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Ferns PDF
Ferns PDF
Atlas of Ferns
of the BritishIsles
.Ina re-drafting
of theIntroduction
an explan-
i
htionof thesignificance
of theopencircle was
inadvertentl omit
y ted.Unless
otherwis expla
e ined,
suchopenCircles arethosesquares
in whichte
.epecies
hasbeenrecordedonlyPRIORTO 1950(nineteen
hundredandfifty).
On page95thespellingof "DrToRteria
xembrosiae"
phnuld be "Dryopteris
x:embroseae"It.is
*similarly miapelt in the Index, (p. 100)0
MM7 i978.
ATLAS OF FERNS
OF THE BRITISH ISLES
edited by
A.C. Jermy
British Museum (Natural History)
The Botanical Society of the British Isles + The British Pteridological Society
London
1978
Botanical Society of the British Isles
and
British Pteridological Society
c/o British Museum (Natural History)
Cromwell Road, London 5W7 5BD
ISBN 0 901158 01 1
INSTITUTE OF
TERRESTRIAL
ECOLOGY
LIBRARY
SERVICE
5JU11978
REF
osH I (40
It is now 16 years since the publication of the Atlas of the determination of hybrids although it is appreciated
the British Flora (Perring & Walters, 1962). Though it that the maps of these can only be regarded as
has proved valuable it was inevitably an imperfect work. preliminary; for detailed information on the
Conditions of the grants under which the work was distribution of hybrids the reader should consult
carried out meant that the first maps had to go to C.A.Stace (ed.), Hybridization and the Flora of the
printers within six years of the start of the scheme and British Isles, London, 1975. As the data were marshalled
the maps of the Pteridophyta were the first to be sent. the B.R.C. staff, under the direction of F.H.Perring, set
Some of these maps e.g. Dryopteris carthusiana (as to work fo'produce the maps and it was Lynne Farrell on
D.lanceolatocristata), D.oreades (as D.abbreviata), whom fell the major task of checking the maps against
D.pseudomas (as D.borrer0, were very unsatisfactory the data cards.
because of difficulties of identification and the map of
the first mentioned species was accordingly marked P, Nomenclature and arrangement of the maps
indicating it was only provisional. Furthermore, The nomenclature follows that used in Flora Europaea
taxonomic revisions in Asplenium, Dryopteris and (ed. Tutin et al., Cambridge, 1964) with a few later
Equisetum were producing data on the distribution of changes. These together with the arrangement are to be
new taxa worthy of mapping. A recent reprint of the found in a List of British pteridophytes shortly to be
Atlas (1976) gave the opportunity to revise maps of published (J.E.Dandy & A.C.Jermy, in prep.). The
about 300 of our rarest species including only five ferns arrangement of the genera is based on that proposed by
but production costs ruled out a complete revision, and J.A.Crabbe, A.C.Jermy & J.T.Mickel (Fern Gaz., 11;
this is a situation likely to continue. 141-162; 1975).
The British Pteridological Society had, from 1962, We have adopted the four genera in Lycopodiaceae as
taken a considerable interest in collecting further data in Flora Europaea but we agree with Holub (Preslia,
on ferns for the Map Scheme and these had over the Praha, 47: 97-110; 1975) that Diphasium should be
years been passed to the Biological Records Centre to be restricted to a few South American species and that
added to the mass of data coming in from the Botanical Diphasiastrum should be used for the European species.
Society of the British Isles. The B.P.S. felt that an Within Dryopteris two major changes on purely
interim publication showing the state of our knowledge nomenclatural grounds have been made. There has been
of the distribution of British ferns, horsetails and club- confusion over the past 150 years with regard to the
mosses, would, apart from removing the anomalies identity of Polystichum abbreviatum DC., the
mentioned above, give a guide-line for further basionym of D.abbreviata (DC.) Newm.; D.oreades
concentrated field work. Therefore when the B.P.S. Fomin should now be taken as the correct name (see
investigated the possibility of B.R.C. revising the maps C.R.Fraser-Jenkins & A .C. Jermy, Taxon, 25: 659-665;
for publication it was very pleased to find the idea 1976). It is also inevitable that we have to change the
already under consideration as part of a scheme to recent name D.assimilis S.Walker; the plant described
revise the Atlas in parts. by Presl from British Columbia as Nephrodium
Discussions were held to see how the two Societies expansum has been shown to be conspecific with the
could best be involved in drawing together all available European plant and that epithet must be taken up.
