wid
Co2
FLOKIS I1C ELERIEN I h 1'4 BKI IAIN AND IREIAhD
Rnnunculur bulbomc
Ranuiiculns circznatus
Ranunculus jcaria
Rnnurirulur jlamniula
Ranuntulur Jluitans
Rrinunculus hederarm
Ranunculus lingua
Rniiunculus omzophyllus
Rnnunculuc ophioqlocsfolius
Ratiunculus paludoJus
RnnunculuJ pany4oruJ
RanuriLulus peltutu.,
Rnriunculus pmzc zllatiis
Rnnunc ulzis repens
Ranunculus reptans
Rniiuriculuc sardous
Rnnunculus Jreleratus
Ranunculus tncfiop/yllu
Rnnunculus tnpartitus
RaphanuJ raphanistium
Rrseda lutea
Reieda luteola
RhamnuJ cnthaitica
Rhinanthus angutlfolius
Rtiinanthu, mznor
RlyiichoJpora alba
RhynchoJpora fusca
Ribus alpinum
9 Ribes nigmm
4 RibeJ mbrum
Rzbec cpzratum
Ribes uua-cnspa
Romulea rolumiiae
Ronppa amphibin
Roiippa zdandzca
Rorqpa microphq lla
Rorzppa nasturtiuniaquatic um
Rorqpa palustns
Rorippa g l r estnr
Rosa flc&VStlS
Rosa aiurn!i$
Rora raesia
RoJa canzna
Rota micranttia
Rota mollzc
RoJa obtustjolza
RoJa pimpznelltjolia
Rosa rubigmosa
Rosa sherardzi
Rosa sglora
Rosa tomentosa
Rubza peregnna
Ruhus airtzcur
RubuJ rausius
Rubus chamaenioiuJ
RubusfilicoJus a g
RiibuJ idneu \
Wid
Sst
83
75
83
73
73
82
74
82
83
91
82
63
73
5.5
46
73
56+
36
RUltlPt' / J U k h f l
\Yid [XI
\$'id
b'id
Tso
81
Rutrirr riipatiir
Runirv Jnngurneu \
Ruppia (irrhom
+
LVid
Wid
\.$'id
tYid
\$'id
h
l
Bo2
Wid
\Yid
Tax
Wid
co2
LVid
Sst
??
Am2 Sst
IVid [56]Sst
\Yid [66]
54
64
r A
-
84
8k
73
45
\ \ i d [86]
\\id [86]
73
73
83:
71
73
615
\Vid
_.
10
66
\Vid
'no
+ 'Im4
Co2 'l'so
Tax
C o 2 -ISO
'32
83
83
83
84
74
54
53
56
52
43
54
72
5373
91
75
44
84
56
73
73
73
73
73
73
53
73
75
73
73
73
73
91
46
74
46
83
56
Rubus sauatilis
Runiev acetota
Rumev acptodla
Ricinr~aquatic ILJ
Riime\ conglonieiatic\
Rimer r n ~ p u ~
Ritmrx l~~~d~olapotliuni
Ruinex loigdohu c
R u n i ~ xmarztmiut
Ricnirr obtnsdolius
Riinirr pnluJttis
117
-3.1
.> .>
83
I6
54
\\id
co 1
Bol
Am1
.54 \'Vid
16
7 :1
54 Sst
53 'Iax
66 c:o2 'l'so ' l a x
7 1 Col 'l'ax
81 cOi'Tax
7 1 C:ol ' l ~ a x
7 1 Col 'I'ax
82 Col Tax
84 \Vid
13 1301 Sst
53
Wid [86]
55
54
Wid
74
\Vid
13
.4ml
Bol
Wid
16
24
44
13
Djt Wid
LYid
Kol Sst
74
54
_16
_
-/I 3-3
84
iVid
'l'so \Vid
Bol
Wid
?I 1
Wid
83
i 3 \\id
~:(i-r\()
