BRITISH FERN GAZETTE
VOL. 9
1959-67
Miss65ur! BoTANICAS
GARDEN LIBRARY
THE
BRITISH FERN GAZETTE
VOL. 9 and supplement 1959—1967
Contents
ALLEN, B. M. Psilotum nudum in Euro
BENOIT, P. M. mor ed gare ira in Wales on the Polypodium
vulgare aggregat
BRUTY, H. J. Cultivation of a under glass
—_____———- Notes from the Royal Botanic Gardens, Kew
CORLEY, H. V. ‘“‘Lastrea dilatata var. alpina
CRABBE, J. A. Herbarium specimens of fect and their ‘aberra-
Herbarium specimens of ferns Il
Plea for historical data on the B.P.
Cultivated ferns
CRANE, F. W. Is ita hybrid ora species?
DRING, M. J. The influence of shaded conditions on fertility of
‘ i ae 2
Bracken
DYCE, J. W. Transplanting ferns (note)
Polystichum aculeat m ‘Pulcherrimum Bevis’ (note).
Variation in Polystchum in the British Isles
Certificates of Meri :
Distribution of British “ferns and their allies (note)
te B.P.S. Austrian Excursion
——____—_— The B.P.S. 1966 Excursion to Bovey Tracey
EVANS, G. B. ‘Hymenopyam transplants in Kent & Sussex
Identification of British Hymenophyllum species -
& Sano A. C. Notes on the history of British
fern species I: Confusion in the aon ees ;
GREENFIELD, P. "Woodsia alpina (Bolton) Gray (no
i lala alpina and C. pn May : ‘the rock
FUCHS, H.P. The genus : Cheilanthes Sw. and its European species
GILBERT, G. Ferns in a Wardia
GILBERT, O. L. Dryopteris villarit if Britain Ks
HOLTTUM, R. E. Some notes on vegetative characters of ferns ..
British ferns in a hitiaies perspective .. a
John Smith of K
JACKSON, F. The rediscovery of Asplentuar < murbeckii (note) ce
A shelter for small ferns in pots (note : ve
Bolivian pteridophytes collected Sais Winifred
Brooke
JERMY, A. C. A preliminary census list of British pteridophytes :
A revised preliminary census list of British pteri-
dophytes .. be - od .. Supplement
Name changes in the British fern list (note) . ie
Two species of Diplazium in Europe 1
See EVANS, G. B.; and W.
JERMY, A. C. & LOVIS, J. D. aided majoricum Litard. 163
JONCHEERE, G. J. DE Ferns of Sardini 114
KAYE, R. Leaf-base propagation of Phyllitis scolopendriuom (note) 120
——__—— ariation in Athyrium in the British Isles 197
LANG, F. A. See LLOYD, R. M.
LLOYD, R. M. & LANG, F. A. a lg ones Mops complex is
. Am
LOVIS, J. D. ae geographical affinities of Ve New Zealand
eridophyte flor 16
Meiosis 1 in g fastest x murbeckii 110
The taxonomy of Asplenium trichomanes in Europe. 147
Fern hybridists and fern hybridising I. The work of
paward J jon Lowe (1825-1900). . 301
See JERMY, A. C.
& REICHSTEIN, T. A diploid form of Asplenium
ruta-muraria .. co ei kee os 141
MORGAN, D. Psilotum triquetrum in Basutoland (note) .. 120
McCLINTOCK, D. The ferns of the Channel Islands. 34
MELVILLE, J. D. ae Bracken on Littleworth Common on
PAGE, C. N. A hybrid horsetail from the Hebrides 117
Sporelings of Equisetum arvense in the wild — 335
PROCTOR, G. R. - Sakon list of pteridophytes from St Kitts . 71
ne — on Jamaican ferns I 13
REICHSTEIN, T. Sec ° LOVI J. D.
ROBERTS, R. H. Some its aat che on Polypodium australe .. 283
SCHELPE, E. A. C. L. E. The identity of two rheueuven® Erne? of
Blechnum 348
SCHULZE, G. The European Chainfern in ‘Corsic 162
SHIVAS, M. G. (Mrs M. G. eee The e Polypodium vulgare
complex .. 65
SLEDGE, W. A. Grammitis in Samoa 339
SLEEP, ANNE A rasonee nig to ihe cytotaxonomy of Asplenium ee
TRYON, A. F. Origin of the fern flora of Tristan da Cunha 269
WALKER, M. G. See SHIVAS, M. G.
WALKER, S. & TERMY. A A.C. Dryopteris assimilis in Britain .. 137
WILLIAMS, S. L. Variation in Adiantum in the British Isles 350
TEE
BRITISH FERN
GAZEA-EE
VOLUME NINE PART ONE
DECEMBER: 1959
EDITED BY A. ©. JERMY
1 Bo oTANICee
wrssoon!
GARDEN LIBRARY
THE BRITISH
PTERIDOLOGICAL SOCIETY
BRITISH PTERIDOLOGICAL SOCIETY
Officers and Committee 1959-60
PRESIDENT
T. H. Bolton, F.R.H.s
VICE-F KRESIVENIS
Rev. E. A. Elliot, M.a.
P. Greenfield, Esq.
Professor R. E. Holttum, M.A., SC.D., F.L.S
Reginald Kaye, Esq.
Dr. T. D. V. Swinscow, M.B., B.S., F.L.S
R. Whiteside, Esq.
SECRETARY AND TREASURER
J. W. Dyce, Esq.
* Hilltop,” 46, Sedley Rise,
Loughton, Essex
EDITOR
A. C, Jermy, B.sc.
Department of Botany,
British Museum (Natural History),
Cromwell Road, London, S.W.7.
COMMITTEE
J. A. Crabbe, Dr. J. Davidson, T. A. Dyer
B. Hayhurst, Mrs. J. R. Healey, F. J. Healey,
F. Jackson, N. Robinson,
H. Wainwright
AUDITOR
Peter Temple, L.R.1.B.A., M.INST. R.A
The SOCIETY is open to all interested in any way in FERNS and FERN ALLIES
a ee Full details will be
sent upon application to the SECRETARY.
THE
BRITISH FERN GAZETTE
NEW SERIES
YOu... 9 DECEMBER 1959 PART 1
Editorial
An interest in fern variation has been foremost in this Society from its
beginning and this interest has in latter years resulted in two factions. One
is the fern grower who is anxious to find new and attractive varieties for his
collection; the other is the taxonomist, equally interested in finding varieties,
but who is interested also in the conditions under which the plant is growing
and its possible relationships to other varieties and other species. It is
obvious that these interests have more in common than is at first supposed
and it is hoped that this will be realised in future activities of the Society.
The Gazette, on the other hand, should reflect the aims of the Society
and, as the Committee agreed to a widening of these to include fern allies
and ferns from all parts of the world, an attempt to do this is made in this
issue. The Gazette is also, however, the organ of the Society through which
members may make contact with each other and I shall be pleased to hear
of notes, views, or requests for information for inclusion in the next issue.
The previous editor, the Rev. E. A. Elliot, has emphasised this in the past
and the Society is indebted to the .vork he has put into the Gazette during
the last ten years. We trust his illness will be short and that shortly he may
again contribute to the Gazette.
SOME NOTES ON VEGETATIVE CHARACTERS OF FERNS
E. HOLTTUM
Kew, Surrey
The classification of ferns on the sole basis of shape of sori fedutien in
an arrangement which in many cases was clearly unnatural.. For example,
the h Fern and the Oak Fern were placed in the genus Polypodium,
though they show little other resemblance to Polypodium vulgare. Such
cases indicate that sori of similar shape have arisen on different evolutionary
lines among ferns. If we are to discover a natural classification, we must
look at other characters besides the shape of the sorus and of the indusium.
The detailed structure of a single sporangium may provide important
evidence, but to observe this requires fairly high magnification and much
practice in learning to see clearly the significant details. There are other
characters more easy to observe which may be equally significant; and these
characters also afford means of distinguishing between genera when plants
lack sporangia. The object of the present article is to call attention to some
of these characters which are not normally mentioned in works on the
‘taxonomy of ferns. The parts concerned are: vascular strands in the leaf-
stalk or stipe; shape of rachis and the way branches and leaflets are attached
to it; hairs and scales. In the discussion which follows, es names use
are those found in Hyde & Wade’s book on Welsh fern
Vascular bundles in the stipe. In the genera Dryopteris and Polynttehige
there are several bundles throughout the stipe (at least three, even in small
plants) and each bundle is more or less circular in section; there are always
two, on the side towards the apex of the rootstock, which are larger and more
complex in structure than the rest.
In the genera Athyrium, Asplenium, Gymnocarpium* and Thelypteris
thére are two vascular bundles at the base of the stipe, and in most cases
these unite in the upper part of the stipe. In Asplenium the united strand so
formed is X-shaped in section; in Athyrium and Thelypteris it is U-shaped;
in Gymnocarpium the two bundles are separate throughout the stipe. This
‘similarity between Athyrium and Thelypteris is one reason why these two
genera have been regarded by some botanists as closely allied; but in my
view other characters showing differences between the two genera outweigh
this similarity, and I do not think Athyrium and Thelypteris are at all near
to each other (cytological evidence supports this opinion).
Asplenium and Athyrium were formerly thought closely related, because
of the tropical genus Diplazium which appears to bridge the gap between
them, and Hooker united all in one genus Asplenium. The arrangement of
the vascular bundles in the stipe is one constant difference between the
genera, and there are others; I believe that the linear sorus in Asplenium
and Athyrium (with Diplazium) evolved from different origins. A difference
tween the two genera given in the Flora of the British Isles, by Clapham,
Tutin and Warburg appears to me however of seat ent, Under
* This is included in Thelypteris in Dandy, List of British Vascular Plants,
2
Athyrium we read “ sori borne on receptacles with a vascular strand branched
off from the vein’; under Asplenium ‘** sori borne direct on the vein.”’ This
statement about Athyrium was due originally to Mettenius, who wrote it
just a century ago, but we have no illustration to show exactly what he meant.
I have examined a leaflet of Athyrium cleared to show the vascular tissue,
and I cannot see the structure mentioned by Mettenius; at any rate it is
not one which can be easily observed.
hape of rachis and of rachis-junctions. In Athyrium filix-femina all
branches of the rachis are grooved on the upper surface, and the midrib of
each leaflet is also grooved; where a smaller rachis joins a larger one, the
groove of the larger is opened to admit the smaller, and this also occurs where
the groove of a midrib joins the rachis bearing the leaflet. This is true also
of Dryopteris and Polystichum (and of Bracken); but Thelypteris has quite
a different structure. In Thelypteris the rachises and midribs of leaflets are not
deeply grooved on the upper surface, and even if they are slightly grooved,
the groove of a main rachis is not interrupted to admit that of a branch.
In Gymnocarpium the fact that the branches are all jointed to the main rachis
complicates the situation; but at least one can see that all rachises are grooved
in much the same way as in Athyrium, Dryopteris and Polystichum. This is
a small piece of evidence pointing to a relationship of Gymnocarpium to
Dryopteris rather than to Thelypteris. In the genus Asplenium, of which there
is an enormous number of tropical species, the shape of the rachises and
their branches is more variable, but in the other genera — the shape
is very constant and is, I believe, an important generic char
Scale and hairs. The British species of Thelypteris have oe
hairs on their fronds (T. palustris also on the rhizome); these hairs each consist
of a single cell. T. oreopteris has also unicellular yellow glandular hairs on
the lower surface of its leaflets; it shares this character with some species of
the mountains of the Malayan region (e.g., 7. viscosa). All species of
Thelypteris, and of the allied genus Cyclosorus (mainly tropical) have uni-
cellular hairs of one form or another, and in this they differ from Dryopteris
and Athyrium. The Thelypteris group of ferns have scales which protect the
growing tip of the rhizome or rootstock, and these scales also bear uni-
cellular hairs, Such hairs can easily be seen on the edges of the scales of the
Beech Fern, T. phegopteris; but if the scales of 7. palustris and T. oreopteris
are examined with a hand lens, they appear to lack hairs. Microscopic
examination shows that there are small round unicellular outgrowths from
these scales, comparable to the glandular hairs on T. oreopteris. The scales
of Gymnocarpium differ from those of Thelypteris in lacking superficial
hairs
The scales of Asplenium, when seen under a low power of the microscope,
show an elegant lattice-like pattern. Like nearly all scales of ferns, these
scales consist of a single layer of cells. The walls of contact between adjacent
cells of this layer are thick and dark, but the upper and lower walls of the
cells are thin and transparent, so that each cell is like a little window-pane
set in a dark frame. Athyrium scales have all walls of about the same thick-
3
ness and colour, so that they show no lattice-pattern; this is another difference
between all species of Athyrium and Asplenium
The scales of Dryopteris, Polystichum nal Gymnocarpium are alike in
essentials. In some species of Dryopteris and Polystichum there are hairs on
the edges of some scales, but these hairs are not needle-like nor round as in
Thelypteris, nor are any of them borne on the surface of the scales.
Polypodium vulgare is so different from all other British ferns that it
needs a separate note. The vascular strands in the stipe are three, or some-
times more (two larger than the rest, as in Dryopteris), each strand more or
less circular in section. The frond has no separate leaflets, as in the other
genera; its midrib is raised, not grooved, on the upper surface. At the base
of the frond, the edges of the leafy part run down each side of the stipe as a
little ridge. This arrangement is also found in ferns of some other (non-
British) genera which are not closely related to Polypodium. The veins in the
frond of our British Polypodium are free; but in almost all the many tropical
ferns to which it is allied the veins form an elaborate network; our species is
a reduced outlier of a mainly tropical family, almost all members of which
are epiphytes. The rhizome of P. vulgare, as in all its family, is creeping and
is dorsiventral in structure (that is, its vascular anatomy has definite upper
and lower sides); our other ferns which have slender creeping rhizomes
(Thelypteris palustris, T. phegopteris and the two species of Gymnocarpium)
have an internal structure which is not dorsiventral. The scales of P. vulgare
are uniformly light brown, the cells not forming a clear lattice-work. There
are no unicellular hairs on either frond or scales
A PRELIMINARY CENSUS LIST OF
BRITISH PTERIDOPHYTES
A. C. JERMY
British Museum (Natural History)
In 1932 George Claridge Druce published his Comital Flora of the British
Isles in which 70 species of Pteridophytes were included together with
their distribution known at that time. In order briefly to summarize this
distribution Druce used the counties and vice-counties drawn up for England,
Wales and Scotland by H. C. Watson“) and for Ireland by R. L. Praeger@.
These divisions do not always agree exactly with present administrative
county boundaries. Further particulars as to the divisions of the larger
counties can be obtained from the references above or from the Census
Catalogue of British Hepatics, Ed. 3.°®) The advantages and difficulties
in this system have been discussed by Dandy‘ and Lousley“® and will
not be taken up here.
The present paper uses the vice-county system and is not intended to be
final in anyway as it is hoped that the end of 1961 will see published an Atlas
showing the distribution of British vascular plants on a 10-kilometre grid-
square basis. This will be the result of a scheme launched by the Botanical
Society of the British Isles to which many members of the B.P.S. will have
4
contributed. However, records are still needed and there are many records in
literature and in herbaria which need confirming in the field; there are
vice-counties from which some ferns have not been seen in the last 30 years
yet their habitats remain, although decreased in area. Such vice-counties
are put in ( ). In other cases the identification or the locality is dubious
and the record of a species in a vice-county in [ ] needs confirming. Lastly
there are vice-counties where the introduction of a species has been affected
by man either by planting or by providing artificial habitats; such vice-
counties are denoted by an *. The arrangement and the nomenclature of
species follow that of Dandy‘® with two changes approved by him; a key to
the vice-county numbers is appended at the end of this paper.
I would like to thank all who have helped in this compilation, in par-
ticular Dr, K, L, Alvin for the Isoetaceae records, and Mr. D. H. Kent and
Dr, F. H. aimee Director of the B.S.B.I. Maps Distribution Scheme, for
their co-operatio
LYCOPODIACEAE
1 LYCOPODIUM L.
I L. selago L. Fir Clubmoss
Native. Heaths. moors, mountain grassland and rock ledges; rarely lowland.
1-6, [7], (8), 9-14, [15], 17, 22, 23, [24], 27. 33, 34, 36, [37, 38], 39, 40, [41],
42-50, 52, [54], 55-60, 62-83, 85-1 12; H1-40.
2 L. inundatum L. Marsh Clubmoss
Native. Wet heaths in lowland areas, loca
1-5, 8, 9, 11-19, 21, 22, 24-31, 37-39, 48, i), Vet 61-63, 69, 70, 75, 85,
87-92, 95-97, 99, 100, 104-106, 108, 111: H2, 3 18, 20, 2 27.
3 L. annotinum L. Interrupted Clubmoss
Native. oors in mountain areas, local.
Oe aL 59], 62. [66], 69, 70, 72, [78], 88-92, 94-98, [100, 103}, 104-107,
4 L. clavatum L. Stag’s Horn Moss, Common Clubmoss
Native. Heaths, moors and grassland in mountain areas; rarely eters
1-6, [7], 9-18, 20-24, 26-30, 32, 34-52, 54-81, 83-100, 103-106, 108-112
H2-6, 10, 11, 13, 15-18, 20, 21, 24, pn Daye
t 1 & S KS 64-66, (67),
(71, 77, 82), 85, (86), 8-90, (91), 92, 93, 98, [100], 104, 106, 108, 110; Hl,
(2, 8), 9, 13, 14, (15, 1. 17, (18), ins, (26), 28, (29), 33, 36, 40.
5 E. fluviatile L. (£. limosum L.) Water Horsetail
Native. In shallow water at the edges of lakes, ponds and in ditches.
In all vice-counties except Monmouth, v.c. 35 and Mo ontgomery, v.c.47.
6
6 E. palustre L. Marsh Horsetail
Native. In most wet habitats from open shallow water to wet woods and
mountain flushes
In all vice-counties except Montgomery, v.c.47.
7 E. gs ks Wood Horsetail
Native. Dam s and moors, usually on acid soils.
1, 2, 4-6, (7), 8 i os] (26), 27, 28, (29, 30), 32-51, 53-60, 62-112; H1-18,
20, 21, 24-40.
8 E. pratense Ehrh. Shady Horsetail
Native. Wet grassy stream banks in hilly district:
[59], 65, 66, (68, 69, 72), 75, (77), Ue), (3.87), "88-90, (91), 92, [94], 95,
96, 98, [104], 107-110, (i11); H33, (34, 35), 3
9 E. arvense L. Common or Field Horsetail
Native. Fields, in sandy = eye in wet areas e.g., flushes, etc.
Widespread in ‘all vice-cou
E. x litorale Kiihlew. ex ee upr. « ed x E. fluviatile)
Native. Occurs with parents quite frequently.
2, 5, 6,8, 12, 14, 16, 17, 22, 48, 53, 54, [59], 69, 88, 90, 107; H. 1-3, 35, 38.
JO E. elmateia Ehrh. (E. maximum auct.) Giant Horsetail
Native. On shady banks often in heavy so
0, [1, 2], 3, ); 5-46, 46-71. 73-77, 81, 83, rea 85, 90-92], 98-102, 104. 105,
108, [111]: H
OSMUNDACEAE
5 OSMUNDA L.
7 O. regalis L. Royal Fern
Native. Fens, bogs, wet heaths and wet woods (carr) on peaty soils.
0, 1-7, (8), 9-19, 21-28, (29, 30, 32), [30], Se 3p ae By 4, >), 56-78, 81, 82,
84-88, 91, 93-95, 97-105, 108-110, 112; H1-3
HYMENOPHYLLACEAE
6 TRICHOMANES L.
ie? ‘cw _ ps andrewsii Sea Bi treed Saw Bristle-fern
Native. In dar revices, and un
48, (49), 63*, (64, ®) 98, 100. [101]; 7 >, : (4, S oe : (9, 10), 13, (20),
28, (33), 35, (36), 3
7 ee Sm.
Ii. ogee . 2 et Tunbridge Filmy-fern
Native. On r runks in moist and ere places
(1, 2), 3, (4), 5, Tel ia, 09, BB6. 39], 41, (42), 44, 46, 48, 49, [52, 57, 58, 59],
60, Bes 1 SE ie Be | 8 71), [78], ey (97), 98-102, (103), 104,
[105]; Hi- 3, (4-6), 8, 1D), 16, (24, 25), 27, (28), 29-31, 33, (35) 36, 37, (39).
2 H. wilsonii Hook. (H. peltatum auct.; Wilson’s Filmy-fern
H. erimwyanteand peg |
Native. Habita nbridgense but often in more exposed ons.
1, (2, 3), 4, [9], G9), (40), “tL, (42), 43-49, o). De, 59], Or 50-65, 6 613,
(74), 75, 76, 78, [80], (85), 86-90, 92, 97- 08, [109], 110-112; HI-3,
7, (8, 9, 12-15), 16, 20, (21, 25, - 27, (28-31), 33-36, 9 39 (40).
DENNSTAEDTIACEAE
8 PTERIDIUM Scop.
J P. aquilinum (L.) Kuhn (Pteris aquilina L.) Bracken
Native. Woods, heaths, mountain grassland, etc., usually on light acid soils.
Widespread in all vice-counties
ADIANTACEAE
9 CRYPTOGRAMMA R.Br.
1 C. crispa (L.) R. Br. ex Hook. (Allosorus crispus (L.) R6Ohl.) Parsley Fern
ative. Screes, rocky slopes and walls; in hilly areas on acid soils.
S*. 37), (4 1), 42-44, 46-50, (57, 58), [59], 60, (62, 63), 64-73, [74], 75,
(76, 77), 78, 79, (80), 81, [83], 85- 88, (89), 90, [91], ee ie 96-99, (100),
101, 103— 106, 108, [109j, 110; H20, "30, 313: 33,.3%:
10 ANOGRAMMA Link
I A. leptophylla (L.) Link (Gymnogramma leptophylla (L.) Desv.)
Native. Shaded banks. Only in Jersey and Guernsey, v.c.0.
11 ADIANTUM L.
J A. capillus-veneris L. Maidenhair Fern
gain Peart crevices in calcareous rocks usually near the
0, O 1117), 137 th A7t, i: 367 3S". (39, 40), 41, [48, 49],
53, 8H, Sol 64*, (69), 71, [91, 100]; H9, 16, [17], 27-29, A
BLECHNACEAE
12 ONOCLEA L.
I O. sensibilis L. Bead Fern, Sensitive Fern
An alien. Occasionally grown in bog gardens and has been reported from
swampy ground in N. Hants., v.c.12, and on the edge of Rydal Water, v.c.69.
13 BLECHNUM L.
J B. spicant (L.) Roth (Lomaria Spicant (L.) Desv.) Hard Fern
Native. A calcifuge of woods, heaths, moors, mountain grassland, etc.
Found in all vice-counties
ASPLENIACEAE
14 PHYLLITIS Hill
7 P. scolopendrium (L.) New Hart’s-tongue Fern
(Scolopendrium vulgar Sh )
Native. in auns hederbatie and walls and in rocky woods.
In all vice-counties except 82, 84, 106, 107.
15 ASPLENIUM L.
J A, adiantum-nigrum L. Black Spleenwort
a subsp. adiantum-nigrum
Native. On rocky banks, ledges and walls.
Found in all vice-counties but less common in E. England.
8
5 subsp. onopteris (L.) Luerss.
Native. On rocky outcrops more often of we ey rocks.
1, 90, 92, 93; H1-S, [8], 9, 11, [16], 20, [26], 27, 3
2 A. obovatum Mas ae lanceolatum Huds. 8 art) Lanceolate Spleenwort
Native. Rocky p and walls usually nea
0, 1-5, [9], 14, fish 16. 23, tae [40], 41, a2 ree [45], 48, 49, 59, 62, 63,
[64], 70, (71), 101; Hl, 4; 4,42;33,2
A. refractum T. Moore (A. obovatum x A, trichomanes)
Native. Found by Moore at Pepperharrow Park, Surrey, v.c.17.
3 A. fontanum (L.) Bernh.
An escape, reed found on walls.
11, DD [17], 24,
4 A. marinum i: Sea Spleenwort
Native. Crevices and ledges of cliffs and pg by th
0, 1-5, (6), 9, (10, 14), [15], 41, 44-46, 48, 49, [50], SI. 32. 58, 59, (60), 62,
66-76, 1, 82), 85, 90-92, (93), 94, 95, 97, 08, [99], 100-112; H1, (2), 3-6,
(8), 9, fl, 12, 16, (20), 21, 27-29, 34, 35, 38-40.
5 A. trichomanes L. Maidenhair Spleenwort
Native. On walls and rock crevices of usually basic rocks.
In all vice-counties but the Orkney (v.c.111) record needs more recent
confirmation
6 A. viride Shar Lge Spleenwort
Native. As trichomanes but prefers more bas
£3, 13, 15, 17, aL 1,27], 35, 36, (37), [38], 39, 41, 42, 44, "18 30, 5), “ue i 9)
60, 52), [63], 64 7, (68), 69, 70, 72, 73, 75- -78, 80, BF 4], 8
(6
94-101, ade 10s, 105, [106], 107-110, 112; H (1-3, 6), 7, 8, 16, on ¢
(29), 33, (34
7 A, Fe Ariat a Wall Rue
Native. Walls and rag ie pa rocks.
Throughout the British Isle
A x clermontiae Syme (A. ruta-muraria X A. trichomanes)
Found once on a wall in Louth, v.c.
A. X P ypmbagie Dorfl. (A. ruta-muraria < pie septentrionale;
uta-muraria var. cuneatum T.
Fe found near Dunkeld, E. Perth, v.c. 30.
8 aS aerate (L.) Hoffm. Forked Spleenwort
Native. Crevices in rocks and occasionally on wa
vi “5, 5, 17), [45], 47-49, “150, 67-70, [78], 80, 83, [85], 89, [90], 92, 97,
A. X alternifolium Wulf. (A. septentrionale x A. trichomanes;
A esate auct.; A. breynii auct.)
Native. Crevices in rocks with the pata.
5, [42], 48, 49, 68-70, 80, 83, 85, 89, [90].
meet oft the Plants of the London Area, London.
unty system in practice, B.S.B.I. 1950. Conf. Rep.: 30-42.
1 , R. L. 1901, Irish Focaar Botany, Dublin.
12 eee a4 Bg The distribution of Dryopteris borreri Newm. in the British Isles, Watsonia,
13 Watson, a C. 1883 (Edited, Baker, J. G. & Newbould, W. W.), Topographical Botany, Ed. 2.
14 Webb, D. ‘A. Fa Irish Plant Records, Watsonia, vol. 2 : 217-236.
15 Wilson, A. 1930, A Census Catalogue of British Hepatics, Ed. 3, Berwick-on-Tweed.
14
VICE-COUNTY NAMES AND NUMBERS
West Cornwall
East Corn
Z
°
S
fe
south Somerset
Wilts
wh
ee:
a
6 West Kent
urrey
South Essex
? North Essex
e
Middlesex
Berks
Oxford
Bucks
25 East Suffolk
KAD AD
Linlithgow
85 Fife (with Kinross)
South Kerry
Waterford
South Tipperary
Limerick
—
orth Tipperary
Kilkenn
Wexfor
Carlow
Leix
jo of vhs via vie oe sks shocks nie ste sfeohs hea
N=Svarnd Auaun=
Sele dlendiea tad
ew
CHANNEL ISL
0 yl eg
ENGLAND AND WALES
=
g
wn
=)
z.
*
KAD AD A
@)
5
os
o
ogg
Ou.
g9
a
aay
=
g
onal
20
£0
o
=
Hereford
Worcester
rwic
Stafford
Ow! COnIN' t
alop
Glamorgan
Brecon
Carmarthen
Cardigan
Montgomery
Merioneth
Caernarvon
IsLE OF MAN
71 Isle of Man
vi gece
86 Stirlin
87 West Perth (with
Clackmannan)
88 Mid Perth
Eas
a a a ae ae a ee
t
CONIA
238
92 South Aberdeen -
3 North Aberdeen
anff
\O \o
BS
95 Elgin
96 Easterness (w. Nairn)
Westerness
98 Main Argyll
IRELAND
’ South-east Galway
West Galway
Si agg ae
Offa
DOD DO ee et et et et
WOO~AIAUN
= xa
g c ogy
AS
oa
=
fiefs ois fe a}e ofa ofan jc spa nge ge spangeo)
Ne
>
= a-%
SREB
=
West Mayo
15
50 Denbigh
51 Flint
n
53 South Lincoln
55 Leices
(vith Rutland)
Nottingham
op
South Lancaster
urham
67 Northumberland
South
68 Cheviotland
69 Westmorland
(with N. Lancaster)
70 Cumberland
99 Dunbarton
100 sts Isles
Kin
South ibialiad
Mid Ebudes
North Ebudes
West Ross
—
East Sutherland
West Sutherland
Caithness
Outer Hebrides
rkn
Zetland
H 28
2
H 30 Cavan
H 31
A
Londonderry
THE GEOGRAPHICAL AFFINITIES OF THE NEW ZEALAND
PTERIDOPHYTE FLORA
a Se LOVIS
Botany Department, Leeds University
The problems of plant geography presented by the plants of the Southern
emisphere are by their very nature more spectacular than those which
exercise students of plant geography who are concerned solely with the
distribution of plants in the Northern Hemisphere. In the Northern
Hemisphere there is only one major barrier to plant migration, the Atlantic
Ocean. The Bering Straits are relatively narrow and have certainly been
in the relatively recent past. In the Southern Hemisphere, however, there are
four major land masses — South America, Africa, Australia, and Antarctica,
all separated from one another by substantial expanses of ocean. The
islands of New Zealand, with which we are primarily concerned here, are.
similarly extremely isolated, there being some thousand miles of the Tasman
Sea between New Zealand and Australia, the nearest continental land mass.
It is therefore not surprising that although the major problems connected
with the origin and relationships of the British Flora — such as the presence
of certain species of Mediterranean affinity, and some of North American
affinity, in isolated localities along the western coasts of the British Isles —
have been the subject of plausible explanations, the remarkable patterns
of plant detitianion in the Southern Hemisphere are still extraordinarily
difficult to explain in an entirely satisfactory manner. It is intended to
demonstrate in this article the peculiar interest of these problems of southern
plant distribution, with particular reference to the pteridophytes of New
Zealand
We must first emphasise that, in relation to the pteridophytes, such
problems of great difficulty really do in fact exist. It is sometimes somewhat
facilely supposed that on account of their copious production of spores,
which being of very light weight are wind-borne, pteridophytes must be more
widely diffused and distributed than are flowering plants and that oceanic
arriers, for instance, must be of little consequence. Study of actual plant
distributions shows this generalisation to be manifestly unjustifiable. Thus
it is known that not only do pteridophytes, and bryophytes and orchids as
well, have distributions which by and large show the same characteristics of
interruption, and apparent inconsistency as do flowering
plants w possess relatively heavy disseminules, but moreover these
former ial include a comparable proportion of endemic species. Long
distance dispersal seems to be as ineffective an agent for explaining the
presumed past migrations of pteridophytes as it is for flowering plants.
It is true that such studies as those made on the recolonisation of the volcanic
island of Krakatoa ®) clearly indicate that forty miles of sea is no obstacle
to plant dispersal, but even to the pteridophytes a thousand miles of ocean
appears to be in general an insuperable barrier.
It has been recognised for some time) that within the New Zealand
flora elements with certain geographical affinities can be detected. It is
16
possible to classify the flora into palaeoendemic, cosmopolitan, palaeo-
tropical, Australian, and Antarctic or South American elements. At the
outset, it should be pointed out that these categories are not all mutually
exclusive. For instance, the same species can be Australian and palaeotropical
in affinity.
With relation to the vascular flora as a whole, the endemic element in
the New Zealand flora is extremely large and by far the most important,
including some 80°% of the whole, an exceptionally high figure. The degree
of endemism in the pteridophyta is much less though still very important,
being of the order of 40%. Some outstanding and distinctive plants come
into this category, including Cardiomanes (Trichomanes) reniforme (Forst.)
Presl, Ap Abed superba (Col.) Pres], Mecodium (Hymenophyllum) dilatatum
(Forst.) Copel., and Loxsoma, a monotypic and endemic genus. A significant
fact is that ie tee-fern genera, Dicksonia and Cyathea, with their huge spore
production, nevertheless show a much higher degree of endemism than the
New Zealand pteridophytes as a whole, some 75% of the New Zealand
species of these two genera being endemic.
Recent researches have clearly shown that the cosmopolitan element in
the New Zealand pteridophyte flora is eauch smaller than was previously
considered to be the case. This fact lier ar ticle, but
the recent publication of cytological investigations by Brownlie, makes it
now possible to give fuller details. Thus H) (Poir) v.
in New Zealand is found to have a chromosome number ofr = 11, im
comparison with m = 18 in Britain (Manton,“” as H. unilaterale Bory).
Equally striking is the case of the former Hymenophyllum tunbrigense of
New Zealand, now cited as H. revolutum Colenso, which has n = 22, as
against n = 13 for H. tunbrigense (L.) Sm. in Britain. In each of these two
cases, the chromosome numbers found are without any numerical relation-
ship to one another. Brownlie,“ was also responsible for the discovery
that Asplenium trichomanes L. in New Zealand is hexaploid, with n = ite
whereas only diploid and tetraploid forms are known elsewhere. How
in this case, the author’s own investigations in New Zealand, still wiskalanod.
show that although this endemic hexaploid is by far the commonest form of
truly co to the British Isles and New Zealand is Anogramma
lestopasite .) Link. Cystopteris fragilis (L.) Bernh. is tetraploid in New
Zealand, with n = 84,‘ but in this case the plant is not identical with its
British tetraploid equivalent, being morphologically quite distinct, its frond
being less dissected, and the pinnules themselves larger and more yar
The appropriate name for the New Zealand plant would seem to
Cystopteris tasmanica Hooker, since the New Zealand plant is esis
with its Tasmanian and Victorian counterpart.
The element of palaeotropical affinity in the New Zealand pteridophyte
flora is not inconsiderable. A number of predominantly tropical genera are
17
represented. These include such genera as Cyclosorus, Nephrolepis, Sticherus,
Dicranopteris, Lygodium, Pteris, Lindsaea, Adiantum, and Marattia, and it is
no accident that most of the New Zealand species of these genera are either
restricted to the sub-tropical north of the North Island or are more
abundant in this region than elsewhere in New Zealand.
The strongest affinity of the New Zealand pteridophyte flora is with
the pteridophyte flora of the temperate regions of Australia. About 45%
of the New Zealand pteridophytes are also present in eastern Australia.
From the Australian point of view the affinity is even more marked, for some
two-thirds of the pteridophyte flora of Victoria and Tasmania are also
found in New Zealand.“*) This common element in the Australian and
New Zealand floras is far more significant amongst the pteridophytes than
the flowering plants. Thus although 45 °% of the New Zealand pteridophytes
occur in temperate Australia, this is true of rather less than 20% of New
Zealand flowering plants. With regard to the Australian flora, the contrast
is even more marked. Some 2,250 species of flowering plants are known to
occur in Victoria and Tasmania, but only fewer than 250 of them occur in —
New Zealand, little more than 10°, in comparison with 60-70% of the
pteridophytes.
This last point is well illustrated in the sub-tropical rain forest which is
developed in suitable locations all the way down the coast of New South —
Wales from the Queensland border region. Although occurrences of this
vegetation type become increasingly less frequent south of Newcastle, there
are still isolated small pockets of this sub-tropical rain forest or ** jungle ”
as far south as eastern Victoria. The general appearance of this forest type
is much like the sub-tropical rain forest of New Zealand, but the trees and
other flowering plants show little relationship with those of the New Zealand
rain forest. However, there is a striking similarity between the fern flora of
the New South Wales and Victorian rain forest and that of northern New
Zealand forests.
Nearly all of the pteridophytes common to New Zealand and eastern
Australia can be placed into one of two geographical groups. There are
firstly such species as Adiantum diaphanum Bl., Diplazium japonicum (Thbg.) —
Bedd., Pellaea falcata (R.Br.) Fée, Pteris comans Forst., Sticherus flabellatus
(R. Br.) H St. John, and Todea barbara (L.) Moore, all of which are in New
Zealand dither almost or completely confined to the far north. In Australia,
these species all extend south throughout New South Wales, and most also
occur in Victoria, although generally much rarer in that state than further
north. This first group of species are clearly of tropical origin, and indeed
Adiantum diaphanum and Diplazium japonicum have wide ranges in the
tropics today. The second group of species, which is the larger, is of mixed
origin. These species are for the most part more or less completely distributed
in New Zealand, while in Australia they are more abundant in Victoria and
Tasmania than in New South Wales. Of some sixty species having this
southern type of distribution in eastern Australia, more than fifty occur
also in New Zealand.“* Examples are Blechnum fluviatile (R.Br.) Lowe, —
18
te etd heterophylla (Labill.) M. Tindale, Hypolepis rugosula (Lab.) J.Sm.,
Asplenium flaccidum en - Mecodium rarum (R.Br.) Copel., and Phomatedie
veri (Willd.) C
also fists e compare the pteridophyte flora of New Zealand
with sis of the far south-west of Australia. The south-west corner of
Australia by a thousand miles of desert. Although the genera are nearly all
the same as in eastern Australia, the vast majority of south-west Australian
For
comparison with the flowering plants, the pteridophyte flora of south-west
Australia is exceedingly poor, there being only twenty-four species fe)
less than sixteen of these also occur in New Zealand, although oe an
insignificant number of species of flowering plants is common to the two
regions. The reason for this paradox is that the impoverished pteridophyte
flora of south-west Australia is composed of species of general wate
over the favourable regions of Australia, and which thus also occur
eastern Australia.
come now to the Antarctic or South American element in the
New Zealand flora. There is, rather surprisingly, a small number of New
Zealand ferns which are also found in the extreme south of South America.
Nearly all of vera also occur on one or more of the remote islands of the
sub-antarctic ocea
‘ileal: (Hymenophyllum) ferrugineum (Colla) Copel., a filmy
fern found in the darkest parts of the New Zealand rain forest, is also found
not only in Chile but also on the island of Juan Fernandez, 400 miles out
into the Pacific off the coast of Chile.
Grammitis billardieri Willd. is a common fern in New Zealand,
Particularly in the South Island. Outside New Zealand, it not only occurs
in eastern Australia, in southern Chile, and in the Falkland Islands, but
also on the isolated island of Kerguelen. The nearest continental mass to
naglecte Island, which lies almost on the 50th parallel of southern latitude,
is Antarctica, some twelve hundred miles distant. Otherwise, Kerguelen
lies nearly equidistant from South Africa and Western Australia. The flora
of Kerguelen is of surpassing importance and interest for the southern
return, but it is first necessary to mention two more species of fern which
have distinctive circumantarctic distributions.
Blechnum penna-marina (Poir.) Kuhn is a common fern in New Zealand,
though most abundant at montane levels in the Southern Alps. Outside
New Zealand, it occurs in eastern Australia and southern Chile. It is also
found on Tristan da Cunha and on Kerguelen, and also on the smaller Crozet
19
and Marion Islands, which lie between Kerguelen and South Africa, and on
the Islands of St. Paul and Amsterdam, north of Kerguelen.
The distribution of Polystichum mohrioides (Bory) Presl is perhaps most
curious of all. The only sizeable land mass where this fern occurs is southern
Chile. It is also known from the Falkland Islands, South Georgia, from two
small islands off the main coast of New Zealand, and from Marion Island
(west of Kerguelen), and Amsterdam Island (north of Kerguelen), though
not, apparently, from Kerguelen itself.
Having now briefly described the geographical affinities of the New
Zealand pteridophyte flora, the problem remains how did these plants of
various distribution all come to be present in New Zealand, separated as it
is from its nearest neighbour of any substance, Australia, by a thousand
miles of ocean.
We have already indicated that the long-distance dispersal of spores
must be rejected as an effective means to explain the origin of the existing
distributions of pteridophytes. There is no doubt that on account of their
light spores pteridophytes can readily be dispersed from one ecologically
suitable habitat to another some tens of miles distant, by-passing unsuitable
terrain in between, but clearly the wind dispersal of spores must generally
be an ineffective agent for securing the successful migration of pteridophytes
across thousands of miles of ocean. There are exceptions to every rule, an
it is quite possible that some orchids and pteridophytes have become
established in New Zealand owing to the transport of their seeds and spores
by the strong prevailing wind across the Tasman Sea from Australia. But
even if long-distance dispersal of this kind has occasionally in such exceptional
cases been successfully accomplished, we are still left without any explanation
of the entirely comparable distributions found amongst the flowering plants,
many of which have seeds or fruits which are so heavy that their natural
long-distance dispersal is clearly impossible.
The earliest rh tainetee studies of any consequence in these
southern regions are those of Sir J. D. Hooker, whose introductory essays
to his Floras of Antarctica, New ak and Tasmania, although written
a century ago, are classical studies of this subject, and are still very well
worth attention today for their scientific content as well as their historical
interest. These, with others of Hooker’s writings, have been ewer
extracted and copiously annotated in relation to subsequent research by
Turrill.“® Hooker, who described the enigmas of southern plant aisiribie
almost as we know them to day, came to the conclusion in relation to the
New Zealand flora that the various migrations of floras which must have
taken place in this region have most likely been effected through land
communications across the ocean barriers, the intervening land having since
disappeared.
Subsequent writers on the New Zealand flora, such as Cockayne,“
have also felt compelled to rely on such hypothetical land bridges, even
though some of these, on the basis of present continental distribution,
20
must have been very long, and even though in many cases studies of ocean
levels seem to render it most unlikely that such a land connection ever
existed.
In this connection the case of the island of Kerguelen, already mentioned,
is instructive. The entire native vascular flora of Kerguelen is extremely
flowering plants and nine are pteridophytes. Although situated between
South Africa and Australia, the flora of Kerguelen shows its greatest affinity
with the flora of the tip of South America, the region known as Fuegia,
seventeen of the thirty Kerguelen plants being found there also. Next in
degree of affinity, and again in spite of the enormous separation in distance,
is the New Zealand flora, for eleven of the flowering plants and probably
four of the pteridophytes of Kerguelen are also found in New Zealand. The
impoverished flora of Kerguelen cannot be accounted for by casual long-
distance dispersal, for eight species are endemic*, including two which belong
to endemic monotypic genera, and one of these, Pringlea antiscorbutica R.Br.
ex Hook., has no known near relative. The existence of a land bridge from
South America is clearly equally unlikely.
There remains the hypothesis of continental drift, the idea that the
major continents were at one time far closer together and have subsequently
drifted apart into their present positions. Ever since this hypothesis was
first advanced in a comprehensive form by Wegener many biologists have
pointed out its great potentialities as a means of explaining various problems
of plant and animal distribution. However, notwithstanding the fact that
Du Toit produced a modified form of the hypothesis, removing the
anomalies present in Wegener’s original concept, continental drift remained
subject to vigorous papain aiigecagii from geophysicists who were
unable to conceive of a mec m by which the wholesale movement of
continents might have been nected In recent years the development of
the study of rock magnetism has produced powerful evidence in support
of the concept of continental drift. There is no space here to describe this
new evidence, but the reader is referred to two admirable popular summaries
which have recently appeared.“*-1*) In consequence of this new development
it can now be said that the total evidence, biological and geological, in
favour of the hypothesis of drift has almost lifted the concept from a state
of possibility to one of probability. What, then, is the oe of this
hypothesis in relation to the problems being discussed here
According to Du Toit there were originally two great initial land
Masses, a northern mass named by him Laurasia and a southern mass
oO
further. Included in Gondwanaland were South America, Africa, India,
Australasia and Antarctica. Significantly, most proponents of continental
drift consider that the disruption of the masses took place relatively late
in geological time, the major part of the movement taking place in
* This is not strictly true, for all of these species occur 0 aie oe ee eS ee
Kerguelen archipelago, namely Heard Island and the Crosst-and Marion
Tertiary period, in which the flowering plants were already the dominant
vegetation type. It is further considered that the separation of South Africa
and India took place before the splitting away of South America and
Australasia from Antarctica. Kerguelen Island became detached directly
from Antarctica. New Zealand and New Guinea became detached from the
eastern coast of Australia and were subsequently displaced in position.
The separation of New Zealand must have occurred before the develop-
ment of the characteristic Australian flowering plant flora as we know it
today, for there are, for instance, no Eucalyptus or Acacia species native
to New Zealand. What affinity does exist between the flowering plant floras
of Australia and New Zealand is predominantly composed of species and
genera of southern or Antarctic affinity and which are today in Australia
mainly concentrated in the mountains of Victoria and in Tasmania. We
have already seen that the affinity between the pteridophyte floras of Australia
dissociated from Australia. Equally, the general relationship found to
exist between the pteridophyte floras of New Zealand, of both east and
Island is entirely consistent with the belief that the circumantarctic element
in this flora had largely developed before the disruption of the great
Gondwanaland continent had proceeded very far. There is therefore no
need for the postulation of extended land bridges to explain the isolated
and anomalous position of the Kerguelen flora which, instead, is seen to be
entirely consistent with the presumed geophysical history of the region
according to the concept of continental drift.
The apparent centring of the southern hemisphere pteridophyte floras,
with the partial exception of Africa, upon Antarctica has intrigued many
students of plant geography. The most extreme attitude is that of Copeland
(5,8) who strongly supports the concept of an Antarctic origin for the
majority of fern genera. His views have been strongly criticised by many
but, if we were to substitute Gondwanaland for Antarctica, perhaps his
hypothesis would come to be regarded as more acceptable.