data that was not already in the data bank. A proforma Furthermore our British D.villarii is tetraploid and
(the 'Green Book') containing the species to be mapped sufficiently distinct from the diploid alpine material
was circulated to B.S.B.I. Recorders and to other described by Bellardi to warrant subspecific separation
interested workers studying regional or vice-county (see C.R.Fraser-Jenkins & A.C.Jermy, Fern Gaz., 11:
Floras. This data was transcribed by A.J.Worland 338-340; 1977). Lastly,European Thelypteris palustris
(B.P.S. Recorder) to single species cards to facilitate Schott has been shown to differ from American plants,
their transfer to the data-bank and to manuscript maps originally called T. thelypteroides Michx, solely in the
from which we could query any outstanding or absence of hairs on the lamina and rachis, a character
anomalous record. A.C.Jermy arranged for the loan of which we agree warrants its separation at subspecific
critical material from the following university or level only. Unfortunately Michaux's epithet has priority
municipal herbaria: Aberdeen (ABD), Aberystwyth over that of Schott thus removing a well-known name
(ABS), Bangor (UCNW), Belfast (BEL), Cambridge from the British list.
(CGE), Cardiff (NMW), Carlisle (CLE), Dublin (DBN), Comments by A.C.Jermy on the distribution of
Edinburgh (E), Glasgow (GL), Kew (K), Liverpool species are given beneath the maps. The
(LIV), Maidstone (MNE), Manchester (MANCH), phytogeographical elements referred to relate to the
Newcastle upon Tyne (HAMU), Oxford (OXF), Perth species distribution in Europe as a whole; the concepts
(PTH) and the South London Botanical Institute as applied to the Atlantic seaboard are those of
(SLBI), and, with the help of the following specialists, D.A.Ratcliffe (New Phytol., 67: 365-371; 1968). The
many new records were added to the data bank: distribution of pteridophytes in Europe is to be found in
H.V.Corley (Dryopteris), J .A.Crabbe (Polypodium), the Atlas Florae Europaeae 1, Pteridophyta (Psilotaceae
C.R.Fraser-Jenkins (Dryopteris), M.Gibby to Azollaceae), ed. J.Jalas & J.Suominen; Helsinki,
(Dryopteris), J.D.Lovis (Asplenium), C.N.Page 1972. For a discussion on the distribution of European
(Equisetum), R.H.Roberts (Polypodium) and A.Sleep pteridophytes using numerical analyses of those data see
(Polystichum). Particular attention has been given to H.J.B.Birks (New Phytol., 77: 257-287; 1976).
Species omitted R.C.L.Howitt, t S.T.Jermyn, Q.O.N.Kay, D.L.Kelly,
Only three aliens have been mapped: Azolla flliculoides A.G.Kenneth, D.H.Kent, M.P.H.Kertland,
Lam. a species so well established that its status is often J.E.D.Lamb, P.J.Lambley, H.Lefevre, F.LeSueur,
forgotten; Selaginella kraussiana A.Br., a native of A.G.Long, tJ.E.Lousley, J.D. Lovis, R.McBeath,
southern Africa, introduced into conservatories from D.McClintock, D.J.McCosh, B.M. Mack,
which it has spread considerably; and Equisetum R.Mackecknie, tV.J.Macnair, L.J.Margetts,
ramosissimum Desf., a single population, of interest M.E.Martin, T.F.Medd, K.G.Messenger, tH.Milne-
because it is the putative parent of E. moorei Newm. In Redhead, R.J.Murphy, A.Newton, A.M.O'Sullivan,
addition to these there are four species grown as C.N.Page, R.C.Palmer, R.J.Pankhurst, C.P.Petch,
ornamentals which have escaped and established E.G.Philp, M.Porter, A.C.Powell, A.L.Primavesi,
themselves in our natural vegetation; none have spread M.C.F.Proctor, A.W.Punter, D.A.Ratcliffe, J.E.
further other than by vegetative means. They are Raven, B.W.Ribbons, M.H.Rickard, R.H.Roberts,
Onoclea sensibilis L., Matteuccia struthiopteris (L.) C.A.Robinson, R.G.B.Roe, J.Rogerson, F.Rose,
Tod., Blechnum chilense (Klf.) Mett. and B.penna- A.Rutherford, C.D.Sayers, M.J.P.Scannell, M.R.D.
marina (Poir.) Kuhn; Dicksonia antarctica Labill. may Seaward, B.Seddon, S.Segal, B.Shepard,F.W.Simpson,
also be included in this category. Three house ferns have A.A.Slack, W.A.Sledge, A.Sleep, J.E.Smith,
spread to warm locations on walls in various parts of A.H.Sommerville, 0.M.Stewart, A .McG.Stirling,
south and west Britain namely, Cyrtomium falcatum K.M.Stevens, R.Stokoe, G.A.Swan, E.L.Swann, D.M.