73
I18
C. 1). PRES'I'ON :LVD 11. 0. H I I L
84 \\'id
56+ .\1ii4 \Vid
73 As1 Sat
7 3 62'id
91
15
1 3 Am 1
16
l(i
Rol
73
26
71
41
I6
16
16
42
73
83
72
73
\Vid IS61
Col Sst
-
sst
+
71
71
13
84
33
Tso
+ 'hll
As2
\Vid
As1
\Vid
\t'id [St;]
\\id
92
84
73
Bol
71
1 :i ,\mI
$33
\Vid
83
82
\Vid 1861
\Vid
74
\\id
sst
Am5 C:u2
!I 2
83
(il
\Vid [8G]
hi5
85
73
43
c:o2 'I'so
Tso
71
85
85
\\id 1861
\\id
X4
91
Anil
71
84
Co2
74
SSl
71
73
7 :3
73
\\id
73
41
73
84
4(i -
64
73
r ,I
0
74
55
71
82
74
81
74
71
83
71
71
71
71
71
71
~
54
82
A \ t ~1i
73
92
71
l\'irl
l\id
81
16
71
41
-/&I
- \\id
23
.\Ill3
46
7:3
7 :3
71
71
74
7-1
73
\ \ i d [76]
\\id [86]
\Vid [8C;]
74
83
83
~
.Is2
St
42
KO1
71
7 :i
71
71
43 + .an1I ;\s2
,Xi
7 6 Xm4 Tao
+
FLORISTIC ELEXIEN'I'S IN BKI I'AIN .AND IRELAND
Spartina mantima
Spegula aruensa
Speplana bocconei
Ypegulana manna
Speguaria media
Spegulana rubra
Spergulana rupicola
Spiranthes aestiualis
Spiranthes romanzojiana
Spiianthes spiralis
Spirodela polyrhiza
Stachys alpina
Stachy c aruensis
Stacks gmanica
Stachys ojicinalis
stacty \ palustns
Stachys Jylvatzca
Stellana graminea
Stellana holostea
Stellana media
Stellana negleita
Stellana nemomrn
Stellana pallida
Stellana palustiu
Stellana uliginosa
Stratiotes aloides
Suaeda mantima
Suaeda Vera
Siibularia aquatica
Surcisa pratensis
Sqrnphytum o@cinale
S~mphytumtuberosum
Tamus communir
Tanacetum vulgare
Tamxarum ojicinale a#
TaxuJ barcata
Eesdalia nudicaulis
5phroseru intepfoha
EfihrosenJ paluJtns
Eucnum batpis
Eucrium chamaedTs
5 u n z u m scordium
Eucnum st orodonia
'(halzctrum alpinum
nalictrum flavuni
Thalictrum minus
'The!vptensp a l u ~tris
77iesiuni humfusum
ThlaJpi arvcnse
IhlaJpi caeruleven,
'Thlaspi pe foliaturu
'Thymu pobtnrfius
n
j
r
n
u
s pulegioides
ThymuJ serpyllum
filia tordata
7i11a p l f l ~ p h l l l o s
Efzeldia pusilia
82
64
91
86
84
83
71
73
41
83
86
73
82
73
73
56
74
54
74
65
73
53
84
55
73
Col
Wid [66]
h
Co2 Tso Wid
Co2 Wid
Am3 Wid
Col
Am5
Tso
Wid
As1
Wid
Wid [66]
As1 As2
LYid
Wid
Am3 As1 As2
Tso Wid
54- Wid
85 Co2 Tso M'id
91 Co2 Tso
46
74
73 As1 Wid
73
92
55 Wid [56]
66 Tso
73 Sst
73
36 f
36
73 83 84
82
16+
+
55
76+
71
75
43
84
53
73
5374
73
16
.