P.S. This article was written before the Royal Society Discussion Meeting
n “The Biology of the Southern Cold Temperate Zone” was held on
December 10th and 11th, 1959. Several important papers relating to problems
discussed in this article were read on this occasion and these will soon
appear in print in the Proceedings of the Royal Society. J.D.L.
vol. 82 :
REFERENCES
i mehde ci G. 1954, 2 EF OEENY note to Cyto-taxonomic Studies of N.Z. Ferns, Trans. Roy. Soc.
2 Brownlie, G. 1958, Chromosome Deeers
Z. Ferns, Trans. Roy. Soc. N.Z., vol. 85 : 213-6.
3 Chaloner, - G. 1959, Continental Drift, New B Bio. lology 29: 7-30.
4 Cockayne 1928, The Vegetation of New Zeal, (Die Vegetation der Erde, 14) 2nd Edition.
rig.
5 Copeland, E. B. 1939, Fern Evolution in Antarctica, Philippine J. Sci., vol. 70 (2) : 157-88.
6 Copeland, E. B. 494i, Bg ed Filicum. vane Mass.
7 Du Toit, A. L. 193 Wandering Continents. London.
”
8 Gams, ne 1938, — der Extratropischen Pteridophyten, Verdoorn F., Manual of Pteridology.
e Hague
9 Harris, te F. 19 1 of the S; f the N.Z. Pteridophyta, Bull. N.Z. Dep. Sci. Industr. Res.
No 116.
td a s, J. D. 1957, Fer n New Zealand, . Fern Gat., vol,
: 184-8.
Manin I. 1950, lvohleint of C grr oe acsiaion in the Pieridophyta Cambridge.
12 Opdyke, N. D. & Runcorn, S.K. 1959, Palaeomagnetism and Ancient Wind Directions, End22vour
vol. 18:
13 Turrill, W.B 1953, Pioneer Plant Geography, (Lotsya, vol. 4.) The Hague.
14 Wake field, Ri A. 1955, Ferns of Victoria and Tasmania. Melbourne.
CULTIVATION OF FERNS UNDER GLASS
R. BRUTY
Kew, Surrey
Ferns grown under glass can be discussed under two main headings, tropical
and temperate. The tropical fern house needs to be maintained at 80°F.
with high humidity from March till September; from October to February
the temperature may be lowered to 65-70°F. The temperate fern house on
the other hand, should be maintained at 50-60°F. throughout the year.
They are not affected by the shorter days and growth may continue.
Light is the next most important factor. Tropical ferns are not used to
such wide seasonal fluctuations as in our own climate, especially in day-
length, but lowering the winter temperatures slows down growth and prevents.
them from becoming etiolated. Summer light is often too intense and the
ferns must be shaded. This is best done by roller blinds, so as to allow the
maximum amount of light in dull weather. The green shading “‘ Summer
Cloud” can be used; if it is a dull summer this makes the plants draw up
vert weak and there is more risk of the plants getting fungal disease.
pagation. The easiest and most interesting way to propagate ferns
is by spores. The fertile fronds should firstly be inspected to make sure the
spores are ready for collecting. With a lens or pocket microscope one can
see if the sporangia have begun to open; when they are just beginning to
open at the base of the frond that frond should be cut and placed in a large
envelope and left in a dry room for two or three days, after which time the
spores will have been shed and can be shaken out of the packet. Such spores
may be stored in a dry packet for some months or may be sown immediately.
Th n be sown in small pots or pans, the latter being far the best as
the prothallus has more room to expand. A compost of two parts peat,
one part loam and half part sand, sifted through a quarter-inch sieve, should
be sterilised together with the pans and crocks. Whilst the pans can be
immersed in boiling water to sterilise them, the soil cannot. A simple and
efficient method is to place the soil in a thin sack and steam in a bucket over
the gas: the soil should be prevented from touching the water by a brick or
similar object and should remain at 180°F. for ten minutes. Only one piece
of crock is necessary at the base of the pan and as the pans are to be stood in
a saucer of water a thin layer of rough peat should cover the bottom. (Instead
of water a weak solution of potassium permanganate may be used to keep.
down algae.) All water is taken up in this way and overhead watering is.
Ot necessary at any time. The pans are filled to within an half inch of the
23
top and a piece of glass cut to shape, or a clock-glass, if pots are used, is
placed on the pan. (If a clock-glass is used it is important that the convex
side is upper-most). These pans must be kept covered until the spores are
sown.
The name of the fern should be written on a label before sowing. The
glass is then taken off and the spores sprinkled on the surface of the soil;
the label is placed in the pan and the glass cover replaced before sowing the
next species to prevent contamination and the mixing of spores.
These pans are stood in saucers of water and put under the staging
where it is moderately light but shaded from the sun. They can be left here
until fertilisation has taken place and the young sporelings appear. From
now on they need more light and must be stood on the staging but the cover
glasses should remain over the pans for a few more days. The young spore-
lings are now large enough to handle and should be pricked out one dozen
or so to a four inch pan and from then into small pots.
Propagation can be done vegetatively from bulbils (really buds) that
arise on the back of the rachis. in, for example, Asplenium bulbiferum. If
such a frond is taken off and pegged onto the surface of a pan filled with
potting soil and placed in a closed frame the bulbils will quickly root. Other
ferns form young plants at the axils of the pinna rachis, e.g., Hemionitis
palmata and Doryopteris pedata and these can be propagated in a similar
way. Rhizomatous species of Polypodium and Davallia can be started by
cutting off a piece of rhizome with a growing point and planting this.
Composts. Peat is preferred to leaf soil when potting ferns because
leaves carry eel-worm and this is not always destroyed when sterilising the
soil. Tropical ferns grown much better in a light compost: one part fibrous
loam, three parts granulated peat, half part sand. To each bushel of compost
add 2 oz. super-phosphate, 2 oz. hoof and horn or bone meal, and | 0z.
potash. For the Maidenhair ferns (Adiantum) add some old mortar rubble.
Angiopteris and Acrostichum danaeifolium being swamp species need @
much heavier compost: one loam, one peat and half sand. A general compost
for temperate ferns is one loam, two peat and half sand with the usual
fertilisers, but for Pteris and Osmunda reduce the peat by half.
There is no need to put this soil through a sieve unless it is for small
pots, just break the loam up with the fingers, and the peat which is purchased
in a bale must be well soaked a week before use. Ferns should not be potted
firmly; there is no need for a potting stick until the ferns are put into five-
inch pots, and then only lightly to make sure the soil goes down between
the plant and pot.
Ferns need watering at least once a day during the summer months and
if it is very warm it is best to check that they are not too dry in the afternoon,
water should be poured into the top of the pot until it is full; never half fill
the pot and do not spray ferns overhead. The water should be allowed to
drain onto the shingle or ash. Never have a pot standing in water. From
October till February watering three times a week will be sufficient for most
terrestial ferns but large tree-ferns (Cyathea) and Osmunda if in pots and not
—_—
recently potted are best watered every day. Species with rhizomes such as
Polypodium and Davallia and epiphytes such as Platycerium need water only
once a Alay in the winter months.
. The chief pest
be een meer As most insecticides are harmful to ferns it is worth 1 mention-
ing a few which have been found to be effective. ‘* Diazitol ” can be used
against mealybug and aphids with very little damage being done at a strength
half oz. to two gallons of water. ‘‘Malathion’”’ or ‘Kill’ can be used for scale-
insects at the strength stated on the bottle but more care should be taken
with this as it may cause damage to the very young fronds. Fumigating
with “* Nicotine Shreds ”’ as stated on the packet, is good for aphids. A
whole house of ferns can be ruined by the eel-worm, and prevention by
keeping the foliage dry is the best answer. If a plant is attacked, the fronds
should be cut off and the roots placed in a container of warm water (at
110°F.) for 10-15 minutes. Badly infected plants should be thrown out com-
pletely once spores have been taken for the propagation of the plant.
kK y 1 4 + lh := hi:
SHORTER NOTES
WOODSIA ALPINA (Bolton) Gray
Woodsia alpina is one of our rarest ferns. It does not grow below at least
2,000 ft. in Britain — it is better suited at 3,000 ft. — and the number of
stations probably does not now exceed six on the mountains of Wales and
Scotla he appearance of the fern in a natural habitat is well shown in
. Swinscow’s photograph reproduced in the 1955 issue of the
Gazette (VII, No. 5). It will be noticed that the fern has some overhead
protection; and this is no doubt important because it gives shade and prevents
damage by heavy mountain rains. The view that some such protection is
needed is supported by the late Professor Bower’s observation that alpine
plants in Scotland are usually to be found on rocks facing N.E. There may,
however, be further advantage in a covering of snow in the winter which at
the top of some Scottish mountains, Ben Lawers, for instance, may persist
until the late Spring; and the grimmer N.E. aspect may be helpful to high
alpines, which are not well suited by our atlantic climate. It is well that this
fern grows in places not accessible to the ordinary rambler — there was
some reprehensible collection of it in the last century.
Woodsia ilvensis (L.) R.Br., almost equally rare, was at one time con-
fused with W. alpina. It is slightly less exacting in its choice of habitats,
both as regards altitude and shelter. It is rather larger, and the pinnae long
as compared with the almost triangular pinnae of W. alpina. In course of
time the two ferns came to be rightly regarded as distinct, and in recent
years this view has been supported by a count of chromosome numbers:
W. alpina has double the number of W. ilvensis.
The writer has himself never seen W. alpina in the wild in this country,
but on a visit to Dauphiné in 1937, by a most extraordinary piece of good
luck, came on the fern in the course of a stroll to fill in an hour or so when
detained en route at a mountain village. From this village at about 5,000 ft.
was a track rising gently along an upper valley watered by a stream fed by
a distant glacier. On a flattish area of this valley, at an altitude of probably
rather less than 6,000 ft. were strewn large boulders which had obviously
fallen at some time from crags high above. On one of the nearer boulders,
more or less flat on the top, some vegetation was noticed which looked like
Lousewort and proved to be so. On closer inspection it was seen that with
the Lousewort were two or three ferns, and these were at once identified as
Woodsia alpina — the peculiar indusium is unmistakable. There was no
time to examine boulders further away and a little higher up, otherwise more
plants of the Woodsia would probably have been found.
A year or two later some detailed information about Dauphiné plants
was found in the Abbé Ravaud’s “‘ Guide du Botaniste dans le Dauphine,”
published at Grenoble about 1890, The Abbé appears to have been a com-
petent field botanist and he covers not only flowers but mosses and lichens,
and ferns and fern allies. The book is schemed as thirteen excursions penetra-
ting the mountains in different directions. _Woodsia alpina is given as occut-
ring in eight different places, but the habitat is described in one case only —
which Woodsia would be growing in the other places for which it was recorded
is about 7,000 ft.
The proximity of the fern to the Lousewort was perhaps merely a coin-
cidence, though the association may have been helpful to the fern. The
species of the Pedicularis cannot be stated. The Abbé lists nine species for
Dauphiné but not P. sylvatica L. The total number of species in the Central
European mountains is approximately twenty. In this country we have two
species only, P. palustris L. and P. sylvatica L.; and P. sylvatica, which grows
among the British mountains, has often been mistaken for a fern; indeed, at
a little distance it does bear some resemblance to Woodsia alpina. Edward
Newman in his History of British Ferns quotes from Ray’s Synopsis 4
comparison of the general appearance of Woodsia with that of Pedicularis.
Woodsia alpina falls within the category of artic-alpine plants. It 1s
characteristic of such plants, suited by alpine conditions in central Europe,
to tolerate lower altitudes in more northerly situations. Woodsia alpina is
a straightforward example as growing at about 7,000 ft. in the Alps and 3,000
ft. in Britain. In northern Europe, e.g. Scandinavia, there is scope for it
to descend nearer to sea level.
P. GREENFIELD
HERBARIUM SPECIMENS OF FERNS AND THEIR ABERRATIONS
Discussions in Ambleside and London at this year’s annual meetings of the
Society prompted the formation of a scheme for the preservation of herbarium
specimens of ferns and their aberrations. Although the original concept was
restricted to varieties (horticulturally desirable aberrants), it is felt that we
should seek to embrace the subject completely in its relation to the botany
26
of ferns and fern-allies. This means that specimens should be preserved for
the following main lines of approach:—
1) to illustrate the range of aberration, not only from separate
localities but also within micro-populations; juvenile and secondary growth
etc. should be included;
2) to build up a reference collection of authentic specimens of
named varieties, bearing in mind the existence of a few herbaria such as those
of Cranfield, T. Moore and Padley. Although the original type-specimens
would be best for this purpose there are very few, or adequate pictures of
them, in existence. Vegetative progeny from the original type would be
the next best choice; it may of course be very difficult to be certain that
what we have is undoubtedly clonal (uniform material derived from a single
individual and propagated entirely by vegetative means). The selection and
preservation of the most representative specimen from the authentic material
available will, consequently, often be necessary. After these three methods
of representation the most useful procedure is to collect together as many
apparently significant specimens as possible;
3) to fill in vice-county distribution gaps for species in the
Preliminary Census List on p.4.
The basic principles to be observed in making a dried specimen are three.
A) The simplest way is to place the fresh frond into a folder of
unglazed paper (e.g., newspaper) in which it will be kept until finally mounted,
immediately laying folder and specimen between several sheets of botanical
drying-paper (or any porous paper — blotting-paper and the majority of
ewspapers are satisfactory) with a weight on top. To keep good colour
and to minimise risk of moulding, change the drying-paper each day for the
first few days. Gentle heat (e.g., beside a fire or on top of.a hot-tank) may
be used to hasten drying, in which case use the alternative method of applying
pressure, viz., place the bundle between wooden lattices or similar stiffeners
as ina botanical press, and tie tightly.
Uniformity of sheet size is desirable for herbarium curation.
The International Code of Nomenclature for Cultivated Plants, ae 1958,
recommends that specimens of nomenclatural importance be in a
public herbarium. If, as seems generally acceptable, the Beith Museum
(Natural History) is to be used, the sheet size would be 13 x 21 inches.
C) Adequate labelling is vital. For specimens other than varieties
the basic details should be:— locality (with vice-county number and grid-
reference, if possible), altitude, date, habitat, collector. For varieties in
cultivation the following points should be covered:— full name of variety
and name of original describer or finder; name and address of present
Owner; previous history of ownership; whether designated as being from the
original (if so, whether clonal or from spore progeny); where grown (pot,
greenhouse, garden, etc.); date of preservation; number (it is a good safe-
guard against possible ambiguity if each plant and dried frond(s) therefrom
bears a corresponding reference number); any other useful information.
J. A. CRABBE
27
THE ANNUAL EXCURSION 1959
The Excursion was held this year from the 5th to 13th September in
conditions ideal for collecting. Blue skies and sunshine every day made the
week an outstanding one in the annals of the Society, for very seldom indeed
can our fern hunters have enjoyed such weather on the annual excursions.
The ferns on the other hand were obviously not enjoying it, and were suffering
badly from the drought, particularly those on walls, most of which were
dried up. Everywhere we went, walls were draped with shrivelled fronds of
Polypodium vulgare and Asplenium trichomanes, and in their more exposed
habitats the Lady-fern and the Beech Fern also were looking in a sorry state.
Our party of 13 members and one visitor was the strongest we have
mustered for very many years, although not all of them attended for the
whole week. We were very pleased to welcome Professor R. E. Holttum to
the Excursion, but our outings must have seemed very tame to him after his
expeditions in Malaya, Indonesia and Borneo. The White Lion Hotel,
Ambleside, was our headquarters, and made a very convenient centre for
exploring in all directions without the necessity for going far afield. The
terrain was a varied one, from cliffs, screes and ghylls on the mountains,
to woodlands, road-sides, and the banks of streams in the valleys. Dried up
streams and bogs allowed easy access to places difficult to approach in a
normal season, and we were able to explore many interesting corners in
consequence.
Two days were spent on the high hills rising from the Kirkstone Pass.
There are old records for Woodsia ilvensis and Asplenium septentrionale
from Red Screes, and the more active members of the party climbed this
hill and concentrated their search on the cliffs and screes near the summit.
Many ferns and flowering plants were seen, but the objects of our search
eluded us. We did, however, find Hymenophyllum wilsonii in a deep ghyll on
the mountainside, our only glimpse of this filmy fern during the week. Our
other climbing expedition was in Deepdale on the north side of the Pass.
A long walk up the dale led to an amphitheatre of precipitious hills. at its
head, where our party split into groups and worked up the slopes in different
directions. Ferns were not plentiful here we found, and the finds. of the day
were not pteridophytes, but the alpine plant Silene acaulis and the creeping
willow, Salix herbacea
One more day was devoted to high ground — on Hutton Roof. This
hill is 25 miles from Ambleside and outside the area we intended to covet,
but as several members who had not been before expressed a wish to 80
there we were not averse to including it in the itinerary, for we all had 4
sneaking desire to revisit the limestone pavements with their fern-filled
grikes. Nothing new was discovered but we all enjoyed the day.
The rest of the week was spent on low ground, one day at Ullswater
and the others in the area lying on the west side of Lake Windermere.
Ullswater visit provided an enjoyable scramble up the bed: of the Aira
Beck where 14 fern species were noted. Among them were the Oak and Beech
28
Ferns, Cystopteris fragilis, Polystichum aculeatum, and many fine specimens.
of Phyllitis scolopendrium some of which were multifids; one good plant
was collected. Later in the day at Glencoyne some crested Lady-ferns were.
found
Ferns were plentiful around Skelwith Bridge and up Little Langdale.
A rough road up Yewdale led us to Tilberthwaite Ghyll, where ferns were
abundant in the slate screes and on the rock faces. During the short time
we were there 16 species were recorded. Asplenium trichomanes grew to a
very large size in the screes and there was a fine little colony of A. viride ona
rock face rising from the beck. A few plants of Polystichum aculeatum were.
also noted. We had one of our best days here, and another, our last day, was.
enjoyed hunting in the country between Esthwaite Water and Newby Bridge.
In Dale Park a scramble along the rocky bed of the stream led us through a
fern paradise where the pene rable at with a tropical luxuriance. Beech
Fern cascading down the slopes had exceptionally large fronds, and.
Dryopteris borreri, Athyrium ited -femina, and Polystichum aculeatum all
grew to a large size. We finished our day —and the week — with the
discovery of a fine Royal Fern.
In all we found 25 species of ferns.
Osmunda regalis L. — Ambleside, Dale Park Beck.
Hymenophyllum wilsonii Hook.— Red Screes.
Pteridium aquilinum (L.) Kuhn — in all areas.
Cryptogramma crispa (L.) R.Br. ex Hook.— Ambleside, Colwith, Deepdale,.
Glencoyne, Hawkshead, Red Screes, Tilberthwaite Ghyll.
Blechnum spicant (L.) Roth — Dale Park, Deepdale, Hawkshead, Red Screes,.
Skelwith Bridge, Tilberthwaite Ghyll.
Phyllitis scolopendrium (L.) Newm.— Aira Beck, Ambleside, Hutton Roof.
Asplenium adiantum-nigrum L.— Aira Beck, Tilberthwaite Ghyll.
A. ruta-muraria L. — Colwith, Dale Park, Deepdale, Glencoyne, Hutton Roof,.
Red Screes, Tilberthwaite Ghyll
A, trichomanes L. — Aira Beck, Ambleside, Dale Park, Deepdale, Hawkshead,.
Skelwith Bridge, Tilberthwaite Ghyll.
A. viride Huds.— Hutton Roof, Red Screes, Tilberthwaite Ghyll.
Ceterach officinarum DC. — 1 mile S. of Skelwith Bridge.
Athyrium filix-femina (L.) Roth — in all areas.
Cystopteris fragilis (L.) Bernh.— Aira Beck, Hawkshead, Hutton Roof, Red Screes,.
Tilberthwaite Ghyll.
Dryopteris filix-mas (L.) Schott — in all areas.
D. borreri Newm.— in all areas, EXCEPT Deepdale and Red Screes.
29
D. abbreviata (DC.) Newm.— Colwith, Red Screes, Tilberthwaite Ghyll.
D. villarii (Bellardi) Woynar — Hutton Roof.
D. lanceolatocristata (Hoffm.) Alston — Dale Park, Hawkshead, Red Screes,
Skelwith Bridge.
D. dilatata (Hoffm.) A. Gray — in all areas, EXCEPT Tilberthwaite Ghyll.
Polystichum aculeatum (L.) Roth — Aira Beck, Dale Park, Hutton Roof, Tilberth-
waite Ghyll.
Thelypteris limbosperma (All.) H. P. Fuchs — Aira Bedk, Dale Park, Deepdale,
Hawkshead, Red Screes, Skelwith Bridge, Tilberthwaite Ghyll.
fs peers (L.) Slosson — Aira Beck, Ambleside, Dale Park, Deepdale, Hawks-
head, Red Screes, Skelwith Bridge, Tilberthwaite Ghyll.
T. dryopteris (L.) Slosson — Aira Beck, Dale Park, Red Screes, Skelwith Bridge,
Tilberthwaite Ghyll.
T. robertiana (Hoffm.) Slosson.— Hutton Roof.
gage tcmer vulgare L. — Ambleside, Dale Park, Hawkshead, Hutton Roof,
elwith Bridge, Tilberthwaite Ghyll.
The vice-county numbers and grid-references of the localities quoted
are as follows: Aira Beck, v.c.70, 353/4020; Ambleside, v.c.69, 35/3704;
Colwith, v.c.69, 35/3303; Dale Park Beck, v.c.69, 34/3592; Deepdale, v.c.69,
35/3913; Glencoyne, v.c.70, 35/3818; Hawkshead, v.c.69, 34/3598; Hutton
Roof, v.c.69, 34/5578; Red Screes, v.c.69, 35/3909; Skelwith Bridge, v.c.69,
35/3403; Tilberthwaite Ghyll, v.c.69, 34/3000.
Once again varieties found were very few, and not of great interest.
Within an hour of our arrival at Ambleside, many crested Lady-ferns were
seen growing on walls and in the roadside ditches. Subsequent observation
in the town confirmed the fact that this type of variation is very common,
due no doubt to the spores that scatter far and wide from the many similar
varieties growing in gardens. It was noted however that the variation never
exceeded terminal crests and crested pinnae — the pinnules were always
normal. A few similar finds were made at Ullswater and in Dale Park.
Two crested Male Ferns were noted by the writer, but again in circumstances
Park some very large-growing Beech Ferns with crenate pinnules were found.
On a wall at Ullswater a colony of Asplenium ruta-muraria contained many
plants with large rotund pinnules, and some of these were collected. The
multifid Hart’s-tongues found at Aira complete a very meagre list of finds.
Evening discussions added to the interest of the Excursion which was
very much enjoyed by all and came to an end all too soon.
J. W. DYCE
30
THE SECRETARY’S REPORT
The past year has seen many changes in the Society. We have welcomed
our new President into office, but have had to accept with great regret the
resignation of our Secretary and Editor, the Rev. E. A. Elliot, a step forced
on him by ill-health. The award to him of the Stansfield Memorial Medal
by the Society is an expression of our thanks for his invaluable services in
the post-war years.
With this Gazette we introduce our new Editor, Mr. A. C. Jerm
who has been appointed to fill the vacancy at the British Museum (Natural
History) created by the death of Mr. A. H. G. Alston. Mr. J. W. Dyce has
been elected to the joint post of Secretary and Treasurer.
The Committee during the year discussed suggestions for developing
the usefulness of the Society and plans for the future of the Gazette which
is to be published annually in December. We are widening our scope to
include not only British pteridophytes but world ferns and their allies,
and our Aims have been amended accordingly. Our Constitution is in need
of. revision and a new one is being drawn up and will be submitted for
approaval at the next A.G.M. after copies have been sent to all members.
More meetings are planned — indoor and outdoor. It is proposed to
form two areas, one in the South and one in the North and day or weekend
meetings can be arranged at convenient centres in both areas. Details
will be advised to members in due course.
A fern exhibit at the Royal Horticultural Society’s show on 11/12th
August was proposed, but unfortunately had to be cancelled because sufficient
ferns up to the required standard were unavailable due to the dry hot summer.
Next year we hope to put up an exhibit on the date of the Alpine Garden
Society’s show in September. Full information will be sent out later but in the
meantime will members willing to participate please give some thought to
the selection of suitable plants and the attention necessary to achieve the
desired high standard.
Another service to members, the correct naming of fronds sent to the
Officers, requires the formation of a pressed collection of the varieties in
existence today. This is discussed on another page and I hope all members in
possession of good varieties will co-operate.
The Financial Statement for the year ended 30th June, 1959, is appended.
It requires no comment — BUT, increased activities mean increased expendi
ture and we need more members. Prompt payment of subscriptions also
helps, so will those members who have not yet found the time to put 10s.
into an envelope and address it to me, please do so right away. Thank you.
31
FINANCIAL STATEMENT at 30th JUNE 1959
as presented and accepted at the A.G.M. on 26th September 1959
1958 Es. od.
30th June
To Balance Sr FT
ubscriptions 58.0.0
Donations s 29k 3 ey
le of uted a9. Ss
£121 3 10 310
59
30th June
To Balance
(signed)
Gazette Vol. 8, No. 10
Subscription R.H.S.
Postages and Incidental Expense
Secretary .
Treasurer
Balance
87
iat J. W. DYCE, Hon. Treasur
PETER TEMPLE, “sat po
NEWS OF MEMBERS
While at Ambleside, one of our party, Mr.
well for many more years to come.
£121
T. A. Dyer, paid a visit to More-
cambe and conveyed our greetings to Mr. Robert Whiteside and Mr. J. D. Dixon.
Mr. Whiteside, aged 93, was one of the group of Lakeland fern hunters who founded
our Society in 1891. He and Mr. Dixon, aged 83, still retain a keen interest in the
Society and enjoyed a chat with Mr. Dyer on our activities. We wish them both
ADDITIONS TO MEMBERSHIP LIST
Bioom, J., Garden News, 62, High Street, Stamford, Lincs.
Burnett, L., 3, Beech Hill Road, Sutton Coldfield, Warwicks.
CARTWRIGHT, R. F., 143, Pitmansfield, Harlow, Essex.
Fucns, H. P., pH.D., Pilgerstrasse 6, Basle, Switzerland.
Jackson, J. F., Knotts View, Stonethwaite, Borrowdale, Keswick, Cumberland.
Jermy, A. C., , Department of Botany, British Museum (Natural History),
Cromwell ad. London, S.W.7
Mor ey, R., Moneyash Close, Chasen: Derby.
Taytor, H., 8, Cotterill Road, Surbiton, Surrey.
Taytor, P. G., The Herbarium, Royal Botanic Garden, Kew, Richmond, Surrey.
We have a very comprehensive
collection of
BRITISH FERNS & THEIR VARIETIES |
We also specialise in
Alpine and Herbaceous Plants
Dwarf Trees, Flowering Shrubs
House Plants, Water Lilies
and Greenhouse Plants.
CATALOGUE ON REQUEST
REGINALD KAYE LTD.
SILVERDALE, LANCASHIRE
We grow
HARDY FERNS
British and exotic, of garden value
We should like to send you our catalogue
and we are always ready to consider
buying good garden varieties which
we do not already possess.
TAYLOR’S NURSERY
BRACKNELL, BERKS
THE
BRITISH FERNGAZETTE
VOL. 9 PART 1! DECEMBER, 1959
Contents
Some notes on vegetative characters of ferns ... R. E. HOLTTUM 2
A preliminary census list of British pteridophytes A.C. JERMY o
The geographical affinities of the New Zealand eee cag 3 eis a
Cultivation of ferns under glass BRUTY 23
Shorter Notes
Woodsia sila (Bolton) Gray . P. GREENFIELD 25
Herbarium specimens of ferns aad their a
A. CRABBE 26
The British Pteridological Society:
Annual Excursion 1959 oe
Secretary’s and Treasurer’s Repodi
News of Members: additions to eacberhio list
ee ee
Courier Co. Ltd., Tunbridge Wells
Eb
THE
BRITISH FERN
GAZETTE
VOLUME NINE PART TWO ~
APRIL 1961
EDITED BY A. ©. . JERMY
% 4
THE BRITISH
Wreecom, Bored ERIDOLOGICAL SOCIETY
SEP 19 1966
BRITISH PTERIDOLOGICAL SOCIETY
Officers and Committee 1960-61
PRESIDENT
Professor R. E. Holttum, M.A., SC.D., F.L.S.
VICE-PRESIDENTS
. H. Bolton, Esq.
DrcS. poe M.B., CH.B., F.R.C.P. (Edin.)
Greenfield, Esq.
Reginald Kaye, Esq.
Dr. T. D. V. Swinscow, M.B., B.S., F.L.S.
R. Whiteside, Esq.
SECRETARY AND TREASURER
J. W. Dyce, Esq.
“ Hilltop,” 46, Sedley Rise,
Loughton, Essex
EDITOR
A.
Gazettes ... nae £442 Council for Nature 2..0..0
ks a 13 0 Stansfie a Medal to Rev. ‘E. A:
Advertisements in| ‘Gazette se a 0:0 2:10:20
Statione nosis oe
Advertisement in A.G.S. “Bulletin 110 0
Postages and —- eer ae
Editor 110 0
Balance axe se oe $2: 3: §
£157 -0 7 $157) OF
1960
30th June
To Balance
o8
(signed) J. W. DYCE, Hon. Treasurer
(signed) PETER TEMPLE, Hon. Auditor
REQUESTS FROM MEMBERS
ur member, Mr. C. J. Taylor, 211 Arundel vee Landport, Portsmouth, Hants,
is anxious to obtain spores of the Mexican Tree “apa ai schiedei) and would
be pleased to hear from any one knowing a sca sou
Mr. Neill D. Hall, 1225E., 95 Street, Seattle 15, Washington, U.S.A., would like to
hear from any members interested in exchanging fe fern spores
LIST OF MEMBERS
HONORARY MEMBERS
MANTON, Professor Irene, Ph.D., F.R.S.
Depart ment of Botan y, The University, Leeds 2.
PRESTON, F. G., M.A., VMH A.H.R.HLS.,
92. Hinton Way, Great Skeitond, Cambridge.
RAMS r. PL
c/o British — Vers History), Cromwell Road, London, S.W.7.
WARBURG, Dr. E. F.,
South Hayes, disor Hill, Cad.
61
SUBSCRIBING MEMBERS
ALEXANDER, T. M. W., The Rose Garden, West Huntspill, Nr. rar eo ete
ALLEN, Mrs. B. E. G., P.O. Box 107, Ipoh, Perak, Federation of Ma
ALLEN, Walter S., 168, Cedar Hill Avenue, Belleville, New Jersey, US. A.
ALLISON, Dr. B. R., M.D., 26, Ives Road, Hewlett, Long Island, New York, U.S.A.
ASHWooD, Mrs. F. M. W., 356, Newbold Road, Newbold-on-Avon, Rugby,
BALLARD, F., a Herbarium, Royal Botanic Gardens, Kew, Surrey.
BANGERTER, B., Department of Botany, British Museum (Natural History),
Cr jie Road, London, S.W.7.
BARLTROP, Seg 23; Brook Street, Nelson, New Zealand.
BASSN s. M.C.L., Laund, Hesketh Lane, Tarleton, Nr. Preston, Lancs.
BELCHER, D. z ma 1h ee 71, Chaffinch Road, Beckenham, Kent.
BENEDICT, Dr. Ralph. C., Ph.D., 185, Hall Street, Brooklyn 5, New York, U.S.A.
BLOOM, John, “‘ Garden News ” , 62, High Street, Stamford, Lincs.
BOLTON, Thomas Henry, F RHS., Birdbrook, Halstead, Essex.
ahi ean! GARTEN UND BOTANISCHES MUSEUM, K6nigin-Luise Strasse, 6-8, Berlin-
a
BRIGHTMAN, F. H., B.Sc., 2, Red Oak Close, Orpington, Ken
BRITISH MUSEUM (NATURAL HISTORY), Cromwell Road, pace S.W.7.
oberg Noel, 102, Stamford Road, Birkdale, Southport, ag
BRUTY, H. J., 23, Dagmar Road, Kingston-upon-Thames, Surr
BUCHANAN, W. C., Douglas Bank, Drumchapel Road, hcieien. Dumbartonshire.
BURNETT, Leslie, 3, Beech Hill Road, Sutton Coldfield, Warwickshire
Us UNIVERSITY, Serials Dept., General Library, Berkeley 4, California,
CAPPER, V., 148, Claughton Avenue, Crewe, Cheshire.
CARTWRIGHT, R. F., 143, Pittmansfield, Harlow, Essex
COBB, pasties 25, East End Avenue, New York, 28.
COCHRAN, John, 30, High Glencairn Street, Kilmarnock, Ayrshire
Hon. T. C. A., 61, East Church Street, Buckie, Banffshire.
COULTHARD, Charles E., 143, Musters Road, West Bridgford, Nottingham
CRABBE, J. A. , Department of Botany, British Museum (Natural History), Cromwell
Road, London, -W.7
CRANE, Mrs. Fern W., 174, Somat Avenue, Summit, New Jersey, U.S.A.
DARTMOUTH COLLEGE, Baker Library, Hanover, New Hampshire, U.S.A.
PAVE James, M.B., Ch.B., F.R.C.P.Edin., Linton Muir, West Linton,
purrFy, E. A. J., 8, Birdwood Close, Selsdon, sop
pyce, James W., Hilltop, 46, Sedley Rise, Loughton,
pYeER, Mrs. E. M., Rockfield, Alrewas, Nr. Burton- crs, Staffs.
DYER, T. A., Rockfield, Alrewas, Nr. Burton-on-Trent, Staffs.
FISHER, C. H., M.A., Lower Drabbington, Thornbury, nigendbed Hereford.
FRENCH, J., 28, Salcot Crescent, New Addington, Croydon,
FUCHS-ECKERT, Dr. Hans P., Dr. phil. 2, Clematisiaan 6l, Wemuniat (ZH), The
Netherlands
GILmMourR, J. S. L., M.A., F.L.S., cima University Botanic Gardens, Cambridge.
GLASNEVIN BOTANIC GARDEN, Dublin,
GREENFIELD, H. F., 41, Brittain’s Lane, Sotedaiek Kent
GREENFIELD, Percy, cy, Beech Bank, 18, Stuart Road, Warlingham, Surrey.
HALL, Neill D., 1225,E. 95 Street, Seattle 15, Washington, U.S.A
HALL, Peter C., 6, Johns Close, Gorsewood Road, Hartley, Se Kent.
62
HALLIGEY, ne P., Officers’ Mess R.A.F., Swanton Morley, Dereham, Norfolk.
HART, D. J., B.Sc., 9, Ridge Road, Sutton, Surrey.
HAYHURST, ‘Fiernard. 91, Windermere Road, Freehold, Lancaster.
HEALEY, F. J., Lily Hill Nursery, Bracknell, Berks.
HEALEY, Mrs. J. R., Lily Hill Nursery, Bracknell, Berks.
HEWITT, A. H., 29, Balfour Crescent, Wolverhampton, Staffs.
HINCKS, W. D., D.Sc., 19, Whitefield, Heaton Norris, Stockport, Cheshire
HOLTTUM, Professor R. E., 80, Mortlake Road, Kew Gardens, Richmond, Surrey.
HOME, Miss H. M. Logan, Edrom Nurseries, rpia gees Berwickshire.
HULL UNIVERSITY, Dept. of Botany, Hull, Yor
IRONSIDE, E. T., The Orchard, 27, Slade Road, Hewtba: Mumbles, Swansea, Glam.
JERMY, A. B.Sc., F.L.S., Department of Botany, British Museum (Natural
eniens ‘Cromwell Road, London, S.W.7.
KAYE, Reginald, Waithman Nurseries, Silverdale, rpiiigsp Lancs.
KIPPAX, Miss Anne, 119, Rosehill Road, Burnley
KNOX, “Sir Alfred, K.C.B., C.M.G., Binfield House, Binfield, Bracknell, Berks.
KNOX, Mrs. Wm. C., 649, Isle of Palms, Fort Lauderdale, Florida, U.S.A.
LANGSTON, Ww. G;; 98, Tom Morris Drive, St. Andrews, Fife
LAWALREE, André, Directeur de Laboratoire au athe Botanique de LEtat, 3;
Avenue Van Elderen, Bruxelles 16.
Lovis, Dr. John D., B.Sc., Ph.D., Department of Botany, The oo Leeds 2.
MARCHANT, H. J., Cardiff RDI, Stratford, Taranaki, New Zea
MASHITER, John, Roseneath, Harewoo: Road, Skipton, Salida
MIDWEST INTER LIBRARY CENTER (C.A.), 5721, Cottage Grove Avenue, Chicago
37, Illinois, U.S.A.
ait a REDHEAD, E., T.D., M.A., F.L.S., 7, Ashley Gardens, Petersham, Richmond,
MORLEY, r V. , D’Arches, Stolford, Nr, Bridgwater, Somerset.
MORLEY, R. 4, Monyash Close, Chaddesden, Derby.
ae cy — Division of Ferns, Smithsonian Institution, Washington,
NATIONAL MUSEUM OF WALES, Department of Botany, Cardiff.
NEILSON, Mrs. J. G., 5, Lady Road, Edinburgh, 9.
NEW ZEALAND GOVERNMENT, National Library Service, Christchurch, New Zealand.
NEWBOULD, Mrs. J. P., B.Sc., Botany Dept., University College, Gower Street,
London, W.C.1.
OWEN, Owen, Orient Cottage, revo yacht
PAYNE, R. M., 8, Hill Top, Loughton, E
PERRY'S HARDY PLANT FARM, Holtwhites Hill, Enfield, Middx.
PICHI-SERMOLLI, Professor Dr. R. E. G., Istituto Botanico “‘ Hanbury ’’, Universita
degli Studi, Via Balbi 5, Genova, Italy.
Sean R. S. W., L.A.M.T.P.L, J.P., 127, Park Road, Chiswick, London, W.4.
RA mF, Wiggin Hill, St. Ives, Hants.
nicleaey R. L., 1030, Stratford Drive, Encinitas, on ya U.S.A.
ROBINSON, N., 27, Yewlands Drive, Fulwood, Pres see Ss.
SAWFORD ., 28, Ermine Street, Huntingdon, sein
— Wilfred, 18, Crakegarth, Dalston, Carlisle, Canibedieid:
w, Rev. C. E., Holy Trinity Vicarage, Waterhead, Oldham, Lancs.
se; LATER, W. P., 100, Ivy Park Road, Sheffield.
SMITH, Ewart G., 147-151, High Street, Christchurch, New Zealand.
ee P. H. B. Otway, V.R.D., D.Phil., M.P., Colebrook House, Winchester,
ants.
63
‘SOUTH LONDON BOTANICAL INSTITUTE, 323, Norwood Road, London, 8.E.24
ST. CLAIR-MORFORD, E., c/o Hongkong & Shanghai Bank, 22, Place Vendéme, Paris.
STERN, Sir Frederick, M.C., O.B.E., V.M.H., F.L.S., Highdown, Goring-by-Sea,
Worthing, Sussex.
Co ie — D.V., M.B., B.S., F.L.S., Everley, London Road, Kneb-
orth,
TAyLOR. C.J,. ane Arundel Street, a Portsmouth, Hants.
TAYLOR, H., g, Cotterill Road, Surbiton,
TAYLOR, , The Herbarium, Royal pteeed Gardens, Kew, Surrey.
TEMPLE, Max, 3 Woodlands Close, Hendon, London, N.W.
TEMPLE, Peter, L.R.I.B.A., M.Inst.R.A., 42, Holly Park, Finchley London, N.3.
TETLEY, Miss M. I., High Gable, Grange-over-San
THOMPSON, Donald H., 180, Wakefield Road, Lightcliffe, Half, Yorks.
THOMPSON, E., St. Davids, Lydwell Road, Torquay, Dev
TINNE, santa A. M., Lochwood Cottage, ee Gartcosh, By ‘Giese
TROTTER, R. D., Leith Vale, Ockley, Surr
tle ‘Dr Rolla , Gray Herbarium, fod University, 22, Divinity Avenue,
Cambridge 38, Mass., S.A.
VERSITY COLLEGE OF WALES, Botany Dept., Penglais, Aberystwyth, Cardiganshire.
ba yk a Pierre, Curator, Musée Botanique Cantonal, Palais de Rumine,
Lausanne, Switzerland.
WAINWRIGHT, H., Rockwood, Buntingsdale Road, Market Drayton, Shropshire.
WALKER, Dr. S., Sub- Dept. of Genetics, The University, Liverpool.
WALKER, Dr. Trevor G., Botany Dept., ge id s bt Reason tee 2.
WATIS. 1774. Starvecrow Place, Tonbridge,
WEBSTER, W. S., The Pharmacy, 7, Station aay acta Nr. Keighley, Yorks.
WHITE, Miss D. M., Birch Cottage, Waenfawr, Nr. i N. Wales
“WRIGHT, W., 145, Monmouth Road, Edmonton, London, N.9
We have a very comprehensive
collection of
BRITISH FERNS & THEIR VARIETIES
We also specialise in
Alpine and Herbaceous Plants
Dwarf Trees, Flowering Shrubs
House Plants, Water Lilies
and Greenhouse Plants.
CATALOGUE ON REQUEST
REGINALD KAYE LTD.
SILVERDALE, LANCASHIRE
We grow
HARDY FERNS
British and exotic, of garden value
We should like to send you our catalogue
and we are always ready to consider
buying good garden varieties which
we do not already possess.
TAYLOR’S NURSERY
BRACKNELL, BERKS
THE
BRITISH FERN GAZETTE
NEW SERIES
VOL. 9 PART 2 APRIL, 1961
Contents
The ferns of the Channel Islands fee ... D. MCCLINTOCK 34
Is it a hybrid or a species? ae — es F. W. CRANE 37
The genus Cheilanthes Swartz and its European species H.P. FUCHS 38
The Southport Spring Flower Show _... ss pa eee.
Shorter Notes:
The rediscovery of Asplenium x murbeckii Dérfl. F. JACKSON 49
Transplanting ferns ... - 3 ae J. W. DYCE ‘50
Polystichum aculeatum ans Bevis Jc W. DYCE 50
Herbarium specimens of ferns. II ... = 5A. CRABRE = oF
Book Notes ae se tee ae es a ea
The British Pteridological Society:
Annual Excursion 1960 54
Reports of Meetings 1960 ... 57
Obituaries Le 58
Secretary’s and Treasurer’s ‘Repad 60
Requests from members =) my we ee
List of Members __... Mop os oF ae de eae
Courier Co. Ltd., Tunbridge Wells
Pat
BRITISH FERN
GAZETIG
VOLUME NINE PART THREE =
MAY 1962
EDITED BY A. £. JERMY
Whssour! BoTANicee
SEP 19 1966
GARDEN LIBRARY
THE BRITISH
p, q PTERIDOLOGICAL SOCIETY
4
BRITISH PTERIDOLOGICAL SOCIETY
Officers and Committee 1961-62
PRESIDENT
Professor R. E. Holttum, M.A., $C.D., F.L.S.
VICE-PRESIDENTS
T. H. Bolton, Esq.
Dr. J. Davidson, M.B., CH.B., F.R.C.P. (Edin.)
¥. Greenfield, Esq.
Reginald Kaye, Esq.
Dr. T. D. V. Swinscow, M.B., B.S., F.L.S.
SECRETARY AND TREASURER
J. W. Dyce, Esq.
ey ~— 46, Sedley Rise,
Loughton, Essex
EDITOR
A. C. Jermy, B.SC., F.L.S.
Cromwell Road, London, S.W.7.
COMMITTEE
R, F. Cartwright, J. A. Crabbe, T. A. Dyer,
B, Hayhurst, Mrs. J. R. Healey, F. J. Healey,
F. Jackson, R. M. Payne, R. S, W. Pollard, N. Robinson
AUDITOR
Peter Temple, L.R.1.B.A., M.INST.R.A.
The SOCIETY ie open. to ofl intetosted fn Katy wity Ss FIRS sal EES AEE
of S$
upon paying an annual
SIXPENCE. pment rane aEbigit tet omaha mae OE
THE
BRITISH FERN GAZETTE
VOL. 9 MAY 1962 PART 3
THE POLYPODIUM VULGARE COMPLEX
M. G. SHIVAS (Mrs. TREVOR WALKER) _ Botany Dept., King’s College, Newcastle
Polypodium vulgare sensu lato is one of the most familiar of British ferns. It
as been described and illustrated in many of our old herbals and was locally
prized for its medicinal properties in the treatment of various ailments.
Britten (1882) for example quotes its cure in several parts of Great Britain
and Europe as a purgative and as a cure for various lung disorders, colds,
whooping cough etc.
In more recent times P. vulgare has been of considerable interest to fern
collectors in this country as a source of morphological variations of horti-
cultural and decorative value. Many forms and varieties have been named,
most of which have been based on some unusual feature such as the presence
of very deep serrations, pronounced auricles etc. An extensive collection of
examples of this kind may be seen in Thomas Moore’s herbarium at Kew.
Such forms are genetically monstrous types and are of little or no biological
importance.
During the past few years P. vulgare has been studied cytologically and
experimentally (Manton, 1950; Shivas, 1961 a and b) and it has been shown
that it is not a single entity but is in fact a complex of three different species.