(L.f.) C.Presl, Pteris cretica L. and P.vittata L. The Synnott, M.A.Turner, I.M.Vaughan, tW.E.Warren,
latter is also established on a warm slag-heap in the M.McC.Webster, T.C.E.Wells, D.J.B.White, L.E.
Forest of Dean, v.c. 34 (see S. Holland, Whitehead, A.J.Willis, A.Willmot, S.R.J.Woodell,
R. N. Glouc. Nat. Soc., 19: 318; 1968).Although these P.Yule.
produce spores their presence is usually ephemeral. A considerable amount of general but often difficult
material, sent in by recorders, was identified by
Acknowledgements J.A.Crabbe and J.M.Mullin, who also handled some
We should like to thank all those, too numerous to thousands of specimens sent on loan from other British
mention individually, who have over the years sent in Herbaria, filling in 'pink cards' and searching for grid
records to B.R.C. Similarly we thank also those many references when new records were found; such routine
people who will no doubt find gaps in our maps and work, so often boring, is gratefully acknowledged.
rectify the situation by sending in further records. We We also thank those curators and staff of Herbaria
are especially grateful to A.J.Worland, for transcribing who have cooperated by sending their material of critical
all incoming data from the 'Green books' to single- groups to the British Museum (Natural History) or to
species cards and to manuscript maps. J.W.Dyce, as Leeds University for checking. Of these we would like to
Secretary of the B.P.S., was alone responsible for mention the following whose knowledge of their own
collating the records made on the meetings of that region, both floristically and geographically was placed
Society. Liaison with the vice-county Recorders of the continually at our disposal: S.G.Harrison,
B.S.B.I.. was initially through the Records Committee and
colleagues at the National Museum of Wales, and
(secretary, F.H.Perring). The following botanists, M.J.P.Scannell and D.M.Synnott at the National
either as B.S.B.I. Recorders or as members of B.S.B.I. Herbarium, Glasnevin, Ireland. Funds to enable
or B.P.S., have made a special effort to cooperate with students to make out 'pink cards' of critical groups at
this revision: Oxford University herbarium were provided by the
D.E.Allen, G.H.Ballantyne, M.Barron, E.P.Beattie,
P.M.Benoit, J.Bevan, Druce Fund.
J.H.Bevis, H.J.B.Birks,
T.L.Blockeel, I.R.Bonner, H.W.Boon, E.H.Booth, Last and by no means least, we thank the staff at
R.P.Bowman, M.Briggs, B.R.C. especially D.W.Scott, and R.A.Cooke, a
A.B.M.Brewis,
J.M.Brummitt, K.E.Bull, E.R.Bullard, A.R.Busby, voluntary worker, who prepared the maps on which the
J.K.Butler, M.S.Campbell, J.F.M.Cannon, J.M.Castle- final production of this Atlas was based.
Smith, E.Chicken, E.R.T. Conacher, P.Copson, Further recording
H.V.Corley, R.W.M.Corner, F.E.Crackles, The British Pteridological Society Recorder will continue
G.Crompton, T.A.W.Davis, J.G.Dony, D.A.Doogue, to monitor additions or amendments to these maps. All
U.K.Duncan, E.S.Edees, P.J. Edwards, T.Edmondson, such records will be passed to the Biological Records
E.A.Ellis, G.Ellis, T.G.Evans, I.K.Ferguson, Centre at I.T.E., Monk's Wood Experimental Station,
F.Fincher, B.E.M.Garratt, G.M.Gent, E.J.Gibbons, on the appropriate record card, and the relevant B.S.B.I.
M.Gibby, V.Gordon, G.G.Graham, I.F.Gravestock, Recorder will be notified. New records may be sent to the
E.F.Greenwood, J.M.Gunn, P.Hackney, P.C.Hall, B.P.S. Recorder, c/o Botany Department,British
A.D.Hallam, G.Halliday, S.G.Harrison, C.C.Haworth, Museum (Natural History), Cromwell Road, London
J.G.Hodgson, S.C.Holland, K.M.Hollick, SW7 5BD.