7
54
9
Tso
Wid [76]
Bo2
\.$'id
Tax
Tax
\+'id Tax
Toidis aruensiJ
Torilisjaponica
Tonhs nodosa
Tragopogon pratensir
Zichomanes speciomm
Xirhophorum alpmum
Tn'thophorum respitosum
TrientaliJ europaea
Tnjolium arvense
Trzfolium boctonez
'Trfohum campertre
Tnfolium dubium
irifolium jiaGfimm
7rfolzum glomeratum
ZIJOliurn incnmotum
Tnfolium medium
Trfolzum micianthum
'Tnjolium orridentale
Tnzhurn ochroleuron
'fiqolium ornithopodioida
irilfolium pratense
Tnjolium repens
Tnfolium scabrum
7 ,folium squnmosum
7itjolzum striaturn
Xfolzum Jtni turn
Xgoliurn subterranrum
Xfoliuni suffot ntum
iriglochin mantimuin
Eiglorhiii palustie
Tiinin glauca
Tnnpleuiosnpemium inodomnr
Trtpleurospemiuvi
inantimuin
Tri vtum flaiiewn \
TrolliuJ cumpa~us
Tuberaria guttata
TuJsilago fadara
q p h a argustlfoha
Gpha lattfolui
I 'lex europaeus
['let gflllli
I 'lei minor
l % n u glabra
~
I%nus minor
17rnuJ plotii
I lmus procera
I ir,bi hcu \ 1zip 5 tns
rtita diozta
l i-tic a uens
I h i r ularia australi,
1 'titi ulana intermedia
I ' t m itlana niirioi
1't)u ularia o( hrokiua
I 'tricrilana stvgia
I tiiczilaria i ~ ~ ( q a i i \
Tati inium mirio(arpum
liir i m u m rrg itillu \
1
119
84
7 i
91
74
Tso
Wid
As1 Mid [92]
\Yld
7I
4b 40
rso
46$ Am4
84 \$'id
(1 1
84 Wid
73 \$'id
84 \.$'id
91
9 1 LVid
54 Wld
92 b'id
71 C a l
73
82
74
54
92
91
83
92
92
91
56
56
83
74
36
73
43
\.\'id [76]
Wid [56]
Wid
Wid
C02
Tso
Wid [76]
Col
\Vid
Sst
c02
%54 Wid
91
74
86
71
71
81
73
73
71
73
I)
1
54
tl4
Am1 Sst
Tso
M'id [72]
LYid
Tax
IVid
M Y [XS]
53
Tso
46
.Xi
Tso
73
J?
71
46
41
Sst
120
L'acrznzum oxycoccos
Vaccznium uliginomm
Vnccinzum vztu-idaea
Valenana dzozc a
Valmana officznalw
Valenanrlla rannata
Vnlmanella dentata
Valenanella locusta
btzlerianella nmosa
Verbascum Iyrhnztzs
Verbasrum n z p m
Vmbasium pulverulentum
Verbascum thaprus
Verbascum vilgatum
$ Verbena o@cinalis
Veronzra agrestzs
Eronica alpina
Vmonzca anagallzs-aquatira
Veronica amrnsis
Veronzca bercabunga
Veronzra ratenata
Ihonira chamaedp
Vernnzcajiuticanr
Veronzca hedmfohn
Veronzca montana
Veranzca o$icinalzt
Veronica polita
4 Veronzca praecox
Veronica scutellata
Veronzca seriyllfolia
Vmonzca spzcata
Vmonzca tnphyllos
VeroniLa uerna
Rburnum lantana
Rburnum opulus
k a bittynica
C . D. PRESION AND hl. 0. HILL
46
26
26
73
55
83
73
73
73
73
74
92
74
82
85
73
14+
85
83
74
76
54
13
83
73
53
84
83 54
56
74 73 74 73
76
K z a crarra
Ikza hzrtuta
Krza lattyoidet
Am1 As1
Wid
As1
Wid
\Yid
Wid
\Vid
Am1 As1
\.$'id
'4sl N'id [86]
\$'id
+ Tax
91
Wid
,41112
Wid
Wid
Am1 Wid
Wid
Kcza htea
Kcza orobus
Rria panitpara
K i n satiua
65cza wpiiim
Kria syltlatira
L h a tetrasfienna
Hola aruenszr
1ioli ranznn
ITola hzrta
T.iola kitaibeliana
Kola lactea
?$ola htea
L h odnrata
Kola palurtru
Rola persicfolza
Kola reichenbachzana
liola nvzniana
Viola rupestrzs
?Sola trzrolor
L k u r i album
Ihlpia bromoidec
k'ulpia d a t a
Vulpiafasciczilata
Vulfna myuior
Vulpia unzlateralzs
H'ahlenbega hederaiea
W'oljia arrhzza
L2'oodJia alpina
llbodsza iluenizs
zannirhellza palustn i
<oh tern angustfolza
<o\tera manna
ZoJtria noltiz
55 Wid [56]
73 Wid 1761
73
92 Wid
72
92
83 Wid
54 Wid
54
73 Wid [76]
74 Wid
54 Am2
74
83
71
43 Bo2
73 \$'id
53 Am3
74- Tax
73
73
75 73 \Yid
73 As1 As2
92 M'id
91
91 c o 2
84 Wid
92 As1
81
84 Tso
26
26
86 Tso
22
..
66
85
Col
Col