All three members of the complex differ from one another in chromosome
hybrids between them are not fertile, hence they are separated from one
another by sterility barriers. All these features, taken in conjunction with the
results of experimental work, have led to the recognition of the members of
the complex as distinct species. These have been named P. australe Fée,
P. vulgare L. and P. interjectum Shivas.
Polypodium australe is a diploid species having 37 gametophytic (i.e.,
prothallial) chromosomes and 74 in the sporophyte (i.e., the dominant stage
of the life cycle of the fern). It is the most southerly member of the complex
in its geographical distribution, being found mainly in southern Europe and
the Mediterranean region. It does, however, extend into parts of England,
The BRITISH FERN GAZETTE Vol. 9 Part 2 (pp. 33-64) was published 24th April, 1961.
65
Wales and Ireland — this representing its most northerly limits. In Britain it
frequently occurs in limestone habitats, as is the case with many other plants
near the limits of their geographical range.
P.vulgare sensu stricto is a tetraploid species with 74 gametophytic and
148 sporophytic chromosomes. It has the widest European distribution of
all the members of the complex and is undoubtedly the plant that Linnaeus
himself knew since it is the commonest (and most probably the 2 ew
Polypodium in Sweden. It is both common and widespread in Britain
P. interjectum is a hexaploid with a gametophytic chromosome number
of 111 and a sporophytic number of 222. Its European range is intermediate
between those of the other two species. It is common in Britain and appears
to favour rather damper habitats, often showing luxuriant growth under wet
conditions.
By making crosses involving all possible combinations of these species
and then analysing the behaviour of the chromosomes at meiosis in the
hybrids one can make deductions concerning the interrelationships of the
parental species. Such techniques have shown that P. vulgare and P. australe
are not very closely related, despite their superficial similarity. In the past
ese two species have crossed, however, to give rise to a sterile triploid
hybrid which completely fails to pair its chromosomes at meiosis. The
doubling of the chromosome complement of such a hybrid leads to the
formation of a fertile hexaploid in which meiosis is normal. This hexaploid
is P. interjectum.
In North America there are several species of Polypodium, some of
which belong to the P. virginianum complex, and others which are closer
to the P, vulgare in morphology. It is now known (from the type of experiment
described above) that the European P. vulgare sensu stricto has at least one,
and most probably two, of these American species in its parentage. Thus one
has the rather curious situation that the European P. vulgare is derived from
American diploid ancestors, despite that fact that there is a diploid species
in Europe in the form of P. australe. Since P. interjectum is a hexaploid
derived from P. australe and P. vulgare it follows that this species has a mixed
European and American ancestry
P. vulgare is the easiest member of the complex to identify in the field,
differing markedly from the other two species in frond and soral characters.
The fronds of this species are lanceolate, bearing pinnae which are all more
or less of equal length from the base to near the tip of the frond. The lamina
is completely flat. The sori are round and the sporangia mature in the
summer.
Both P. australe and P. interjectum have sori which are oval in outline
when young, but the sporangia of P. interjectum mature in summer and
autumn whilst those of P. australe mature in the autumn or the following
spring.
66
Frond pee [x 4] and sporangia [x 60] of Polypodium.
Fic. 1: rale Fée (with paraphysis, 1b) ; Fic. 2: P. vulgare L. ;
Fic. a P. pers bt Shivas.
The fronds of P. australe are produced in the autumn and are broadly
ovate, almost triangular. In this species it is usually the second pair of
pinnae from the base which is the longest. In P. interjectum the new fronds
are produced in both summer and autumn and are ovate — the longest pair
of pinnae being the 4th, 5th or 6th from the base. In both species the lowest
pair of pinnae are markedly inflexed — a very conspicuous character in the
field but which, however, tends to be lost on herbarium specimens.
In addition to the macroscopic characters mentioned above which can
be used both in the field and in the herbarium there are several other features
which require the use of a microscope of low magnification. These features
provide very useful checks on identifications made in the field. :
Martens (1950) described the presence of branched paraphyses inter-
mixed with the sporangia of some European plants of Polypodium and it is
now known that these are diagnostic of P. australe, the other two species
lacking them (see fig 1b)
The number of the veigeaaten, thickened cells of the sporangial
annulus — the indurated cells — varies from one species to the next, as has
been noted by Futo (1905) and Nsdios (1950), and therefore provides a good
identification character. P. australe has an average of 4-7 indurated cells
in the annulus, P. vulgare has 10 - 14, whilst P. interjectum is intermediate
with 6- 10. In addition to this Mr. P. Benoit has drawn my attention to the
fact that the indurated cells of P. vulgare turn a much darker brown and are
more conspicuous than is the case in P. interjectum. The contrast in colour
tween the indurated cells and those of the rest of the sporangium is so
marked that the dark brown line of the annulus in P. vulgare is easily seen
with a x 10 lens or even with the naked eye. The reasons for this lies not only
in the greater number of indurated cells in P. vulgare but also in the fact that
after dehiscence the sporangia gape open and expose the whole of the annuli
to view. This is in contrast to the sporangia of P. interjectum, which not only
have fewer indurated cells per annulus but also do not open widely at
dehiscence and hence part of the annulus is concealed. I am very grateful to
Mr. Benoit for these observations and for allowing me to quote him.
Rhizome scales are often used as specific characters in ferns and an
examination of these have been made in each of these three species. The
variation in shape of the scales around the base of the youngest frond has been
found to be so wide that without a statistical analysis this character would
not be useful. However, there is a strong tendency for P. australe to have a
long tapering, almost linear, scale whereas P. vulgare has a narrow triangular
scale drawn out to an accuminate tip which is rarely longer than the triangular
area itself. The scales of P. interjectum are, as one might expect, intermediate
between the ae agers _ ho drawn out tip being at least 13 times as long 4! as
the triangular port J
character on sessile 5 olendis and for that reason is included here.
68
Finally, spore size also varies, being on average greater than 74, in
length in P. interjectum, less than 74, in P. australe and less than 70, in
P. vulgare.
One final advantage in the use of a microscope should be noted and
this is in the diagnosis of hybrids. Where any two species of Polypodium grow
together hybrids may be formed and all three possible types are known from
. vu
australe X P. interjectum and the pentaploid P. vulgare x P. interjectum.
These may be rather difficult to distinguish from the true species with certainty
as they combine the characters of the parents but all of them have irregular
meiosis and as a consequence produce spores which are irregular in shape
and size. A glance at mature spores of a suspected hybrid is sufficient to
settle any doubts as to its nature. Triploid and tetraploid hybrids are
infrequent because of the limited distribution of P. australe; the pentaploid
more frequently occurs in mixed populations of the parental species
These characters may be summarised in the form of two keys which
readers may find easier in practice to use than a single combined key.
KEY TO MACROSCOPIC CHARACTERS
1 Sori round; fronds lanceolate, pinnae more or less equal in length to near the tip
mS 6 frond; lowest pair of pinnae not projecting forward; new fronds
i pe
than otha cells of the sporangium and very conspicuous : . vulgare
Sori oval; lowest pair of pinnae projecting forward See 2
> ae ovate, longest pair of pinnae being the 4th, Sth or 6th from the base;
w fr onds p roduced in summer and autumn P. interjectum
ae broadly ovate or triangular, longest pair of pinnae being the xe’ from the
base; new fronds produced in the autumn ustrale
KEY TO MICROSCOPIC CHARACTERS
1 Paraphyses present: number of indurated cells eel ca! 4-7; Baja spo
length less than 74 ustrale
Paraphyses absent .. ae
2 Number of indurated cells averaging 10 - 14; mean spore length hss than React
Number of indurated annular cells averaing 7-10; mean og length greater
than 74, . P. interjectum
At present an attempt is being made to record the occurrence of the
three species of Polypodium in the British Isles and a table is given at the end
of this paper summarizing our present dts by listing the vice-counties
in which each of the species has been foun
As is the case with most common species of plants, however, we tend to
have far less information concerning the detailed distribution and ecological
69
requirements of Polypodium than is the case for very rare plants. It is hoped
that the descriptions and keys given above will be helpful to readers and
encourage them to collect specimens and compile appropriate information
on habitat etc., in an attempt to increase our knowledge of those ferns.
My grateful thanks are due to Mr. A. C. Jermy in preparing the
distribution table and illustrations.
LITERATURE CITED
BRITTEN, J., 1882. European ferns. Cassell. London.
cial 4 son: Se Polypodium vulgare L. and Polypodium vulgare subsp. serratum Willd. Hedwigia
MANTON, I., 1950. Probl f cytol d evolution in the Pteridophyta. Cambridge.
MARTENS, P., 1950. Les paraphyses de Polypodium vulgare et la sous-éspéce serratum. Bull.Soc.
Bot. Belg. $2:225 - 261. . 34
peptic M. G., 1961a. Contributions to the cytology and ected of species of Polypodium in
urope and ‘America: I Cytology. J.Linn.Soc.(Bot.)58 :13 -
1961b. Contributions to the cytology and taxonomy a species of Polypodium in Europe
and America: II Taxonomy. J.Linn.Soc.(Bot.)58:27 - 38.
DISTRIBUTION OF THE THREE POLYPODIUM SPECIES
IN THE BRITISH ISLES
The numbers used re the Watsonian vice-counties as adapted for A
Preliminary Census List te British Pteridophytes [Brit. Fern Gazette 9(1) page: 15
POLYPODIUM AUSTRALE Fée: 2, 3, 5, 6, 9, 35, 41, 45—47, 69; H3, 4, 7, 9, 12, 28, 29.
POLYPODIUM VULGARE Ls. i—6, 8-12, 14—23, 24, 25, 27—29, 31—33, 33,
x | 42, 45, 4750, re 54, 57, 58, 61, 64—67, 69—72, 74, 78, 80, 81, 83,
87—93, 9%6, 97, 100—1 102, 104-106, 108, 110—112; HiI—4, 16, 29, 33, 35.
CTUM Shivas: 0, 1—6, 9—11, 13—18, 22—25, 27—29, 32, 34,
"35, "38, 4147, 49, 50 50, 52, 58, 64, 67, 69—T71, 83, 92, 98, 99, 101, 102, 104, 108,
110; H2—6, 8, 12, 23.
N.B. The apparent scarcity of these species in oo is due to the fact that little
Irish aggregate material has been re-determined. will
be welcomed from all areas either on a Sav eoeity ue 10-km. grid square basis.
A REVISED LIST OF PTERIDOPHYTES FROM ST. KITTS
G. R. PROCTOR
Kingston, Jamaica, W.1I.
The Lesser Antillean island of St. Kitts (St. Christopher), one of the verdant
chain of tropical volcanic peaks that rim the Caribbean Sea along its
eastern end, has a long but intermittent history of botanical investigation.
From the time of Sir Hans Sloane, who collected the first ferns there in 1687,
relatively few botanists have visited St. Kitts, and these for the most part at
long intervals. It is not surprising, therefore, that today a collector can still
make significant contributions to our knowledge of the flora, in spite of the
small area involved.
Until the present century, there was little systematic search for ferns.
Until recently, our knowledge was chiefly based on the collections made by
N. L. Britton & J. F. Cowell in 1901, and H. E. Box in 1931-32. The
present writer was able to make several visits to the island in 1958 and 1959,
during the course of field work! connected with the preparation of a “ Flora
of the Lesser Antilles,” for which the volume on ferns is nearly ready for
publication.
In 1937 (Journ. Bot. 75:241-258 t. 6/2), H. E. Box and the late A. H. G.
Alston published the first comprehensive list of the pteridophytes known
from St. Kitts. A total of 97 species were enumerated. Most of these had
been collected by Mr. Box (whose specimens are deposited at the British
Museum), but a few other records of earlier collectors were also included.
The present writer’s work has resulted in the addition of 32 species not
specifically listed or authenticated before, including two that have proven
new to science. It therefore seems desirable at this time to present a new
augmented St. Kitts’ list, not only for the sake of completeness, but also to
record peony — changes that will be adopted for “* Flora of the
Lesser Anti
The present eer cannot presume to improve on Mr. Box’s excellent
account of the St. Kitts’ vegetation in relation to the fern flora, but a few
minor discrepancies of detail which appear therein can be mentioned for the
Sake of accuracy
Firstly, there seems to have been inadequate liaison between Box’s
descriptive section and Alston’s list, as several names of ferns mentioned in
the former either do not appear at all in the latter, or else in the form of
synonyms. Anemia adiantifolia, Diplazium striatum, Dryopteris megalodus,
and Hymenophyllum hirtellum are examples of the first category; Dryopteris
mollis (=D. quadrangularis), Diplazium arboreum (=D. cristatum), and
Olfersia cervina (=Polybotrya cervina) are examples of the second. It is
difficult to understand how these were editorially passed without correction.
* Supported by National Science Foundation Grant No. G 4441, in collaboration with Dr. R. A.
Howard, Arnold Arboretum of Harvard University.
71
In addition, there are several references to a palm called “ ? Acrista
monticola”’ which apparently allude to the well-known Euterpe globosa
Gaertn., a common species of the moist forests at middle to higher elevations.
Finally, the plant called Columnea hirsuta cannot be that species, which
is endemic to Jamaica. No Columnea is in fact recorded for St. Kitts; Box’s
plant was probably the common and striking Alloplectus cristatus (L.)Mart.
var. brevicalyx Morton, which resembles a Columnea in habit
A further descriptive point is that following a small earthquake during
a period of heavy rains in 1960, a great landslide occurred in the crater below
Mt. Misery; this completely filled up the lake which formerly occupied part
of the floor of the crater. The vegetation inside the crater has therefore
suffered considerable destruction and change since so ably described by Mr.
Box.
In the present list, the erect is given in accordance with the
treatment in the forthcoming ‘“‘ Flora of the Lesser Antilles,” and therefore
indicates the writer’s present views, isda on the names and delimitation
of genera. Within the families, the genera and species are listed i ical
order. In each instance, the accepted name is given first in small capitals
(new names or combinations in bold face); if the name was not the same in
Alston’s list, the one he used follows parenthetically in italics. Box’s localities
and numbers are then cited (or others if any), followed by those of the
the British Museum (Natural History) for his help in preparing the Latin
description, and to Dr. K. U. Kramer and Mr. Wessels Boer of the State
University of Utrecht for several useful comments and determinations.
© specimens collected by the writer are deposited in duplicate at the
Harvard University Herbarium and the Institute of Jamaica. Further sets
have been sent to the State University of Utrecht and the U.S. National
Herbarium.
The total number of pteridophyte species now known from St. Kitts
is 129, all occurring in an area roughly fourteen miles long and five miles
wide, of which more than half, being cultivated with sugar cane, is nearly
devoid of ferns. With the exception of the peculiar Asplenium described here
as new, there appear to be no species endemic to this island, though several
are shared only with others of the Lesser Antilles.
PSILOTACEAE
PsiILOTUM NUDUM (L.)Griseb. Wingfield, Box 367; Wingfield Ravine, on
shaded vertical cliff of volcanic conglomerate, Proctor 19242.
LYCOPODIACEAE
LYCOPODIUM CERNUUM L. Masson (BM); floor of The Crater, Proctor 19238
(growing beside L. tortum rele perfectly distinct from it; Alston’s suggestion that
the two fine merely eco-variants will not hold).
72
*L. CLAVATUM L. Floor of The Crater, Proctor 19239; upper slopes and summit
of Mt. Misery, Proctor 19617.
L. DICHOTOMUM Jacq. Britton & Cowell 304 (not seen).
L. SIEBERIANUM Spring. Mt. Misery, Britton & Cowell 547 (not seen).
L. TAXIFOLIUM Sw. Olivee’s Mt., Box 401; Nine Turn Gut, Proctor 19569.
L. ToRTUM Sieber ex sya i & Lloyd. Rim of The Crater, Box 269, Proctor
{9217 ° Olivee’s Mt., Box 3
SELAGINELLACEAE
SELAGINELLA FLABELLATA (L.)Spring. Mt. Misery, Box 258; Molyneux Water
Source, Proctor 19278.
*S. PLANA (Desv. pr 3 Extensively naturalized on moist banks, Molyneux
Water Pat Proctor 19287.
S. ROTUNDIFOLIA Boring. Upper slopes of Mt. Misery, Box s.n
S. SUBSTIPITATA Spring. Mt. Misery, Box 292 and s.n.; - Olive’ s Mt., Box 394;
NW rim of The Crater, Prottoe 19496.
S. TENELLA (Beauv.)Spring. Nine Turn Gut, Box 35/; Wingfield Ravine,
Feoctar F 19241; Molyneux Water Source, Proctor 19277.
*MARATTIACEAE
*DANAEA Noposa (L.)J.E.Sm. Outer NW slopes of The Crater, Proctor 19535.
*SCHIZAEACEAE
*ANEMIA ADIANTIFOLIA (L.) Sw. Mentioned in the descriptive section (p. 246
Box & Alston’s paper, but not included in the annotated list. Phillips Level, on a
ruined overgrown wall, Proctor 19546.
GLEICHENIACEAE!
GLEICHENIA BIFIDA (Willd. pies ae MS ye sid bifida (Willd.) Maxon)
Edge of = _ Lake, Box 257; NW rim of The Crater, — 19222.
G. ATA (L.)S (Diranoteri facet (L.)Underw.). Mt. Misery,
Britton & Conch (not Seahes 290; NW rim of The ee pabbed 19224.
HYMENOPHYLLACEAE
*HYMENOPHYLLUM ELEGANS Spreng. NW rim of The Crater, Proctor 19515.
*H. rucomes (Sw.)Sw. NW rim of The Crater, Proctor 19512.
*H. HIRSUTUM (L.)Sw. var. HIRSUTUM (the typical form not listed by Alston).
Outer NW wl of The one EProeior Lae
7H. UTUM var. tum (Fée)Proc based o ae —
Fée, Mém. hee 12:118, 2. ; © f. 1, but te " 866) iy oon ling” um by
Alston). “* Near the crater,” Box 253; oute ter NW slo opes of The Crater, f bas 10523.
[H. hirtellum Sw. was listed by Box in his descriptive section (p. 249), but oh
ar on a Kitts, as there are authentic records from and Guadel ‘
tum Fée. Candle Walk Ridge, along track to Dodans (Dos d’Ans)
Pond,” proce te 19581.
1 The generi Sa eS a eer igs fae i nie s. wat jet 4(2) :257- 280
(1957), + : :
73
*H. LINEARE (Sw.)Sw. Vicinity of Dodans (Dos d’Ans) Pond, Verchild’s Mt.,
Proctor 19592.
*H. MACROTHECUM Fée. Candle Walk Ridge, Proctor 19571.
POLYANTHOS (Sw.)Sw. Olivee’s Mt., Box 392-A; NW rim of The Crater,
Proctor 19511.
*H. SIEBERI (C.B.Presl)v.d.Bosch. NW rim of The Crater, Proctor 19514.
TRICHOMANES ALATUM Sw. “ Near the crater,” Box 254; Molyneux Water
ary Proctor 19275,
Ridge above the Mescie) Box 271; outer NW slopes of The
Crates Poca. 19524; ive s Mt., Box 39].
. HYMENOP v.d.Bo ne Belmont, Britton & Cowell 410 (not cr
Wingfield — s icieces & Cowell 452 (not seen); outer NW slopes of The Crater.
Procto
ng KAPPLERIANUM Sturm. Wingfield or yo Proctor 19269. 1 am indebted to
Mr. Wessels Boer, at the State University of Utrecht, for this determination. This
species is similar to, and often confused w ay the Greater ~ illean 7. hookeri
CB Presi, tae which name the present collects was distri
i SII . & Grev. (Alston, like some other recent authors el this
eg - Tras Hie ; however, the plant was gees for a Dr. Kraus, who sent
specim s from iominica, and the original publication uses bat one 5").
Olives s yer Box 388; W Wingfield Ravine, Procor 19247.
T. MEMBRANACEUM L. Nine Turn Gut, Box 25]; interior slopes of The Crater,
Proctor 19229.
T. PoLypopioiwes L. Nine Turn Gut, Box 252.
*T. PUNCTATUM Poir. pants Water Source, Proctor 19276.
ANGUSTIFRONS (Fée) Wessels Boer (7. ” sais Sw.). Olivee’s Mt., Box 402;
outer NW slopes of The Crater, P tector 1952
T. RIGIDUM Sw. Ridge above the ae Box 280; NW rim of The Crater,
Proctor 79495.
T. TRIGONUM Desv. Ridge above the crater, Box 279; outer NW slopes of
The Chater: F Procior 19205; E spur of Mt. Misery, "Proctor 19613.
CYATHEACEAE!
GRANDIFOLIA (Willd.)Proctor. a cote" (Willd.)
Seek Ridge above the crater, Box 283; NW rim of The Crater, Proctor 19501.
Bo der ARBOREA (L.)J.E.Sm. Wingfield omeirehy Box ped Olivee’s Mt.,
x
C. A Willd. (Hemitelia muricata (Willd.)Fée). Above Belmont,
Box 281; Oliver’ s Me Box 398; Wingfield Ravine, Proctor 19642.
POLYPODIACEAE sens. lat.
vegan TENERUM Sw. Wingfield Ravine, Box 3
76.
APHYLLUM Humb. & Bonpl. ex Willd. Nine Turn Gut, Box 362;
Wingfield avid; Box B72.
A. VILLOsuM L. Canada Hills, Box 348.
ANETIUM CITRIFOLIUM (L.)Splitg. Nine Turn Gut, Box 260, 353; Wingfield
Ravine, Proctor 19258.
* Asp
LENIUM ABSCISSUM Willd. Wingfield re Proctor 19248.
A. AURITUM Sw. var. BIPINNATISECTUM Mett. hg ado the common variety of
the species in sg let Lesser Antilles, aed the typical form (var. auritum) does not
1 The generi pts recognized in this family follow those of Copeland, Gen. Fil: 94-99 (1947).
74
B ances listed as A. auritum [«] var. rigidum (Sw.) Hook. (1860),
It
butt this epithet } is antedated in
rate? the application of Mette
Nhe oa
Wingfield Ravine, Procior 19352.
Im-smithii Proctor, sp.noy.!
varietal status by on - Mettenius (1859). In order
rom his numerous
name.
leloupe is naiies: aaactell as lectotype. (Listed
Nine Turn Gut, Box 259; lower (E) slopes t
rater, Proctor 19537.
re)
ingfield Ravine, Proctor 19265.
zome erect or oblique, 5 - 6 mm. — clothed at apex with dusky-brown,
long, 0.5- 0.9 m
Sicheste mona = Itate scales
Fronds erect,
crenatedentate, the teeth chiefl
m interior slo
TYPE fro pes of The
a 19234, collected 22nd Februa
t the Bri tish Museum (Natural
2.5-4 m
m. long; stipes 8.5 - Ide em. ‘ine: minutely ee
rondie: puiciae above and n
1,2- 1.7 mm. broad at
the lowest acroscopic one 2-forked, the middle ones ag both sides 1-fork
toward the apex simple. Sori 6 - 11
hyaline, entire, hee the sporangia; vai a
irs, 2.5 - 5.5 m
a iso
Hale ceca -~ the Institute of Jamai
r the late Mr.
n-marginate be
mm. broad at base.
se. Veins oblique,
those
ag inframedial; indusium
Crater, St. Sica elev. 2100 - contd ft.,
x 1959, mn moist sha
ypes a
ledges. Holotype
t the Harvard University
aces Smith of Basseterre, St. Kitts, whose keen
N
interest in mountir-limbing and in natural history have substantially nirsponsire
th direct
to our know
ledge of his native isla
through observations and
pment and indirectly thr rough his hospitality to. and piatt toaed 1a th oF ae
natu
Coe crater he loved so much.
ms possible that A. malcolm-smithii may be a hybrid between A. abscissum
and A. petro soc both of w
abortive spores su
mediate nature of the new species
ich occur in St. Kitts.
The irregular pinnae and
ggest hybrid origin, and the following table illustrates the inter-
ween its possible parents
abscissum malcolm-smithii pseuderectum
scales: 1.5 mm. long 2.5-4 mm. to 7 mm. long.
Sti 7.5 - 20 cm. long 8.5- 14cm - 10 cm.
rachis: green-marginate cone green-winged green-winged.
pwardly :
blade: deltate- eens to linceclete a narrowly _ linear.
: eo ance-oblo
pinnae: sis 25 - 40 pai
margins: dou eng or triply obliquely agp poe obliquely crenate-den-
dentate-serrate = Nees 1.2 - 1.7 mm. soot , teeth 1.5-2.5mm
veins: 1-3-forked 2-forked at base, I- cate simple.
forkedin middle simple
: toward a
sori: to 9 mm. long 2.5 - 5.5 mm. 2-4 mm. long.
* Rhizo rect 1 , S<6 , paleis fusco-brunneis clathratis anguste
deltatis 2.5 -~4 ma eee basi 0. 5 ans mm, latis fo eo vestitum. Folia erecta, 30-45 cm.
longa; stipes 8.5 - 14 cm longus, basim versus minute gg alg , superne late ee
is tis viridi inato; lamina pinnata, + lata, anguste eae AtO-
oblonga vel anguste elliptico-oblo! 25 - 28 cm. longa, 6-11. oe . lata, yan |
angustata, acuminata; rhachis m viridi-alata; pr ome ard gae, oppositae
alternae, horizontales vel inferiores deflexae, trapezio-oblongae cm. longae, basim
Sessilem inaequilateralem versus 1- 1.6 cm. latae, acuminatae Peaudatae, margine
crenato-dentatae, rere basi 1.2 - 1.7 mm. latis; venae obliquae, infimo acroscopico copico bifurcato,
mediis ig me tthe sei unifurcatis, eis apicem versus simplicibus. Sori pike My inframediales,
5.5 mm. longi; indusia Seaman, integra, sporangia occultantia; sporae
75
*A. PSEUDERECTUM Hieron. Interior slopes of The Crater, Proctor 19231.
A. PUMILUM Sw. Canada Hills. Box 349.
A. SALICIFOLIUM L. Nine Turn Gut, Box 265; Wingfield Ravine, Proctor 19257.
BLECHNUM DIVERGENS (Kunze)Mett. (Struthiopteris pf Bars Broadh.).
Mt. inaty; Box 311; outer NW slopes of The Crater, Proctor
B. NESIOTICUM edie ec iy striata (Sw.) Broadh. i “Ridge above the
crater, Box neg NW rim of The Crater, Proctor 19223; Mt. Misery, B
B. oc NTALE 9 gpetiiette i Box 370; Wingfield, Box an Phillips
Level, Pibetie 19553.
*B. PLUMIERI (Desv.)Diels. NW rim of The Crater, Proctor 19507.
*B. UNDERWOODIANUM (Broadh.)C.Chr. E spur of Mt. Misery, Proctor 19611.
BOLBITIS CLADORRHIZANS (Maxon)Ching. Olivee’s Ravine, Box 381; Wingfield
Ravine, pone 19262.
CHEILANTHES MICROPHYLLA Sw. Canada Hills, Box 350.
DIUM SEMINUDUM (Willd.)Maxon. Mt. Misery, Box 31/2; Verchild’s
Mt. near Dodans Pond, Box 359; NW rim of The Crater, Me 19503, 19520.
ONALIS (Poir.)Ching site ge tog sme (Poir.)C.Chr.).
Wig Ravine. B Box 364; The Hole, West Farm, Pro
DIPLAZIUM Pay gc oie )Alston. a in ws desert re by
Box (p. 247) 4 nder the name D. arboreum. Nine Turn Gut, Box Wingfie: id
or Box 373, Proctor 7: 19261; interior slopes of The Eater, peo 19 923
IATUM (L.)C.B.Presl. Mentioned in the descriptive section he Box
(p. Ay, tat not listed “a Alston. Outer NW slopes of The Crater, Proctor 79198;
Win rc Ravine, Proctor 19263.
ELAPHOGLOSSUM APODUM (Kaulf.)Schott. NE slopes of Mt. Misery, Box 278;
outer NW slopes of The Crater, Pricer 19197.
19 te. BORYANUM (Fée)T. Moore. Outer NW slopes of The Crater, Proctor 19202,
E. crinitum (L.)Christ (Hymenodium crinitum (L.)Fée). NE slopes of Mt.
Misery, Box 274; outer NW slopes of The Crater, Proctor 19528.
E. pussa Underw. NE slopes of Mt. Misery, Box 275; NW rim of The Crater,
ieee 19218.
*E. HERMINIERI (Bory & Fée)T. Moore. Phillips Level, Proctor 19550.
*E. IMPRESSUM (Fée) T. Moore. NW rim of The Crater, Proctor 19210.
= LONGIFOLIUM (Jacq.)J.Sm. NW rim of The Crater, Proctor 19225.
TINICENSE (Desv.)T. Moore (E. underwoodianum Maxon). Olivee’s
Mt., a 306; Phillips Level, Proctor 19532.
*E. PLUMIERI T. Moore. NW rim of The Crater, Proctor 19503.
E, UNDULATUM (Willd.)T. Moore. Mt. ea Box 298.
GRAM asplenifolia (L.)Proctor, comb. noy., based on Polypodium aspleni-
folium L. 54 Sp. PL. 71084, excl. 8 (1753). This epties has often thoes confused with the
plant named Polypodium suspensum by Linnaeus, but the two are entirely distinct,
s ident t one who takes the tro o compare the Linnaean
descriptions and the Plumier plates on which they are ultimately based r present
species is the abundantly hairy one illustrated by (Tr.Fil.Amer. t. 102 - A)
under the name Aspleniu tius & villosum, an etiver copied as
his Polypodium asplenii folio villoso (Fil. 26, t. 7-f). Petiver’s copy 1s actual
Linna obtained his material from ebasse,
Martinique, which must be considered the type locality. The present oe ha
recent este this species at the same place in Martinique, where
common. St. a ee — the crater, Box 272; NW rim of The Crater, Proctor
19216; Cities Mt Box
76
"sg, eggersii (Baker)Proctor, comb. . based on Polypodium eggersii Baker
in Hook., Ic., Pl. t. 1671 (1886). ry oe ig related to G. asplenifolia, and might be
treated as a vari ety of it, gt ai that poe, it mai rage identity without inter-
gradation throughout its range. It is certainly not a mere juvenile state of G.
asplenifolia, as considered by Copeland (Phil. Journ. Sci. 84(4) :443.(1956)). NW
rim of The Crater, Proctor 19226.
G. MOLLISSIMA (Fée)Proctor cag cg mollissimum Fée). Mt. Misery,
Box 291; Candle Walk Ridge, Proctor 19577
*G. SERRICULA A et adeca pee Vicinity of Dodans (Dos d’Ans) Pond, Verchild’s
Mt. , Proctor 19593
G.s ATA (Sw) Sw. (Polypodium duale Maxon). Mt. Misery, Box 293 &
CA, pat ‘of Dodans ae d’Ans) Pond, Verchild’s Mt., Proctor 19594.
G. sa (L.)Pr Ves ae on Polypodium suspensum L.,
Sp. he “1084. (1754): Polypodiim ‘sabtieloriatie aulf.; Grammitis pee (Kaulf)
Proc Although it is perfectly clear that the glabrous species named P. suspensum
by ena (founded on Plumier, Tr.Fil.Amer. t. 87) ane be synonymous
Calebasse, Martinque, has produced Aeon more das one glabrous-fronded
species of this affinity, and until recently the writer was of the opinion that an
unusually large plant oe * P. flabelli agg viet could have the
ean species. A paper was actually n (but _ oh poe an = which this
view was expounded at some length, ee fio cwnthoonk me private mi
In December, 1960, an opportunity arose to meni nearly a week cuhating on
Morne [de la] Calebasse, and a special effort was made to chin turn the several
one of his ae which co reasonably be equated with this species is ‘* Polypodium
pendulum glabrum ” (Tr. Foug. t. 87, Halt haod figure), upon which Linnaeus much
later founded his | i ef en This seemed to be quite convincing, though perhaps
not conclusive, eviden
mewhat later, Mr. C. V. Morton (who sabe had read the writer’s
; lypodium um
the Bibliot Génerale, Muséum d’Histoire Naturelle at
ray Sichaal still not ee does give a somewhat clearer idea of the similarity
eforme; at least, it does not suggest any veces alternative.
pet me seems, therefore, to decide definitely that P. s um L. must reasonably
be assumed the same as the later P. jubaeforme Kaulf, “of which the type also came
from Martinque!) and it is seorcntie treat faces dingly.
St. Kitts: Verchild’s Mt., Box 356; E spur of Mt. Misery, Proctor 19614.
G. LIA (Jenm.)Proctor (Polypodium taenifolium Jenm.). Near the
crater, PP 2553 OTS MW cant , The Crater, Proctor 19227; Olivee’s Mt., Box 391.
TAXIFOLIA (L.)Proctor (P; taxifolium L.). Ridge above the crater,
G. T ) (Polypodium taxi
Box 270; outer NW slopes of The Crater, Proctor 19201.
77
it TRIFURCATA (L.)Copel. NW rim “ The Crater, Proctor 19513.
MIDICTYUM MARGINATUM (L.)Presl. Not yin by Box or Proctor; listed
by ales on the basis of Hook., Exot. Ferns t. 73 (1858). The presence of this
species in St. Rate is altogether probable; it has is collected recently as near as
adjacent Nevi e genus Hemidictyum seems doubtfully distinct from Diplazium.
HYPOLEPIS REPENS (L.)C. a Presl. Phillips Level, Box 264; Molyneux Water
Source, Proctor 19280.
*LIND: UADRANGULARIS Raddi, ssp. ANTILLENSIS Kramer. NW rim of
The Ornter, Proctor 19499,
LOMARIOPSIS SORBIFOLIA (L.)Fée (Stenochlaena sorbifolia 33, wp Nine
Turn Gut, ‘Box 261: Wi as Ravine, Proctor 19246; Olivee’s Mt., Box
Lonc L. (Anisosorus Beso: (L.)Underw. & Maxon). ae
Mt., res 382: Wingfield Ravitie: Proctor 1925
NNEPHROLEPIS BISERRATA (Sw. ch: Winget Ravine, Box 365.
N. RIVULARIS (Vahl)Mett. Olivee’s Mt., Box 392; E. spur of Mt. Misery,
Proctor 19608.
a Scares CALOMELANOS (L. YLink. Wingfield, Box 407; Hope Estate,
x
num r Antillean islands (St. Barthelemy, tigu edonda, an
Montserrat, in addition to St. Kitts) on the basis of specimens chiefly collected on
cliffs, old walls, and other exposed places ey r from typical P. calomelanos
c The
in havi shorter, more deltate pa tii which tend to be more fine
(or rather, the divisions smaller and denser Ja nd have the “basal basiscopic pinnules
TS mi expec O appear in plants of
under difficult conditions. It s be remembere at the latter is a prot
species capable of great be benigee it has a deceptive plasticity of form, offering
temptations which a e fern taxonomist u s ;
ically tg: oh co and Central bts
pastes oa ge aulf. Retired’ to by Box as Olfersia ieee
( p. 247). N of t thie mater Box 288; Nine Turn Gut, Box 361; Wingfield Ravin
spapyss i esis
OLEPIS Liebm. Trail from Belmont to the crater, Box 286;
Phillipe Level. Pr Price 19568.
P. AUREUM L. Box, sight record, not to be doubted.
P. LORICEUM L. Mt. Misery, Box 300; NW rim of The Crater, Proctor 19494.
19 pe Lycopopioies L. By side of Crater Lake, Box 285; Phillips Level, Proctor
P. PECTINATUM L. Nine Turn Gut, Box 267; Phillips Level, Proctor 19556.
P. PHYLLITIDIS L. ho ails Ravine, Box 385; Phillips Level, Proctor 19567.
P. PILOSELLOIDES from Belmont to the crater, Box 286, in part, not
seen by Anon (det. miaaoaye Phillie Level, " Proctor 19566.
P. poLypopiorpes (L.)Watt. Wingfield Ravine, Box 366.
*P. REPENS Aub., sens. lat. Wingfield Ravine, Proctor 19244.
P. SECTIFRONS Kze. Mt. Misery, Breutel, not seen.
ml TRISERIALE Sw. (P. brasiliense Poir.). Nine Turn Gut, Box 355.
USSIANUM (Benedict)Benedict. Outer NW slopes of The
Fy "Decne 19 19529.
78
; P. Feel (Schaffn.)Maxon. Nine Turn Gut, Box 262; Wingfield Ravine, Proctor
19260.
PTERIS BIAURITA L. Recorded by Urban, nore Ant. 4:48 (1903); along track
between Sicbtions and Phillips Level, Proctor 1
06.
me HERACLEIFOLIA (Willd.)Underw. Wingfield Ravine, Box 369, Proctor
6
T. mnctsA Cay. (7. martinicensis (Spreng.)Copel.). Wingfield Ravine, Box 379,
Proctor 19641.
T. PLANTAGINEA (Jacq.)Maxon. Nine Turn Gut, Box 268, 352; Wingfield
Ravine, Proctor 19249.
T. TRIFOLIATA (L.)Cav. Nine Turn Gut, Box 266, 358; Wingfield Ravine,
Proctor 7 9253.
THEL Proctor, in Rhodora 63:33 (1961) (Dryopteris
oligocarpa sensu Alston, non O. Ktze.). Mt. Misery, Britton & Cowell 529; upper
SW spur of Verchle’s Mt. along track to Dodans (Dos d’Ans) Pond, Proctor
19587, type collec . Known otherwise only from Dominica.
- CLYPEOLUTATA (Desv.)Proctor (Dryopteris I’herminieri(O.Ktze.) C.Chr.).
No precise regi Box 273; outer NW slopes of The Crater, Proctor 19203.
2 ATA (L.)Proctor (Dryopteris decussata (L.)Urban). rded from
7. a by Urban: Symb. Ant. 4:19 (1903); outer NW slopes of The Crater, Proctor
oo DENTATA (Forsk.)E.St.John (Dryopteris quadrangularis (Fée) Alston)
Attempts to apply Alston’s criteria for distinguishing West Indian material from the
Arabian frican T. dentata have not oro a bane , and it does not seem
possible to maintain their separation. Admittedly, however, a great deal of variation
: —— i erage and perhaps statistical techniques may eventually pe
*T. GERMANIANA cieaiecec Outer NW slopes of The Crater, Proctor 19199.
a: A (Desv.)Proctor. Pores brachyodus ( Ktze.)
Britton & ‘Cowell. 407 (not seen); Candle Walk Ridge, Proctor 19580.
T. GONGyLopEs (Schkuhr)Small ees gongylodes (Schkuhr)O. Ktze.).
Edge of Crater Lake, Box 256, Proctor 19236
T. mNvisa (Sw. chy (Dryopteris oligophylla Maxon). Wingfield Ravine,
Box 378; — tor 19286
2. A (Sw ok or Devore hnbaie (Swartz)O. Ktze.). Mt. Misery,
Box 289, 203, 999: Proctor 19625; N of The Crater, Proctor 19505.
[T. megalodus (Schk Tete is tied twice by Box in his ro section
(p. 247, 250, as Dr prophihle megalodus), but it is lacking from Alston’s list. The
writer has not so © far been able to authenticate the record. Box may pomibly have
mistaken T. glandulosa for this species, as they have often been confused in a
T, NEPHRODIOIDES (Klotz.)Proctor et wcacasd Re 8 f. guadalupens
(Fée)C ae Ovects Ravine, Box 387; Olivee x 399; Wingfield Ravine,
ire 1925
OPPOSITA (Vahl)Ching (Depopteris opposita (Vahl)Urban). Wingfield
Sateen — & Cowell 446 (not seen).
T. PATENS (Sw.)Small (Devore bag (Sw.)O. Ktze.). Olivee’s Ravine,
Box 380. Mol Olyneux Water Source, Proctor 19288. ste
T. PorreaNA (Bory)Proctor els poiteana (Bory)Urban). Ollivee’s
Ravine, Box oe
“Tt. s (Gmel.)Morton var. TENERA (Fée)Proctor. Interior slopes of
The Conetyi Pacer 19232.
79
T. RETICULATA os pretex (Dryopteris reticulata (L.)Urban). Phillips Level
Box 263, Proctor 1955.
Las (Ra ron age sprengelii (Kaulf.) O. Ktze.). Wing-
field Ravine, Box 371 x Ghaut, Box 403; Molyneux Water Source, Proctor
19291; Molyneux Estate, ar ve & oned, 312 (not an: Lambert Estate, Britton
& Co well 637 (not s
T. TETRAGONA 7 )Sma ll (Dryopteris mets Se (Link)Maxon). Wingfield
Ravine, Box 374; Millikan Estate, Box 404; The Hole, West Farm, Proctor 19635.
cc FILIFOLIA Fée. Rodger 24 (BM).
V. CL: = E.Sm. ‘ Cheekles ’(acc. Krug); outer NW slopes of The
Crater pecs 4 9527
ABSTRACT
A shor and augmented list of St. Kitts | peel sae is presented, adding
es to the known catalogue for a total o One new species, Asplenium
las oie anal (p. 75) is descri i following are new co ombinations:
Hymenophyllum hirsutum var. gratum (p. 73), Grammitis asplenifolia (p. 76), G.
eggersii (p. 77), and G. suspensa (p. 77). The identity of G. asplenifolia and G. sus-
pensa are discussed.
SHORT NOTES
‘‘LASTREA DILATATA var. ALPINA”’
In 1857 Thomas Moore (Handb.Brit.Ferns, 3 ed.:126) described a small form
of Lastrea dilatata from Ben Lawers under the name var. alpina. Recently
has thought fit to raise it to specific rank and has named it Dryopteris
assimilis S, Walker [Amer.J.Bot.48 :607 (1961)]. It is hoped that more will be
heard of the distribution of this (diploid) close relative of D. dilatata (Hoffm.)
A. Gray.
The purpose of this note, however, is to report on another diploid
(chromosome number counted) Dryopteris of the same complex found by
myself in a wet wood on a steep hillside in Wester Ross. This fern may be
D. assimilis but as we have only Moore’s varietal description to go on I can
only compare it with that. The scales are concolourous and the rootstock
is definitely prostrate, creeping and branching with the fronds arising in
tufts at the ends of the branches. The stalk, blade and even the scales all
have stalked glands yet Moore (who loved his glands) never mentions them.
I think this plant is possibly D. spinulosa (lanceolatacristata) var. decipiens
(Syme) Druce but what relationship it bears to either D. assimilis (apart
from being another diploid) or to D. lanceolatocristata only bree ing
experiments will sh
H. V. CORLEY
NOTES ON THE HISTORY OF BRITISH FERN SPECIES I:
CONFUSION IN THE FILMY FERNS
. B. EVANS,
University College of N. Wales, Bangor
and
A. C. JERMY,
British Museum (Natural History)
In 1686 John Ray, the eminent English natural historian, published the first
volume of Historia Plantarum in which he included (p. 141) an account of a
filmy fern under the descriptive phrase-name sone petraeum perpusillum
Anglicum foliis bifidis vel trifidis Newtoni. Of it he says ‘“* This was first
shown to me by a Mr. [James] Newton, who, in company with Mr. Lawson
found it on Buzzard Rough Crag, near Wrenose [Wrynose] ” in the English
Lake District, in July 1682 and Ray’s description of it makes it clear that
Newton had found the plant we today call Hymenophyllum wilsonii Hooker.
He mentions, however, that he had “‘ heard that this little plant was found a
few years ago by Mr. Daire [a London alchemist], near Tunbridge town in
Kent ”’. In 1688 he published the second volume of his Historia containing an
Appendix to his 1686 account and here (p. 1853) he explained that ‘“ Mr.
Doody has told [and later showed] me this same little fern [i.e. Adiantum
petraeum ...]... at the house of Dr. Plukenet.’’ Ray’s description that
followed and the origin of the material suggests H. Mag shlcos In the same
year Ray published a work entitled Fasciculus Stirpiu which
he again mentions his Adiantum petraeum . gi to the two accounts
mentioned above for a “ fuller description ” Thus, although by chance Ray
had seen both species and published separate descriptions of them, it is clear
that he considered them as one in 16
Ray begins to have doubts in 1689 when, in reply to Edward Lhwyd of
the Ashmolean Museum, Oxford, who had sent him some Irish material,
he writes that he “ inclines towards the opinion of Dr. Plukenet who believes
in two sorts’ [of Adiantum petraeum ...]. A year later in his Synopsis
Methodica Stirpium Britannicarum Ray states that Dr. Plukenet, a doctor in
Westminster, makes two species, one fitting the Newton material, thé other
corresponding to Mr. Dare’s plant, and a written contribution by Plukenet
to this effect was appended at the end of the Stirpium Britannicarum. Plukenet
figures the two species in his Phytographia (1691):t.3,f.5 (which he names
Adiantum radicosum humisparsum Filicula pellucida nostras Coriandri foliolis
pest te globuliferum [ H. tunbrigense]) and f.6 (named Adiantum
osum liolis imis bisectis, caeteris vero integris tenuissime crenatis
‘ H. wilsonii]) and these are the first drawings to be published of these two
species. It should be noted that Plukenet used, in his phrase-name for the
plant we now know as H. wilsonii the diagnostic character erectius and for
that known as tunbrigense, humisparsum and his figures although of
sterile material show well the habit of both species. The reproductive
81
structures of ferns were little understood at that time and it is not surprising
that Plukenet did not look closely at the “‘ Seed vessels ”’ to see the difference
in the indusium.
By 1696 it is apparent that Ray accepted Plukenet’s views that two kinds
of plants were involved, for in the second edition of the Stirpium published
species [i.e. H. wilsonii]. ... Many botanists of the time used Plukenet’s
drawings for identification, thus the Rev. Adam Buddle correctly equated a
specimen of H. tunbrigense (now in the Sloane Herbarium in the Brit.Mus.