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Three subspecies of Asplenium trichomanes are subsp. quadrivalens will have a stout rachis, with oblong,
recognised in Flora Europaea: subsp. trichomanes, thick, convex pinnae, and crowded, more numerous sori.
subsp. quadrivalens D.E.Meyer and subsp. inexpectans The following micro-characters may also be used:
Lovis. Only the two first named subspecies are found in 1. Rhizome scales The scales of subsp. trichomanes
the British Isles; the last is a south-eastern European possess a central red-brown stripe; those of subsp.
taxon of limestone rocks. The absence of subsp. quadrivalens have a dark-brown stripe, although this
inexpectans from Britain makes the separation of the difference is only readily observed in a liquid medium.
other two often possible on ecological characters alone as There is also a difference in the maximum size of the
subsp. trichomanes is a calcifuge and subsp. scales (3.5mm in subsp. trichomanes, 5mm in subsp.
quadrivalens predominantly a calcicole. The following quadrivalens). This character must be observed with
notes on identification of these subspecies have been some care since both bear scales with a wide range in size,
provided by Dr .J.D.Lovis. and only the largest provide usable characters. A fair
The distinctions between the two subspecies are sample of scales must be examined from each specimen,
particularly apparent in the upper half of the frond; the and in practice ,the scales do not provide a convenient
most evident field character lies in the aspect of the upper character for the determination of a large number of
pinnae, in which the lamina is conspicuously concave specimens.
(i.e. the margins turning upwards) in subsp. trichomanes 2. Spores These provide the most suitable micro-
but convex with inrolled margins, or less commonly, flat, character, , where confirmation is necessary. The
in subsp. quadrivalens. Both subspecies are very plastic sculpture of the perispore is highly variable in both
and present a very different appearence when growing in species, and of no value as a criterion, but the spores of
exposed as opposed to sheltered conditions. subsp. trichomanes are paler than those of quadrivalens.
Plants of subsp. trichomanes from sheltered sites may Spore size is often diagnostic and it is usual to measure
be distinguished by the following combination of only the exospore*. For reasons which are not yet
characters: stipe thin, wiry, red-brown; pinnae distant, satisfactorily explained, different workers have reported
arrangement mostly alternate, obliquely inserted, with a different size ranges. Ranges of exospore length
distinct stalk, asymmetric (oval to rhombic) up to 8mm measurements (in gum chloral) recorded by J.D.Lovis
long, often with a perceptible basiscopic auricle; lamina are as follows (mean values in bold): trichomanes 23-29-
36-42pim; subsp. quadrivalens 27 - 34 - 43 - 50pm.
delicate, that of upper pinnae distinctly concave with
upturned margin; soli small, short (up to 2mm), P.M.Benoit (Nature in Wales 9: 75-79; 1964) has found
relatively few in number (4-6 (-9)); indusia narrow and the .smean size ranges (measured in air) of spores of
delicate. Merioneth plants to be quite discrete: subsp. trichomanes
In contrast, plants of subsp. quadrivalens from similar 27-34gm, subsp. quadrivalens 38-48pm. It is necessary to
sites have: stipe thick, often dark brown or blackish bear in mind that different mounting media have
brown, pinnae more crowded, mostly opposite, with different effects on spore size. It would be prudent for
approximately transverse (square) insertion, almost new workers each to establish his or her own standards
sessile, symmetrical in shape, usually oblong, rarely from material of confirmed identity.
auriculate, larger (up to llmm long); lamina thicker, flat 3. Stomata The length of the guard cells can provide a
or convex, margin ± inrolled, with often crowded, more valuable confirmatory character: subsp. trichomanes 31-
numerous sori (4-9 (-12)), longer (up to 3mm long); 38-43-52pm; subsp. quadrivalens 35-41 -49-571.un.
indusia conspicuous. 4. Chromosome number The two British subspecies also
In very shaded situations, the lamina of subsp. differ in chromosome number: subsp. trichomanes is
quadrivalens is delicate and the pinnae are distinct, as in diploid, with n = 36, whereas quadrivalens is tetraploid,
subsp. trichomanes. Moreover the lamina of the upper with n = 72 chromosomes.
pinnae is flat, not convex but the main pinnae are large, An excellent photograph displaying the general
with long sofi and broad indusia, unless the shade is too morphology of characteristic examples of fronds of the
.extreme to permit normal growth, when only a few two subspecies is given in Welsh Ferns (ed. 5); pl. 10
scattered sori of variable size are produced. (Hyde, Wade & Harrison, 1969). The frond of subsp.