(Nat.Hist.)) with fig. 5 of Plukenet’s tab. 3. Ray published a Supplement to
his Historia in 1704 in which he equates Plukenet’s second species (=
wilsonii) with his Adiantum petraeum . . . and agrees that it is different from
Dare’s Tunbridge plant.
In 1700 James Petiver published a series of writings entitled Musei
Petiveriani and listed (p. 73) Ray’s second plant naming it Darea Tunbrigensis
minor possibly being unaware of Plukenet’s name for it. There is no doubt
as to which plant he is referring as he says, “ I gathered this elegant Plant
A.D. 1698 from the Rocks near Tunbridge-Wells, in the same Bese where
Mr. George Dare, Apothecary, had long before observed it ’’; a specimen is
in his collections in Herb. Sloane (H.S.190f.96). A year lates ange name
appeared in the literature namely in Morison’s Historia Plantarum (3:627,
s.15,t.7,f.49 [referred by the author as f.50, in error]). Morison quotes Ray’s
description of the Lakeland plant but only Plukenet’s fig. 5 [=tunbrigense]
but considers the plant to be a moss (a suggestion which Ray had made
some years earlier) and named it accordingly1. His figure appears to
represent two fronds of H. wilsonii and one of H. tunbrigense, all infertile.
Sherard, an English business man, had, in 1721, called over to England
the German botanist Johann Dillenius who, amongst other tasks, was set
to work on revising Ray’s Stirpium Britanicum. The resulting book did not
have Dillenius’ name associated with it, however, and was published in 1724
as a third edition. In this, a book with a large and wide sale, Dillenius,
under Ray’s original name says, “‘ whether it is truly the same and not such a
distinct species it is at least a variety owing its distinction to age, place and
circumstances thus arising?. He adds new localities: ig Pet Yorks;
near Llanberis, Caerns.; and Cockbush,* near Chichester, Susse:
In 1753 Carl Linnaeus published his Siesccat’ Sediés pistioe
where, by inserting epithets in the margin of this work, he produced the basis
for all modern (binomial) plant nomenclature. Under the name of Tricho-
manes ae ae he gives a brief arene (which could be either species)
i his).
‘ : ie ee. : a:
. Utrumque enim idem est ¢ et non tam _distincta species, quam varietas saltem est aetati, loco et
* A Settle locality reported for H. wilsonii in 1911.
4 Cockbush: “ cain also plectifully by ed Rand, in Company with Mr. Sherard, amongst Pebbles
at Cockbush . on the of Susse: This almos certainly refers to a misidenti
ck
(possibl the alga Pelvetia prone Bae fe (ef. Wolley-Dod, FL Sussex, 937, who suggests the pebble
velered to could only have been marine
82
and refers in synonymy to Ray (1704) and (1724) (i.e. to Dillenius for whom
Linnaeus had a great regard), to Plukenet (1696) quoting both figures, to
Petiver (1695) and to Boccone! (1697); for localities he gives “ Anglia;
Italia’’1. Linnaeus accepts the view that there is one species and bases his
concept (as does Dillenius) on H. tunbrigense as he quite likely never saw
H. wilsonii; there is in fact one sheet of Trichomanes [Hymenophyllum]
tunbrigense in the Linnaean Herbarium in London, annotated by J
Smith “* Ex Anglia”? and presumably from Tunbridge Wells. In his own
copy of Plukenet, Linnaeus has written in the margin of t.3 “* 5, 6, [bracketed
together] Adiantum ” and in a later pen has crossed this out and written
“Trichomanes tunbrigense”’. Similarly this name is written against
Adiantum petraeum ...in Linnaeus’ copy of the Stirpium Britannicarum
3rd ed.
Between the years 1785 and 1790 James Bolton of Halifax published a
set of plates and text entitled Filices Britannicae. On page 58 he gives a
description under Trichomanes tunbrigense of a plant which is more likely
H. wilsonii [cf. “* pinnae 2 or 3 lobed ” and “ seed-vessels, globular, size of a
small mustard seed ’’]; his figure, a bad one, could be either species. Bolton
adds to the confusion by adding the following comment ‘I suspect the
ee pyxidiferum [=T. speciosum Willd., the Bristle-fern] of Bell
Bank [near Bingley, Yorks.] to be a luxuriant variety of this plant; their
figure, texture, colour and whole habit are the same, only differing in
magnitude . . .” [in this he is incorrect for the Bell Bank plant collected by
Dr. Richardson is in the Sloane Herb. (H.S.302f.66) and is T. tape
Furthermore he writes in the Introduction (p. xii), “‘ There is a variety of
Trichomanes tunbrigense which grows in little caverns under moist rocks
where the sun is excluded, and where the water, dripping from the points of
the leaves, enlarges them greatly; in this state the plant has been taken
notice of, and called Trichomanes pyxidiferum”’. The concept at that time
may well have been based on a luxuriant habitat form of H. wilsonii.
J. E. Smith was the first to divide the genus Trichomanes as conceived
by Linnaeus and in 1793 published the genus Hymenophyllum®, on the basis
of the enclosed columella and bivalved involucre, wed (Trichomanes)
tunbrigense L. as the first, now regarded as the type, species. The British
Floras of the time, e.g. Withering, Botanical Arrangement 03 British Plants
(1792), Hull, British Flora (1799) and J. E. Smith, Flora Britanica (1804)
omit the true Bristle-fern and include the Bell Bank plant (v.s.) and other
records of T. pyxidiferum L. as varieties of H. (or 7.) tunbrigense. Smith
went further in 1805 when he described the Bell Bank plant as a new species
Hymenophyllum alatum in an attempt to show that Trichomanes pyxidiferum
of many British authors was not the same as the American plant of that
name (i.e. true T. speciosum Willd.). Sowerby’s drawing of H. allatum
J.E.Sm. in English Botany 20:t.1417, whilst omitting the exserted columella,
shows a typical narrowly campanulate indusium of a true Trichomanes!
. Seat ot plante rare della Sicilia, ete... « 224 V2 f.1 [a poor drawing of sterile H. tunbrigense ;
s is the first record from the Continent).
° = Mem. ‘Acad.Roy.Sci:Turin $:418, t.9, fig. 83
Twenty-five years later, in 1830, W. J. Hooker cleared the matter up
in his British Flora: he included Trichomanes speciosum (as brevisetum) in
the British Isles and besides Hymenophyllum tunbrigense described a new
species — H. wilsonii, as ‘‘ abundant in the Highlands of Scotland and many
parts of Ireland ’’, and named after Mr. W. Wilson, the bryologist, who
“found them both growing at the Lakes of Killarny and distinguished them
specifically”. Later in the Synopsis Filicum (1868) Hooker had second
thoughts and wondered if wilsonii may be of subspecific rank This later
opinion was formed after he had seen much more material of this species —
complex from various parts of the world — material that appeared inter-
mediate between tunbrigense and wilsonii. Hooker, by widening his concept
of the species brought about nomenclatural problems as to the earliest name
and whilst H. unilaterale Bory ex Willd. is common in the literature, H.
peltatum (Poir.)Desy. the legitimate name (if regarded as a single species) is
also in the Floras. Today, however, H. wilsonii Hook. is accepted as a
separate species confined to N.W. Europe and as such the name is legitimate.
We have tried to show in this short paper that whilst the older botanists
discerned two species of filmy fern on vegetative characters alone Linnaeus
followed Dillenius and regarded them as one, basing his concept (luckily)
on an English plant of H. tunbrigense. Confusion continued with tunbrigense,
wilsonii and Trichomanes speciosum being taken as a cline in the one species-
complex. Hooker eventually redescribed the original taxon as H. wilsonii,
thinking later that the differences may only be infra-specific. Recently new
criteria (such as cell-size, chromosome number and ecological behaviour)
have removed the apparent similarity of much foreign material and
emphasised the distinction in Britain so that specific rank may now be given
to our species with some certainty. We should like to thank Dr. W. T. Stearn
of the British Museum (Natural History) for much helpful discussion.
SHORT NOTES (cont.)
HYMENOPHYLLUM TRANSPLANTS IN KENT AND SUSSEX
As part of researches into the autecology of British Hymenophylla the
following transplants were made Nov. 4, 1961: at Eridge Rocks, Sussex,
two mats, (approx. 10 sq. cms. of H. wilsonii (from Tryfan Mnt., Caerns.)
were planted in patches of H. tunbrigense, another on a bare rock-cleft wall
and a fourth near the site of the Dryopteris aemula. At Hungershall Rocks,
Kent, two similar mats one of wilsonii and one of tunbrigense (from Eridge
Rocks) were planted a yard apart on a peaty ledge with a N. aspect. No
Hymenophyllum has ever been recorded specifically from Hungershall Rocks,
to my knowledge. It is planned to keep these experiments under complete
control and the wilsonii will eventually be removed from the locality. I shall
be pleased to hear of any observations on the gametophytes or young
sporophytes of these two species.
GARETH B. EVANS
84
CYSTOPTERIS ALPINA AND C,. DICKIEANA
IN THE ROCK GARDEN
Cystopteris alpina and C. dickieana have both been growing for more than
twenty years in the writer’s garden in Surrey. They have been given the
minimum attention: none at all for five years during the last war and it
seems worthwhile to describe the rather unorthodox method of cultivation.
The small garden in approximately 400 ft. above sea-level, on the sides of
the Downs, sloping south and west; the soil consists almost entirely of
disintegrated chalk. The ferns were planted in a small rockery of irregular
form made of rough-hewn sandstone which is hard but retentive of moisture.
Part of this rock-work faces north and forms a shady “ cliff ’’ about 15 inches
high, the spaces between the horizontally laid stones being filled with garden
soil. A good deal of moss forms on this cliff-face but does not damage the
Cystopteris as these ferns have the alpine characteristic of dying down
completely after a short period of growth in the summer. In the early spring
whilst the fern is still below ground it is desirable to clear away the remains
of fronds and to remove the moss. Grown in this way the ferns look much
the same as they do in their native haunts; C. dickieana happily works its
way along the horizontal crevices in the rock-work. It is constantly small in
stature, averaging perhaps about 4 or 5 inches, the fronds tend to be broad
and rather foliose and the pinnae slope down from back to front suggestive
of the laths in a Venetian blind — this is a useful identification character in
a living specimen not so evident in a pressed frond. A further distinctive
character of C. dickieana as compared with alpina is the production of a few,
and usually less vigorous, fronds in the late autumn. On the face of it it is
difficult to see why C. dickieana should be considered a variety of the
polymorphic Cystopteris fragilis.
P. GREENFIELD
85
BRITISH PTERIDOLOGICAL SOCIETY
FIELD MEETINGS FOR 1961
KEW GARDENS, Saturday, 29th April. This visit to the Fern Houses at Kew
Gardens, arranged by our member Mr. H. J. Bruty who is in charge of the
Houses, was attended by eleven members and one visitor. The ferns all
belong to warmer climates and our President was able to impart much
interesting information on the many plants which he sent home to Kew from
Malaya, Borneo and elsewhere. Mr. Bruty was kept busy answering questions
on the methods of growing these ferns, particularly in his propagating
houses where we saw plants in all stages of growth from the prothallus
onwards.
We have to thank Mr. Bruty for a most enjoyable day, and the
opportunity given for getting to know some of our newer members, whom
we were very glad to welcome.
SHAFTESBURY, DORSET, 9th- 11th June. The headquarters of this meeting
was at the St. Edward Hotel, Shaftesbury where the party met on the
Friday evening. The following morning the party of ten members and
friends went under the direction of Mr. Peter F. Hunt to Asham Wood, an
ancient ashwood on limestone. Here the ground flora over the large boulders
that formed the sides of this wooded valley consisted of many Phyilitis plants
(each of which were carefully examined —but in vain) Asplenium
trichomanes, Dryopteris borreri and Polystichum aculeatum and the occasional
plant of Cystopteris fragilis. After lunch in Nunney Castle the Polystichum
hunt was continued at Railford Bottom and from there to Shepton Forest on
the Old Red Sandstone. Here in the boggy lags around the plantations
Dryopteris lanceolatocristata was found along with dilatata and Athyrium
filix-femina.
Sunday was spent in the woods, mainly of birch, on the Greensands
around Stourton. The usual ferns were noted and it was interesting to see
the not too common Egquisetum telmateia and the very uncommon E. sylvatica
in King’s Wood Warren.
EAST GRINSTEAD, Sunday, 16th July. Persistent rain could not discourage
our party of six which met at East Grinstead Station and set off for a locality
near Turners Hill where we hoped to find Hymenophyllum tunbrigense and
Dryopteris aemula. We found neither, but had good hunting in wet woodland
which produced Dryopteris lanceolatocristata in quantity. Working in an
easterly direction towards Tunbridge Wells after lunch we hunted Lycopodium
inundatum in vain, and we finished our day near Newbridge where our finds
included Thelypteris palustris. In all eleven species were found.
VISIT TO MR. T. H. BOLTON, BIRDBROOK, ESSEX, Saturday, 12th August.
This outing will long be remembered by the participating members 4s
one of the most enjoyable ever held by the Society. Sixteen members and
friends were entertained to lunch by Mr. Bolton and his family, and later
86
were let loose among his ferns. It was typical of our host’s generosity that he
had garden forks and plastic bags all ready, and ferns coveted by members,
provided they were not the one and only specimen of their kind in the
collection, were dug up and bagged. The number of ferns leaving Birdbrook
that day had to be seen to be believed, but made little impression on the
magnificent Bolton collection. After tea we went our several ways home and
our thanks to Mr. and Mrs. Bolton and their family, though profuse, could
not adequately express our appreciation of a happy day and their warm
liberal hospitality.
THE ANNUAL EXCURSION, 23rd - 30th September. Snowdonia proved to be a
popular choice for the Annual Excursion, which was attended by a record
number of members. We were very pleased to have our President with us,
together with Mrs. Holttum, who joined in all our expeditions. In addition,
the London area was represented by J. A. Crabbe, P. Halligey, A. C. Jermy,
R. M. Payne, R. S. W. Pollard, Mrs. Pollard, W. Wright and the Secretary.
From the North came Miss A. Sleep, Messrs. F. Jackson, R. Kaye, N.
Robinson and W. S. Webster. Later in the week we were joined by our
Midland members, Mr. and Mrs. T. A. Dyer and L. Burnett. We were
glad too, to welcome Mr. Proctor who is in England from Kingston, Jamaica
to work on a fern flora of Jamaica, and who took this opportunity to have
a look at our ferns. It was a great disappointment not to have with us Mr.
P. Greenfield who for so many years has been the mainspring of the Excur-
sions. It was a bitter disappointment for him, as at the last moment he had
to cancel his plans because of serious illness in his family. We mi i
Our headquarters was the Aberdunant Country Club, Prenteg, near
Portmadoc in Caernarvonshire, and here we had an ideal centre for our
purpose, in an old manor house set in extensive wooded grounds with
glorious views of the mountains to the North. The half-mile entrance drive
was up a dark ravine rich in ferns, including Polystichum setiferum and
Hymenophyllum tunbrigense. Naturally, we commenced our hunting here on
the Sunday, and recorded in all 12 species. Later in the day we wandered
further afield along the Caernarvon road, and explored up the valley of the
Afon Dwyfor which was not particularly rich in ferns, and only the more
common species, 12 in number, were found.
Snowdonia is one of the richest areas in plant life in Britain, and it is
satisfactory to know that this has been recognised and steps taken to preserve
the region, which in 1951 became a huge National Park. Protection does not
end here however, for within the Park, preserving its riches for posterity, two
other powerful bodies are active, the National Trust which owns seve
properties, and — what concerns us more — the Nature Conservancy which
selects and maintains areas of outstanding interest. There are several of these
Nature Reserves in the Park, and one of the objects of our Excursion was to
visit as many of them as possible. In this we were greatly helped by Dr.
Brian Seddon from the Nature Conservancy Headquarters for Wales at
87
Bangor, and by Mr. Evan Roberts, the well-known naturalist and Warden
of the Cwm Idwal Reserve, and their co-operation was very much appreciated.
We visited four of the Nature Reserves. The Coed Tremadoc Reserve
is near Prenteg and we visited this on our first day. It is a precipitous area
of dense oak woodland with ferns dominating the undergrowth on the wet
rocky slopes. We recorded ten species, with Polystichum setiferum the
common fern growing strongly in great abundance, very reminiscent of its
extensive growth in the West Country. A very similar place is the Dolgarrog
Reserve, with dense woodland clinging to a steep rocky hillside. We visited
this on Thursday accompanied by Dr. Seddon, and again ferns were dominant
in the undergrowth. The same ones were here, the large-growing common
species and some of the smaller rock ferns, with the addition of Thelypteris
limbosperma, but P. setiferum was not so abundant and occupied but one
small area.
Of much more interest to us was the small Reserve of Cwm Glas
Crafnant at the head of the Afon Crafnant valley, visited on the same day in the
company of Dr. Seddon. This is a precipitous headland on the hillside
facing north-east, closely covered with stunted trees. The high humidity
and shade foster the development of a great variety of mosses, which de-
manded more than passing attention. Here too grows Sedum telephium and
S. forsterianum, and many other flowering plants to which we could have
devoted much time, and we had to remind ourselves that it was the ferns we
had come to study. They too are there in variety, and in this small reserve,
only an acre or two in extent, we recorded 15 species. Among them we
noted a few plants of Asplenium trichomanes, A. adiantum-nigrum, A. viride,
and Cystopteris fragilis. A fine colony of Hymenophyllum wilsonii was
found on wet rock, and P. setiferum and P. aculeatum were both seen, but
only in small numbers.
Our visit to Cwm Idwal Nature Reserve was a much more strenuous
expedition which occupied a whole day — the Wednesday, and was led by
the Warden of the Reserve, Mr. Evan Roberts, accompanied by Dr. Seddon
and Mr. G. B. Evans from the University College of North Wales, Bangor,
who is working on the filmy ferns. The going was hard and not all of us
completed the course. Lunch was taken in pouring rain, sheltering as best
we could under great boulders below the Devil’s Kitchen, and trampling
H., wilsonii underfoot. Afterwards only the more energetic members of the
party continued upwards, and the others retired to hunt in the Cwm until
the deteriorating weather with high wind and driving rain forced them to
retreat. Woodsia ilvensis grows on rocks in this vicinity and this was a magnet
to attract us upwards through any weather. Soaked through and cold we
plodded on led by Mr. Roberts, and we got our reward. In some rock
crevices grow a few insignificant plants of the rare fern, and about 15 minutes
were spent photographing them in the most impossible conditions — cold
lashing rain driven by the wind on the exposed hill-top. Then on once more
to a higher top below which, in the steep stony screes, grows Polystichum
88
lonchitis. Again there was a division in our party, and only a few enthusiasts
traversed the loose screes to see the holly fern. Some of us had already seen
it a few days earlier on Snowdon, and by this time our chief desire was to
get back to warmer and drier conditions. This was the only wet day we had
during the whole week, and although it would have been pleasanter without
it, still it did not detract from the enjoyment of the day. Not very many ferns
were seen, and in addition to those already mentioned the most noteworthy
is the Quillwort, Isoetes echinospora, found in the Lyn Idwal.
A day was spent on Snowdon. Walking from the Gorphwysfa Hotel,
Pen-y-pass, we climbed by a track past the two lakes, Llyn Teyrn and Llyn
Llydaw up to Glaslyn at the foot of the great precipice leading to the summit
of the mountain. Hunting as we walked, the number of species mounted
quickly, and we finished the day with a total of 20. Nowhere were ferns to
be seen in great abundance, but the types of habitat are varied and each
fosters a selection of suitable species. On the screes, Cryptogramma crispa
flourishes happily, and in odd places grow the occasional Devons filix-mas
D. borreri, D. dilatata, Athyrium filix-femina, Polypodium vulgare, and
Blechnum spicant. The ruined buildings of the long disused copper mines
below the precipice of Clogwyn-y-Garnedd we found Cystopteris fragilis,
Thelypteris Phegopteris, T. Teabiperiie and D. abbreviata. On the cliffs
above grows Woodsia alpina and a few of us climbing in search of it, had the
good fortune to find one plant on the rock face about ten feet above a ledge
on which we were standing. Near here too we found the rare Polystichum
lonchitis, and also H ‘'ymenophyllum wilsonii. Our twentieth find was Jsoetes
echinospora, growing in one of the lakes. Three club-mosses were also
noted, Lycopodium selago, L. clavatum, and L. alpinum.
Again in pursuit of Woodsias, this time W. ilvensis, we spent a day on
the rocky slopes of Moel Hebog above Cwm Meillionen near Beddgelert.
The approach was through Forestry Commission plantations, and having
missed the path, we had to struggle upwards through the dense growth of
young conifers until we emerged on the open hillside. Hidden among the
trees we passed a low cliff, its wet surface covered with a luxuriant growth of
H. tunbrigense. 1n the lower and more open part of the wood a colony of
Thelypteris phegopteris spread itself happily through the sedges and grass
on the wet hillside. Climbing and scrambling on the cliffs, we were again
fortunate in finding our Woodsia. On the sheer wall of a perpendicular
chimney, we counted in all nine plants of W. ilvensis, and with some difficulty
were able to take close-up photographs of one of them. Other ferns on the
rocks were A. trichomanes, A. viride, A. ruta-muraria and C. fragilis, while on
the screes some Cryptogramma crispa was seen. The club mosses L. seago,
L. clavatum and L. alpinum were abundant on the hillside, particularly
L. alpinum which was present in great quantity.
89
We found so much to occupy us in the Snowdon area that we were able
to turn to the south for one day only. Two very rare ferns were our objectives,
Trichomanes speciosum which has been found on the south side of the Afon
Estuary, and the rare hybrid Asplenium alternifolium which grows
on the hills bith of Llanelltyd. The habitats were known to a few of us and
we were successful in finding two plants of the Asplenium, one growing in a
rock crevice and the other on scree. The latter was ideally positioned for
some successful photography. Both parents of the hybrid, A. vcr
and A. trichomanes were growing in numbers nearby. The Killarney
was not so obliging unfortunately, and after a wet fruitless hunt up ee
course of a mountain torrent exploring every nook and cranny and behind
waterfalls, we reluctantly gave up the search. In spite of this disappointment,
we were very satisfied with the day’s hunting. In addition to A. alternifolium,
we found Dryopteris aemula near a large waterfall, growing vigorously on a
narrow ledge above the waters, and presenting a very tricky problem for our
photographers. Here too, the two Hymenophyllums were abundant on the
rocks, as was A. obovatum on stone walls by the roadside on our way up the
coast from Barmouth. These ferns with D. abbreviata and others, totalling
21 species, made the day outstanding.
At the end of the week we were very conscious of all we had not done,
and in a vain attempt to put some of this right, Saturday developed into a
scramble to visit a multitude of places. Our first stop was to see Asplenium
marinum on the sea-cliffs near Criccieth. Thereafter each car went its own
themselves with exploring odd corners near Prenteg. Rushing torrents
cascading down the steep hillsides in deep ravines are a feature of this
countryside and ferns revel in the humid conditions created by the drifting
spray. On the rocks high above one very deep and spectacular ravine,
A. septentrionale was plentiful in the crevices. Today too, after searching in
vain in all likely places during the week, we were able to add D. lanceolato-
cristata to our list.
This excursion was outstanding — from the number of members
attending which compares with the attendances during the early days of the
Society when the fern cult was in its heyday, and from the number of species
found, 34 in all, which is a pretty impressive figure for one small corner of the
country when we consider that there are only about 50 fern species in Britain.
Many of our rare ferns were seen for the first time by most of us, and it was
very satisfying to see the two Woodsias, P. lonchitis, and A. alternifolium.
We should include A. septentrionale among our rare finds, but we saw sO
much of it that we ceased to think of it as rare. Dr. Seddon introduced us to
one large colony numbering hundreds of large plants growing in the
stone wall by a roadside in the Conway valley. This road is scheduled for
widening in the near future, and the wall is to be pulled down. It is a tragedy
to think that what must be one of the finest colonies—if not the finest—of this
fern in Britain is going to be destroyed. The Nature Conservancy at Bangor
90
is very conscious of this, and it is planned to move the ferns before the wall
is pulled down, and establish them on other walls in the vicinity. We
appreciate very much the thought and care which is being given to their
preservation although we cannot help feeling pessimistic about the success
if the operation. It deserves to succeed and we sincerely hope it will. Along
the same road, the sight of a fine colony of Ceterach officinarum covering an
old high wall gave us great pleasure, and many fine plants were collected
without making any impression on the numbers there.
Although Snowdonia can offer a wealth of fern species, variation is
absent. Those of us who are variety hunters kept our eyes open and two
finds only were made, one a C. officinarum with crenate fronds, and the other
a promising small plant of A. filix-femina growing on high rocks. Although
small it was not a young plant and consisted of two crowns with fronds of a
very unusual character which however is difficult to describe as they were
so small and past their best. Mr. Jackson and the Secretary each have a
crown and we look forward to seeing what the fern will do next year.
A list of the species recorded is appended, and the numbers against each
indicate the localities where found as shown in the list of place names also
appended. Vice-county numbers and grid-references are included.
Isoe pora Durieu — 1
Hymenophyllum : tunbrigense (Ly ee i) 92-99.
H. wilsonii Hook.— 11, 13, 17, 19.
Pteridium aquilinum (L ') Kuhn — 1, 3, 6, 10, 12, 15, 16, 17, 18, 19, 20, 22.
Cryptogramma crisp: a Br.ex Hook.— 11, 12, 13, 16, 20.
Blechnum spicant (L.) Roth — i. 5, 6, 9, 10, il, 12, 13, 15, 16, 19, 20.
Phyllitis scolopendrium (L.) Newm. — 7, 11, 18, 22.
As losin betectuannigrom L—A, 3) 6:7, 10; 85.17, 38, 20 20
A. oboyat v. — 1, 4, 8 21.
co
93 3
—
Po
—
2
A Seatindaiy
A. trichomanes L. — 2:3 6 7: 10; 11, 12,13; 15) 16, 17, 18,
A. viride Hu ven lis ef 13, i7.
A Saat S 10, il, on Ls 20.
A septentrionale (L.) Hofim 4, 20.
A, enw
yoshi
Cystopteris fam ih )E Bernh. — 11, eb 1B.
i r.— 12, 13.
j — 1, 3,7, 9, 10, 11, 12, 15, 17, 18, 19, 20, 22.
. borreri Newm. — 1, 3, 6, 7, 9, 10, 11, 12, 15, 16, 17, 18, 19, 20.
1 ye 20.
Alston — 9.
. dilatata (Ho ng ies Gray + & 3. 6, 7, 9, 10, 11, 12, 13, 15, 16, 17, 18, 19, 20.
) u sia €'8
‘olystichum setiferush (Forsk.) lah — 1, 7, 17, 18, 22.
> lonchitis (L-) Roth 11s 13.
itis (L.) Roth — 11,
Thelypteris limbosperma (Ail.) H. P. Fuchs — 1, 3, 6, 9, 10, 11, 12, 13, 16, 17, 18, 19.
: JH
7. phegopteris (L.) Sioson — 10 i, 12, 17, 19.
13
Polypodium vulgare L.1, 3,6, 7,9, 10, 12, 15, 16, 17, 18, 19, 20, 22.
93.
>
P. interjectum Shiver 24; 11
91
Prenteg — V. CG 49, 23/5841.
Portmadoc V.C. 49, 23/5738.
Cerrig-y-rhwy: wydw — V.C.49, 2 ia
Paavacaiceenn — “V. C.48 23/6 143
Afon Maesgwm, near Croesor — V.C. 48, 23/6344.
Aberglaslyn — Vv. C. 49, 23/5946.
Coed Fremadoe Nature Reserve — V.C. 49, 23/5640.
5237.
WwW :
Llanfihangel-y-pennant — ey C.49, 23/5244.
Snowdon — V.C.49, ais 154.
23/5647.
Cwm Idwal Nature nS V.C. 49, 23/6459.
Near Llanrwst — V.C.49, ince
9
wm Glas Nature Reserve, Crafnant — V.C. 49, 23/7360.
econ Nature ge — V.C.49, 23/7766.
Ganllwyd — V.C.48, 2
Near Llanelltyd — V. a a, 23) 7221.
Llanfair — V.C.48, 23
Near Penmynydd, se ibook — V.C.52, 23/5174.
PESSSIAARGN=SemraAMAwN—
x6
°
ge
E
>
LOWViS:
ASPLENIUM
x
MURBECKII
(see page 110)
LOVIS: ASPLENIUM x MURBECKII 111
tinuous smog at the end of November 1962. It has therefore proved to be
very fortunate that Mr Clive Jermy had sent me in May 1961 a vied sage
from the plant at Chelsea. This fixing yielded only a single sporang nan
informative stage but some of the spore mother cells from this sporangium
gave clear analyses of chromosome pairing which were both unexpected and
interesting. Since it now appears rather unlikely that more fertile fronds will
become available in the immediate future and in view of the fact that there is
no previous account of chromosome pairing in A. X murbeckii, it has been
decided to present a report on this limited cytological information
CYTOLOGICAL RESULTS
Preparations were made by the aceto-carmine squash method.
stated above, only one sporangium was found in a stage suitable for study of
the pairing of the chromosomes in meiosis. The spore mother cells from this
sporangium were in diakinesis. Fourteen cells in all were available for study.
Of these, only two have yielded confident analyses. One of these two cells
(illustrated on plate I, fig. 1, with explanatory diagram on text-fig. 2), shows
60 bivalents and 24 univalents, the other (plate I, fig. 2, text-fig. 3), shows 62
bivalents and 20 univalents. Though the remaining cells have not yielded
accurate analyses, they clearly show the same order of pairing.
INTERPRETATION
The basic chromosome number (x) in Asplenium is 36. Both A.
septentrionale and A. ruta-muraria are tetraploid species with perfectly
gular meiosis, constantly forming 72 bivalents at meiosis. The analyses
obtained from the Castle Crag hybrid indicate that it is tetraploid, thus
giving cytological confirmation of the correctness of identification as distinct
from A. x alternifolium which is a triploid.
Studies have been made on the chromosome pairing found in the
hybrids formed between A. septentrionale and the diploid and tetraploid
cytotypes of A. trichomanes L. and it has been shown that the analyses
obtained can only be consistently aes if it is assumed that the two
genomes contributed by the A. septentrionale gamete pair together in the
hybrids formed with A. trichomanes. From this conclusion it can be inf
that A. septentrionale is of effectively autopolyploid origin. In other words,
A. septentrionale has arisen from within a single ancestral diploid species by
doubling of the chromosome complement (Lovis, in preparation).
There are two possible alternative simple interpretations of the chromo-
chromosomes donated ae the A. — parent. Either all the bivalents
are formed by pairing between chromosomes contributed by different
parents (i.e., all the Fase: are : SR), or else the two genomes contributed
BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963
Biaacer,
Silhouettes (natural size) of fertile fronds of Asplenium x murbeckii
Text-fi
oie Ch eo Crag taken from Pg division now in cultivation at Leeds on its arrival
n the autumn of 1961.
2, NS
ee
i
ase Of
y
) fe)
3 % 2 OR, ro
S BY ge ~ Os
4,° ee
Text-fig. 2. Explanatory diagram to cell illustrated on plate I, fig. I De ment
x 1250. Bivalents in solid black, univalents in outlin
ta
Text-fig. 3. Explanatory diagram & @ > af
to cell illustrated on plate I, fig. 2. 2 ah 2
50. Bivalents a &
in solid — univalents in Bg w ome 4
utline. eas 9
208 |
LOVIS: ASPLENIUM x MURBECKII 113
by each parent are pairing between themselves, (i.e., all the bivalents are
SS or RR).
The first interpretation is less likely since, from other information
(Lovis loc.cit.), it has already been inferred that the two genomes present in
gametes formed by A. septentrionale are pape of pairing with one another
at meiosis. Morever, d A. ruta-muraria
to be separated from one another by cay a limited number of gene differences,
as is probably the case if their chromosomes are capable of pairing together
to form as many as 62 of a possible 72 bivalents. If correct, this interpretation
would involve a further rather curious inference since, if the assertion that
A. septentrionale is of effectively autopolyploid origin is accepted and if in
fact A. ruta-muraria were separated from A. septentrionale by only a limited
number of gene differences, then it would evidently follow that A. ruta-muraria
must ultimately be derived from the same origin as A. septentrionale, in other
words it must have as its ancestor the same single diploid species as does A.
Septentrionale,
The alternative interpretation of the chromosome pairing also leads us
to an unexpected conclusion regarding the origin of A. ruta-muraria. If the
chromosomes of the two genomes from A. septentrionale are pairing together
iv:
the A. ruta-muraria parent (RR), indicating a high degree of homology
between the two genomes present in the gamete of A. ruta-muraria. This
Conclusion would indicate that A. ruta-muraria is either a segmental
allopolyploid or an ancient autopolyploid. Such a situation, notwithstanding
the claim that A. septentrionale is effectively an ancient autopolyploid, is
nevertheless believed to be of generally rare occurrence amongst ferns,
wherein nearly all of the polyploids so far investigated have proved to be
typical allopolyploids (Manton 1950, 1961).
It is not possible to arrive at a firm decision here as to which of these
two suggested interpretations of the chromosome pairing found in the
Castle Crag example of A. x murbeckii is correct, Equally, it is not possible
on this evidence alone to determine just how closely related or otherwise are
A. septentrionale and A. ruta-muraria. Cytogenetic study of hybrids formed
7 es septentrionale and A. ruta-muraria and other species of Asplenium
It isa blair to record my grateful thanks to both Mr Fred Jackson
and Mr Clive Jermy for their kind and generous co-operation.
may m
REFERENCES
JACKSON, F, (1961). The sre of Asplenium x murbeckii Dérfl. British Fern Gazette 9:49.
LOVIS, J. D. J. Linn. Soc. (Bot.), in preparation.
MANTON, I. (1950). Probleme = cytology and evolution ies a gees Cambridge.
MANTON, I. Gest). Type y Mg Pteridophyta. Conference Report 6, A Darwin
Centenary, edit. P.J. Wanstall:
114 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963)
FERNS OF SARDINIA
by
G. J. pe JONCHEERE
Antwerp
In 1960 it was my good fortune to visit Sardinia with my friends Professor
and Mrs van Steenis. Botany and holiday were agreeably mixed.
When flying in to Alghero from Rome the fernhunter’s eye is struck by
the wild uncultivated landscape and the vast expanse of forests still covering
the northern part of the beautiful island. The expectation which I felt was
certainly confirmed by our subsequent journey. We crossed from Alghero
in the west to Santa Teresa in the extreme northeast via Sassari and Tempio,
then driving southwards via Monti, Nuori and Fonni to Desulo where we
climbed Gennargentu (1830 metres) the highest mountain of Sardinia. Our
faithful little Fiat carried us back to Alghero via Abbasanta and Santa
Lussurgio. We did not visit southern Sardinia which, being drier and less
wooded than the north, is probably less interesting to a pteridologist. Before
getting down to botany we saw the Ascension Day pageant at Sassari,
representatives from all parts of Sardinia, each in their own local costume, ,
march in procession through the sun-bathed piazza of the old town making
a magnificent display for which the island is famous.
From Sassari we went through Tempio to Aggius for our first real
botanising. A wild jumble of basaltic rocks hides a wealth of ferns tucked
away in cracks of huge boulders shaded by small copses of mediterranean
oak and chestnut. Asplenium adiantum-nigrum ssp. onopteris mingled with
‘A. obovatum and although these ferns are both well known they still offer
problems.
A. adiantum-nigrum ssp. onopteris is quite different from the North
European form of A. adiantum-nigrum ssp. adiantum-nigrum, yet there 1s a
complete sequence of forms without a clear demarcation line right down to
the extreme forms in the Canary Islands and Palestine; it seems therefore
and tetraploid) and ssp. onopteris (diploid). However, if it can be proved
DE JONCHEERE: FERNS OF SARDINIA 115
that the hybrid between diploid adiantum-nigrum and onopteris is also
sterile the case would be reversed and I would agree with Dr Meyer that two
real species are involved.
The case of A. obovatum is different. Viviani described it from a
mediterranean plant, small with rounded segments and diploid. The atlantic
forms which Hudson described as A. Janceolatum are larger, with pointed
segments and atinsgr: The differences are small and it takes a lot of
experience to separate them. They were both growing together in Sardinia.
Manton & Reichstein (1962) in Bauhinia 2(1) state that A. obovatum is
diploid and A. billotii is eo et also, that according to unpublished
investigations by Dr M. G. Shivas (Mrs T. G. Walker), A. billotii is an
alloploid from A. obovatum sv A, onopteris; pending further substantiation
I cannot accept that the two plants be treated as different species and prefer
to regard them as A. obovatum sspp. obovatum and lanceolatum.
Other ferns seen at Aggius were Bracken and Ladyfern, both ubiquitous.
Huge clumps of Osmunda regalis in wet places were most impressive and the
little annual Anogramma ie i on earthy banks was a reminder of
links with more tropical flor
scenery around Santa Teresa is strongly reminiscent of Cornwall,
Sardinia is one of its most easterly localities; it is always worthwhile to look
further above it for Asplenium obovatum as the two often grow together,
albeit on different levels. It is curious that in Santa Teresa, so far away
from the British and French coasts, the same association was found; if one
had not seen this before one would be tempted to regard it as a very curious
habitat for A. obovatum which otherwise grows everywhere in Sardinia in
rockfissures and hedges, if not too dry and high.
We climbed Monte Corrasi (1460 metres) from the old village of Oliena
and we drove by car practically to the top of Monte Limbara (1360 metres).
Their dissimilarity is striking. Corrasi is limestone with a large bare plateau
on the summit and treeless through generations of herdsmen grazing their
goats and sheep; Limbara is basalt and the vegetation resembles North
European heathland. Limbara was not very exciting pteridologically but
Athyrium filix-femina, Dryopteris borreri, eget see Asplenium
trichomanes and Polystichum setiferum again showed European ferns
finding suitable terrain in a mediterranean locality. It was ays surprising to
nd Asplenium ruta-muraria on the top of Monte Corrasi; this is rare in
Sardinia and not recently reported as far as I know; other limelovers found
there included Ceterach officinarum, Adiantum capillus-veneris and Dryopteris
villarii which latter, as found in Sardinia and southern Italy (var. australe
TEN.), is different from the well known fern of the Alps which is also fou
rarely in Great Britain on limestone pavements; var. australe is distinct -
116 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963
ts ecology, appearance and morphology and there is maybe a good case for
elevating it to specific rank, as was done by Guadagno. I did not find
Phyllitis hemionitis on Monte Corrasi but it was growing in small pits eroded
out of a huge limestone boulder near Ittiri; very small plants of Asplenium
— had also settled there and as the habitat was so uncommon I
thought at first I had found A. petrarchae, up till now not recorded from
Sardinia; alas, no, but it could be found there as it is known from Sicily and
the coast near Salerno.
The piéce-de-résistance of our trip, the climbing of Monte Gennargentu,
proved to be arduous and long. We were guided by an old herbalist from
Desulo, where the women still wear their vivid red and yellow costumes in
ordinary life. Blechnun spicant and Asplenium septentrionale emphasised the
northerly atmosphere of this old volcanic mountain but the most striking
feature was Athyrium filix-femina forming broad vivid-green splashes in the
shade of alders along streamlets rushing down to warmer places.
For me it is always a thrill to find Jsoetes. Sardinia is a well known
locality for these plants and we found two amphibious and one aquatic
species. Between Monti and Nuero, shallow hollows in the meagre pastures,
still damp in May but certainly dry in midsummer, produced J. hystrix and
an unidentified species. The only freshwater pool we saw north of Alghero
was fringed with numerous thriving plants of J. velata among lush water-
plants and algae; in Northern Europe Isoetes is strictly confined to oligo-
trophic waters but this pool was strongly polluted with cow and sheep dung;
the amphibious species were found in heavily grazed land but Professor van
Steenis assured me that the vegetation was typical Jsoetes-association.
Apart from Isoetes, Selaginella denticulata which is very common and a
few plants of Equisetum ramosissimum, no other fern allies were seen.
Polypodium australe was common, P. vulgare grew high up on Monte
Gennargentu and Pteridium aquilinum was reminiscent of landscapes in Great
ritain.
In conclusion I thank Mr Clive Jermy (British Museum, Natural
History) and Dr D. E. Meyer (Botanical Museum, Berlin-Dahlem) for pes
help with Isoetes and Asplenium respectively and Professor and Mrs V:
Steenis for organising the trip and for their invaluable botanical advice bs
friendship
PAGE: HYBRID HEBRIDEAN HORSETAIL 117
BRITISH FERN NOTES
A HYBRID HORSETAIL FROM THE HEBRIDES
While collecting plants in Harris in the Outer Hebrides during the summer
of 1962 I had the opportunity of obtaining a number of horsetails, many of
which grew in profusion in certain localities in the area, mainly on roadside
verges and in ditches. The rainfall is sufficient in Harris to maintain over
most of the area a permanently waterlogged soil and the gradients steep
enough for there to be a continual run-off of peaty water, the streamlets and
roadside ditches seldom, if ever, running dry.
In a small area of one such ditch grew several dozen specimens of a
horsetail which appeared at first sight much like Equisetum variegatum. It
consisted of simple or sparingly branched stems (text-fig. 1) arising out of the
mud and growing prostrate over it for about half their length and then
curving upwards. All the stems were sterile, dark green and 6 to 18 inches
long. Closer inspection of the stem revealed a greater likeness to E. arvense
or E. palustre.
This small colony grew about 300 feet away from a larger colony of
E. palustre which had upright-growing stems mostly regularly branched and
which at this time of the year (late August) still bore withered cones. Within
a mile radius of this spot were also to be found Equisetum fluviatile, E.
sylvaticum and E. arvense and slightly further away E. telmateia, though only
arvense and palustre grew in any number.
appear almost identical with those of palustre. The first internode of the
branch is slightly longer than the stem sheaths. It appears to resemble
fluviatile, however, in the fact that the endodermis, which in palustre and
pani + i the whole set of vascular bundles, here surrounds
Iw OWL UE
each vascular bundle separately, there being no link between the bundles.
This condition is also reported in E. x litorale (arvense x fluviatile). The
ratio of the size of the central hollow to the diameter of the stem appears to
be intermediate between that of palustre and fluviatile (more nearly like
arvense). The branches are 4-5 grooved and hollow. The arrangement of
the silicified sclerenchymatous tissue differs from that of palustre, arvense OF
fluviatile. There is a large amount of this tissue at the ridges linking with a
smaller amount in the furrows between.
4 (1963)
GAZETTE: VOLUME 9 PART
BRITISH FERN
fi
} " 2b
i \
-sections of inter-
E Fig. - bing palustre X 7 eg Seine. shoots
| uvia (all at same ma tion)
\ t thes shoots (x 4) aE fluviatile
c: rhizome (x 2/3) b: E. palustr
E. palustre x fluviatile
d:n me send sheath. ry
Pe Gay as
PAGE: HYBRID HEBRIDEAN HORSETAIL 119
These features appear not to tally with those described for any species
and appear to be somewhat intermediate between palustre and fluviatile. It
may therefore be a hybrid, growing as a separate colony, its parents being
absent in the immediate vicinity. This plant clearly needs some further
investigation.
KING’S COLLEGE, NEWCASTLE C. N. PAGE
NAME CHANGES IN THE BRITISH FERN LIST
14.2 Asplenium billotii F. Schultz in Flora 27:807; 28:738 (1845). Manton
and Reichstein in Bauhinia 2:79-91 (1962) have shown that the
complex known lately as A. obovatum Viv. (incl. A. lanceolatum
Huds.) in Europe consists of two species, a diploid and a tetraploid.
The diploid has a mediterranean distribution and is the plant
described by Viviani from Corsica as A. obovatum. The tetraploid
is A. billotii, found in the British Isles and over a wider area on the
Continent, described from a French (Vosges) specimen; Manton
suggests it is an allotetraploid with A. onopteris as one parent.
True obovatum differs from billotii in having shorter, more tri-
angular pinnae with fewer and less distinct pinnules; it may turn
up in the Channel Isles or in the SW peninsula, but so far has not
been recorded from the British Isles.
18.2 Athyrium distentifolium Tausch ex Opiz in Kratos (Zeitschr.f.Gymnasien.
Prag.) 2(1):14 (1820) must we A. alpestre Clairv. (1811).
Clairville’s type specimen at Ziirich has been examined and
identified as A. filix-femina a. pics
21.6 Dryopteris carthusiana (Vill.) H. P. Fuchs in Bull.Soc.Bot.France
105:339 (1958). D. lanceolatocristata (Hoffm.) Alston is now
considered to be a possible hybrid between D. carthusiana and D.
cristata. The type specimen of Polypodium lanceolatocristatum
Hoffmann cannot be traced. Polypodium carthusianum Villars
Hist.Pl. Dauph. 1:292 (1786) is the next available name.
24.4,5 Gymnocarpium dryopteris (L.) Newman and G. robertianum (Hoffm.)
Newman in The Phytologist 4:371 (1851). Stipe anatomy, scale
and frond structure and chromosome number are distinct from
Thelypteris. Most pteridologists regard these species as probably
more correctly placed in Aspidiaceae.
BRITISH MUSEUM (NATURAL HISTORY) A. C. JERMY
120 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963
SHORT NOTES
PSILOTUM TRIQUETRUM IN BASUTOLAND
Psilotum is 7 a tropical epiphyte. Sometimes it grows wpa titty bie
and, as Manton ! says, “‘they are familiar objects in most botanic gardens
since they are gah easy to grow and, in addition, reproduce accede by
bulbils”’. This record of Psilotum triquetrum (P. nudum) growing non-
epiphytically in Basutoland, whose climate cannot be described as tropical,
noteworthy.
a Re mention by the Rev. J. Gomes, O.P., that P. triquetrum had
been seen growing in Roma Valley, Basutoland, possible locations on the
eastern escarpment of the Cave Sandstones flanking Roma Valley were
searched in November 1961; it was found in only one place (recorded on a
grid-map) at about 5650 feet altitude as a colony of eighteen stems arising
from deeply implanted mycorhizal rhizome in a cleft in a rockface in running
water. The infertile stems were more divided and had more rigid pointed
leaves than the seven fertile stems of the colony. Basutoland specimens of
P. triquetrum are in the herbaria of University College, Roma, Basutoland
and the British Museum (Natural History).
Other records of P. triquetrum in southern Africa include Port St. John’s
(Cape Province) and Pondoland 2, Natal 3, Southern Rhodesia and Ghana.
UNIVERSITY COLLEGE, BASUTOLAND D. MORGAN
LEAF-BASE PROPAGATION OF PHYLLITIS SCOLOPENDRIUM
C. T. Druery, in the third chapter of his British ferns and their varieties,
described leaf-base vegetative propagation of ferns, an invaluable meth
when for any reason division of crowns or growing from spores is impractical
or impossible. Phyllitis scolopendrium and its varieties are particularly
adaptable to this method which, over many years, I have used most
successfully.
Lift the parent plant, preferably in spring, shake off the soil, divide into
single crowns, trim each to three inches of stem and replant. The severed
portions of stem provide the material for propagation. Wash them
thoroughly. Starting at the bottom, cut away until plump frond bases about
an inch long are reached; detach them by carefully snapping off or by under-
8 eee
1 Manton, L., Problems of cytology and evolution in the Pteridophyta. 1950.
2 Jacot-Guillarmod, A., personal communication. 1961.
3 Sim, T. R., Ferns of pe Africa, ed. 2. 1915.
KAYE: LEAF-BASE PROPAGATION 121
cutting with a sharp knife. Cut off any roots, trim neatly down to green
tissue, wash thoroughly in cold water and spread them half-an-inch apart
over clean wet sand one inch deep in a clean lidded plastic box or a glass jar
covered with polythene tied on with rot-proof material (not rubber bands
as these soon perish and snap). Place the box or jar in a light but shady
position away from direct sunlight. After a month or six weeks up to about
twenty pimples should appear on each leaf-base, to develop into tiny plants.
When the plantlets are well rooted, open up the container, detach them and
prick out into boxes or pans of open compost; keep covered for a few days
before gradually hardening off; grow on until large enough to prick out
again further apart or into thumb pots. At no stage must the plants be
allowed to get dry. The following year the plants should be big enough to
put into a shady border and in two years may be handsome subjects.
From one original plant I have been able, in one operation, to raise
hundreds of progeny which, being vegetative fragments, truly bear all its
characters.
SILVERDALE, LANCS, REGINALD KAYE
BOOK NOTES
WELSH FERNS — A DESCRIPTIVE HANDBOOK by H. A. Hyde and
A. E. Wade; 4th ogra pp. ix, 122, 12 plates. National Museum of Wales,
1962. Price 15 shillings
Since the publication of the first edition of Welsh ferns in 1940 this work
has been a handbook for all interested in finding, identifying and growing
ferns. This latest edition has been fully revised and rearranged, with two
exceptions, according to J. E. Dandy, List of British vascular plants which
incorporates the views of the late A. H. G. Alston; the authors recognise
Gymnocarpium as a genus distinct from Thelypteris on both cytological and
morphological grounds, although they do retain it in Thelypteridaceae
despite other opinions of closer affinity to Aspidiaceae; the Polypodium
vulgare complex has been treated according to the work of Manton and
Shivas, published after Dandy’s List.
As in previous editions the Introduction (pp. 1 - 28) includes an excellent
account of the life-history of the Malefern and of fern morphology and
geography. In the remaining pages 23 genera in 13 families are treated
species by species. There is a key to genera and there are keys to species ;
122 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963)
y 4 - | 1 W 1 + 1 oon + + mh 4 iL. rtare
of most species; the text gives a full description of the species, its habitat
and distribution, first and foremost in Wales but also in the British Isles and
the World. In spite of the title the book covers the fern flora of the British
Isles; those species not occurring in Wales are described in smaller type and
are included in the keys but are not illustrated; however, Wales has such a
good fern flora that this means only 6 species out of the 51 described have
had to be dealt with in this way.
The biggest disappointment is that the publishers have not seen fit to
include horsetails, quillworts and clubmosses in this new edition, to make
the work a concise treatment on Pteridophyta. The price has been doubled,
mainly due to large parts that needed re-setting; the binding is the stouter
standard round spine type, but there was something to say for the flush cut
boards and cloth spine of the third edition when it came to packing it into a
rucksack. Nevertheless it is very good value and its attractive dustcover
and crisp typography throughout make it a neat book and once bought I
doubt whether it will spend much time on the shelf.
THE FERN GUIDE by Edgar T. Wherry; 318 pp. Doubleday Nature
Guide No. 9, New York 1961. Price 3.95 dollars.
This book by a former president of the American Fern Society covers those
ferns and fern allies found in NE and midland U.S.A. and adjacent Canada;
135 species are described and illustrated with line drawings. I thoroughly
recommend this to British fern enthusiasts; Professor Wherry, also an
expert gardener, includes cultivation notes for each species; there are many
that would be worth growing in the British Isles.
HOW TO KNOW THE FERNS by Frances T. Parsons; “* second edition i
215 pp. Dover Publications, New York; Constable, London, 1961. Price
1.25 dollars.
With the exception of the omission of 6 photographs, this is a reprint of the
1899 edition published by Scribner & Sons. 57 North American species,
ouped into 6 categories based on position of sporangia, are described and
illustrated with 103 line drawings and sketches. In spite of some out-of-date
nomenclature and a restricted use in the British Isles, this book by a fern
lover of older days makes interesting reading with its many observations
from living and dried specimens and from literature.
A. C. JERMY
CRABBE: REQUESTS 123
REQUESTS
PILULARIA GLOBULIFERA
Pillwort or Peppergrass is sporadic in its known distribution although, being
easily overlooked because of superficial resemblance to small grass or rush,
it may in fact be more evenly distributed than the records show. It is
spasmodic in its growth, often perennating for years by means of a tough
impervious indusium around each of the four sori within each sporocarp
* pill ”’
It is usually found in soft sandy acid soil at the margin of standing
water, when it usually bears one-eighth-inch spherical hairy sporocarps, at
ground level. Sometimes it is submerged and is then mostly without fruit, but
€ circinnate vernation of the unrolling threadlike leaves distinguishes it
with watchspringlike leaftips.
Fresh material, packed in plastic with no loose water and sturdily
wrapped, together with details of locality, habitat, etc., is required for a study
of the biology of this species. If requested, I would gladly refund any postage.
BRITISH PTERIDOLOGICAL SOCIETY
Although the bones of the Society’s history are reasonably well known, finer
details of flesh and feature are obscure. Apart from several facts scattered
through the British Fern Gazette, some of which are mentioned on pages 135-6
of this number, there are many episodes which must have been interesting.
For example, CHARLES THOMAS DRUERY (1843-1917) crossed swords
somewhat with the Linnean Fern Chapter (later the American Fern Society)
whose Fern Bulletin (later the American Fern Journal) was virtually the
progenitor of the British Fern Gazette. Who was WHILHARRIS who col-
near Glastonbury in the 1890s (Gazette 5:93;7: 33)?
How and where did the West of Rapid Pteridological Society operate ?; was
it centred partly at Chislehurst with GEORGE BUCHANAN WOLLASTON
(1814-99) and partly at Clifton, Bristol with Colonel ARTHUR MOWBRAY
JONES (1826-89)? Is there any detailed record of Colonel Jones’s method of
natureprinting? What sort of people were the well known names of the
Society, several of whom are mentioned on pages 97-108 of this Gazette?
Biographical and genealogical clues must surely lie in such places as local
newspapers and flyleaves of family bibles and fern books.
Any information, cuttings, photographs, etc., would be most useful to
me for compiling a history of our Society.
BRITISH MUSEUM (NATURAL HISTORY) LONDON SW7 J. A. CRABBE
124 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963)
THE SOUTHPORT FLOWER SHOW 22-24 August 1962
Professor Holttum writes:
This was my first visit to Southport so I have no previous experience
from which to make comparisons. The day was sunny but with a cool and
blustery sea breeze which I met as I walked along the esplanade admiring
the spacious and varied gardens and recreation areas into which the coastal
belt of sand-dunes has been transformed. At the Show I soon found my way
to the Society’s stand in tent No. 2 where I received a warm welcome from
Messrs Hayhurst, Jackson (senior and junior) and Robinson, who had
brought a varied selection of plants to attract visitors. Another of our
members, Mr R. Kaye, had his own exhibit, including ferns, in another part
of the Show; he was one of the judges of the fern classes.
Messrs Brookfield’s plants in Class 8 were suitably varied and of a high
general standard, but I felt that the whole presented the effect of a collection
of plants rather than a group arranged to give unity of design, admittedly
a difficult effect to achieve and perhaps I am unduly critical.
In general, I was impressed with the excellence of the plants and by the
fact that they were collectively so representative. The following notes on
individual species and varieties may reflect my own particular interests and
prejudices. Perhaps I should preface them by a word about names. Most
members of the Society are probably better versed than I am in the
intricacies of varietal nomenclature, but I doubt if anyone now remains who
has a knowledge comparable with that of Charles Druery and other founder
members of our Society or whether it is really possible to name every plant
precisely in accordance with Druery’s list; it needs experience to know how
much variation may occur in a single plant. After returning from Southport
I spent a morning looking over the books of Moore and Druery and also the
earlier issues of the British Fern Gazette, supplemented ig consulting Thomas
Moore’s fern herbarium which, of course, does not contain some later
varietal names listed by Druery. I found almost all a names given to
exhibits at Southport, but I am not sure that all were properly applied; and
there were some exhibits which had no varietal nam
The plants which I found most attractive were ses Polystichums, which
included a fine example of what I regard as the most elaborately elegant
V:
deltoideum (see frontispiece of Gazette vol. 5, no. 12, June 1929). There
were others of the divisilobum group which were apparently divisilobum
densum. There were also two good examples of both pulcherrimums, na
* Moly’s Green’ and ‘ Bevis’, and one named Wakeleyanum. Contrasting
with these, the rather sak open branching of a Polystichum labelled
gracillimum (is this P. aculeatum pulcherrimum gracillimum of the Gazette,
vol. 1, no. 10, Dec. 1911?) was very pleasing.
SOUTHPORT FLOWER SHOW 1962 125
There was also much excellent variety in the Polypodium exhibits,
though the much-dissected var. cornubiense (or elegantissimum) did not
come near the size of specimens in Moore’s herbarium. The largest Poly-
podium plant (from Brookfield) had no varietal name; it was like a large
P. interjectum (see Gazette vol. 9, part 3) with rather strongly crenate seg-
ments. There were also crested varieties and some fine forms of var.
cambricum.
Of Scolopendrium there were several varieties, but these all seem to me
to distort the form of the original species without adding anything in
elegance. Mr Jackson had one which showed repeated forking in a very
interesting way; it was perhaps var. conglomeratum.
There were several beautiful exhibits of the smaller Aspleniums, includ-
ing a remarkably well-grown plant of A. septentrionale shown by Mr
Jackson, a result of our excursion to North Wales in 1961. Mr Jackson
showed also a plant of the hybrid alternifolium.
Among many Athyriums I was struck by the open branching of A.
filix-femina var. Clarissimum and especially by A.f.-f. var. plumosum Drueryi
which to my mind is the most beautiful form of this astonishingly poly-
morphous species. There were a number of crested forms which seem to me
to distort the natural form of this species, destroying completely that graceful
elegance which is its prime characteristic.
Other native British species exhibited were Dryopteris villarii, Cystopteris
fragilis and C. dickieana, Woodsia ilvensis, Cryptogramma crispa and an
unusual form of Blechnum spicant with little difference between fertile and
Sterile fronds.
There were some fine plants among the greenhouse ferns and I was glad
to see these; in a cool greenhouse one can grow a large number of species
from warmer parts of the earth, some of great beauty and interest. I would
like to suggest especially that there are some other species of Pellaea which
are extremely elegant and not too large which would repay the attention of
members.
In addition to Adiantum cuneatum from tropical America and our
native A. capillus-veneris there were fine plants of an unnamed maidenhair
which was said to be ‘ Kensington Gem’ but I have not found that name
in the R.H.S. Dictionary of Gardening.
Pteris cretica was represented by an excellent plant of the typical form of
the species and one of a fine crested variety. There were single plants of
Cyrtomium falcatum, Asplenium bulbiferum, Onychium japonicum and
Phlebodium (or Polypodium) aureum and a very pleasing one of Davallia
canariensis. Nephrolepis was represented by typical N. exaltata and two or
three varieties, but none of the more finely dissected ones.
Altogether, a few of our members had put up a quite impressive show.
126 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963)
I hope that more will be fired by their enthusiasm and that this section may
once again grow to something like its former dimensions. Members from the
south of Britain would find Southport well worth a visit and they would
certainly be welcomed by our northern stalwarts.
N. Robinson our Northern Meetings Secretary reports:
From the interest shown by many of the visitors, their questions and
discussion, there is no doubt that the decision was a wise one to obtain space
for a British Pteridological Society bureau to display various items of litera-
ture, application forms for membership and copies of the British Fern
Gazette. Messrs N. H. Almond, N. Brookfield, B. Hayhurst, F. Jackson,
J. Jackson, R. Kaye and N. Robinson provided constant attendance at the
bureau in the marquee and at the exhibits in the fern tent and also supplied
ferns to decorate the bureau tables. The Society, which benefited by
applications for membership and by £1 5s. 6d. from the sale of ferns, ack-
knowledges with gratitude the support given by this group of enthusiasts.
A thunderstorm with high winds and torrential rain on the second day
b
exhibitors had performed a magnificent job in cultivating their plants to
exhibition standard. Except for a few spelling corrections and abbreviations,
fern names in the following list are as given on the labels.
CLass 8 = group of hardy British ferns, with accent on diversity and variation, arranged in a natural
anner = a decors 13 feet by 8 a and not higher than 6 feet above the ground: L
this class there has been only the most attractive entry of Messrs
F
Brookfield & So on apa a t o hold the Silver Newscunrt Challenge Trophy presented by
British Pteridological Socie:
9 Six dissimilar hardy British ferns with accent on variation:
lineata,
Ist Messrs Brookfield; Athyrium filix-femina ‘ Druery’, A. f.-f. fieldiae, Fas: ate ae
phan scolopendrium crispum, Polypodium cambricum hadwinii,
2nd =s«éB. Hayhurst; A.f.-f.cristata, A.f.-f.frizelliae capitatum, Phyllitis scolopendrium crispum,
Polypodium cambricum, "Polystichum setiferum imbricatum cristatum, P.s.iveryanum.
£ Dhwlliei dvi:
cLass 10 Three
Ist Messrs ‘Recckiadl: crispum, cristatum, winoiaeed
2nd N. H. Almond; crispum, cristatum, hebetatum
3rd B. Hayhurst; eS cristatum, ramocristatum.
CLass 11 Three distinct varieties of Polypodium vulgare:
Ist Messrs Brookfield; bifidomultifidum, cambricum hadwinii, ‘ Oxford’
2nd B. Hayhurst; cambricum hadwinii, ‘ Oxford’, pulcherrimum.
cLass 12. Three distinct varieties of Polystichum:
Ist B. Hayhurst; densum, divisilobum densum, gracillimum
2nd Messrs Brookfield; ‘ Bevis’, congestum, wakeleyanum
3rd N. H Almond wletmtisen tmehwlords, Baath saad " * Fi 1°
cLass 13 Three distinct varieties ° ee oa
1st B. Hayhurst; cris: *, percris
2nd N. H. aera ond; fe, frizelliae lig ace
3rd Messrs Brookfield; c stum, ‘ Druery ’, percristatum.
SOUTHPORT FLOWER SHOW 1962 127
cLass 14 Three distinct varieties of Dryopteris geal Thelypteris:
Ist B. Hayhurst; angustata, cristata, ramosissima
2nd Messrs Broo id; angustata, lineata cristata
3rd F. Jackson; cristata, linearicristata, ramocristata.
cLass 15 Three distinct species and/or varieties of Asplenium
Ist F. Jackson; alternifolium, ruta-muraria, septentrionale
2nd N. plain ruta-muraria, septentrionale, trichomanes
3rd Messrs Brookfield; trichomanes, trichomanes bipinnatum, viride.
CLass 16 Three distinct normal species of hardy British ferns
Ist N. Robinson; Blechnum spicant, Cryptogramma crispa, Report vulgare
2nd__—sCF. Jackson; Cystopteris fragilis, Dryopteris villarii,
3rd Messrs Brookfield; Adiantum capillus-veneris, Phyllitis ARNE Ss Polypodium vulgare,
CLASS 17 One British fern:
Ist Dr A. A. Wilkinson; Polystichum setiferum divisilobum densum
2nd B. Hayhurst; P.s.divisilobum densum
3rd F, Jackson; Athyrium f.-f.cristatum.
CLass 18 Six dissimilar greenhouse ferns:
Ist B ay gi Adiantum, C. ium fal Nephrolepis, N.exaltata, Onychium japoni
phe
2nd Messrs id; Asplenium _bulbiferum, Devaliia canariensis, Nephrolepis, Pellaea
sinter my Poinian lium aureum areolatum.
CLass 19 Three dissimilar greenhouse ferns:
Ist B. Hayhurst; Adiantum ‘ Kensington Gem’, oe asin N.plumosum
2nd = Messrs Brookfield; Davallia canariensis, Nephrolepis, Pte
greenhouse fern:
Ist Messrs Brookfield; Adiantum ‘ Kensington Gem’
2nd = M.. Plowman; Preris cretica
3rd B. Hayhurst; Adiantum ‘ Kensington Gem’.
REQUEST
SOUTHPORT FLOWER SHOW
Thanks to the help of our President and of the Northern Meetings Secretary,
it has been possible to present the foregoing account of last year’s Southport
Flower Show. The article on variation in polystichums on pages 97 - 108
emphasises how important are records of the ferns exhibited. A list of winners
can be obtained from the Show Committee, but details of exhibits means
Painstaking work by private individuals.
The Southport Show is one of the major events in the world of boris
culture and it is a source of profound regret that so few of us get to see it;
we have to rae on other eyes and pens.
In r that a similarly ample and accurate account may be given of
future “irs Shows, anyone who is fortunate enough to attend and can
offer help is asked to contact the British Pteridological Society’s bureau.
128 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963)
BRITISH PTERIDOLOGICAL SOCIETY
MEETINGS IN 1962
BRITISH MUSEUM (NATURAL HISTORY), LONDON, 17 March. 13 members
attended. Among many exhibits and informal talks H. J. Bruty spoke on
growing ferns from spores, illustrated with living specimens of young ferns
in stages from the prothallus.
SAVILL GARDENS, WINDSOR and TAYLOR’S NURSERY, BRACKNELL, BERKSHIRE
26 May. Led by F. J. Healey on a bitterly cold day 7 members visited the
Savill Gardens. Amid the colour of rhododendrons and other early-
flowering subjects were seen many British and foreign ferns growing happily
in the borders. In the afternoon we were most hospitably entertained by
Mr and Mrs Healey at their Bracknell Nursery where we were especially
attracted by two specialities, waterlilies in numerous pools and ferns in
extensive borders well protected by bamboo hedges.
SOMERSET and WILTSHIRE, 15 - 18 June. This joint meeting with the Botanical
Society of the British Isles was based on Wells in the Mendips. The first day
was spent anes the rich limestone gorge | at Ebbor and Leigh Woods in
the Avon gorge a the doubt
fully recorded Pobisstcliah braunii. Shapwick Heath National Nature Reserve
was visited on the second day and among the many ferns seen at this classic
locality was an enormous Osmunda with a circumference of fourteen feet.
South Somerset was the venue for the third day when Hamdon (Ham) Hill
near Montacute was visited. This rarely botanised area yielded all the usual
species, often in great abundance. The last day was devoted to the woods on
the Greensand at Stourton on the Somerset - Wiltshire border. Specially
noteworthy finds were the rediscovery of Thelypteris limbosperma in an old
locality and a fine colony of Equisetum sylvaticum at the edge of a plantation.
WHITBARROW and LONG SLEDDALE, WESTMORLAND, 14 and 15 July. F. Jackson,
J. Jackson, R. Kaye and N. Robinson spent the Saturday on Whitbarrow, the
long limestone hill running parallel to the Lyth Valley, an area noted for Red-
and Roe-deer and Damson blossom. Most of the lower slopes are boulder-
strewn woodland, much of the top is open moorland; there are limestone
pavements and evidences of past quarrying. Among the profusion of ferns
were extensive stands of luxuriant Gymnocarpium robertianum and Ceterach
officinarum. The weather was fine and 18 species of ferns were seen. On the
Sunday N. H. Almond and B. Hayhurst joined the party which visited Long
Sleddale, a long narrow valley running into the heart of the hills; many
hedgerows and walls were encountered before reaching fells and rocks; in
spite of a less favourable terrain and a blustery cold rainy day, 18 species of
ferns were again seen including some magnificent clumps of Cryptogramma
crispa. N. Robinson, the Northern Meetings Secretary, has also made
several excursions during the year, including day visits to Whitbarrow an and
BRITISH PTERIDOLOGICAL SOCIETY 129
Warton Crag and a long weekend at Troutbeck near Windermere; 14 species
of ferns were noted at Troutbeck
ROYAL BOTANIC GARDENS, KEW, SURREY, 21 July. Last year’s visit to the Kew
Ferneries proved so interesting that we included another for this year.
Despite reluctant holiday absences, a small party toured the Houses under
the expert guidance of H. J. Bruty (Foreman-gardener of the Ferneries),
enjoying the sight of so many varied ferns from so many parts of the world.
Mr Bruty, an experienced grower with a wide knowledge of ere —
ferns, answered our many questions with hi
MILL GREEN WOODS, INGATESTONE, ESSEX, 12 August. A party fot 15, including
members of the Essex Field Club, spent the day in this extensive woodland.
7 species of ferns were recorded in addition to plants of the hybrid Dryopteris
carthusiana x dilatata. Blechnum spicant, now in very few places in Essex,
was plentiful and some fine plants of Thelypteris limbosperma were growing
with it. Dryopteris carthusiana grew happily in wet places. Other species
noted were Athyrium filix-femina, Dryopteris dilatata, D. filix-mas and
stoennoes wa toroset Later i in the day i in ” an wood near Kelvedon Hatch we
foun dilatata, filix-mas and carthusiana.
AL EXCURSION, 8-15 September. The Cruachan Private Hotel,
Aberfeldy, Perthshire provided an excellent headquarters for visiting the
central scottish highlands. Our President and Mrs Holttum took an active
part in the daily expeditions. Dr J. Davidson’s intimate knowledge of the
district was of great help. Other members present were H. J. Bruty, L
Burnett, J. A. Crabbe, Mrs E. M. Dyer, T. A. Dyer, F/Lt P. Halligey,
A. C. Jermy, R. Kaye, Miss A. Sleep and Mrs M. R. Stuart. Welcome guests
were Mrs R. Kaye and their son Jeremy, Miss M. J. P. Scannell of the
National Museum of Ireland and Miss P. A. Sims of the British Museum
(Natural History). We were sorry our Secretary was not with us and were
grateful to him for making many prior arrangements. Other contributions
to the success of the excursion were the hiring of a minibus which provided
long distance and local transport, the efficient drying by the Proprietor of
masses of wet clothing and, for evening work, the free use of a large room
with ay tablespace and fanheaters for warming ourselves and drying our
specimen
The — expedition, on the Sunday, is memorable as a day of record
rainfall during a particularly wet summer. It was decided to inspect the
famous Birks of Aberfeldy and its waterfalls; perhaps no better description
can be given of the Glen of Moness than the poem which gives it its modern
name; to quote one verse: ‘* The hoary cliffs are crowned wi’ flowers, while
o’er the linns [waterfalls] joo burnie pours and, rising, weets wi’ misty showers
the birks of Aberfeldy’; Robert Burns did not mention ferns but they can
hardly have passed ri os to such a keen observer of nature. We set off
at 10 and by lunchtime had worked up the mile or so from the town to the
bridge over the main falls. 15 species of ferns had already been found includ-
130 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963)
ing Phyllitis scolopendrium on the Crieff Road bridge, Asplenium viride,
A. trichomanes less frequently on wet overhanging rocks and plentiful
Thelypteris phegopteris and Gymnocarpium dryopteris; in higher and drier
parts T. limbosperma was dominant. Also of interest were a few plants of
Wild Basil (Clinopodium vulgare); bushes of Sambucus racemosa among
groves of raspberry canes contrasted with the common elders and black-
berries of farther south. It was agreed that the linns were well worth the
climb; after lunch most of the party were glad to make their way back to the
hotel and dry clothes.
Better weather prevailed on Monday for our first mountain trip. The
mountains of the district are composed of mica schist which weathers to a
soil of high mineral content and largely accounts for the richness of the flora;
a most helpful chapter on this subject is that by John Raven entitled “* The
mica-schist mountains of the Central Highlands ” in Collins’ New Naturalist
Mountain Flowers 1956. The main road runs for 15 miles along the north
side of Loch Tay and at Edramucky a road runs up to a carpark at about
1500 feet. From there we walked up to the slopes of the west side of Lochan
na Lairige which, in recent years, has been considerably enlarged by damming
by the North of Scotland Hydro-electric Board. The activities of the Board
are rapidly putting maps out of date; not only have numerous similar large
reservoirs been formed but they are augmented by diverting streams in
neighbouring catchment areas through tunnels in the mountainsides;
interference on this scale must inevitably react on wild life and it will be
interesting to learn what changes occur. On the lower slopes Bracken and
Malefern soon gave way to Thelypteris limbosperma in sheltered positions
and small plants of Blechnum spicant in the turf, its fronds pressed close to the
ground. From about 2000 feet Lycopodium and Selaginella began to appear.
Piles of boulders frequently formed miniature caves whose shelter and
moisture had established a microhabitat similar to the damp woodlands
below; in such hollows were found Adoxa moschatellina, Asplenium viride,
Chrysosplenium alternifolium, C. oppositifolium, Cystopteris fragilis, Dryopteris
dilatata (montane, diploid-looking forms) and Oxalis acetosella. On exposed
rockfaces was Polypodium vulgare. Two plants of Botrychium lunaria were
seen in a small crevice. Polystichum lonchitis was fairly frequent and on one
ledge was a community of P. lonchitis, P. aculeatum and young plants; in
vain did we search for adult plants of the hybrid P. illyricum; material for
further investigation at Leeds University was collected. At this time of year
most alpine flowers were over and we were not at a station of the real rarities.
Nevertheless, Alchemilla alpina was so abundant that it often formed a turf,
Saxifraga aizoides was still in bloom and S. oppositifolia, Sedum rosea,
Silene acaulis, Thalictrum alpinum and Cerastium alpinum were plentiful.
Dryas octopetala was also found
On Tuesday we explored a similar mountain area in Glen Lochay.
The Hydro-electric Board had provided a road (from grid reference 27/
BRITISH PTERIDOLOGICAL SOCIETY 131
500370) up which the minibus climbed halfway to the summit, to a small
dam which diverts the Allt Dubhchlair. Above this dam the burn cuts
through a small ravine where birches have gained a foothold and in whose
shelter flourishes an astonishing range of plants in this otherwise barren spot;
thus, Pteridium aquilinum, Dryopteris borreri, Thelypteris phegopteris and
Gymnocarpium dryopteris were found alongside ferns of higher altitudes. After
lunch on the open moor the party split into two, particular note being taken by
one of them of Saxifraga stellaris, 4 alpine Salix, Draba rupestris, Triglochin
palustris, List data, A ia maritima, Poa alpina, Carex saxatilis and by
the other of both large and small forms of Dryopteris dilatata.
Wednesday was fine and was spent exploring woodlands along Glen
Lyon; two of the party went further to an alpine site on Meall Ghaordie
where Salix myrsinites was seen. The first stop was at Fortingall with its
famous churchyard yew said to be 3000 years old; the churchyard wall
provided our first Asplenium adiantum-nigrum of the day. Reaching the
Glen we stopped at Macgregor’s Leap where the river falls through a
narrow gorge; 12 ferns and 2 horsetails were found. Six localities were
visited along the Glen in one of which Dryopteris abbreviata was found.
At Carnbane Castle Malefern and Ladyfern were growing vigorously and
with variation and Corydalis claviculata was seen. On returning to Aberfeldy
we stopped to inspect the Tay Bridge built in 1733 by General Wade as part
of his military road through the highlands; Asplenium ruta-muraria and
A. trichomanes beautify the bridge and also the rugged stone pedestal of the
nearby Black Watch monument.
On Thursday we were joined by a local member, Mrs M. R. Stuart.
As the day boded fine it was decided to tackle Ben Lawers itself. Most of the
mountain and the area on the Loch Tay side is protected by the National
Trust for Scotland who have produced an excellent account entitled Ben
Lawers and its alpine flowers. The route taken, up the Lawers Burn, was
long and only the two youngest reached the summit; the first half mile
Parnassia palustris and Drosera rotundifolia. On reaching the National
Trust boundary the woods ceased and a long bare valley stretched ahead
with few ferns except Thelypteris limbosperma. At the head of this valley is a
small dam and from there to Lochan nan Cat is a boggy heath where were
Whortleberry and Crowberry, laden with ripe fruit, Cottongrass and other
marsh plants. Lochan nan Cat lies at 2300 feet and is surrounded on three
sides by a vast amphitheatre of cliffs rising a further 1000 feet and more;
these cliffs were our main objective; alpine plants abounded, Hollyfern
flourished, but we failed to find Woodsia alpina, On returning to the main
road tea was taken at the Lawers Hotel which has catered for generations of
botanists and has interesting mementos of some of the most famous; Urtica
urens was recorded right outside the hotel. In the evening, thanks to a local
enthusiast and to the local Camera Club in loaning a projector,
132 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963)
show of lanternslides including fern propagation by Bert Bruty, botanising
in Austria and Greece by Anne Sleep and Irish High Crosses by Maura
Scannell
was mainly wet and we went further afield to Glen Clova.
From the youth hostel at the head of the glen we were permitted to take the
Forestry Commission road up to the foot of the Fee Burn but, in spite of this
time-saving, only an hour-and-a-half was left to explore cliffs. and loose
screes; 13 fern species were found, including Athyrium distentifolium and
Dryopteris abbreviata. On the way back Polypodium interjectum was co
by the roadside. Before joining us, Mrs Stuart had found Hart’stonguefern
on a wall near Pitlochry.
On Saturday, the day of farewells and departures, the minibus party for
the South decided to detour through Aberdeen to see Cystopteris dickieana
still growing happily in its type-locality.
This was the first major excursion of the Society since the publication
of the Atlas of the British flora to which we provided 13 unpublished records |
(7 ferns, 2 fern allies, 4 flowering plants). For another society we recorded
frogs and toads, which were frequently encountered. In spite of intensive
search in known stations we failed to find Woodsia or Cystopteris montana.
A list is appended of the 23 ferns and 8 fern allies which were found; the
numbers refer to localities, details of which follow.
WG: we id Perth Date National
(O.S. 1” map, ser. 7, sheet 47) visited Grid Ref.
September /
1 MEALL NASAMHNA, 10 miles WofTyndrum 11 4833, 4933
2 ALLT DUBHCHLAIR, 10 miles NofTyndrum 11 5034
3 MEALL GHAORDIE, 12 miles NE of Tyndrum _—12 5141
(O.S. sheet 48)
4 Slopes on the W side of LOCHAN NA LAIRIGE,
5 miles N of Killin... EA 10 5939
5 Around GALLIN, GlenLyon ... ... ... 12 5345, 5445
6 INNERWICK, Glen Lyon TORIES 5847
7 LOCHAN NAN CAT area, Ben Lawers. ws. —Ss 13 6442, 6542
8 LAWERS BURN, Ben Lawers ey ee 13 6741, 6742
LAWERS BURN, lower wooded area ... son 13 6740
10 INVERVAR, Glen Lyo ee 12
1 CARNBANE CASTLE, GlenLyon . nore OEE 6748
12 MACGREGOR’S LEAP, Glen Lyon is 12 7247
13 FORTINGALL, 7 miles W of Aberfeldy in wna 7447
14 BIRKS OF ABERFELDY it aha 9 8547, 8548
15 ABERFELDY os 1a Sel ae CS peas $548, 8549
V.C. 89 East Perth
(0. : sheet 49)
MOULIN, near Pitlochry et Li. det LBS Cie
17 By A.923 road, 3 miles Eof Blairgowrie... ... 14 1244
90 Angus (Forfar)
xo :: sheet 41)
FEE BURN corrie, Gla Clova . ii ivd 14 2574
BRITISH PTERIDOLOGICAL SOCIETY 133
Lycopodium selago — 1, 4, 7, 8, 13, 18
L. clavatum — 1, 4, 7, & 13
L. alpinum — 1, "4, ’s,
Selaginella utuplasideis | 4, 8, 13, 18
Equisetum fluviatile — 4
E, palustre — 1, 8
BE. — 8, 12
Pteridium aquilin — 2, 3, 5, 6, 8, 9, 10, 12, 14
Blechnum spicant — 1, 2, 3, 4, 5, 6, 7, 8, 10, 12, 14, 18
Phyllitis scolopendrium — 14, s.
-Asp! source nigrum — 42,23
A. trichomanes — 4, 6, 10, 11, 2 14
A. viride — 1, Ne 4, 14, 18
. Tuta. ep
Athyrium ot aH 6 *, 10, 11, 12, 14, 15, 18
e um —
ieana — y aan
Dryopteris filix-mas — ewe 10. it. “ 13, 14, 15, 18
D. borreri — 2, 5, 8, 9, 10,'11, 12, 14, 18
4 abbreviata — 10, 18
D. dilatai e. 4, 5, 7, 8, 9, 10, 1 i, 13, 14, 18
Polyatichum a Pinar ede 4,9, 12, 14
P. lonchi st, 3, 4,7, 18
Thelypteris i bosr — 1, 2, 4, 5, 6, 7, 8, 10, 12, 14, 18
T. phegopteris — 1, 2, 3, 4, 7; 9, 12, 14, 18
Gymnocarpiu: mew h 2) 3 S548
Polypodium vulgare — 1, 2, 4, 7, 8, 9, 10, 11, 12, 14
P interjectum —
ANNUAL GENERAL MEETING, 22 September, at the British Museum (Natural
History), —— A report has already been circulated and the financial
8 appear in a Newsletter. After business had ae
Professor et spoke on treeferns; we hope a presidential address ma
me a custom.
TUNBRIDGE WELLS, Kent, 23 September. Four people had a good day’s
botanising around Frant and Eridge Park. 12 fern species were found, the
Outstanding ones being Dryopteris aemula (one plant) and Hymenophyllum
Junbrigense (several colonies, one of them extensive and luxuriant).
ROBERT WHITESIDE
Robert Whiteside 1 who died in November 1960 aged 94 was one of the
founders of the Northern British Pteridological Society ne tonoreREE
Lakeland in 1891 2%. In his late teens he was handed his father’s garden
at Lancaster to look after and being anxious to have ferns in it he visited the
» Autobiographic details in British F. Gazette 7:227-230 (1949).
but it
® There British Pterido Society holding mectings in Kent in in the 1870s but it presumably
became defunct, ges nero de Wollaston) joined the 1891 Society.
134 BRITISH FERN GAZETTE: VOLUME 9 PART 4 (1963)
from the title of the Society which soon became a focal point for fern.
enthusiasts throughout the British Isles.
Whiteside in his younger days had great physical energy and he led a
busy life; he owned and managed a laundry at Lancaster, took part in local
affairs and was several times president of the local horticultural society. A
serious heart attack in 1941 led to his retirement from business and his
moving house to Morecambe whence he had an uninterrupted view across.
the bay of some of the Lake District mountains he knew so well.
I gratefully acknowledge Whiteside’s strong support given to me and to
the Society when I had to take over the Secretaryship in disturbed conditions
on the death of Dr Stansfield in 1937.
P. GREENFIELD
SPORE AND PLANT EXCHANGE
AMERICAN FERN SOCIETY. Neill D. Hall, 1225 East 95 Street, Seattle 13;
Washington, U.S.A. has been appointed National Director of the Spore
Exchange of the American Fern Society which invites botanic gardens,
universities and individuals throughout the world to participate. The
current list of available spores includes many rare and choice ferns. Our
Secretary (J. W. Dyce) will be pleased to supply a copy of this list to
anyone interested.
UNIVERSITY OF HULL. The Department of Botany, The University, Hull,
VY 1 M4 ps 4 e
+
CACHALIEL.
UNIVERSITY OF MALAYA. Edmund A. Turnau, Department of Botany,
University of Malaya, Pantai Valley, Kuala Lumpur, Malaya will be
happy to supply local material to those interested in the ferns of Malaya-
STUART L. WILLIAMS, 664 Rochester Way, Eltham Park, London SE9 would
like spores of the Japanese hardy form of Adiantum flabellulatum (sy2-
A. amoenum).
ie Sasi
* T. Bolton, J. Garnett, J. Gott, J. J. Smithies, J. Stewardson, R. Whiteside, G. Whitwell, J. A- Wilson,
W. Wilson, J. Wiper.
BRITISH PTERIDOLOGICAL SOCIETY 135
ADDITIONS TO MEMBERSHIP LIST
JULIAN J. BRIGHTMAN, 2 Red Oak Close, Orpington, Kent
Prof. J. COLHOUN, Dept. of Botany, The University, Manchester 13, Lancs.
Mrs G. CROMPTON, Thriplow Farm, Thriplow, Royston, Herts.
EKKER EN DEMANN’S WETENSCHAPPELIJKE BOEKHANDEL, Voorburgwal 243,
Amste ase C, Holland
Mrs J. D. EVERARD, 152 Sheaveshill Avenue, London NW9
F. FINCHER, Randan Wood, Woodcote, Bromsgrove, Worcs.
J. C. GARDINER F.C.A., 7 Stanhope Terrace, London W2
ERIC GOWING-SCOPES, Rosewood, Stonehouse Road, Halstead, Kent
GRAY HERBARIUM Library, Harvard University, 22 Divinity Avenue, Cambridge 38,
Mass., U.S.A.
Mrs J. F. HALL, 6 John’s Close, Gorsewood Road, Hartley, Longfield, Kent
P. F. HUNT M.Sc., F.L.S., The Herbarium, Royal Botanic Gardens, Kew, Surrey
INDIAN BOTANICAL SURVEY, The Chief Botanist, 14 Madan Street, Calcutta, India
A. J. KELLY, 7 Tudor Road, Wheathampstead, Herts.
Mrs E. M. LLOYD, 47 pes Street North, Hamilton, Ontario, Canada
LONDO socieTy, The Botanical Reading Circle Secretary, 28
Sietherinatoet Road, ppc tim Sw4
Prof. J. A. MACDONALD, Botany Dept., The University, St. Andrews, Fifeshire
R. MAUDE, 72 Hampton Road, Southport, Lancs.
J. N. MILSUM O.B.E., F.L.S., Grays, Tilford, Farnham, Surrey
Dr ALols PATZAK, Botanische Abteilung, Naturhistorisches Museum, Burgring 7,
Vienna I, Austria
G. R. PROCTOR B.A., Botany Dept., Institute of Jamaica, Kingston, Jamaica
Miss T. M. RICHMOND, Richmond Hill, Silverdale, Carnforth, Lancs
Dr ALBERT SCHUMACHER, 8 Eichbornweg, 522 Waldbréll, Germany
WALTER SCOTT junior, Easterhoull, Scalloway, Lerwick, Shetland
4. E. M. SHAKESHAFT B.Sc., 26 Moon Street, Wolverton, Bucks.
P. W. STRACHAN B.A. (Cantab.), 71 The Mall, Southgate, London N14
JOHN SUMMERTON, 81 Leeds Road, Tadcaster, Yorks.
KENNETH TREWREN, 22 St. Barnabas’ Road, Linthorpe, Middlesbrough, Yorks.
EDMUND A. TURNAU, Dept. of Botany, University of Malaya, Pantai Valley, Kuala
“poco Malaya
Mrs B. WELCH, 203 Lichfield Court, Richmond, Surrey
P. W. WOOD, 9 Slievemoyne Park, Newcastle, Co. Down, Northern Ireland
£) an
os aN
tf pate Lt! ise eeve
near
We have a very comprehensive
collection of
BRITISH FERNS & THEIR VARIETIES
We also specialise in
Alpine and Herbaceous Plants
Dwarf Trees, Flowering Shrubs
House Plants, Water Lilies
and Greenhouse Plants.
CATALOGUE ON REQUEST
REGINALD KAYE LTD.
SILVERDALE, LANCASHIRE
We grow
HARDY FERNS
British and exotic, of garden value
We should like to send you our catalogue
and we are always ready to consider
buying good garden varieties which
we do not already possess.
TAYLOR’S NURSERY
BRACKNELL, BERKS
THE
BRITISH FERN GAZETTE
VOL.9 PART 4 1963
Contents
Variation in Polystichum in the British Isles ... 3. W. DYCE 97
Meiosis in Asplenium X murbeckii ... eee J. D. LOVIS 110
Ferns of Sardinia ie a .. G.J. DE JONCHEERE 114
British fern notes:
A hybrid horsetail from the Hebrides... ... C. N. PAGE 117
Name changes in the British fern list... A. C. JERMY 119
Short notes:
Psilotum triquetrum in Basutoland aK D. MORGAN 120
Leaf-base propagation of Phyllitis scolopendrium R. KAYE 120
Book notes ... 121
Requests:
Pilularia globulifera aa = ae J. A. CRABBE 123
British Pteridological Society... .... J. A.CRABBE 123
Soutlipecst Flower Show: -35 oe ae ee 127
Polystichum varieties oS “Se a = we he 109
‘The Sduthport Flower Soe Sw a ae 124
British Pteridological Society:
Met W196) a ee oe 128
Obituary: Robext Whiteside. = Sie a ae oe 133
Sporé and plant ceachange.c = GS a a 134
Additions to membership list > ..5. ss She we 135
Courier Co. Ltd., Tunbridge Wells, Kent
iV
THE
BRITISH FERN
GAZETA2
ee
VOLUME NINE PART FIVE
1964
EDITED BY A; C-FERMY
ASSISTED BY J. A. CRABBE
Missour, BOTANIcar
SEP 19 1966
GARDEN LiBRARY
THE BRITISH
PTERIDOLOGICAL SOCIETY
4
BRITISH PTERIDOLOGICAL SOCIETY
Officers and Committee 1963-64
PRESIDENT
Reginald Kaye Esq.
VICE-PRESIDENTS
T. H. Bolton Esq.
Dr J. Davidson M.B., CH.B., F.R.c.P. (Edin.)
. Greenfield Esq.
Professor R. E. Holttum M.A., SC.D., F.L.S.
T. D. V. Swinscow M.B., B.S., M.SC., F.L.S.
SECRETARY AND TREASURER
J. W. Dyce
Hilltop, 46 Sedley Rise
Loughton, Essex
EDITOR
A. C. Jermy B.SC., F.L.S.
Department of Botany
British Museum (Natural History)
Cromwell Road, London SW7
COMMITTEE
. F. Cartwright, J. A. Crabbe, T. A. Dyer, P. Halligey,
B. Hayhurst, Mrs J. R. Healey, F. J. Healey,
F. Jackson, R. M. Payne, R. S. W. Pollard, N. Robinson
AUDITOR
Peter Temple L.R.1.B.A., M.INST.R.A.
The SOCIETY is open to all interested in any way in FERNS and FERN ALLIES”
upon paying an annual morn of SEVENTEEN SHILLIN'
Full details will be sent upon application to the SECRETARY: d :
BRIT. FERN GAZ, 9(5), 1964 137
DRYOPTERIS ASSIMILIS s.wALKER IN BRITAIN
S. WALKER and A. C. JERMY
Dept. of Genetics, University of Liverpool British Museum (Natural History)
The second edition of the British Flora (Clapham, Tutin and Warburg, 1962)
contains the description of a species of Dryopteris (viz. Dryopteris assimilis
S. Walker) that may on first sight be new to many British botanists. It is
however the description of a plant known to fern enthusiasts as Dryopteris
dilatata var. alpina, first described under Lastrea by Thomas Moore in 1856
from specimens he had collected on Ben Lawers, Perthshire, a year earlier.
These specimens Moore placed in his Herbarium (now at the Royal Botanic
Gardens, Kew) on three sheets labelled in his hand “ Ben Lawers, Perthshire
1855 T.M.”; a photograph of one of these sheets is shown on PLATE | ee
typical mature frond from one of these sheets has been designated lectotype
by one of us (A.C.J.); a silhouette of that frond, and a drawing of a spore
from it is shown in Fic. la & b.