In exposed sites both subspecies are much reduced in trichomanes is very typical of a luxuriant specimen from
size. Subsp. trichomanes may still be distinguished by a a rock-cleft in deep shade. See also J.D.Lovis, Br. Fern
delicate rachis and orbicular, flat or concave pinnae with Gaz. 9: 147-160; 1964.
small sori, which are relatively few in number; in contrast *the outer wall beneath the perispore
ASPLENIACEAE 61
0
1
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Dryopteris filix-mas (L.) Schott, D.pseudomas (Woll.) usually concave, inclined to apex; pinnules with an
Holub & Pouzar and D.oreades Fomin have, since their obtuse or rounded apex with blunt teeth arranged
inception, been confused by botanists both in England palmately or like a fan, basal pair adnate or slightly
and abroad (see C.R.Fraser-Jenkins & A.C.Jermy, stipitate, longer than the pair above; basal lobes forming
Taxon, 25: 659-665; 1976). The following descriptions of auricles. Indusia 0.5-1mm, highly convex, thick,
these three species have been drawn up with the help of glandular, green when young, fitting closely round
C. R .Fraser-Jenkins . sporangia, scarcely shrinking at maturity, becoming
D.filix-mas Rhizome little branched. Fronds grey-brown when old. A plant of open screes, rocky
spreading, semi-persistent (although in very clement banks and open hillside.
western areas they may be persistent) petiole about + D.pseudomas Rhizome little branched. Fronds ±
length of the frond, ± densely clothed with pale coloured upright, persistently green throughout winter; petiole
scales; lamina ovate-lanceolate, truncate at base, ± variable in length, densely clothed with gingery scales,
herbaceous, mid-green. Pinnae flat, ± horizontal; many of which are narrow and with a dark base which
pinnules toothed, teeth ± acute, curved towards the acute remains as a blackish speck when the scale drops; lamina
apex, basal pair of each pinna ± stalked, usually longer variable in shape ovate to lanceolate, leathery, yellow-
than those above, lobes at base of pinnules usually green when young becoming blue-green when old.
forming auricles. Indusia 0.5-2mm, convex, thin Pinnae flat, ± horizontal; pinnules with few or no teeth
sometimes glandular, white or translucent when young, on parallel side, teeth at apex acute; basal segments not
at least part of the margin adpressed to lamina surface, obvious; junctions of pinna rhachis with main rhachis
amply covering sporangia but shrinking at maturity, darkly coloured. Indusia 1-2mm, highly convex, thick,
becoming brown when old. A plant of forest, hedgerow margins inrolled, often glandular, brown when young,
and open places on rocks usually on lighter soils. scarcely shrinking at maturity, becoming grey when old.
D.oreades Rhizome much branched. Fronds upright, A plant of woodland, hedgerows, especially on clay soil
frost sensitive; petiole about length of frond, densely and on open hillsides, often replacing D.oreades in
clothed with pale scales; lamina lanceolate tapering to maritime situations.
base, very slightly coriaceous, pale grey-green. Pinnae
1. Frond texture thick, somewhat glossy and dark 2. Frond mid-green; teeth of pinnules with acute
green on top (yellow-green when young); stipe and tips, converging towards the apex; immature
rhachis bearing long narrow scales with a dark base indusium thin, margins not involute and
and usually a dark centre, junction of rhachis and usually extended over the lamina.
pinna axis darkly coloured; lower margins of D. fdix-mas
pinnules (pinna-segments) parallel, with few teeth.
D. pseudomas NB: Hybrids between these species are intermediate in
their morphology but it is likely that D. x tavelii
1. Frond texture thin, mid- or grey-green on top; stipe will key out as D. pseudomas and D. x mantoniae
and rhachis bearing both narrow and wide con- as D. fdix-mas.
colorous scales, junction of rhachis and pinna-axis
not darkly coloured; pinnule-shape tapering, lower
margins usually bearing teeth or lobes.
ASPIDIACEAE 83
0 3 0
0
27.2
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Index
Synonyms are given in italics, map pages in roman