In the course of a cytogenetic investigation of the Dryopteris spinulosa
complex in Europe, the results of which are published elsewhere’, Dryopteris
dilatata var. alpina from Ben Lawers (see Fic. 1c) was examined and found to
be a diploid (2n=82). Furthermore, artificial hybridisation and the study
of chromosome pairing at meiosis showed this diploid to be one of the
parents of the allotetraploid D. dilatata var. dilatata, the more common
form. A frond silhouette and a drawing of the spore of the tetraploid is shown
in Fic. 2. It will be seen that the basal pinnule on the lowermost pinna Is
shorter in proportion to the length of that pinna than the corresponding
pinnule of assimilis (cf. Fic. 1c). The difference in spore outer-wall morpho-
logy between diploid and tetraploid D. dilatata agg. was pointed out by
Fern Ward Crane?; those of the tetraploid (D. dilatata sensu stricto) appear
darker in colour because of the thicker and denser spinules. Hybrids between
the two cytotypes are sterile. Because of these findings it was decided to
treat the diploid as a separate species and the name D. assimilis was given to
the plant originally called Lastrea dilatata var. alpina by Moore. The
description (with diagnostic characters in italics) that follows enlarges upon
that given by Moore:
* WALKER, S., 1955, Watsonia 3: 193-209.
* CRANE, F. W., 1955, Watsonia 3: 168-9.
Thé BRITISH FERN GAZETTE Vol. 9 Part 4 (pp. 97-135) was published 26 September 1963.
138 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
it % ‘oe
> “set
natty e yet
tw g (er
Ma: + Ae
i" ft!
t
a
pm)
Ic
FIGURE 1 Dryopteris assimilis s:.wALKER
la, b: Lectotype; ic: Cytologically d ined ial fi
Ben Lawer
WALKER & JERMY: DRYOPTERIS ASSIMILIS PLATEI
PEMEAREEM BOARS ARTES NECA
Bh a CLD FERRE
— MUSSEL BRIT ANS i
3
Lit wlan
CpLaerin
Lastrea dijatata var. al
an
Nakare Printed Ferns Gt Brit. B irel.
Sub tad. 12 (1886)
ee
A9 at fire st
le
PLATE I Drypteris assimilis Ss. WALKER Type Coll. Hb. Kew
WALKER & JERMY: DRYOPTERIS ASSIMILIS 139
Dryopteris assimilis S. Walker, in Amer.Journ.Bot. 48:607, footnote
(9 Aug. 1961).
Lastrea dilatata var. alpina T. Moore, Nature Printed Ferns of Great Britain
and Ireland, sub tab. 22 (sine figura) (1856); Handbook Brit. Ferns
3rd ed.:126 (1857); Nature Printed Brit. Ferns: 225 (sine fig.) (1859).
Lectotype and syntypes, Ben Lawers, 1855, T. Moore (K). Rhizome
short, erect or ascending, often with several crowns. Fronds 7-60cm
long; stipe pale brown darkening only at the base, scaly below, scales
ovate-subdeltate abruptly accuminate into long, often falcate, apex 4-4
of total length, light brown usually with well-defined dark brown central
stripe; blade 1-14 times length of stipe, light green, thin, linear-lanceolate,
often almost parallel-sided below, apex acute, bi-tripinnate; pinnae
narrowly triangular to linear, the lowermost one having a markedly
longer basal pinnule on the lower side which is about half as long as that
pinna; pinnules finely pinnatisect, with mucronate teeth. Sori situated in
two rows near the pinnule midrib, c. Imm in diam., with small indusia
which are glandular on the margin and soon becoming evanescent.
Spores 45-50n x 30-37, with thin pale brown perispore with a few
trow wings and widely spaced acute spinnules up to Ip in length
(cf. Fics. 1b & 2b).
Whilst Ben Lawers is the only locality from which material has been
cytologically examined, a morphological investigation of herbarium material
at the British Museum (BM) and Kew (K) revealed the following other
probable localities for D. assimilis:
v-c 98 Argyll, Oban, D. Clarke, , Her oore
v-c 100 Clyde Isles, Arran, E. Greatorex 1861, Herb. T. Moore (K)
Other material which was immature, but which on scale and frond characters
may possibly be D. assimilis came from:
v-c 88 Mid Perth, Aberfeldy, E. C. Tollemache, 1870 (K)
v-c 89 E. Perth, Glen Beg, 7. J. Foggitt, 15:7,
v-c 99 Dunbarton, Ben Voirlich, Balfour, 1845; (BM) ‘ane
v-c 104 N. Ebudes, Hallaig, Raasay, M. McCallum Webster, 20.6.
(BM
Plants from all these localities require further investigation and we should
be pleased to receive material from any botanist who visits them. Livil
specimens are best sent with the rhizome wrapped in polythene together with
damp moss. If, however, the number of plants in the locality is small, only
pressed fronds, preferably fertile with mature spores, should be sent.
Mr H. V. Corley has provided some interesting plants from a number of
localities in the country and has so far revealed a diploid in at least one new
locality. Following up reports from Mr Corley, one of us (S.W.) has recently
140 BRITISH FERN GAZETTE: VOLUMES PART 5 (1964)
cm. 7
\ A.
Brit 4 w
Py j
+. eth
3 f I
reo eg,
,
fe Re os - ?
b ay AA
rn =<
\
. \ 14 ; ~
_ p ae - ?
= if
es ah ap. ee
FIGURE 2 Dryopteris dilatata (Hoffm.) A
Tetraploid plant from Bowland Knotts, Yorks,
10
|
Gray
visited areas in Ross-shire, Argyll-
shire and Perthshire and sampled
a number of populations. Plants
have been brought back for
further study.
An initial reaction to a more
concentrated search for the diploid
D. assimilis is that it may be more
variable in morphology than at
first thought. This is of course the
situation found in tetraploid D.
dilatata where variation is extreme
in different habitats and often
within the same population. The
diagnostic and especially the spore
characters of D. assimilis are seen
in other varieties of Lastrea
(Dryopteris) dilatata described or
determined by Moore and now in
his herbarium (viz. ornamentata,
lawersensis and micromera) and it
is possible that these may prove to
be variations within assimilis.
The re-discovery of these plants
in the wild and their subsequent
cytological examination can be the
only answer.
We should like to thank the
Director of the Royal Botanic
Gardens, Kew, for allowing us to
examine the Moore specimens.
BRIT. FERN GAZ. 9(5), 1964 141
A DIPLOID FORM OF ASPLENIUM RUTA-MURARIA
J. D. LOVIS
Department of Botany, University of Leeds
T. REICHSTEIN
Institut fir organische Chemie der Universitdt, Basel
In the last issue of this journal!, an account was given of meiosis in the
example of Asplenium x murbeckii found by F. Jackson on Castle Crag,
Borrowdale®. A. x murbeckii Dérfler is the hybrid between A. ruta-muraria
L. and A. septentrionale (L.) Hoffman. At the time that note was written,
only tetraploid chromosome numbers were known for either A. ruta-muraria
or A. septentrionale in any part of their geographical ranges*. An unex-
pectedly high degree of chromosome pairing was found in the Castle Crag
hybrid, and the possibility was considered that all the bivalents present were
in fact produced by autosyndesis, in other words, that the bivalents formed
in this hybrid are all produced by chromosome pairing between genomes
contributed by the same parent, and not by pairing of chromosomes con-
tributed by different parents. If this were true, it would imply that ct
forms referable to A. ruta-muraria and A st certain
existed in the past and might indeed still be living today. Most remarkably,
as has already been briefly reported elsewhere*, in rapid succession diploid
forms of these two species have now been discovered.
The diploid form of Asplenium septentrionale is known at present only
from the Caucasus, and has not yet been studied so sufficiently as to justify
taxonomic description at this time. No specimens of this plant have yet been
detected from the area of Europe as defined for the purposes of Flora
Europaea.
The probable existence of an extant diploid form of Asplenium ruta-
muraria became evident to us after a putative hybrid between A. lepidum
C. Presl (another Peg ie species®, *) and A. ruta-muraria detected by one
of us (T.R.), in the possession of Herr Th. Egli of Canton Zurich, had
Proved to be triploid and not tetraploid, as is usual for this hybrid®. This
Plant had originally been collected by Herr Egli from the Buco di Vela near
Trento in N Italy, and subsequent cytological examination of a specimen of
, LOVIS, J. D., 1963, Brit. Fern Gaz. 9: *
3 * TaCKSoN F., io, re Fern Gaz. 9 eee ne ‘ie: wel
‘ON, I, 1950, Pri Cytology and Evolution in the Pterido, MEYER, D ,
_ Deutsch. ie Ges. yr STs le of Coe 2 H., Jr., 1955, Rhodora §7:219 (A. ruta-muraria ia, U.S.A
= 72); BIR, S. S. in MEHRA, P.H., 1961, Res. Bull.Pi anjab Univ., n.s., 12:139 (A. septe
Himeley ya, n=72).
LOVIS, J. D. 1964, Nature, in press.
, LOVIS, 3. D .. and REICHSTEIN, T., unpublished.
MEYER, D. E., 1959, Ber. Deutsch.Bot.Ges. 72;37; 1964, ibid. 77:3.
142 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
A. ruta-muraria from this locality already in cultivation in Basel showed this
to be diploid. Since then we have found diploid plants of A. ruta-muraria in
situ in the Buco di Vela, and in other localities in the Italian Dolomites. At
present, this plant is not known outside the Italian Dolomites, although
it is quite evident that within this region it is rather common, and indeed
appears to be there the prevalent form of A. ruta-muraria, at least on native
rock. The diploid form of Asplenium ruta-muraria is described below as a
distinct subspecies (dolomiticum).
MORPHOLOGICAL COMPARISON OF THE SUBSPECIES OF ASPLENIUM RUTA-MURARIA
1 MACRO-CHARACTERS It is not easy to distinguish the diploid subsp.
ene from the tetraploid subsp. ruta-muraria on the basis of macro-
c ers alone. In both subspecies the shape of the pinnules is extremely
Laney we those of dolomiticum are usually smaller and more regular
in s innules of the latter are certainly more delicate and thin in
texture, and the pares and stipe are usually less stout than in ruta-muraria.
However, the most striking macro-character lies in the margin of the pinnules.
In living plants of dolomiticum the colourless margin of the pinnules is very
obvious. This clear border is of uneven width, being widest on the irregular
teeth at the tips of the pinnules, making these teeth relatively conspicuous.
e rhizome scales do not provide a reliable character, being peculiarly
inconsistent in both subspecies. Many collections of ruta-muraria Possess
markedly larger scales than those of any of the collections of dolomiticum yet
examined, but in some collections of both subspecies the rhizome scales are
quite minute and difficult to examine.
2 MICRO-CHARACTERS Under the microscope, the two subspecies may
be distinguished with relative ease by examination of either spores or stomata.
In both subspecies the length/breadth ratio of the stomata is variable, the
stomata being longer in etiolated fronds produced in deep shade conditions.
Consideration of stomatal length alone is consequently inadequate to
distinguish shade forms of subsp. dolomiticum from exposed forms of subsp.
ruta-muraria, In such circumstances there is, however, a marked difference
in the breadth of the stomata, sufficient to distinguish the two subspecies.
The figures obtained from measurements of stomata and spores are
given below; in each case, fifty spores or stomata were measured. Stomata
were measured in water, the pinnae having first been cleared with a hypo-
chlorite agent. The figures given for spore size are from spores mounted in
De Faure’s gum chloral. Only the exospore was measured, the perispore
being ignored. All the measurements given are obtained from material of
known chromosome number. Since the number of cytologically known
samples at present avatiabis i is very limited, it will be evident that the ranges
Pe ete entually
of size for the two P y eventua
LOVIS & REICHSTEIN: ASPLENIUM RUTA-MURARIA PLATE: £2
crevice of perpen
oe II Holotype of Asplenium ruta-muraria subsp. dolomiticum in situ in shaded
dicular dolomite vock wall tn V. Val di Uda above Mazzin, E of Bolzano, N Italy. Photo: JDL10.9. 1963.
LOVIS & REICHSTEIN: ASPLENIUM RUTA-MURARIA 143
require amendment when more cytologically determined material has been
examined.
TYPIFICATION OF ASPLENIUM RUTA-MURARIA L. AND DESCRIPTIONS OF SUB-
SPECIES Although the question of the typification of Asplenium ruta-
muraria L. has not been investigated in detail, it does not appear to present
the same difficulty as the case of Asplenium trichomanes L.?. It is already
evident that the diploid subspecies of A. ruta-muraria is very much rarer and
more local in distribution than the tetraploid. Very probably only the
tetraploid is found in Sweden. Consequently, in the absence of any contrary
evidence, it is a reasonable assumption that it is the tetraploid plant with
which Linnaeus would have been familiar, and which therefore best deserves
to be regarded as the type subspecies.
Asplenium ruta-muraria L., Sp.Plant. 2:1081 (1753), subsp. ruta-muraria.
Fronds relatively robust; rhachis comparatively stout; pinnules variable in |
shape, but usually somewhat - more angular, texture relatively thick, colourless
margin rarely conspicuous. Stomata 37-66» long (range of means 50-57,),
31-46 broad (range of means 35-42). Spores rg of exospore only in
chloral medium) 41-60» long (range of means 49-54), 33-46» broad
(range of means 37-42). Chromosome number n=72, 2n=144. Habitat in
fissures of basic rocks throughout Europe. Also in North America and Asia.
Asplenium ruta-muraria L. subsp. dolomiticum Lovis & Reichstein subsp.nov.
(vide figs. 1-9 et tab. 2).
Frondes delicatae, rhachibus B cesprovie” —— plerumque constanter
rotundis-rhomboidis, texturae conspicuibus pellucidibus
cicatricibus membranibus iets. ‘Stomata 35- 56 longee (ordo Send BE.
27-38 u latae Ser mediis 30-344)". Sporae (magnitudo exospori s
chlorale), 33-54 longae (ordo mediis 39-46 »), 25-42 latae (ordo mediis 29-39x)f.
romosomata gesnetophiyti 36, sporophyti
sea N ITALY, Trentino Alto Adige: Val di Uda, above and ca, 1km
NW of Mazzin (Val di Fassa), 25km ESE of Bolzano, ca. 1850m in crevices
of shaded perpendicular dolomitic rock, E. Hauser, J. D. Lovis & T. Reich-
stein JDL1963:87T, 10 Sept. 1963. Holotype plant (PL. II & Fics. 7-9) divided,
one half deposited in the herbarium of the British Museum (Nat. Hist.),
other half in cultivation in T. Reichstein’s garden in Basel, sub TR950.
Paratypes: N ITALY, Trentino Alto Adige: Buco di Vela, ca. 3km W of
Trento, ca. 450m on dolomitic rock, Reichstein TR19, summer 1956, cult. in
’ LOVIS, J. D., 1964, Brit. Fern Gaz. 9: 147
* Holotype 37°5-55:5u (mean 46-1) long, 29: 5-37-54 (mean 32-4y) broad
+ Holotype 39-5-53-5u (mean 45-64) long, 31-5-41-Su (mean 38-64) broad.
144 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
Basel; ca. 410m on same rock, Hause- Lovis & Reichstein JDL1963:97,
pa 11 Sept. 1963; Val d’Adige ca. 3km SW of Salorno, ENE of Trento,
. 230m on dolomitic rock-wall, Hauser, Lovis & Reichstein JDL1963:91,
Pied 964, 11 Sept. 1963; Monte Bondone, ca. 5km SW of Trento, ca.
1700m on dolomitic rock Hauser, Lovis & Reichstein JDL1963:99, TR966,
11 Sept. 1963. N ITALY, Venezia Giulia: N of Chiusaforte, Canale del Ferro
N of Udine ca. 450m on dolomitic rock, Lovis & Reichstein JDL1963:83,
ae 9 Sept. 1963.
Fronds delicate, with a slender rhachis; pinnules usually regularly round-
aaa texture thin, margin with conspicuous colourless membraneous
order. Stomata 35-56p long (range of means 41-49), 27-38» broad
(range of means 30-34). Spores (size of exospore only in chloral medium),
33-54 u long (range of means 39-46), 25-42, broad (range of means 29-394).
Chromosome number n=36, 2n=72. Habitat in fissures of dolomitic rock in
N Italy
TABLE I SPoRE MEASUREMENTS
Range Mean
subsp. dolom Length» Breadth » LxB
Mazzin, E of Bolsane, N tly wee 39-5—53°5 31-5-41-5 45-6 X 386
nr. Salorno, N of Trent hes 2 33-5-47°5 29-5-37°5 41-0X 33-1
nr. Chiusaforte ,Nof Udine, N taly ... 37°5-45°5 29-5-35-5 41-7 x 32:3
Buco di Vela, nr. Trento, N fo Ws 33-5-45-5 25-5-33°5 39:7 X 29-7
33 -— 54u 25 - 42 39-46 X 29-39
subsp. ruta-muraria
Chapel-le-Dale, Yorkshire, England ... 49-5-59-5 37-5-43-5 53:3 X 41:3
Kraubath, Steiermark, Austria 41-5-57-5 335-455 49-4 x 37-4
Convento’ Bigorio Tessin, Switzerland 41-5-57-5 33-5-41°5 49-5 x 373
pag Roof, Westinerta nd, England .. 45-5-55:5 37-5-41°5 50: ee
Varese, Nw of Milan, N Italy nee ie ay
41 - 60u 33 - 46u 49-54 x 37-42
TABLE II StoMA MEASUREMENTS
Range Mean
subsp. dolomiticum Length» Breadth» LxB
Mazzin, E of Bolzano, N Italy (TYPUS) 37:5-55-5 29-5-37°5 46-1X 32-4
nr. Salorno, N of Trento, N Italy 37:5-49-5 29-5-35-5 41-9X 33-2
nr. Chiusaforte, N of Udine, N Italy .. 37-5-53°5 = 27-5-33-5 48:2 x 30°9
Buco di Vela, nr. Trento, N Italy 33-5-53:5 29-5-37°5 47:7 X 33-9
35 - S6u 27 - 384 41-49 x 30-34
subsp. ruta-muraria
Chapel- o-Dale. Yorkshire, England .. 51:5-65°5 33-5-43-5 56-9 X 39-3
Kraubath, Steiermark, Austria 41-5-595 35-5-45-5 53-2x 41-1
Convento Bigorio, Tessin, Switzerland 41-5-59-5 31-5-39-5 54:2 x 35-4
Hutton Roof, Welmitent England .. ee see evs
, NW of Milan, N Italy |. 37-5615 33-$-43-5 50-8 x 39°8
37 — 664 31 - 46 50-57 x 35-42
FIGS. 1-9 Photo-copy silhouettes (reproduced —— size) of fertile fronds of Asplenium ag pig
subsp. ee, all pressed fom wild material. Fic. 1 nr. Chiusaforte, N of Udine. Fis. 2,
> nr. Trento. Fic. 6 Monte Scodoun SW of Trento. Fics. 7, 8 and 9 HOLOT YPE,
bg th worl Reichst tein SDLIs6s :87T.
LOVIS & REICHSTEIN: ASPLENIUM RUTA-MURARIA
145
FIGURES 1-9
146 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
SYNONYMY Correlation of Asplenium ruta-muraria ssp. dolomiticum and
the numerous minor taxa already described within A. ruta-muraria is a very
difficult task, one which will require much more investigation. Because of
the uncertainties that remain unresolved, no attempt is made here to indicate
synonymy. Thus, it is very likely that one or more of the varieties recognised
by Christ’, may refer to subsp. dolomiticum, but without reference to Christ’s
herbarium, which at present remains inaccessible, any attributions not based
on very careful study would be open to question.
Asplenium ruta-muraria is divided into three subspecies by Grintescu in
Savulesci’s flora of Roumania®. One of these, subsp. typicum, by application
of Articles 24 and 26 of the current International Code of Botanical Nomen-
clature, should be more correctly known as subsp. ruta-muraria. No
specimens or illustrations have been seen, but from the descriptions given, it
is clearly extremely improbable that either of his two new subspecies, subsp.
ponticum Grintescu, and subsp. grammitioides Grintescu, corresponds to
subsp. dolomiticum. In the case of subsp. ponticum, A. haussknechtii
God. & Reut. is cited in synonymy. Unless this attribution is entirely
erroneous, it is most interesting, for A. haussknechtii is a rather obscure
taxon described from Asian Turkey, certainly quite distinct from A. rufa-
muraria, but very close to A. samarcandense Koss. and having some affinity
with A. lepidum C. Presl. If A. haussknechtii is really concerned, then we have
here an indication of an addition to the Asplenium flora of Europe. What 1s
perhaps more likely is that the Roumanian plant really refers to A. lepidum,
not recognised for itself. It is relevant to note that A. /epidum has only very
recently been detected in Hungary!°, and thus might well grow unrecognised
in Roum
THE RAD - THE SUBSPECIES OF ASPLENTOM giedease ties The
nature t ies of Asplenium
peti 8: recognised i in this paper is a subject of obvious interest and is
now under active investigation.
It is a great pleasure to record here our grateful thanks to Professor I.
Manton, for her advice and encouragement. We also thank Herr Egli for
his kind co-operation.
$ anes H., 1903, ego 42:153.
GRINTESCU, G., in SAVULESCI, T., au -~ Repub.Popul.Romane 1 Add.: 634.
1° VIDA, G., 1963, Acta raf Acad.Sci.Hung. 9
BRIT. FERN GAZ. 9(5), 1964 147
THE TAXONOMY OF ASPLENIUM TRICHOMANES IN EUROPE
J. D. LOVIS
Department of Botany, University of Leeds
The existence of a species complex within Asplenium trichomanes L.
has been known ever since the publication in 1950 of Manton’s magnum opus
on the cytology of the pteridophyta, in which the discovery of diploid and
tetraploid chromosome numbers in European A. trichomanes is described.
For the last ten years, I have been engaged on an intensive biosystematic
Study of the complex, a study which is far from complete, and is still in
progress,
Preparation of the account of Asplenium for Flora Europaea made it very
desirable that an adequate taxonomic treatment of the complex be formu-
lated, representing as accurately as possible our present state of knowledge of
this complex as found in Europe.
In 1962 Meyer? described the two European cytotypes (diploid and
tetraploid), as distinct subspecies, and this treatment has been followed in
the Flora, as being consistent with the concept of subspecies as generally
applied therein. However, some amendments of Meyer’s nomenclature and
diagnoses are required, and are given in this paper. Very recently, too
recently for inclusion within Flora Europaea, another diploid taxon within
Asplenium trichomanes has been discovered in Europe. In direct contrast to
the strictly calcifuge character of the more familiar diploid, which is particu-
larly frequent in central and northern Europe, the new taxon appears to be
confined to limestone in central and south-eastern Europe. This plant is
described below as a new subspecies.
t +ha
In the final section of this paper th fused situatio:
recent publication of three new specific names tein the Asplenium
trichomanes complex is discussed.
THE TYPIFICATION OF ASPLENIUM TRICHOMANES L.
In describing the two cytotypes found in Europe as distinct subspecies,
Meyer? circumvented the very difficult problem of the typification of
Asplenium trichomanes L. by the expedient of giving new names to both
subspecies, a procedure directly contrary to Article 26 of the International
MANTON, I., 1950. Problems of Cytology and Evolution in the Preridophyta, Cambridge.
* MEYER, D. E., 1962. Ber. Deutsch.Bot.Ges, 74:455-6.
148 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
Code of Botanical Nomenclature. In so doing, Meyer in effect left the now
vexed question of the typification of A. trichomanes for subsequent authors
to decide, since one of these two subspecies must include the nomenclatural
type, and must therefore be regarded as the type subspecies. This subspecies
should carry the trivial (specific) epithet in repetition, i.e., it must be called
subspecies trichomanes.
Already, since the appearance of Meyer’s paper, contrary interpretations
gi
Fuchs produced a later homonym, and in an addendum to Fuchs’ paper
(.c., p.20), Sod (the editor of the journal Secee provided another
name for the diploid, adding the explanatory comment “ (A. trichomanes
auct.p.p. non L. herb.)””. As a reason for seers the choice of type
subspecies, this statement alone without any elaboration is not acceptable.
It is relevant to quote W.T. Stearn, an acknowledged authority on Linnean
specimens present cannot always be accepted as nomenclatural types; indeed
it would be highly misleading to rt uncritically a certain specimen in the
Linnean Herbarium as a type .
It is not my intention to get involved any further in arguments concerning
the typification of Asplenium trichomanes L. in this paper. To do so would
greatly increase its length, and distract from its main purpose. The typifica-
tion of A. trichomanes is a most complex issue, difficult to resolve. It is my
intention to consider this problem and to explain my opinion regarding the
typification of this species in a separate paper. For our present purposes I
think it is sufficient to state that I base my own interpretation on the guiding
principle that where a Linnean name covers a species complex, this name
should be retained by that segregate with which Linnaeus would have been
most familiar, and which would therefore most likely have represented to
* LANJOUW, J. (edit.), 1961. snlecsinayad Code of Botanical Nomenclature, Utrec
in
rk eS (l 436) j Hi aoe
vorkommt.. .”’ (l.c., p. |. He not appear to ha lised the / al co!
his argument would be that our A. viride must become i waumean can benioneeke te Spleenwort
must take another name! Fortunately, as I shall show elsewhere, there are good reasons for believing
Meyer’s basic premise unjustified,
*# ROTHMALER, W., 1963. Exkursionflora von sepin IV, Kritischer Erganzungsband, p. 5.
pci HY. 1963. ‘Acta Bot. ne Sci.Hungaricae 9;1
* STEARN, W. T., 1961, Taxon 10:1
LOVIS: ASPLENIUM TRICHOMANES 149
him the most typical form of his species. In the case of A. trichomanes, my
investigations have led me to conclude that this would be the familiar
iploid taxon widespread in Scandinavia.
THE SUBSPECIES OF ASPLENIUM TRICHOMANES RECOGNISED BY MEYER
Meyer? divided Asplenium trichomanes L. into two subspecies, subsp.
bivalens D. E. Meyer, and subsp. quadrivalens D. E. Meyer, corresponding to
the diploid and tetraploid cytotypes respectively. In order to produce an
acceptable taxonomic treatment of Meyer’s taxa, it is not sufficient merely to
substitute the name trichomanes for Meyer’s bivalens. In his diagnoses and
descriptions, Meyer gives only size, sculpture, and colour of spores, chromo-
some number and ecological preference for each subspecies. Thus, the only
characters available from his diagnoses for determination of herbarium
material are those of the spores. The qualitative characters of the spores in
my experience much too finely balanced and variable to be of practical value.
One is left with spore size alone, which will serve to identify positively a
considerable proportion of specimens, but there is an inconvenient overlap
between the ranges of mean length determinations for diploids and
tetraploids’.
The situation is aggravated by the fact that the ranges of spore size
given by Meyer and myself® do not agree. The degree of discrepancy is very
great, of the order of 30%. I would like to record here my gratitude to Dr
Meyer for his co-operation in investigation of this problem. I am most
indebted to him for sending me mature fronds from his cultivated type
specimens. Without this material it would not have _— Possible for me to
exclude beyond doubt the p lay in the
Plants themselves.
In the cir t ,Ih checked very carefully indeed the calieetos
of the particular micrometer .'
I use in making my measurements, and satisfied myself of ‘the accuracy of the
Calibration. Another obvious possible source of discrepancy lies in the
Medium used in making spore mounts. Measurements were made of spores
from the same frond of Asplenium trichomanes mounted in different media.
The results indicate that spores mounted in gum chloral or in glycerine jelly
give measurements of the order of 6% greater than spores mounted dry
(i.e., in air), and of the order of 7% greater than spores mounted in
Pure xylol or in balsam in xylol®. My own spore measurements are usually
" LOVIS, J. D., 19st. As evolutionary study of the fern Asplenium trichomanes. Ph.D. Thesis, Univ.
of ag un:
* LOVIS, J. D., 1955. 1 Bot,Soc.Brit,Isles pg ats 4:102.
: E was terme oes don Bad we There has not not yet been opportunity to discover
whether these di would be repeated if the spores of other er species and genera savemianees
but in the light of this experience, it is evidently very a ot tant ee farare descriptions of spore size
sheidd stwete eden a Dek ‘wood.
150 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
made in De Faure’s gum chloral, whereas Dr Meyer has informed me that
his measurements were made from dry-mounted spores. A further source
of possible discrepancy lies in different selection procedures used in making
measurements. I myself, when measuring spores in gum chloral, always
ignore spores that are obviously end-ways on or obliquely aligned in the
medium. Such a selective procedure is less easy to carry out on spores
mounted dry, because of the optical effect of the different refractive indices,
and in consequence I determined what difference results from measurement
of a sample using the selective procedure described above, as compared with
measurements derived from all spores, irrespective of orientation, only mis-
shapen or shrunken spores being excluded. My small pilot experiment
indicates a difference in mean of approximately 1-5-3-5%. It is clear that
neither difference in medium nor possible difference in selective procedure are
adequate to explain the discrepancy between Meyer's results and my own.
I can therefore only ¢ come to the conclusion that for some other reason Dr
The discrepancy between the published figures for spore size given by
Meyer and myself is of real practical importance, because Meyer’s figures for
the length of tetraploid spores correspond more or less to my own figures for
the length of diploid spores! It is clear that reliance on the figures for spore
size given in Meyer’s diagnoses can only result in confusion.
It is therefore necessary to provide an amended diagnosis for subsp.
quadrivalens, and the opportunity is taken to amplify the diagnosis at the
same time, since it is in my opinion undesirable to characterise a plant on
spore characters and chromosome number alone.
actual fact, it is usually possible to distinguish subsp. trichomanes and
sitbene quadrivalens without recourse to micro-characters. This is particularly
true of living plants, herbarium specimens being admittedly more difficult to
determine, especially if of indifferent quality. In subsp. trichomanes the pinnae
are usually small, suborbicular, distant, and delicate, in contrast to the
pinnae of subsp. quadrivalens, which are usually larger, oblong, relatively
crowded, and more massive except in very shaded situations, when the pinna
size and shape alone provide an adequate distinction. The suborbicular form
of the pinnae is obscured in luxuriant forms of subsp. trichomanes (see TEXT-
Fics. 1:3 and 2:6) by the development of a very characteristic auriculate or
subauriculate condition not found in subsp. quadrivalens. A further criterion,
particularly evident in living plants, is found in the insertion of the upper
pinnae, which is always oblique in trichomanes, but most often more or less
square in guadrivalens.
Rhizome scale characters can be useful, but only if applied with care.
os both subspecies, the rhizome scales display a wide range of development,
d useful comparisons are obtained only if all but the largest scales are
wesc It is then found that the scales of quadrivalens are longer, narrower,
LOVIS: ASPLENIUM TRICHOMANES
EEXT -FIGURE 1 Photo-c
Ps es 0 in cultivation at. y= “8 ss. Peas. 2 and 3
Pia. =e onshire, N Wales. Fic. 4 subsp. PEE Mes mt
subsp. quadrivalens, ex moesiey Glen, Ingleton, N
f Asplenium
a ph romnai ex Pam of Li pore
ex Aber Falls, es eg
W Yorkshire, Englan
rvonshire, N Wales
152 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
with the sige gripe | of dense cells darker in colour than in subsp. tricho-
manes, and g further into the tip of the scale?°.
Asplenium trichomanes L., Sp.Plant.2:1080 (1753), subsp. trichomanes.
A, melanocaulon Willd., Enum.pl.hort.reg. pts :1072 (1809).
A. trichomanes L. subsp. bivalens D. E. Meyer in Ber.Deutsch.Bot.Ges.
74:456 (1962).
A, trichomanes L., emend.Rothm., Exkursionfl.Deutschld. IV, Kritisch.
Erganz. :5 (1963).
A. trichomaniforme H. a ove in Acta Bot.Acad.Sci.Hungaricae 9:19
(1963) non Woynar (19
A. linnaei So6é ex H. P Packs loc. cit.:20 adnot (1963).
Rhizome scales >3-5mm, lanceolate, with red-brown central stripe.
Vien 8—18(-25) cm long, tapering towards apex, apex acute, with (10-)
pland regions hroughout Europe. "Also in North America and
[Melanesia pie eh 2]
Asplenium trichomanes L. subsp. quadrivalens D. E. Meyer emend. Lovis.
A, trichomanes L. subsp. ear lat D. E. Meyer in Ber.Deutsch.Bot.Ges.
74:456 (1963). nomen ambigu
A. lovisii Rothm., Exkursi on Deutechid. IV Kritisch.Erganz.: 5 (1963)
nomen nudum
A, fioen ae sensu H. P. Fuchs in Acta Bot.Acad.Sci. Hungaricae 9:20
1
P. rhizomati_
SYDOI sNOsIvI|ed-UOU
tL
19§-67
(8-) 9-4
nEy-8E
snonsidsuooul ‘moieu
enbijgo
yuej}sIp
jUBUNXN]
Udy dETNOLIN ‘ye;NdIqJOgGns
Sutiode}
87-FI OD
wd ($Z-) 8I-8
SOUDUIOYIIAJ
adOUNA NI NOLLAGRLLSIG
LVLIGvH
(UZ) YaaIWAN ANOSOWOUHD
HLONT NVAW Sauods
(A4VNNId ATGCIW) Twos
HLONAT NVAW VLVWOLS
VNNId TVNIWUGLL
(@VNNId 4addn) NOLLUASNI
XadvV LV NOLLVUVdas
adVHS
Sulvd VNNId
HLONGT
adIaLS TVULNAD
SGNOW
SATVOS AWOZIHY
adOund NI SANVWOHDIAL WAINTTdHS¥ AO SAIDAdSHNS AHL AO NOSIYVIWOO
158 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
certain other authors (taxonomists of more conventional approach), have
almost immediately accorded them specific status
I am not myself unduly concerned whether these segregates be given
specific or omit oe rank. In the treatment of Asplen um trichomanes in Flora
Europaea | have been content to accept for the segregates that rank which
has been ioe editorially as most consistent with the concept of species as
fication, and inevitably whatever rank is selected is subject to one objection
or another. Thus the selection of specific rank for segregates of this sort has
the real disadvantage of obscuring their close mutual relationship. On the
other hand, the choice of subspecific status is equally objectionable on the
grounds that it may conceal the existence of real genetic discontinuities. In
advocate the recognition of the species aggregate as a formal taxonomic
category, coupled with the use of some type of trinomial system to indicate
gregates equivalent to specific rank, e.g., Asplenium trichomanes: tricho-
manes L. However, until the unlikely day when a system of this general type
gains formal acceptance, one must continue to apply the existing categories
as best one can.
2 THE SPECIFIC NAMES RECENTLY GIVEN TO SEGREGATES OF THE ASPLENIUM
TRICHOMANES COMPLEX These are three in number. In order of publica-
tion, they are A. lJovisii Rothm., corresponding to the tetraploid cytotype,
A. trichomaniforme H. P. Fuchs, and A. linnaei Sod, the latter two corres-
ponding to the diploid cytotype.
Rothmaler’s work‘ was first circulated privately in page proof. It is of
course most undesirable to quote names used only in an author’s proofs and
not appearing in the published work, since such names are unpublished and
have no validity 15, However, since Fuchs has seen fit to quote in synonymy
names used by Rothmaler only in proof, the harm has already been done.
In the proofs of Rothmaler’s flora, the cytotypes of Asplenium trichomanes
were given subspecific rank, the diploid as subsp. trichomanes, the tetraploid
as subsp. Jovisii. Only in the posthumously published volume is specific
status given to the cytotypes. In view of the tragic death of Werner Roth-
maler prior to the appearance of his flora, it is very unlikely that we shall ever
know the reasons for the decision to promote to specific rank, and further
discussion is not desirable. It is obvious that Rothmaler, a very competent
taxonomist, must have intended to publish more formally on the complex,
since his new name is not validly published in his flora, and indeed now
remains a nomen nudum
45 Cf, Recommendation 34A of the Int.Code Bot. Nomenclature.
LOVIS: ASPLENIUM TRICHOMANES 159
Subsequently, Fuchs® reversed Rothmaler’s decision as to the type
segregate, and published a new name, A. trichomaniforme H. P. Fuchs, for
the diploid cytotype. This name is illegitimate, being a later piri of
A. X trichomaniforme Woynar}®, given to a hybrid between A. trichomanes
and A. adulterinum Milde. The most remarkable feature of Fuch’s i
tion is that he should consider that there was no necessity to give any reason
e
Purpose. Unconsidered or arbitrary decisions are in fact valueless and may
indeed obscure the issue, since a later worker must start from the beginning,
all the real work involved in typification, including elucidation of all the
circumstances dh ees the original application of the name, still remain-
ing to be don
It is regrettable that Sod5, in perceiving that Fuchs had made one major
mistake, did not consider more carefully whether or not he may have com-
mitted others, before substituting another new name, A. linnaei Soo, for
Fuchs’ illegitimate name A. tri igdiinge fehl Both Fuchs and Sod have
entirely overlooked the necessity to respect the principle of priority of
publication, and the need therefore ‘cs ‘irene the existing valid names
Synonymous to A. trichomanes s.|., in spite of the fact that no less
twelve synonyms (exclusive of attributions to genera other than Asplenium),
are given by Christensen!7, One of the oldest and most important of
synonyms is A. melanocaulon Willdenow. Owing to the co-operation (in the
absence of Dr Meyer), of Dr Buchheim of the Botanisches Museum, Berlin-
spore length i “7 By in the air). - dens eoucheeer tide is based ona North American
specimen, b North American
diploid plants and type material of Meyer’s subsp. bivalens (on which
A. linnaei So6 is based), A. melanocaulon Willd. must take precedence over
A. linnaei So6, which lapses into synonymy as a superfluous name,
It must surely be evident that in a group so difficult as the Asplenium
trichomanes complex, the publication of new names without adequate study
Serves no useful scientific purpose, and can only lead to more confusion
which the Code of Nomenclature enables us to resolve, but which it is
Powerless to prevent. As a botanist primarily concerned with studies on
evolution, I grudge the effort spent on the clarification of tiresome and
unnecessary nomenclatural tangles. e is acutely aware that only by
future authors exercising responsible caution will it be possible to avoid the
a ae aes
‘* WOYNAR, H., 1913. Mitteil.Naturwiss.Ver.Steiermk. 49
" CHRISTENSEN C.., 1906, Index Filicum, and supplements «i913, 1917, and 1934)
160 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
creation of further nomenclatural chaos in this much abused but interesting
group of ferns.
ACKNOWLEDGEMENTS
First of all, I must make it quite clear that I accept total and sole re-
sponsibility for the content of this paper. The opinions expressed herein are
my own, and acknowledgement of assistance from the persons mentioned
below must not be interpreted as implying acceptance by them of my views.
I am very grateful to J. Armitage, Prof. H. Melzer, Prof. T. Reichstein
and Miss A. Sleep for living collections of subsp. inexpectans. My greatful
thanks are due to the Director and Dr G. Buchheim of the Botanisches
Museum, Berlin-Dahlem, for the loan to the British Museum, of spore and
rhizome scale preparations of the holotype specimen of A. melanocaulon
Willd. and to Dr Meyer of the same institution, for his generous co-operation
with regard to the re-investigation of spore sizes and for type material of his
subspecies. I owe a particular debt of gratitude to A. C. Jermy, who has
helped in many ways too numerous to describe, and to Professor Manton, for
her valuable advice and constant encouragement.
TWO SPECIES OF DIPLAZIUM IN EUROPE
A. C. JERMY
British Museum (Natural History), London
It has long been the subject of debate as to whether Athyrium and Diplazium
should be treated as distinct genera and, if so, as to what criteria should be
used to distinguish them or whether, in fact, intermediate genera should be
recognised. The problem will not be solved without intensive monographic
work! and it may therefore be considered presumptuous to transfer Athyrium
crenatum (Sommerf.)Rupr. and A. umbrosum (Ait.) C. Pres] to Diplazium
without considerably wider researches.
However, as is well known, two chromosome base numbers have been
found within the complex: n=40 in Athyrium and n=41 in Diplazium; it is
believed that both the species above have n=41,?, 3. They have oblong or
linear, flap-like (asplenioid) indusia and on numerous fronds examined,
sori have never been found lying back to back on the same vein. Both have
bipinnate-tripinnatisect fronds arising singly or in groups of two or three
from a creeping rhizome. Furthermore, the rounded outline of the base
of the lower-most pinnae (i.e., those pinnules nearest the rhachis are shorter
than the second pair) is distinct from that in Athyrium; in the latter the basal
pair is distinctly longer than the second pair, giving an auriculate outline to
the pinna.
Diplazium sibiricum (Kunze) Jermy comb.nov.
Asplenium sibiricum Kunze, Anal.Pterid. 25, t.15 (1837).
Aspidium sibiricum Turcz. in Flora 1834, Beibl.1:28 (1834) nomen nudum.
JERMY: DIPLAZIUM 161
Aspidium crenatum Sommerf. in fet Akad.Handl. 1834:102 (1835); non
Diplazium crenatum Poir. (18
Cystopteris crenata (Summerf.) E. ie Nov.Fl,Suec.Mant. 3:165 (1842).
ey crenatum (Summerf.) Rupr., in Nyland., Spicil.Pl.Fenn. 2:14
1844).
Asplenium crenatum (Sommerf.) E. Fries, Summa Vegt.Scand. 1:253 (1846).
Athyrium mite Christ in Bull. Acad.Géogr.Bot.Mans 1909:36 (1909).
ial C3 eens Kom. in Bull.Jard.Imp.Bot.Pierre le Grand 16: 148
1916).
Kunze based his name on Turczaninov’s plant from the Baikal region of
Dauria and refers to a specimen in herb. De Candolle. There is also a
specimen labelled “‘Aspidium sibiricum mihi, Trentia 1828. Turcz.” in the
herbarium of Oxford University (OXF) (!) which agrees very well with
Norwegian material. Eastwards the species extends to Japan where Tagawa
has described a variety glabrum*. Christ described a species, Athyrium mite,
from Sagalien which, from an isotype at the British Museum is identical with
D. sibiricum. The range is extended westwards via central Asia, the Urals
and Karelia to Finland and to relict localities in the Gudbransdalen, Norway.
Diplazium caudatum (Cav.)Jermy comb.nov.; non D. caudatum J.Sm.
(nomen nudum
Tectaria ‘Couuhina Cav. in Anal.Cienc.nat.Madrid 4:100 (1801).
Polypodium umbrosum Hr Hort.Kew. 3:466 (1789); non Diplazium
a Willd. Ate ose
Aspidi art Schrader, Journ. Bot. 1800(2) :42 (1801).
Maniodic umbrosa (ait) fur Enum. Fil. 179 (1824).
Asplenium umbrosum (Ait.) J.Sm. is Hook., Journ.Bot. 4:174 (1842);
non Villar (1786) nec RRauit (1824).
Asplenium axillare sensu J.Sm., Bot.Mag. 72, Comp.30 (1846), excl.syn.
Asplenium aitonii Moore, Ind. Fil. xlix (1857).
Cavanilles based his name on a plant collected by Broussonet on Teneriffe,
Is. Canarias; the type is presumably at Madrid. The species is also found on
the Acores, Madeira and the Cape Verde Is.; it has no close relative on the
African mainland. The babi australe tenerum complex of Australasia
has been confused with A. umbrosum in the past. Whilst superficially very
Close, the balloon-like Rseinatice indusia which ati split along the
top, puts the Australasian complex into a different secti
The epithet axillare (Ait.) is often applied to this “iil under various
genera. Polypodium axillare Ait. was published in the same book as P.
umbrosum Ait. and there is a specimen in Herb. BM labelled “‘Hort.Kew. 1780
€ Madeira’ and named Polypodium axillare in Solander’s hand®. We can
only assume that this was a plant sent back by Frances Masson and grown
at Kew under that name; it is however, a form of Athyrium filix-foemina (L. )
Roth. John Smith, on the other hand, on the Garden’s staff at Kew from
1822-64, had a different concept. Although he referred in synonymy to
P. axillare Ait. when publishing Asplenium axillare J. Sm. it is obvious from
162 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
s herbarium (at BM) that the plant, grown at Kew at that time under
Asplenium axillare, was Athyrium umbrosum. Smith’s concept of Asplenium
of seeing living specimens he seldom liked to look at them — if he had done
so he would not have united A. umbrosum and A. axillare’”. So in respect to
this great gardener further study on living plants from Macaronesia may pay
dividends — on the other hand, movable pot-labels may be the source of
error!
REFERENCES
Recently R. C. oaENG a. Acta Phytotax. a 9:41-84 (Jan. 1964), proposed severa] new genera
in ragga The pecies mentioned here fit on vegetative grounds into Allantodia R.Br.
which Ching segrega py istinct from Diplazium, but have an 8 ete of the Springs type
which chatectertecs “ase ttkernent aeong chad ee Diplazium chrysocarpus v.A.v.R. group which
is put into a new genus Rhachidosorus Chi
, 1960, Blyttia 18:33-47 [where the author found n=41 in Norwegian A. crenatum
and Saiheed ~ the affinity of this species to Diplazium].
* Prof. I. ee ANTON counted Maderan material of 4. umbrosum and obtained a very strong indication
of n=41; later pa lhe to establish the exact number have so far vailed due to lack of suitable
ccatriak ” (personal communicati
* TAGAWA, M.,Acta Phytotax. Geobot. 11:238 a
* Aiton took his ae yn to Banks’ Lib where mined by Solander, whose descrip-
tions were utilised by Dryander in the first. ming pry Mets rosie [see Stearn, W.T., Cat.Bot.
Books in the Coll. of R.McM.M.Hunt: cviii (1961)].
THE EUROPEAN CHAINFERN IN CORSICA*
In May 1963, I discovered in Corsica four tufts of Woodwardia radicans
Swartz, a fern not previously known in this area. It was growing in the
upper part of the Sisco valley of Cap Corse, well sheltered by 600-1000
metre mountains against cold, desiccating, north and west winds. This
valley is accessible only to the ‘‘ Levante ’”’ and the ‘* Gregale ”’, both humid
warm easterly winds which periodically give it almost subtropical conditions.
The tufts grow near the river in the midst of a very dense stand of Alnus
glutinosa, A. cordata, Castanea sativa and Juglans regia, and do not receive
direct sunlight at any part of the day. Unlike some Spanish localities for
Woodwardia, the hardly accessible Corsican station seems to be quite natural
and untouched by man. Each tuft was up to one-and-a-half metres high wi
five to seven fertile fronds of the previous year’s gro
LUDWIGSHAFEN-am-RHEIN GERHARD SCHULZE
* SCHULZE 1963. Découverte de la fougére Woodwardia radicans Sw en Corse. Bull.Soc.Sci.Hist.
Nat.Corse 83 (569):55.
BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964) 163
ASPLENIUM MAJORICUM Litardiére
A. C. JERMY
British Museum (Natural History)
J. D. LOVIS
University of Leeds Botany Department
During the course of preparation of the account of the genus Asplenium for
the forthcoming Flora Europaea’, we have had cause to investigate one
species, A. majoricum Litard., which, although described more than fifty years
ago, has remained an obscure taxon. This plant, which is apparently endemic
to the Balearic Islands, has scarcely come to the attention of British botanists,
and was indeed until recently unrepresented in the herbaria at the British
Museum or Kew.
To our great good fortune we have had available to us material of this
species collected by J. Orrell in 1959 from the Jocus classicus, the town of
Soller in Majorca. This gathering is represented by a specimen in the
Herbarium of the Department of Botany at Liverpool University, and by
another specimen in cultivation in the Botanic Garden of the same University.
Subsequently, a number of plants have been raised at Leeds from spores
obtained from the original plant in cultivation at Liverpool and specimens
from these are now in Herb. B.M.
A combination of morphological and cytological studies has left us in
no doubt that 4. majoricum Litard. is in fact a species, fully distinct from
any other species of Asplenium known to us. For a variety of reasons, am
investigation of A. majoricum was not completed in time for this species
receive full treatment in Flora Europea and consequently a description of this
plant, drawn up according to the conventions used in the Flora, is given
below, together with an indication of its position in the key to the genus
Provided in that work.
COMPARISON WITH OTHER EUROPEAN SPECIES OF ASPLENIUM As can be
seen from the silhouettes illustrated (Fics. 1 and 2), Asplenium majoricum,
though variable, is nevertheless a very distinctive species. It has perhaps most
often been confused with A. fontanum (L.) Bernh., but A. majoricum has less
* Flora Europea Vol. 1 (Pteridophyta - Platanaceae) Ed, D. H. VALENTINE et al. In press.
164 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
ze
;
'
TEXT-FIG. 1 Photo-copy silhouettes (reproduced natural size) of fertile fronds of Asplenium majoric
all from plants in cultivation ~ cone peeps — rh potions right, whi “a is one of the fronds sent by
Dr Ball from which the Leeds c cu re wa “eee marks indicate maximum extent of black
colouration on underside of thachi
JERMY & LOVIS: ASPLENIUM MAJORICUM PLATE III
PLATE I Cytolo of Asplenium majoricum. Permanent aceto-carmine preparations of meiosis, photo-
graphed using Leitz 2m 4 m N.A. 1.4 apochcomat, with x 10 ocular. Magnification | Fic. 1
Meta aphase I showing n= 72 bivalents. FiG. 2 Late diplotene, also showing n=72 bivalents. Chiasmata :
clearly displa vad,” reed or more per bivalent. Four the valents associated with nucleolus. Fics. 3 an
Explanatory diagrams to Fics. | and 2 respective
JERMY & LOVIS: ASPLENIUM MAJORICUM 165
finely dissected pinnae, the lobes of which are broader than in A. fontanum
with obtuse, not acute, teeth. A. majoricum may readily be distinguished
from A. obovatum Viv., sensu stricto, by the characters of the pinnae, which
are more triangular in outline in A. obovatum, and with lobes of more
rounded shape than in A. majoricum. The pinnae also differ in their
dissection, being bipinnate below in A. obovatum but never more than
innatifid in A. majoricum. This last character also serves to distinguish
satisfactorily A. majoricum from A. billotii F. Schultz (A. lanceolatum Huds.).
degree of dissection of the frond, A. majoricum most nearly resembles
A. forisiense Le Grand, but the pinnae are shorter and more distant, and the
segment teeth are less mucronate than in A. forisiense.
In A. majoricum the rhachis is thin and wiry, in contrast to the relatively
thick and inflexible rhachis found not only in A. forisiense, but also in A
obovatum, A. billotii, and A. fontanum. A further distinctive feature of the
rhachis, again serving to distinguish A. majoricum from all these four species,
is its colour. On the undersurface of the frond the rhachis and stipe are deep
shining black in colour for about two-thirds of their combined length.
However, when grown in less favourable conditions, the fronds remain small,
and this character is not so evident. The frond is then less divided, with the
upper pinnae even entire. In this condition, which may obtain in nature in
excessively sun-exposed and dry situations, A. majoricum might be mistaken
for stunted forms of A. viride Huds. or A. obovatum, but A. majoricum can
always be readily distinguished from all but aberrant forms of A. viride by the
dissection of the lower pinnae, which are always pinnatifid in A. majoricum,
while even in reduced forms A. obovatum has a massive appearance quite
different from the relatively delicate A. majoricum. In A. trichomanes L. the
thachis is black to the tip, easily distinguishing this species from even dwarf
forms of A. majoricum.
The only other species with which Asplenium i
be confused is A. _petrarchae (Guerin) a, but A morienee is quite
glabrous, totally | dular hairiness of A. petrarchae.
i i
1 nahlyu
Amendment to the key to Asplenium (see Fl.Eur.1:14)
Pinnae adnate to rhachis ne eo hs 8 jahandiezii
10 Pinnae shortly stalked
10a Lower pinnae suborbicular, not deeply cut 7 viride
10a Lower rt co mee:
or deeply cu ai sae 9/1 majoricum
9/1 A. majoricum Litard. in Bull. aye Bot.Mans 1911:28 (1911) (A. lanceo-
latum var. majoricum (Litard.) Sennen & Pau). Rhizome erect; scales without
dark central stripe. Leaves 8-12cm, petiole c. one-third the length of the
lamina, wiry, dark at base, shiny black on the underside (adaxial side)
—
oO
166 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
TEXT ote 2
Photo-copy a (reproduced natural size) of fertile fronds from plants in cultivation at Leeds.
Fic. 1 and 2 Asplenium obovatum, ex Capo di Testa, Serdiois (Fic. 2 diminutive fo rm). a 34. bélioes
ex Ronco, Cant on x THAD. Swivveclant. Fic. 4 A. fontanum, ex Roche, Canton Vaud, S itzerland.
Fic. 5 A. forisiense, ex Peuchot, Dept. Lot, France. Fio. 6 A. viride, ex Saalbach, Germa ny.
JERMY & LOVIS: ASPLENIUM MAJORICUM 167
greenish above; lamina elliptical to oblanceolate, pinnate-pinnatifid; light
green in life, dark green on drying; rhachis green above, black beneath for
} its length; median pinnae 8—10mm, lower decrescent and distant, pinnatifid
or deeply lobed, segments with broad obtuse teeth without a mucro. Sori
small, lying near the midrib of the pinna; indusium persistent, deeply and
conspicuously lobed; spores 35-50 (mean 42-8), perispore irregularly
echinulate, sometimes with cristate wings, surface shallowly reticulate or
wavy cristate. 2n=144. Dry stone walls. @ Soller, Islas Baleares. Bl.
CYTOLOGY Examination of meiosis in plants in cultivation at Leeds has
shown that A. majoricum is tetraploid, with a regular meiosis showing n=72
bivalents. (see PLate III). The provenance of the investigated plants is given
above (p. 163).
This finding of a tetraploid chromosome number in A. majoricum is in
contrast to the diploid chromosome number (n=36, 2n=72), found in
A. fontanum?, *, A. obovatum*, and A. viride®, *. Of the other species disc
in this paper, A. billotii2, +, A. petrarchae®, and A, forisiense’ are tetraploid,
while both diploid and tetraploid: levels are known in European A. tricho-
manes?, ®, 7,
THE AFFINITIES OF ASPLENIUM MAJORICUM The chromosome number of
A. majoricum indicates that it is at least possible that this plant is yet another
allotetraploid species amongst European Asplenium. Nevertheless, even if
this suggestion happens to be correct, the parentage of A. majoricum remains
uncertain, and it is probably wisest that further speculation concerning the
nature and origin of A. majoricum is withheld pending the outcome of an
investigation into the cytogenetic affinities of A. majoricum which is now in
Progress at Leeds University.
Our grateful thanks are due to Dr P. W. Ball who on his own initiative
sent us the pressed mature fronds from which were raised the plants of
A. majoricum used in this investigation.
MANTON, L., 1950, Problems oa Cytology and Evolution in the Pteridophyta, 304.
MEYER, D. E., 1957, Ber. Deuts s. 70:58,
MANTON, |, em 1 REICHSTEIN, T., 1962, Bauhinia 2:81.
MEYER, D. E., sch.Bot.Ges. 73:387.
YER, D eos ‘paaone nese 30 Heft 123:31.
LOVIS, J. D., ‘ass B.S.B.1.Conference Rep. 4:99.
168 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
THE POLYPODIUM VULGARE COMPLEX
IN NORTH AMERICA
ROBERT M. LLOYD
Dept. of Botany, University of California, Berkeley
Dept. of Biology and Botany, University of British Columbia, Vancouver
The Polypodium vulgare complex in North America taken in the broad sense
consists of at least five, extremely variable, distinct species. The majority of
these occur on the western seaboard. Since the massive work by Eaton (1879)
on North American ferns the conceptual limits of these species as related to
P. vulgare s.\. has varied from almost complete synonymy with the latter to
complete taxonomic recognition as independent entities. It has only been
since 1950 that the true evolutionary picture has begun to emerge, primarily
through the efforts of Drs Irene Manton and Molly Shivas. The evolution of
the complex in Europe, clarified by the use of cytology, cytogenetics,
morphology, ecology and geographical distribution, has aided in explaining
some of the perplexities of the complex in North America
ir ae ee foo nator : 1 in America
one centering around P. virginianum L. and the other about P. glycyrrhiza
D. C. Eaton. The cytotaxonomic study of these species has not, as yet
reached the refined state known of European members of the complex. By
utilizing the methods of Manton (1950) and Shivas (1960a, b; 1962) we are
endeavouring to clarify some of the remaining problems.
This paper summarizes the current concepts of the complex in North
America. However, there is still much work to be done before the evolution
and systematics of the species can be clarified.
Polypodium californicum Kaulf.
Polypodium californicum is a complex of two cytological races, @
diploid (n=37) occurring from San Francisco, California, south to Baja
California, Mexico; and a tetraploid (n=74) occurring in California from
Monterey County to Humboldt County, and the foothills of the Sierra
Nevada range (Lloyd, 1963). Hybrids between these races have not been
found. Plants also occur on the Santa Barbara Islands, California, and
Guadalupe Island off the coast of Baja California.
Ficures 1 - 5: Frond silhouettes of North American species of Polypodium (x9:
ie pore a 2 Ste couler. igh coastal form; 3 P. glycyrrhiza with attenuate
ements; 4 P, hesperium; 5 P. virginianum
LLOYD & LANG: POLYPODIUM VULGARE 169
4
170 BRITISH FERN GAZETTE: VOLUME9 PARTS (1964)
This fern is commonly found growing on moss-covered rocks and
bluffs throughout the northern part of its range. Plants of the drier regions,
such as southern California and the foothills of the Sierra Nevada range, are
usually in the rich soil of shaded canyons or form an understory to more
xerophytic vegetation. Occasionally, P. californicum is found growing
epiphytically on moss-covered tree trunks and branches.
The herbaceous deciduous fronds drop during the early part of the
summer dry season. Fronds of tetraploid plants may persist throughout it if
sufficient ground water is available.
Fronds of both cytological races are ovate to oblong-lanceolate (see
Fic. 1) and range up to 590mm long and 395mm wide. Margins of most
fronds are serrate or serrulate. The number of pinnae is not constant and is
related to the length of the frond. As many as 69 pinnae have been observed
on plants growing in the mesophytic habitat of the coastal forest. Pinnae
endings are acute-attenuate to round.
Sori are less than 3mm broad and are round to oval. Annuli have 10-21
indurated cells. Although paraphyses are generally lacking, a plant from
San Clemente Island which resembles P. australe was found to have them in
abundance. These are similar to those illustrated by Martens (1950b) and
Shivas (1960a) for P. australe.
Plants with irregular meiosis and inviable spores have been found
beneath the lower Yosemite Falls i in the central Sierra Nevada range. This
population is ically and its evolution is unknown.
Manton (1951) observed 37 univalents and 37 bivalents in plants from Pt.
Reyes, California. Plants investigated from this locality have less than 10%
spore fertility and are morphologically intermediate between P. californicum
and P. scouleri. Due to these characteristics they are believed to be hybrids
(Lloyd, 1962). All of these plants were found growing among the matted
rhizomes of P. scouleri on the bluffs above the ocean. Additional plants with
intermediate characteristics have been found on Morro Rock, San Luis
Obispo County.
Polypodium californicum and P. glycyrrhiza are difficult to separate
morphologically in areas of sympatric occurrence. As the range of the
tetraploid P. californicum is identical with the overlapping range of the two
species it can be hypothesized that the allotetraploid P. californicum is a
product of hybridization between diploid P. californicum and diploid
P. glycyrrhiza. This would produce plants intermediate between the parents
and account for the difficulty in separating them.
Polypodium californicum var. kaulfussii D. C. Eaton was segregated from
the species by its coriaceous fronds and regular series of areoles on each side
of the midrib. This variety inhabits coastal bluffs and occurs sporadically
throughout southern California in moderately more xeric habitats than var.
californicum. Studies have shown these plants to be morphologically the
LLOYD & LANG: POLYPODIUM VULGARE 171
same as the species in all other characteristics (Kendall, 1923; Lloyd, 1962).
Although frond coriaceousness is ecotypically controlled they grade imper-
ceptibly into the herbaceous leaves of the species.
Chesnut (1902) reported that the juice of the rhizome of the California
Polypody was used by Indians of Mendocino County for treatment of
open sores and rheumatism. Root extracts were sometimes used as an
eyewash.
: Frond aberrations such as lobed or cleft pinnae are somewhat common
in the species. Morton (1961) gave forma status to two such individual plants
emphasizing their lobed or deeply laciniate segments.
Polypodium glycyrrhiza D. C. Eaton
Polypodium glycyrrhiza is a coastal diploid species (n=37) which extends
from Kamtchatka along the Aleutian Islands to Alaska; south through
British Columbia, Washington, and Oregon to central California. In the
Southern half of its range it extends east only to the Cascade Mountains,
where it is found growing in the valleys of the major rivers as far inland as
Yale on the Fraser River and Hood River on the Columbia River. It is also
found at Hazelton on the Skeena River in the Coast Mountains of western
Canada.
The species is apparently uniformly diploid (Manton, 1950, 1951;
Taylor and Lang, 1963). Since Shivas (1960a) has shown Polypodium vulgare
S.S. to be a distinct tetraploid species and has evidence that indicates that
P. glycyrrhiza might be one of the possible ancestors of it, it seems illogical
that the two taxa should be considered conspecific.
The Makah Indian name for this species is translated ‘* crawling root on
trees ” (Gunther, 1945) attesting to the typical habitat of the species on the
Olympic Peninsula of Washington. Throughout most of its range P. glycyr-
rhiza may be found growing as an epiphyte on the trunks and horizontal
branches of trees. In Washington, Oregon, and southern British Columbia
the species is often found growing as an epiphyte on the big leaf maple,
Acer macrophyllum Pursh. It has also been found growing on willow (Salix),
elderberry (Sambucus), alder (Alnus) and spruce (Picea). In addition,
however, it is found almost as frequently on rock surfaces, stumps and logs.
This species can be distinguished in the field on the basis of its thick,
Sweet rhizome, uniformly colored scales, free vein endings, and round sori
located midway between the margin and the costa of the lanceolate, often
attenuated, acute frond segments (see Fic. 3). The rock form often has
shorter fronds with less attenuated segment tips. New fronds appear in the
iy
172 BRTiISH FERN GAZETTE: VOLUMES PART 5 (1964)
The specific epithet g/ycyrrhiza is an allusion to the sweet, licorice taste
of the rhizome (the legume Glycyrrhiza glabra is the source of commercial
licorice).
Fischer and Lynn (1933) investigated P. glycyrrhiza as a possible source
of licorice and found that no glycyrrhizin (a compound found in true licorice
that is fifty times sweeter than sugar) was present. They concluded that the
licorice-like taste was due to a mixture of a glucoside polydin, various
unidentified substances, and sucrose.
The Indians of western Washington used the rhizome either raw or
cooked as a cough medicine. The Cowlitz Indians of the Columbia River
used an infusion of the crushed rhizome boiled with young fir needles as a
treatment for measles (Gunther, 1945).
Polypodium hesperium Maxon
Polypodium hesperium is more continental than P. glycyrrhiza or F.
californicum being found in the montane areas of western North America.
Its distribution is from Alaska, south along the Rocky Mountains to Arizona,
New Mexico, and Baja California; east from the Olympic, Cascade, and
Sierra Nevada ranges to Colorado and North Dakota. It is not present in
the Great Basin region of Utah and Nevada. The range of this species
approaches that of P. californicum in the Sierra Nevada Mts. and overlaps
the range of P. glycyrrhiza in Washington and British Columbia.
This montane polypody is found growing on rocks and in narrow rock
crevices from moderate (500ft. in British Columbia) to high (9,000ft. in
Arizona) elevations. It is rarely if ever found growing as an epiphyte. In
British Columbia it produces new fronds in the spring before the dry season
fei
Polypodium hesperium appears to be a polymorphic species with at least
two cytological races. The diploid (n=37) was first reported from the
Columbia River Gorge (Manton, 1951). Taylor and Lang (1963) found
diploid populations in British Columbia. Both Knobloch (1962) and Lloyd
(1963) report tetraploidy (n=74) for the species in Arizona.
Polypodium hesperium can be distinguished from the other North
American taxa by its relatively small segments with rounded blunt tips (see
Fic. 4). The diploid is similar to “ _virginianum in having characteristic
glandular paraphyses and marginal so
The diploid has been postulated ie A. Murry Evans (pers. comm.) to be
the other ancestor of the tetraploid P. virginianum. He finds that in many
respects (e.g., the shape of epidermal cells) the tetraploid P. virginianum is
intermediate between the diploid P. hesperium and the diploid P. LD
Shivas (1960a) has found by crossing the two North American of
LLOYD & LANG: POLYPODIUM VULGARE 173
P. virginianum with European taxa (P. vulgare s.s.) of higher ploidy level, that
the hybrids produced had no paraphyses. If the character is genetically
recessive, as it appears to be, Evans argues that both parents probably had
paraphyses since they are present in the tetraploid P. virginianum. The
diploid P. hesperium with paraphyses therefore seems to be a likely candidate
for the other parent.
Polypodium scouleri Hook. & Grev.
Polypodium scouleri is a diploid species (n=37) (Taylor and Lang, 1963)
and is the most distinct of the North American P. vulgare complex. The
species is entirely coastal occurring from British Columbia south to Monterey
County, California, and disjunctly on Morro Rock, San Luis Obispo County,
in the Santa Barbara Islands and Guadalupe Island. It can be easily recognized
in the field by its very coriaceous evergreen fronds with crenate margins and
its thick rhizome which is glaucous or pruinose.
The habitat of P. scouleri is in shallow soil or rocky crevices on coastal
bluffs or within forested areas. In the northern part of its range it is frequently
epiphytic on moss-covered trees. The island habitats of the fern are meso-
phytic in nature and tend to indicate a more continuous distribution of the
species throughout coastal California during past periods of wetter climate.
Parish (1901) reported that the original collector of the species on Guadalupe
Island found it “encircling the trunk of a single oak, in a thick mat of moss,
and constantly wet by fogs, covering the tree with a network of its strong,
tough roots to a height of ten feet ’’.
Maximum frond sizes up to 89cm long and 22:5em wide occur in plants
growing in the more mesophytic habitats of the coastal forest. In exposed
coastal bluff habitats plants usually do not have fronds longer than 40cm.
Fronds are deltoid to ovate in shape with 3 to 33 pinnae (see Fic. 2). The
great majority of pinnae have rounded endings. Sori are round to oval and
over 3mm broad. Sporangia have 12 to 20 indurated cells in the annulus,
greatly overlapping the range found in P. californicum.
Polypodium virginianum L.
es found in eastern North
America and eastern Asia. It is a complex of two cytologi
(n=37) and a tetraploid (n=74). Natural triploid hybrid
between these races (Manton, 1957; Evans, pers. comm.) wi
chromosomes.
Polypodium virginianum s.1. is distributed
Arctic Circle south through northeastern
Manitoba to Alabama and Georgia (where its range ;
distinct species of the southeastern United States, Polypodium polypodioides
s have been found
th 111 sporophytic
in North America from the
174 BRITISH FERN GAZETTE: VOLUME9 PART 5 (1964)
(L.) Watt var. michauxianum Weath.) and east from northeastern North
Dakota, Minnesota, and Illinois to the Atlantic Ocean.
Manton (1957) reports that the diploid race is one of more northern
latitudes and montane regions, while the tetraploid race is found in lowland
areas and southern regions. Evans (pers. comm.) has found that the triploid
plants have a somewhat independent and southern range. He suggests that
the triploid has successfully circumvented its sterility barrier by reproducing
apomictically.
Analysis of pairing behavior of the chromosomes at meiosis in
synthesized hybrids between the diploid and tetraploid races has shown that
the diploid is one of the ancestral parents of the tetraploid (Shivas, 1960a).
The habitat and general appearance (see Fic. 5) of this species is
adequately described by its French Canadian name tripe de roche. This name,
according to Cobb (1956), refers to the rich, velvety, mantle-like appearance
of the species as it grows over rocks. The species is commonly found growing
on rocks or rock ledges in shady woodlands. It is not often found as an
epiphyte, although it occasionally grows on trees and stumps. Polypodium
virginianum produces new fronds in the summer with the spores maturing by
August or September.
The polypody of the eastern United States and Canada was for many
years considered to be identical with P. vulgare of Europe. M. L. Fernald
(1922) offered strong morphological evidence that the Polypodium of eastern
North America was a distinct entity. He separated P. virginianum on the
basis of its soft spongy, narrow, somewhat acrid rhizome, with its scales
darkened on the back, and its nearly marginal sori. This is contrasted to
P. vulgare with its sweet, firm rhizome with uniformly pale-cinnamon to
castanaceous scales and medial sori. The two complexes differ also in the
characteristics of the glandular paraphyses among the sporangia (Martens,
1950a).
Meiotic analysis of hybrids between the diploid P. virginianum and the
tetraploid P. vulgare s.s. suggests that the North American diploid race is one
of the probable ancestors of the European tetraploid (see Shivas, 1960a, 1962
for details). This is a particularly interesting situation since P. virginianum
is restricted to North America and eastern Asia.
Some work has been done on the comparative morphology of the two
main cytological races (Manton and Shivas, 1953; Evans, pers. comm.).
Manton and Shivas found that the characteristic paraphyses are present in
both the diploid and tetraploid plants but there were many fewer in the
latter (4 as against c.20 per sorus). They also found the cells of the tetraploid
to be somewhat larger. Evans finds that the tetraploid plants are typically
smaller, narrower, with the lower pinnae shorter rather than longer, more
LLOYD & LANG: POLYPODIUM VULGARE 175
upright, and the pinnae tips rounded rather than angular.
Fernald (1922, 1950) does not recognize any varieties or subspecies of
P. virginianum. He does, however, give the status forma to nine morpho-
logical aberrations which are of local or very exceptional occurrence. These
freaks are of little biological significance but are much sought after by fern
collectors for horticultural purposes.
CONCLUSIONS
From the above discussion several characteristics have emerged which
may be used to delineate the species currently recognized in North America.
These are given below in the form of a key.
Sori marginal, with paraphyses
fuamed . Sane acute to obtuse; mostly eastern North
Ameri P. virginianum
Binnae ZO} to obovate, obtuse to rounded; western North
America P. hesperium (diploid)
Sori medial, without paraphyses
Rhizome glaucous to pruinose; fronds coriaceous ..... P. scouleri
Rhizomes not glaucous to pruinose; fronds usually herbaceous
Pinnae usually less than 3cm long; montane south-
western United States P. hesperium (tetraploid)
Pinnae usually more than 3cm long; coastal western
tth America
rond veins free; apex of aes acute to
attenua glycyrrhiza
Frond veins anastomosing into ae apex of
pinnae acute to round
Fronds herbaceous .........-.- P. californicum
Fronds COTIACCOUS ..vcesscseseorsmncesnecsrinernnmnsneenenenenes
P. californicum var. kaulfussii
The cytological evidence to date has shown that the Polypodium vulgare
complex in North America consists of nine cytological races, more or
distinct but interrelated. Polypodium scouleri is morphologically the most
distinct and seems to have had little or no part in the evolution of the remain-
ing members of the complex. On the other hand it can be shown that the
tetraploids P. californicum and P. virginianum have possibly evolved through
ybridization between closely related members of the complex within the
America.
176 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964
In comparing the distribution of polyploid races with the entire complex
it is interesting to note that P. hesperium and P. virginianum are the only
species with diploids occupying more northern latitudes than tetraploids.
This can be contrasted to the situation found in P. californicum and the
European members of the complex. In addition, the habitat occupied by
tetraploid P. californicum seems to be much more favorable for the species
than that occupied by the diploid.
Of special interest are the isolated populations of Polypodium found on
the Santa Barbara Islands and Guadalupe Island. These plants generally
exhibit characteristics somewhat scent from the mainland populations as
a whole (e.g., spore surface, paraphyses).
In Europe the number of indurated cells in the annulus has proved to be
a valuable tool in delimiting the species. This does not seem to be the case in
North America. The number of indurated cells is quite constant in P.
virginianum, P. glycyrrhiza, and P. hesperium averaging 12-14 per sporangium
(range from 9-24). In P. californicum and P. scouleri the number of cells
overlaps so greatly that it is not a useful character.
Fernald (1922) used the base of the rhizome scale as a means of separating
P. virginianum and P. vulgare s.1. He found that in P. virginianum the scale
had a sinus leading to the point of attachment while in P. vulgare it was
peltately attached. Careful examination of the base of the scale of P.
glycyrrhiza (considered a variety of P. vulgare by Fernald) has shown that it
* not peltately attached but that there is a sinus leading to the point of
tachment. The peltate appearance is caused by an overlapping of the lobes
_ the sinus.
ACKNOWLEDGEMENTS
The authors are indebted to Drs Robert Ornduff and T. M. C. Taylor
for criticisms of the manuscript and Mr Richard Mitchell for help in the
preparation of the figures.
LITERATURE CITED
Chesnut, V.K. 1902. Plants used by the Indians of Mendocino County,
California. Contrib. U.S. Nat.Herb. 7:295-422.
Cobb, B. 1956. A Field Guide to the Ferns. Houghton Mifflin Co. Boston.
Eaton, D. C. 1879. The Ferns of North America. Vol. 1. Cassino Publ.
Salem. 272p.
Fernald, M. L. 1922. Polypodium virginianum and P. vulgare. Rhodora
24:125-142.
LLOYD & LANG: POLYPODIUM VULGARE 177
——. 1950. Gray’s Manual of Botany. American Book Co. New York. 1632p.
Gunther, E. 1945. Ethnobotany of western Washington. Univ. Wash. Publ.
Anthropol. 10:1-62.
Fischer, L. and E. V. Lynn. 1933. Licorice fern and wild licorice as substitutes
for licorice. J. Am. Pharmacol. Assoc. 22:1225.
Kendall, M. Louise. 1923. A study of variation in Polypodium californicum
Kaulf. Am. Fern J. 13:75-87.
Knobloch, I. W. 1962. Tetraploid Polypodium vulgare var. columbianum from
Arizona. Am. Fern J. 52:65-68.
Lloyd, R. M. 1962. Studies in Californian Species of Polypodium (Poly-
podiaceae). Master’s thesis, Claremont Graduate School, on deposit
R na Bot Garden, Claremont.
——. 1963. New chromosome numbers in Polypodium L. Am. Fern J.
53:99-101.
Manton, Irene. 1950. Problems of Cytology and Evolution in the
Pteridophyta. Cambridge. 31 6p.
——. 1951. The cytotology of Polypodium in America. Nature, London
167 :37.
——. 1957. The problem of Polypodium virginianum. Am. Fern J. 47:129-134.
—— and Molly G. Shivas. 1953. Two cytological forms of Polypodium
virginianum in eastern North America. Nature, London 172 :410.
Martens, P. 1950a. Les organes glanduleux de Polypodium virginianum. Ill.
Nouvelles données geographiques, systematiques et histologiques. La
Cellule 53:187-212.
——. 1950b. Les paraphyses de Polypodium vulgare et la sous-espece serratum.
Bull.Soc.Roy.Bot. Belg. 82:225-262.
Morton, C. V. 1961. Some forms of Polypodium californicum. Am. Fern J.
51:73-75
Parish, S. B. 1901. Southern extension of the range of Polypodium scouleri.
Fern Bull. 9:40-42.
Shivas, Molly G. 1960a. Contributions to the cytology and egg
species of Polypodium in Europe and America. I. Cytology. /. 4
Soc.Lond. (Bot.) 58:13-25.
——. 1960b. Contributions to the cytology and taxonomy 0
Polypodium in Europe and America. II. Taxonomy. J. Linn.
(Bot.) 58:27-38. oe
——. 1962. The Polypodium vulgare complex. British Fern = Pb ae
Taylor, T. M. C. and F. A. Lang. 1963. Chromosome counts in some
Columbia ferns. Am. Fern J. 53:123-126.
f species of
Soc.Lond.
178 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
CULTIVATED FERNS
J. A. CRABBE
British Museum (Natural History), London
Ferns in cultivation, like many things, have phases of popularity. A century
— particularly in the British Isles, the collecting of fern varieties! had a
fev of enthusiasm which, within a decade or so, somewhat stifled itself
en collectors were more selective, resulting in the formation of fern
societies, the making of fine collections and a spate of literature. Interest
dwindled again at the beginning of the twentieth century, but today there is
another revival, particularly in this country and the United States of America.
In this short paper I want to consider cultivated ferns from the stand-
point of societies and institutions and on national and international levels;
also the contribution of the British Pteridological Society.
HORTICULTURAL SOCIETIES often have some interest in ferns,
although most of them, including the Royal Horticultural Society of London,
have so far not received enough support from their membership to set up
fern committees. Societies of fernlovers are few and include the American
Fern Soci e British Pteridological Society, and the Nippon Fernist
Club. Their iol contain a wealth of information; the first fifty numbers
of that of the latter are in Japanese and in 1961 they were planning to publish
in English or with English summaries.
BOTANICAL INSTITUTIONS usually try, in one way or another, to
take an interest in cultivated plants. An institution may have no relevant
books or specimens at all, its staff relying on personal experience. the
other hand it may concentrate primarily on cultivated plants, outstanding
examples being the Bailey Hortorium at Ithaca, New York and the Royal
Horticultural Society of London whose Lindley Library is one of the finest
collections of literature on this subject. The majority of institutions fall
between these extremes, some places being more fortunate than others in
possessing the herbaria of famous pteridologists, although few have herbaria
containing important fern varieties such as, for example that of Thomas
Moore at Kew.
At NATIONAL levels cultivated ferns come under review by horti-
cultural societies and they are usually included in encyclopedias of gardening
Laws of copyright and patent apply in most countries but the introduction
1 yariety and cultivar (see p. 180.).
CRABBE: CULTIVATED FERNS 179
of special bills for their application to plant breeders is rare; the United
States of America has had such rights for thirty years. In her speech on the
opening of Parliament in November 1963, Her Majesty the Queen said
** Legislation will be introduced to promote the well-being of horticulture.
Proposals will also be made to enable rights to be conferred on breeders o
new varieties of plants”. She was referring to a Parliamentary Bill2 to create
a system of proprietary rights for breeders and discoverers of new varieties
of plants, to establish a Plant Variety Rights Office with a tribunal (a panel
of biologists and plant specialists under the chairmanship of a lawyer), an
ndex of names of plant varieties and a Gazette. This Bill replaces and
extends in scope the Seeds Act 1920 and relevant parts of other Acts are
restricted. Plantbreeders’ rights, in respect to plant varieties, are available
d or di
knowledge being established by reference to plant varieties already in
cultivation or exploited for commercial purposes, or those included in a
recognised commercial or botanical reference collection, or those of which
there are precise published descriptions.
INTERNATIONALLY there is the International Society for Horti-
cultural Science which publishes Chronica Horticulturae. The desire to
Classify and give names is universal+. It was not until the nineteenth century,
however, that international attempts were made to achieve uniformity in the
naming of plants, although rules of nomenclature for certain groups of
Cultivated plants had been made by societies long before. The first code of
botanical nomenclature was published in 1867 and it has been revised
Several times, the latest being the Botanical Code® of 1961. It was not until
1953 that a separate Code for cultivated plants appeared; it was revised’ in
1956, and the second, latest revision® was published in 1961.
* Plant varieties and seeds bill. Her Majesty's Stationery Office, London, 19th December. 1963.
Price
‘1S.L.G GILMOUR, 1961. A decade ng poe Bull, Alpine Garden Soc, 29:53-61
ELSPETH bee HARRIS, 1963. What’s in a name, Bull.Hardy Plant Soc. 3:44-46.
1964, at’s ina nam me ?, J. Scotti sh Rock Garden Club, 9: 69-80. :
: “i x Kanpur 1953. Historical introduction, International code of nomenclature for cultivate
pla;
bs Amiel code of botanical nomenclature. Adopted by the Ninth International Botanical C my
Montreal August 1959. Published as volume 23 of Regnum vegetabile, Utrecht, December
by the Inte national A. sociation for Pant Texonoaty with financial oy a I UBS. -U. ~ E. - oe oO:
be i sake sacha text of the 1956 Code for .
—— code of nomenclature for cultivated plants. Formulated and adopted Sy the lencmsiont
Commission for the Nomenclature of Cultivated Plants of the In test b Aves > tg
Sciences. ‘Published as volume 22 of Regnum tabile, Utrecht, ae a btainable
Roya! Horticultural Society, Vincent Square, London, S.W.1, price 2s. 6d., post free.
180 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
A cultivated plant may be referred to by using three kinds of names;
e.g., Ladyfern Athyrium filix-femina ‘ Druery’ comprises a common
(vernacular) name, a botanical name, and a varietal name. The common
name in any language is governed by popular usage. The botanical name is
always in latin, consists of a generic name followed by a specific epithet, and
is governed by the Botanical Code. The varietal epithet is governed by the
Code for cultivated plants.
Confusion in the nomenclature of cultivated plants has been caused
mainly by the indiscriminate use of both latin and vernacular varietal names.
In ferns, most varietal names have been composed of several latinised
epithets Which: fortunately, may be adapted for presentday use.
The following points from the Code for cultivated plants seem appropriate
to be mentioned here:
Variety (not to be confused with varietas®) is synonymous and inter-
changeable with cultivar; for ferns, however, it seems desirable to continue
using variety, which can be one of four kinds of unit. Ideally, a variety is a
clone (vegetative progeny from a single individual); it can also be a line
(unifo - Or spore-progeny); or an assemblage involving minor gene
differences; or an assemblage of uniform first generation hybrids.
The name of a variety. From 1 January 1959, a new varietal epithet
should be a fancy name markedly different from a scientific name of latin
form, consist of up to three words, each spelt with a capital initial letter
(except when national custom demands otherwise) and enclosed within
ay quotation marks; it should agree in gender with the genus concerned
can precede or follow the botanical or the common name. Thus,
“dikes filix-femina ‘ Victoriae ’; had this variety been described after 1958
a suitable name could be Aliiriaine filix-femina‘ Queen Victoria’ or ‘ Queen
Victoria ’ Ladyiern, similarly, after 1958, ‘ Frizelliae ’, ‘Padleyi’, ‘Cathedrale’
coul ‘Mrs Charles Frizell’, ‘Reverend Charles Padley’, ‘ Lichfield
Cte: 4
Change of scientific name leaves the varietal epithet unchanged, ¢.g.,
Scolopendrium vulgare ‘ Hookeri’ becomes Phyllitis scolopendrium ‘Hooker?’ ;
likewise variety ‘ Barnesii’ of Lastrea montana, Dryopteris oreopteris OT
Thelypteris limbosperma, etc.
nadmissable names. Since 1958 it has not been permissible to introduce
the following kinds of fancy names for new varieties: the botanical or
llia
combination of parts of the latin names of the parent species when the
* yarietas is a botanical category between species and forma.
CRABBE: CULTIVATED FERNS 181
variety is of hybrid origin; names including the word “ variety ” (or “‘var.”’)
or “ form ”’; translated personal names, e.g., ‘ Charles’ must not be altered
to ‘ Karl’, ‘ Karel’, etc. Also, the avoidance is recommended of nine further
kinds of fancy epithet, such as names beginning with abbreviations (except
Mrs and its equivalent in other languages), those lending themselves to
confusion, exaggeration or vagueness and (unless required by national
custom) names containing numerals, symbols, an initial article (e. g., The),
forms of address (e.g., Miss and Mister and equivalents in other languages
are rejected; Mrs, Madame, Frau and other equivalents are admissible).
The correct name of a variety is, with few exceptions, the earliest available
legitimate name.
Legitimate variety names are those validly published in accordance
with the aries of the Code for cultivated plants. Names contrary to
€commendations of the Code are not on that account illegitimate. Names
given before 1959 must either be validly published or accepted by an
official registration authority; a description or a reference to a previously
Published description is desirable but is not necessary. Names given after
1958 must be validly published and also accompanied by a description
(in any language) or by a reference to a previously published description.
Recommendations are that English, French, German, Russian or Spanish be
used for a description, that an illustration be provided, and that a preserved
specimen and/or illustration pared) coloured) be deposited in a public
herbarium and be cited in the descript
id publication is effected by the distribution to the public of printed
or similarly duplicated matter. Microfilm, microcard and xerograph are
accepted. Handwritten material (even though reproduced by mechanical or
graphic processes) and non-technical newspapers are excluded.
Registration is the acceptance of the name of a cultivated plant or group
of plants by a registration authority and the inclusion of this name in a
register. There is no such register at present for ferns.
A registration authority is any international or national organization or
agency so designated either by appointment or by mutual agreement of
societies, organizations and agencies interested. Its main function is to
compile and publish lists, with descriptions and historical details, of varieties
in cultivation and of defunct but historically important varieties. The British
Pteridological Society is preparing revised lists of the names of cultivated
€rns!°; such work may result in the recognition of an official regi —
ety To ‘help ir in this work we are considering making a survey to en
the utm ted in registration. Published
J
a list compiled by J. W. Dyce of ferns to which Certificates of Merit have been awarded by the
a Pteridological Society is given on page 183.
182 BRITISH FERN GAZETTE: VOLUME9 PART 5 (1964)
treatments of cultivated ferns, such as those in the American Fern Journal
and the British Fern Gazette and lists such as in encyclopedias of gardening
and in Standardized plant names by the American Joint Committee on
Horticultural Nomenclature, are of great help in this work; the Editor
hopes for more contributions similar to the treatment of Polystichum in
the last Gazette. As another aid in the registration project we are preparing
a census of fern varieties growing in the British Isles.
Description. A variety, like other taxonomic groups, should be described
definitively, i.e. so that other plants may be readily compared. Ideally, such
up by preserved material. Seldom do we have this ideal. The British
Pteridological Society is planning to define each fern variety by publishing
illustrated descriptions.
THE BRITISH PTERIDOLOGICAL SOCIETY. In his presidential
address to the sixtieth Annual Meeting of the Society in 1963, Reginald Kaye
asked: ‘“* What is going to happen to all the varieties of ferns in cultivation
today, hundreds less than fifty years ago; what is going to be the position in
the future?”
Our plans include a survey of societies interested in ferns and a census of
ferns cultivated in the British Isles, publishing treatments of relevant genera
and illustrated descriptions of varieties, compiling lists of varieties for registra-
tion and distributing living material to gardens in the British Isles, thereby
seeking to increase the popularity and chances of survival of our heritage 0
fern varieties.
BRIT. FERN GAZ. 9(5), (1964) 183
FERN VARIETIES TO WHICH CERTIFICATES OF MERIT HAVE
BEEN AWARDED BY THE BRITISH PTERIDOLOGICAL SOCIETY
Compiled by J. W. Dyce
The names are those under which the certificates were
awarded (with dates) and are subject to revision
Asplenium trichomanes eens Clement 1921.
thyrium filix- put medusae 1938; cruciato-cristatum baccatum Kingsmill
Moore 1922; plu tieoedclts statum Lovelady 1930; plumosum laxum Sheldon
1931; setigerum percristatum Sheldo on 192 9.
Blechnum spicant bipinnatum Sheldon 1931; percristatum Rowlands 1926; ramo-
cristatum Blow 1927; Smithies seedling 1938; tricapitatum
Lastrea dilatata (=D is dilatata) grandiceps Cranfield 1938.
ean sedis a Dryopteris filix-mas) cristato-tenuis Parker 1932; grandiceps
ae ee se a, limbosperma) formoso-cristata F. W. Stansfield 1922;
ceps Smithies 1903.
Lastrea Seal (= si abbreviata) crispa Cranfield 1939.
Lastrea pseudo-mas (=Dry borreri) fimbriato-cristata angustata Cropper 1922;
polydactyla W Whiteside 1999, Whitwell 1922.
Polypodium vulgare bipinnatum F. W. Stansfield 1926; cambricum Henwood 1922,
cristatum Henwood 1922; omg nates ‘Oxford’ 1921; plumosum Whilharris
1927; pulcherrimum rem 8.
Polystichum aculeatum cristatum Henwood 1927; divisilobum Cranfield rnd
faticiecincicarr Sheldon 1933: gracillimum attenuatum Druery 1938; g. cri
latum Cranfield 1937; g. pulcherrimum Cranfield 1937; grandiceps Sheldon
1937; pulcherrimum Cranfield 1927, Druery 1921, Ki ngsmill Moore 1934;
Ra UO soe Edwards 1922; p. plumosum Green 1921; sinuosum Kingsmill
re 1931.
erie angulare (=P. setiferum) mee oP Stansfield pes a. divisum
W. Stansfield 19303: a. laxum Sheldon 1934 a. macrodon T. Stansfield 1930 1930;
M
d. plumosum resales ldo
1923; foliosum Moly 1927; } grande “Walton 1922; inclan- stone ee ae
1934: lineare caudatum T. Stansfield ag 1. ‘cristatum m Ed. 1906; mult ey
1934: m. rotundatum Be ie Id 1921;
=
nw
eo
“$S
35
ch
&
a
Z
une
a)
[Nw
+i
&
AO
7
'
mill es
38;
Soepenirion. ver (= Phyllitis ie scolgpends ta) crispissimum muricatum 19.
Cris cate gustifrons Sados 1937; cristatum John Co ousins 1927; .
—_ we 1921; cc. f. sagittatum pumeegy toe hes
Sheldon Is 1931: speciosum Moly 1921; ¢. sp ndens : ;
fimbriat wih Sheldon eigen pening grande 1931; ramo-cristatum Cranfield
1937; sngitteto-criapuat tum Perry 1926.
184 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
DISTRIBUTION OF THE BRITISH FERNS AND THEIR ALLIES
During 1962 the Atlas of the British flora* was produced by the Botanical
Society of the British Isles. This contains over 1600 small outline maps of
the British Isles, showing the distribution of about 1700 species of native
or naturalised plants including ferns. The Ordnance Survey grid system was
utilised, and dots indicate the presence of the plants in the relative 10-
kilometre squares. This of course does not show the complete picture, as a
dot recorded on a map cannot show whether the plant exists in a square as
an individual specimen only, or is abundant throughout the area. It is
extremely interesting to study the distribution pictures of the ferns and many
cases show the lack of interest in the Pteridophyta which seems to prevail
among botanists generally. It is very obvious that a lot of work has still to be
done on fern distribution and the Atlas has acted as a spur to do something
about it.
BRITISH PTERIDOLOGICAL SOCIETY
DISTRIBUTION CARD FOR
FERN
S AND FERN ALLIES
OMMO sats la}s{6{7{8{9 ho UNCOMMON | 1/2/3/4[5|6 [7[8| 9/10
Asple adi Adian cap
rut Anogr lep Ll
tri Asple alt =
vir bil —
Athyr fil mar oa
Blech spi sep EA Se
Ceter off Athyr dis feel fel
Crypt cri le id esa
Cysto fra Azoll fil a ee
Dryop abb Botry lun =S
aem Cysto dic |—_}_+-—
bor mon _—+—+
car Dryop cri }+—+—
dil Nib li Sa GE
fil vil ae (es Dae
(agg) fil Hymen tun ia ee
ymno dry il oun be
Ophio lus ag ES oe
Osmun reg vul =
Phyll sco |_| Pilul glo Lf}
Polyp aus Polys lon |}
int Thely pal ae
vul Trich spe +
(agg )vul Woods alp +}
Polys acu ilv ae
set Hed been Sie foe Be? OR
Pteri aqu Ban
Thely lim Sage
phe
od of the British flora. Ed. F. H. Perring & S. M. Walters. Pp. xxiv+432. B.S.B.I. & Thomas
Nelson & Sons Ltd, London & Edinburgh, 1962. Five guineas.
DYCE: DISTRIBUTION RECORDS 185
FERN ALLIES LOCALITIES
(rare)
1| 2! 3/4{ 5/6} 7/ 8] of1 1
Lycop
alp
a (abundant) c (common) £ (frequent) o (occasional) r
I!) gee
|
|
I have devised a card as shown above, and use a separate one for each
10-kilometre square. Filed according to squares this system works admirably
for quick reference and gives a clear picture for any area under review. Cards
pee of course be used for any unit area. Also, instead of a tick in the
Ocality box one could put a symbol indicating the frequency, e.g.
a ppc lias c (common), f (frequent), o (occasional), r alas
It is thought that many members of the Society t
on the fern distribution in their home districts or when on holiday in other
Parts of the country. I shall be pleased to hear from anyone interested in
using these cards and if the demand is sufficient to cover the costs the Society
will consider bulk printing. It is hoped that some will be available at this
year’s Excursion.
J. W. DYCE
186 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964
BRITISH PTERIDOLOGICAL SOCIETY
MEETINGS IN 1963
BRITISH MUSEUM (NATURAL HISTORY), LONDON, 9 February. Conversazione
and exhibition meeting.
ROYAL HORTICULTURAL SOCIETY’S GARDENS, WISLEY, SURREY, 18 May.
excursion to see, amongst others, the remnants of W. B. Cranfield’s British
fern varieties.
BROCKENHURST and LYNDHURST, HAMPSHIRE, 14-16 June. Among 18 species
of ferns seen, of particular note were Ophioglossum vulgatum, Osmunda
regalis and Pilularia globulifera. We hunted in vain for Polystichum setiferum,
although it is recorded for the area and is most abundant a few miles further
south.
NATIONAL PINETUM, BEDGEBURY, KENT, 7 July. A meeting jointly held with the
Kent Field Club.
ROYAL BOTANIC GARDENS, KEW, SURREY, 27 July. A visit to the Herbarium
and Library conducted by Dr Frances M. Jarrett. After lunch a tour of the
Gardens with Mr H. J. Bruty.
THE ANNUAL EXCURSION, 15-21 September. Pencerrig Country House Hotel
a pony-trekking station about two miles north of Builth Wells on the border
of B hire and Radnorshire turned out to be an excellent centre. Mention
has been made before of the added interest the Atlas of the British flora
gives to our records; our Secretary has developed a most efficient system
(see p. 184) whereby he is able to recognise straightaway a new record for a
10 kilometre square (a new dot for the appropriate Atlas map). Permits
had been obtained to enter and collect in six Nature Reserves; new fern
records were made in all of these, and details of our observations have been
sent to the Nature Conservancy.
On Sunday we worked down the Wye valley to Clyro and returned via
Painscastle. The first stop was at Aberedw to inspect a steep wooded bank of
northerly aspect with cliffs rising above a stream, which was typical of good
fern ground in the district; within half-an-hour we had found 15 species
including 2 new records. We then visited a similar site at Erwood where an
additional species, Gymnocarpium dryopteris, was found in plenty. The next
stop was at a bridge with a fine display of Asplenium trichomanes, large
sprays of Cystopteris fragilis, and interesting Polypods, some with forked
pi . A mile north of Glasbury we explored a wooded gorge where
Polystichum setiferum and Phyllitis scolopendrium were the dominant ferns,
which is remarkable since neither of these is plentiful in this part of the
country. Crossing a field on the way we noted silvery bushes of Artemisia
BRITISH PTERIDOLOGICAL SOCIETY 187
absinthium which proved to be a new record. At Clyro we turned off the
main road and went over the hill to a bog known as Rhosgoch Common*.
Mr and Mrs Walter Jones, proprietors of a local guesthouse, told us that
Osmunda grows in a wood at the far end but that it is decreasing; unfortunately
time did not allow us a visit to this remote spot, but it was some compensation
to discover Pilularia globulifera; many other bog plants were found, such as
Potentilla palustris and Senecio aquaticus which was still in flower. The
first day’s hunting had yielded 18 fern species, including 4 new records.
On Monday we went to Llandovery and turned north up the valleys of
the Tywi and Doethie rivers to visit the Nature Reserve known as Allt
Rhyd-y-groes. Up a steep and beautiful mountain stream we found 11 ferns
including 2 new records; this stream is noted for uncommon bryophytes.
North of the stream a Sessile Oak wood, typical of the region, forms part of
the Reserve; the broad dark-green leaves of the Quercus petraea give deep
shade which, where there is ample surface water, induces a most luxuriant
undergrowth of ferns but, where the soil is dry, the undergrowth is mainly
composed of Bilberry. After lunch we went further up the valley to some
cliffs where Cryptogramma crispa had been recorded in 1958 at almost the
extremity of its southern range; since this was no part of the Reserve we
asked permission and were given every assistance to reach our objective; we
failed to find the Parsleyfern but we did discover a plentiful growth of
Hymenophyllum wilsonii by a stream cascading down the steep hil Iside; this
filmy fern was recorded several times during the week and we wil] make no
furihes special mention of it
Tuesday was spent at Craig-y-cilau N.R. near Crickhowell. This is an
extensive line of cliffs and old quarry-workings on Carboniferous limestone
which underlies the South Wales coalfield syncline and outcrops around its
Periphery. The remains of a tramtrack provides a path along the Reserve and
€asy access to the cliffs and screes at any point. Gymnocarpium robertianum,
in varying shades from green to red, was abundant in screes where Oakfern
and Beechfern were also found. In rock crevices were Asplenium ruta-
muraria, A. viride, Ceterach officinarum and Phyllitis scolopendrium. At the
top of the cliffs, in complete contrast, Blechnum spicant thrived on the over-
lying acid moorland.
On Wednesday, the only wet day, we went to some waterfalls two miles
northwest of Rhayader. Access to these was difficult and approach had to be
from above, over a mile of boggy moorland where we found Lycopodium
alpinum and L. clavatum, which are uncommon in this region, and Hypericum
elodes, a new record. The waterfalls were worth the visit and 10 ferns were
found in the wooded oasis surrounding them, but after trudging back across
ee ene eee ern
New Phytolo-
* for nalysis, and stratigraphy of Rhosgoch see D. B. BARTLEY 1960,
gist $9: S627, Bon Ecol, 48:205-214.
188 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
the moor in the rain it was agreed to call it a day. However, a small group
had decided to by-pass the waterfalls and continue along the mountain
road to Devil’s Bridge; they escaped the rain and found plentiful ferns; at
one of their four stops were two plants of Cryptogramma crispa in boulder
The following day we went south to Craig Cerrig Gleisiad N.R. which
lies in the Brecon Beacons National Park. This is on the Old Red sandstone
which gives rise to an extensive area of bleak moorland mountain country.
The Reserve is named after a horseshoe of cliffs some 600 feet high which
form its principal feature. These supported a luxuriant vegetation of interest-
ing plants, including Asplenium and Polystichum, while the screes below were
dominated by Dryopteris borreri among other ferns, including a few
D. abbreviata. Six new records were soon made and it is remarkable that
Thelypteris limbosperma which was the commonest fern over considerable
areas had not been recorded for this well-botanised locality. The site is
noted as the southernmost station of a number of arctic alpine plants; none
of these was in flower at the time but we noted cushion saxifrages and, high
on the cliffs, an old record for Saxifraga oppositifolia was confirmed. After
lunch the main party decided that there was still plenty of ground to explore
around the cliffs, while two small parties set off to other cliffs, one over
moorland to the west and the other down the road to the south; the former
had all the fern species of the main cliffs and at the latter Cryptogramma
crispa was found on scree.
On Friday we went via Llanwrtyd Wells to Abergwesyn where, in the
neglected churchyard wall, were hundreds of plants of Asplenium ruta-
muraria and Ceterach officinarum, one plant of Ceterach being identified as 4
‘Crispum’ form. We then visited Nant Irfon N.R. which is in a similar
situation to Allt Rhyd-y-groes but is more open and varied. After lunch the
main party took the mountain road to Tregaron to visit the extensive and
famous raised bog. A smaller party exploring the valleys north of Beulah
found that if the scenery was less exciting this did not apply to the fern flora.
At the first stop an unexpected find was Equisetum sylvaticum, one of the very
few horsetails found during the week. Then, in a disused slate quarry a fine
colony of Asplenium adiantum-nigrum was found. A bridge on the B4358
road was seen to have iron railings and was nearly passed by; however, on
stopping, no less than 9 species of ferns were found growing on or very close
to the bridge, including luxuriant Nihon fragilis, quantities of Ceterach
and even Oakfern. Finally, a halt was made at a hillside oakwood which,
although festooned with lichens, onsen out to be dry and fernless; however,
in marshy ground at the bottom of the wood the last find was our first
Dryopteris carthusiana.
Saturday was the day of home-goings. Nevertheless one party went to
Cwm Clydach N.R. which is on the Carboniferous limestone and proved
BRITISH PTERIDOLOGICAL SOCIETY 189
most interesting. At the bottom was a dark ravine with luxuriant fern growth,
including Polystichum aculeatum and P. setiferum. Higher up was a disused
railway track in the ballast of which were found plentiful Gymnocarpium
robertianum and Ceterach, while in the brickwork of a tunnel entrance was
a magnificent display of Cystopteris fragilis. In boulder screes were found
more Ceterach and Asplenium trichomanes, and on limestone cliffs were
A. adiantum-nigrum and A. ruta-muraria. Blechnum spicant grew in acid
pockets in the higher gullies. In all, 17 fern species were found by late
afternoon, although only half the area had been covered. Another party
visited two more limestone localities, the first of which was a noted beauty-
spot near Ystradfellte with waterfalls and potholes; some good ferns were
found, including Phyllitis scolopendrium growing in the deep shade of
recorded in full flower beside the river. The second site was Penmoelallt
Forest N.R., which is a wood on a very steep slope topped by cliffs over-
looking the River Taff on the northern outskirts of Merthyr Tydfil; this was
not a particularly ferny spot and the complete absence of Thelypteris
limbosperma may perhaps be attributed to atmospheric pollution; it is of
interest for its rare trees, including Sorbus leyana of which only a few bushes
remain
ing the week a great deal of interesting and varied ground was seen;
103 Polypods were collected for a survey which has segregated them to
86 P. vulgare, 15 P. interjectum, and 2 presumed peataploid hybrids. Our
experience with the Atlas of the British flora has shown, at least for ferns,
that the frequency of dots on the maps may be more indicative of the
distribution of botanists. We should therefore not be deterred from visiting
a region merely because the Atlas shows a poor fern distribution; in fact it is
in such regions that we may be able to contribute most.
Appended is a list of pteridophytes seen during the Excursion; figures
(bold= new record for the Atlas) refer to localities.
copodium clavatum — /9
7 alpinum — /9
Equisetum palustre — ¢
= sylvaticum — 15 ou
ymenophyllum wilsonii 4, 6, 20, 28, 30.
Pteridiurn naekeia Pe a 6, 7, 9, 10, ii. 12, 14, 15, 16, 18, 19, 20, 21, 23, 24,
tdi, 30, 31, oa
a crispa — 4, 33
chien bith cant — 2, 3, 4, 5, 6, 7, 9, 10, Is 19, 20, 21, 22, 28, 29, 30, 33
eee sme rium — 5, 7, 9, 10, 12, 14, 15, 16, 2
: 7,20
ran en yee Le 10, 11, 12, 14, 16, 17, 18, 21, 22, 23, 28, 30
A. ruta-muratia — 1, y 8, 2, ie 13, 16, 18, 21, 22, 23, 32
Coteraeds 9, 11, 12, 22 20, 24, 22,
Asean nen hy 3, 4, , 6, 7, 9, 10, 11, 12, 14, 15, 16, 18, 19,
23, 24, 28, "59, 30, 31, 33, 34
190 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
cane | taygeowalaag we 6,:7,.9, 10, 12; 14, 16, 17,18, 21, 29,30
Dryopteris filix-mas — 1, 2, 3, 4, 7, 9, 10, 11, 12, 14, 15, 16, 18, 21, 22, 23, 24, 28, 29,
D. borreri — a; "4, 5, 6. 7,9, 10, 11, 14, 15, 16, 18, 79, 20, 21, 22, 23, 28, 29, 30, 31, 32,
33, 34
D. abbreviata —4
D. carthus: —3
D Silesia — 2: 3, 4; 5; 6, 7, 9; 10, 11; 42; 14; 15, 16; 19, 20, 21; 22, 23; 24; 28, 29-2
Polystichum setiferum — 9, 10, 14, 21, 22, 29
P. aculeatum — 4, 6, 9, 10, 16, 18, 21, 28,
some ne ag dryopteris — 2, 4, 6, 7 , 10, 12, 19, 20, 30.
roberti
Thelypteris lenboeperass— 1, 2, 3, 4, 5, 6, 7, 9, 10, 11, 15, 16, 19, 20, 21, 22, 28, 29,
ais phegoptcris — 2, 4, 5, 6,7, 11, 19, 20, 28, 29, 30, 33
Polypodium vulgare — 2, 3, 4s 5, 6,7, 9,10, 11, 12, 14, 15, 16, 17, 18,-19, 20, 21,
ea Mo 4. 26, 27, 28, 29, 30, 31, 3.
10, 11, eis “i; 28
Bee
Piluleee globulifera —
Date
V.C. 42 Brecknockshire visited National
(O.S. 1” map ser. 7, sheet 140) September Grid Ref.
: — Church . 20 22/ 8552
IRFON NR, | a NE of Abergwesyn ... 20 8453
oe S. go 141)
3 COED-Y-CILIAU 24 miles NE of Beulah 20 9553
4 CRAIG RIG-GLEISIAD N.R. 6 miles SW of
recon 9621, 9622, 9722
5 CRAIG-Y-FRO 14 miles S of Craig Cerrig- gleisiad 19 9720
6 DU 2 miles W of Craig Cerrig-gleisia 9421
7 CRAIG-Y-CILAU N.R. 2 fodiens SW of Crickhowell 17 32/ 1915, 2015, 2016
8 CRICKHOWELL — Bridge over Usk . 17 2118
9 cwM CLYDACH N.R. 14 miles E of Brynmawr 21 2112, 2212
10 ERWOOD — ravine 15
11 GILWERN ihr miles N of Beulah 20 22/ 9153
12 GLANDULAS Bridge over Dulas 2 miles NE of
Beul i 20 9453
13 LIBANUS 3 miles SW of Brecon |. as 19 9925
14. LLANGOED CASTLE, roadside le at CP me 15 32/ 1139
15 LLWYN cei 1 W of Beulah_... 20 22/ 9052
16 PORTH-Y-OGOF 1 tile S a ¥stradfelite BG 21 9212
17 YSTRADFELLTE =e < oe Pees 21 9213
(O.S. sheet 154)
18 eee 9 ORS E a eric aA Sata aaest as, 21 32/ 0109
V.C. 43 Radnorshire
(O.S. sheet 128)
19 CRAIG-pDu 2 miles NW of Rhayader 18 22/ 9469
20 NANT-Y-SARN WATERFALLS 3 —— NW of
Rhayader 18 9371
— S. te ec 41)
21 Dw — wooded ra 15 32/ 0847
paps. DINGLE 14 et) N of Glasbury ... 15 1840, 1841
23 PENCERRIG COUNTRY HOUSE HOTEL 2 miles N
of Builth Wells as. ‘ha ce oe eee 0453
RHOSGOCH a ge ae 15 1847
25 RHOSGOCH COMMONbog ... tis 15 1948
BRITISH PTERIDOLOGICAL SOCIETY 191
46 Cardiganshire
6. S. sheet 127)
AEN-Y-CwM bridge over river 6 miles E of
Devil’s Bridge ies 18 22/ 8275
27 BRYN 3 miles SE of Devil’s Bridge iis ees 18 7774
28 DEVIL’s ane ynach Falls... ae 18 7477
(0. s. shee 140)
29 seta OES N.R. 64 miles W of
a nwrtyd Wells .. 16 7647, 7747
30 cRIBYN Du 14 miles NE of Allt Rhyd-y-groes NR. 16 7548, 7648
31 JUNCTION OF AFON DOETHIE and A
6 miles W of Llanwrtyd Wells . ai 16 7746
32 TREGARON, railway bridge mileNof _... 20 6861
V.C. 46/47 Cardiganshire/Montgomeryshire border
CRAIG-Y-LLUEST (V.C. 46) 18 8575, 8576
34 CRAIG-y-LLUEST south of stream by ‘roadside
(V.C. 47) 18 8575
ANNUAL GENERAL MEETING, 18 September. Held during the Annual Excursion.
Reported in Newsletter No. 1, November 1963, to which were appended
Minutes of this sixtieth A.G.M. pee ss hisiats sat talk.
BORROWDALE and COCKERMOUTH and RYDAL, WESTMORLAND,
5 and 6 October. Eight veople: ceatia 4 this satiate weekend meeting
during which twenty fern species were recor:
BRITISH MUSEUM (NATURAL HISTORY), LONDON, 19 October. Conversazione
and exhibition meeting.
THE SOUTHPORT FLOWER SHOW 28 - 30 August 1963
Our President, Mr. Reginald Kaye, judged the fern exhibits this year with Mr.
W. G. Mackenzie (Curator of Chelsea Physic Garden). Each fern had to be
assessed carefully for rarity, cultivation, presentation and condition, and it
was exceedingly close between first and second in many cases. There ba
evidence of more care in preparation, one exhibitor in particular having
taken great pains in cleaning his pots and neatly surfacing them with nice
fresh moss. Dressing a plant for exhibition means a good deal and may
make just that little extra when being judged. There were still one or two
specimens marred by not removing weeds when potting them; this should
never hap
In Class 16, several exhibits were noted for having one fine specimen
Supported by two which looked as though they had recently left the Coens
of a hedge. One contained what was probably the best specimen in the
Show, but was supported by two such rough ones that the first had to go to
@ group of more uniform merit. The best possible fern in such a rt
cannot earn more than a third of the points and will not “ pull up ° ” inferi
companions,
192 BRITISH FERN GAZETTE: VOLUME 9 PART 5 (1964)
The ground group (Class 8) as usual was very well done and contained
some magnificent specimens in splendid condition particularly, amongst
many others, Athyrium filix-femina ‘ Clarissima ’ and varieties of Polypodium
vulgare.
The classes for greenhouse ferns seemed to be better than last year and
some really delightful well-grown specimens were shown. Incidentally,
some ferns shown as greenhouse subjects were hardy ones. The Show
Schedule should indicate that ferns shown in these classes should be tender
species requiring greenhouse conditions. In the judges’ view, the inclusion
of hardy species in classes for greenhouse ferns would have to be taken into
account in the event of more numerous entries as, strictly speaking, they are
not “‘ according to schedule”’ and, indeed, we had some discussion as to
whether we could overlook the offenders.
Thanks are due to Messrs Almond, Hayhurst, Jackson and Robinson
for staffing the Society’s Information Bureau. Any assistance members can
give in this direction will be greatly appreciated.
A glance over the following list of prizewinners and their entries shows
the selection of ferns available for visitors to see. Except for a few abbrevia-
tions the fern names are those with which they were exhibited.
CLASS 8 = > of Boma gris ferns, with accent on diversity and variation, arranged in athood
e 13 feet by 8 feet and not higher than 6 feet above the gr
Ist aa re cna wate who continue to _ the Silver Perpetual Challenge Trophy
presented by the British Preridological Socie
CLASS 9 Six dissimilar hardy British ferns with accent on variation:
B. Hayhurst: Athyrium filix-femina victoriae, Dryopteris lineata cristata, | Phyliitis ey
ae y Seah Senge Polypodium pulcherrimum, Polystichum ‘ Hironde lle’,
st ivery
alis
F. perene | Pi oven filix-femina cristatum, Dr ryopteris lineata cristata, Osmunda reg
ristata Moore. Phyllitis a egg marginata, Polypodium cambricum, Poly-
pe a
§
|
s
S
o
S
S
my mind some of the red back-
€ unpleasing. Detailed charts tabulate all
tns dealt with in the previous sections, so that it 1s
very easy to look up descriptions and requirements. A good glossary, a list
of nine commercial sources of supply in U.S.A., and a comprehensive index
complete this very well-presented book.
Waithman Nurseries, Silverdale REGINALD KAYE
BRIT. FERN GAZ. 9 (6) 1965 205
BRITISH FERNS IN A WORLD PERSPECTIVE
R. E. HOLTTUM
Royal Botanic Gardens, Kew, Surrey.
Thirty years ago Carl Christensen, in the introduction to the third
Supplement to his mit Filicum, estimated that almost 10,000 species of
ferns were then known. A good many new species pate been described
since 1934, and most — collections od the wet tropics produce more.
One may thus regard 10,000 species as a conservative estimate of the world
fern flora. The number of species in Britain is about 0. 5% of the whole.
My present object is to try to show how this small fraction is related to the
great mass of tropical ferns.
The great majority of the world’s ferns have sporangia which are basically
Similar in structure, though differing in details; they are usually called
leptosporangiate ferns (/eptos means thin, delicate). The remaining ferns
long to a few rather small and very diverse groups: Ophioglossaceae (the
Adder’s Tongue and the Moonwort in Britain have relatives throughout the
world, but they are nowhere abundant); Marattiaceae (a family of ferns found
only in the wet tropics, many growing to a very large size, probably related
to ancient fossils); and the heterosporous ferns (only Pilularia native in
Britain) which belong to three distinct families the relationship of which to
other groups of ferns is still obscure. The remainder of this discussion
deals only with the leptosporangiate ferns.
The “ primitive ’’ leptosporangiate ferns
The distinctive feature of the sporangium of leptosporangiate ferns is
the annulus, a more or less continuous ring of cells with walls thickened in
such a way that on drying stress develops in the ring, which ultimately breaks
and so scatters the spores. The more highly evolved ferns have smaller
sporangia and a more perfect mechanism for this breaking of the annulus
than the more primitive ones. The fact that the latter are primitive is
demonstrated by their occurrence in fossils. Botanists first began to examine
the annulus critically just before the year 1800, and soon recognised the
families which are usually called primitive because of the primitive structure
of their sporangia; many of these ferns are highly specialized in other ways.
OSMUNDACEAE. The British representative is the Royal Fern
Osmunda regalis. There are a few other Osmunda species throughout north
temperate regions and in the wet tropics; also in south temperate regions the
allied genera Todea and Leptopteris. The family has a more clearly defined
fossil history than most others. The sporangia are large, almost spherical,
with a clear line of dehiscence but no distinct annulus. They are however
very like those of the genus Mohria (Schizaeaceae) and one can see the
relationship between the sporangia of Mohria and of other members of its
family which have adistinct annulus.
206 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
SCHIZAEACEAE. This family is not represented in Britain today, but
there are fossils of the genera Anemia and Lygodium in the London Clay;
these genera occur today in N. America and probably disappeared from
Europe in the glacial period. The other living genera are Schizaea and
Mohria, e four genera are very diverse and all specialized; the most
GLEICHENIACEAE. Ferns of this family are the commonest thicket-
forming sun-ferns in the wet tropics today; they have increased greatly in
recent years owing to the felling of forests by man. The family is repre-
sented by fossils in the Eocene of Britain and by numerous others in north
temperate regions (and Greenland) but its extra-tropical distribution today
is only in the southern hemisphere. Most modern members of the family
have repeatedly forked fronds; the forking has been regarded as a primitive
character, but it is due to dormancy of apices in very precise patterns, and
in my opinion there is nothing primitive about it. It is possible that Poly-
podium and allied genera of today have a common ancestry with this family,
but this is still very uncertain.
OPHYLLACEAE. Filmy-ferns (three British species) are
extremely abundant in the wet tropics, especially on trees in mountain forest
in the cloud zone, where the ferns may be dripping wet a great part of the
time. € number of species now known is about 600. Copeland has
divided them into 34 genera, mostly on minor characters, and in my opinion
two feet or more long; these mostly grow on rocks by streams in deep shade.
CYATHEACEAE. This is the family of the tree-ferns. In recent years
Bower thought to be very divergent; but it seems to me that they are so
Closely related that it is natural to unite them (see Flora Malesiana, Ser. HU,
vol. 1). The genera with a more primitive type of sorus, at the end of a
are on the lower surface of leaflets, not at the ends of veins, and in most
Cases have indusia. The German morphologist K. von Goebel suggested
HOLTUM: BRITISH FERNS IN A WORLD PERSPECTIVE 207
that a simple change in marginal growth would change a Dicksonia sorus
into one like Cyathea, and this seems to me a very reasonable suggestion.
Other characters besides the sori which are important are the structure
of the rhachis and the way in which the bases of leaflets are adjusted to it,
the distribution of hairs (especially on the upper surface of branches) and the
shape of the frond (whether broadest at base or narrowed downwards). In
all these characters one can see close similarities to various members of the
great mass of ‘‘ modern ”’ terrestrial ferns which are now commonly included
in the genera Dryopteris, Thelypteris, Athyrium and their respective allies (see
below). It seems to me also that the indusia of these genera could all have
been evolved from the various types of indusia known in Cyatheaceae. I
suggest therefore that these great groups of ‘‘ modern”’ ferns are derived
from the same stock as Cyatheaceae, though there are considerable differences
in anatomy, scales, sporangia, spores and chromosome numbers whic
indicate that the ‘“‘modern’’ ferns could not be included in a family
yatheaceae
The ‘‘ modern”’ leptosporangiate ferns
Until very recently it has been the custom to include almost all ferns
other than the families mentioned above in a comprehensive family Poly-
podiaceae, which included some 80% of the world’s ferns. But it has
become increasingly evident that such a “ family” included a number of
more or less distinct sub-groups, each of which must have had an indepen-
dent evolution; and sporangia indicate that they all came originally from the
same stock as the “ primitive” families. The only rational solution seems
to be to try to distinguish these different evolutionary lines, and to call each
of them a family. There will inevitably be some difficult and doubtful cases,
but I think the general scheme is now becoming fairly evident. However,
no two pterologists agree on details, and I prefer at present to arrange the
genera in ‘‘ groups,” rather than give these groups definite family names.
BRACKEN (Pteridium aquilinum) belongs to a not very clearly defined
group of genera which all have a hairy creeping rhizome and so are more or
less capable of forming thickets. This group is rather on the border-line
between ‘‘ primitive ’’ and ‘‘ modern ”’ as regards form of sporangia and sori.
The genus Preridium, which is world-wide, is specialized in its sori which
spread from one vein-end to the next, and in the complex anatomy an nd
branching of its rhizome. The important tropical genera are Dennstaedtia,
Hypolepis and Microlepia; 1 think these still need to be more clearly define d.
There is an awkward genus which has sori much like those of Dennstaedtia
y
problems. Chromosome study might help, but not enough species have
yet been investigated. In the new Flora Europaea the Bracken family is
called Hypolepidaceae; Copeland includes it in his too-comprehensive family
Pteridaceae. (For a note on Pteris, see the Adiantum group.
208 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
THE THELYPTERIS GROUP. This is represented in Britain by three
species, now usually known as Thelypteris palustris (type of the genus),
T. limbosperma and T. phegopteris. The whole group includes about 900
species throughout the world. C. V. Morton is so impressed by the common
characters within the group that he would include all in one genus Thelypteris
(Amer. Fern Journ. 53: 149, 1963). R.C. Ching, on the other hand, proposes
17 genera for the species in Asia (Acta Phytotax. Sinica 8, no. 4, 1963).
Certainly sub-groups can be recognized, but not all have yet been clearly
characterized. Of our British species, 7. phegopteris, belongs to a small but
_ rh sub-group which would be called Phegopteris if given the rank
a genus. But there is a closely allied sub-group which Ching calls Pseudo-
ttasoamieg and it is not yet clear to me whether these should be united. It
appears that our other two British species have tropical allies (especially in
mountain forest) but they have not yet been sorted out. The details of the
scales on young fronds give the best means of recognition of genera, and they
investigations are still very incomplete. There is a range of basic numbers
from 30 to 36, contrasting with 40-41 in the Dryopteris group. It seems to
me likely that our British species should rank as members of three genera,
which would be called Thelypteris, Lastrea and Phegopteris. In distribution
of hairs, and in some characters of scales, the Thelypteris group shows
resemblances to Cyathea which seem to me significant; these need to be
more fully investigated.
DRYOPTERIS GROUP. Besides Dryopteris (8 species in
Britain), I would include here Polystichum (3 species in Britain), Gymno-
carpium and Woodsia. These agree in scale-characters, rhachis-structure
and chromosome-number (40 or 41), in all of which they differ from the
Thelypteroid ferns (Gymnocarpium is sometimes included in Thelypteridaceae,
but I see no reason for this). Dryopteris is a very large and world-wide
genus, reaching its maximum in the region from the Himalayas to West
China, where its distinction from Polystichum possibly breaks down. The
solution of recognizing a third genus (wrongly called Rumohra by Ching and
Polystichopsis by myself, correctly Arachniodes) seems to me not yet clearly
established. Gymnocarpium is a very distinct genus of few species in north
temperate regions (extending to the Philippines and New Guinea); it was
ormerly included in Polypodium because of the absence of an indusium, but
in other characters is very unlike Polypodium. In many genera of ferns
(including Cyathea) some species have lost indusia, and one can find indusiate
and exindusiate species which in other respects are extremely alike, so that
the absence of an indusium by itself is not a very important character in
classification. Woodsia is a more isolated genus, of about 38 species, on
mountains in the northern hemisphere and in South America. Its cup-
HOLTUM: BRITISH FERNS IN A WORLD PERSPECTIVE 209
shaped indusium is at first sight very different from the kidney-shaped
indusium of Dryopteris, but within the genus Cyathea differences as great
can be found; I think that both the Dryopteris and the Woodsia form of
The North temperate Ostrich Fern (Matteuccia struthiopteris, hardy in
Britain) is Dryopteris-like in scales and indusia, but has very reduced fertile
pinnae, superficially like those of some species of Blechnum. The genus
Onoclea (now north American, known as a fossil in Britain, often cultivated)
is usually associated with Matteuccia, but has important differences, including
chromosome-number, and its place seems uncertain.
THE ELAPHOGLOSSUM GROUP. This comprises several genera
which appear to be related, and together to form a distinct offshoot from
the same stock as Dryopteris. They have reduced fertile fronds which are
covered with sporangia on the lower surface, and their rhizomes are dorsi-
subtropical. The most aberrant genus is Elaphoglossum, almost all species
of which are epiphytes with simple fronds (one species in Madeira; the
majority in South America but a few through Africa and Asia). The climb-
ing genera are exceedingly interesting, and one of them (Teratophyllum) has
been much confused because young plants bear peculiar fronds which have
not always been correctly associated with fronds of mature plants.
THE TECTARIA GROUP. This is entirely tropical and subtropical.
It has sori much like Dryopteris but a different rhachis-structure and peculiar
multicellular hairs on the upper surface of the rhachis.
THE ATHYRIUM GROUP. In Britain we have three species of
Athyrium; with these I would associate the genus Cystopteris. Alston and
others have recognized a family Athyriaceae which includes these genera and
Woodsia. 1 cannot see why such a family should not be united to Dryopteris;
if it were, it would be called Aspidiaceae. The whole question seems to me
to be still open. But I would not agree to the suggestion of Ching and
Copeland that Athyrium is nearly related to Thelypteris.
Athyrium is a world-wide genus of nearly 200 species; but Copeland has
added to it the genus Diplazium of at least 400 species, stating that no clear
line can be drawn between them. To draw such a line one needs to eliminate
some peculiar groups of species as separate genera, and this has not yet been
satisfactorily accomplished. It is a very complex, worldwide, problem.
tropical species of Diplazium are among the commonest terrestrial forest
fe The sori of some of them are very like Asplenium, for which reason
they were formerly included in Asplenium. But in anatomy, rhachis-form
210 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
and scales they are very different from Asplenium, and I believe that the two
genera have evolved their similar sori from different origins. The genus
Athyrium (sens. str.) shows how the elongate sori of this group have evolved
from the kidney-shape of Dryopteris; all stages can be seen on one pinna of
A, filix-femina.
Cystopteris is a genus of 10 species. Two of these have been separated
as a genus Acystopteris, but this ranks as a subgenus in the latest monograph.
Most species grow in rock-crevices, like our native C. fragilis, which itself is
worldwide apart from the wet tropics.
THE ASPLENIUM GROUP. Asplenium (seven British species) is a
worldwide genus of 700 species, most of them epiphytes or rock-plants.
Copeland includes Phyilitis in Asplenium, and I would agree. It is evident
that the primitive condition in Asplenium is a large much-branched frond (the
largest tropical ones may be six feet long). From this condition, various
evolutionary lines have led to simple fronds, and in species on at least three
of these lines there are pairs of sori facing each other as in Phyllitis scolo-
pendrium. It is clear that a definition based on this single character would
give a quite unnatural genus. It is also clear that Phyllitis scolopendrium is
closely related to some of our native species of Asplenium, as demonstrated
y the occurrence of hybrids. The genus Ceterach is more distant, but still
near enough to Asplenium to permit of intergeneric hybrids; it is a genus of
few species of north temperate regions. The position of Asplenium in
relation to other groups of genera seems to me still uncertain. It is perhaps
nearest to Blechnum, but not very near.
HOLTUM: BRITISH FERNS IN A WORLD PERSPECTIVE 211
THE ADIANTUM GROUP. In Britain we have the genera Adiantum,
Cryptogramma and Anogramma, one species of each. ere is no true
indusium (though the edge of the frond is sometimes reflexed, protecting
the young sporangia) and the sporangia run along the veins. These three
genera represent a very large and diverse group of genera which mostly occur
in the drier and warmer parts of the world; few occur in the wet tropics.
Opinions differ as to the delimitation of families within the group, and more
study is needed. The genus Adiantum includes more than 200 species, most
numerous in South America, whence many have been brought into cultiva-
grow under glass. Cryptogramma is credited with only three species, all
north temperate, but it is very closely allied to some other genera. Ano-
gramma is a small genus of world-wide distribution. It seems probable
that all the genera in the group are allied to Schizaeaceae.
The genus Preris, worldwide and complex, is often included in this group
of genera, though formerly associated with Pteridium. It still seems to me
possible that it is allied to Pteridium rather than to the group of Adiantum
and Cryptogramma.
THE POLYPODIUM GROUP, or Polypodiaceae in the strict sense.
This is a large and very natural family, almost all members of which are
epiphytes (as our British species often is). Until recently all members of
the family which have round sori (a majority) were included in a very large
genus Polypodium (which was also made to include the genus Grammitis and
its allies, now regarded as constituting a distinct though probably allied
family). The present tendency is to recognize a number of distinct genera,
but their precise limits are not agreed; in particular it is not yet clear what
should be the limits of a restricted genus Polypodium (I am doubtful whether
the treatment in my book on Malayan ferns is a good one). Almost all
members of the family have veins in an elaborate network, with free veinlets
in each mesh of the net; Polypodium vulgare with its free veins thus looks like
A similar netted
(some in Britain). It has therefore been suggested that Dipteris represents
the ancestral stock from which modern Polypodiaceae have evolved. But
Dipteris has many differences from Polypodiaceae (in cytology, anatomy,
scales and sporangia) so that it is probably better in a family of its own.
Specialization has occurred in the family Polypodiaceae in various differ-
ent ways. Some species have very thin fronds and are adapted to life in
shady forest near streams (some on wet rocks rather than trees), others are
very tough and grow on exposed tree-branches; in some species of bo
212 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
and tough types, sori have spread along the veins and joined to form lines,
or in some cases have covered the lower surface of fertile fronds completely,
the genus Lecanopteris have a fleshy rhizome, parts of which internally are
soft. The soft parts are eaten away by ants, which make their nests in the
cavities. The ants also bring small seeds, some of which germinate in the
crevices between branches of the Lecanopteris rhizome, and so a mass of
epiphytes (often of large size) starts on the branch of a tree. The most
extreme specialization is found in the genus Platycerium, which has two kinds
of fronds, one growing upwards, collecting dead tree-leaves which rot and
form a compost for the fern’s roots, the other kind growing outwards or
downwards and bearing the sporangia, which are in large fertile areas and
are protected when young by interlacing stellate hairs. There is another
very distinct group of species, now placed in several genera allied to Drynaria,
which have two kinds of fronds serving similar purposes to those of Platy-
cerium but in a simpler way.
much more study before we can have a satisfactory conspectus of them, and
that the whole problem is one of considerable complexity.
REVIEW
COLO URED ILLUSTRATIONS OF THE JAPANESE PTERIDOPHYTA
Motozi Tagawa; (IV) +270 pages, 72 plates; Osaka: Hoikusha, 1959, and
pa Ree reprinted; price about £3.
this eh ag and reated work the entire fern flora of Japan is
illustrated. Although the main text is in Japanese, Latin names are given
and each of the ya (1-399) is numbered so that it is easily identified or
looked up in the INDEX PTERIDOPHYTARUM JAPONICARUM which
occupies pages 173-262. The latter is a complete enumeration of Japanese
ferns under their Latin names, giving referen ynonymy
aw ed species, with all synonyms cross-indexed. Inevitably there will
be s gr over nomenclature, for example in the use of Lastre
; men a
instead of Thelypteris. The figures were prepared from colour-transparencies
and are good, even though there is some loss of detail. For those interested
m mete sig ferns but not acquainted with the Japanese language, the x
the itietentiorss het make this book most valuable. The price, though
high, is not excessive and the standard of production is excellent.
Royal Botanic Gardens, Kew FRANCES M. JARRETT
BRIT. FERN GAZ. 9 (6) 1965 213
TAXONOMIC NOTES ON JAMAICAN FERNS
G. R. PROCTOR
Institute of Jamaica, Kingston, Jamaica
ABSTRACT. New combinations are proposed in Arachniodes, Blechnum
Grammitis, Pityrogramma and Stigmatopteris. In addition are the fo llowing
e cies: Blechnum antillanum (p. , Ctenitis ore
parvula (p. 00), and Pityrogramma williamsii (p. 00). Asplenium sciadophilum
e for a species wi
e
Polypodium microchasmum is transferred to the synonymy of P. /euco-
Sticton, and a few other ferns are also discussed.
There seems to be no end to the problems confronting anyone who attempts
to deal comprehensively with the ferns of Jamaica. The present writer has now
spent more than 15 years trying to learn enough concerning these plants to
publish a book about them, and usd it eee rs admitted that other activities
have taken up a good deal of time, one might suppose the end to be in sight.
problems (not dealt with here) lie in the genera Diplazium, Hypolepis, and Poly-
podium, and also in some general questions of family and generic status. It is
- ;
uture
r is indebted to William T. Stearn of the British Museum (Natural
History) for lacnizing bee descriptions of new species. Also acknowledged with
gratitude is the opportunity to have worked for a time at the Bri tish Museum
(Natural History) and fe Kew ake bs so many types of Jamaican ferns are kept.
nless otherwise noted by the appropriate yibols, cited specimens are in
the herbarium of the Institute of Jamaica (IJ).
ARACHNIODES Blume, Enum. P1. Jav. 241-242 (1828)—Byrsopteris Morton
in Amer. Fern Journ. 50: 149 (1960).
The separation of this taxon from the related genera Dryopteris and
Polystichum has been thoroughly discussed es Tindale (Contr.N.S.Wales
Nat.Herb. 3: 89-90, 1961; op. cit. Flora Ser. nos 208-211: 35-59, 1961).
te
overlooked Blume’s earlier name. This necessitates new combinations again
or the two Jamaican species of the genus
~ ARACHNIODES denticulata (Swartz) fae comb. nov., based on Polypo-
dium denticulatum Swartz, Prodr. 134 (1788). This species has also been
placed in Aspidium, Polystichum, Nephrodium, Lastrea, agi Rumohra,
and Byrsopteris by various authors. TYPE from Jamaica, Swar
CHNIODES rigidissima (Hook.) Proctor, comb. nov., based on Nephrodium
enticulatum var. rigidissimum Hook., Sp. Fil. 4: 148 (1862); Dryopteris
rigidissima (Hook.) C.Chr. in Dan. Vid. Nagress Skr. VII, 6: 118, f.27 (1 cab.
Byrsopteris rigidissima (Hoo k. ) Morton in Amer. Fern Journ. el 152 (19
E from Jamaica; Hooker cited * Purdie, ei sed (who obse ‘I mistook
this at first for the leaf of a carrot *)”, but Morton (under Byrsopteris) wees
i only the Wilson collection (K), ise aionity choosing thi
Otype.
ASPLENIUM sciadophilum Proctor, nom. nov. “‘Caenopteris ogg nie Smith”
sensu Hook, & Grev., Ic. Fil. a ’t.193 (1831), non Smith, Pl. Ic. Ined. .50
214 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
(1790); non Asplenium rhizophyllum L. (1753), nec Kunze o5 Anite:
r.) B
myriophyllum Spreng. (va macraei Griseb., Fl. Br. W. I. 684 (1864), p
ea no f Avene macraei Hook. & Grev. (1831). Asplenium pin 8
ar. aii? Jenman, Ferns Br. W. I. & Guiana 162 (1909), non
A
eieleiinan “een rag in Wilkes (1854). TYPE from Jamaica: Hook .
Grev., Ic. Fil. 2:
NF mith’s gale rhizophylla was based on a specimen allegedly
from Ceshiicn collected by Thierry, which Smith received from Thouin;
this appears to represent the species now called Asplenium conquisitum
derw Maxon.
are
narrower and more attenuate. ut does not appear to be very closely allied
0 the A. myriophyllum group, which it has sometimes been confused in
the past. Hooker and Greville’s pat am type) i wn sys excellent panei
and can be exactly matched by numerous Jam pecimens, which
ong
can be cited Clute 135 (PH), aaa 7166 (JAM), aa P1353 AM), Proctor
a sg 24640 (IJ, etc.), cae ver 100 (UP). Other, mostly unlocalized
ns occur in many her
Max annotated various + avbatick specimens with the unpublished
epithet Wophboin. I have not seen Maxon’s own material or his notes, and
specimens seen, H. H. Smith 2206 (NY, PH) from Santa Marta, Colo mbia mi
pra ng Chad close, Further study is required to establish the npr yenr
0 ese
BLECHNUM antillanum Proctor
I noy.
Rhizoma breviter ee Me ae ens, apice paleis nitide brunneis deltato-
Stheniatie squamatum ndes ascendentes, plerumqu ue 15-30 cm_longae,
fere dimorphae; stipites ent versus sparsim squamati, ei frondium fertilium
PROCTOR: TAXONOMIC NOTES ON JAMAICAN FERNS 215
laminas aequantes vel excedentes. Laminae oblongo-ovatae ad late ovatae,
8-15 cm latae, basi perfecte pinnatae, sursum pinnatisectae; segmenta supera
basi adnata parum dilatata, apice a pinnam magnam integram vel basaliter
auriculata abrupte redacta; pinnae 3-8-jugae, lanceato-lineares vel lanceato-
1
breviora; venae simplices vel 1-2-furcati. Sori continui vel interrupti, ei
divisionum vin ie plerumque basi secus rhachim decurvi; indusium
age _ atu
ema eeping to ascending, scaly at apex (the scales glossy
brown, “detateattenuate), _ akly stoloniferous. eine ascending, mostly
15-30 c ng, subdimorphic; stipes sparsely scal ards base, those of
the fertile ae as long as de cmdian the heer Blades oblong-ovate to
ay ovate, 8-15cm toad "fully p innate at base, pinnatisect upwardly, the
upper segments being adnate and slightly dilated at py abruptly reduced 2
apex to a large entire or basally auriculate apa pinna; pinnae 3-8 pair:
0-15
lance-linear or lance-oblong, falcate, 1 —18) broad (widest eae
middle), the lowest 1-2 pairs equal to or slightly shorter Seed the next upper
ones and often slight! flexed; veins simpl 1- or 2- d. Sori
e
tinuous or interrupted, those of upper divisions usually Socucved at base
along the Monat indusium me aig rors
PE m Jamaica, i eri t. Mary: near the Ugly River Bridge,
J.P.282 (J AM, isotype US). P siahich oe from the same locality, Maxon
ws). akan 21398 (IJ); St. Andrew: Brandon Hill, Harris 7422 (JAM);
: deep ravine above House Hill, Maxon GH, NY, US).
nea Repcblie: Arroyo Loma Bajita, Valeur 891 (IJ, ex x US). What is
apparently the same species has also been recorded fi Se the —_ rsa
Guadeloupe & St. Vincent), and a very similar occu
America (cf. British Honduras: Gentle 5071; and onder: Srandley 72626).
The species therefore probably has a wider distribution than here indicated,
but at the prese che sandal i is not available.
This Seats Aas 5 calied Blechnum lonsifoiam by Jenman and his con-
temporaries, and B. fraxineum “‘sens. lat sit later writers, but differs
having ‘fewer, relatively larger pinnae, terminated abruptly in a large terminal
pinna, byt the lon nger ee y ill sor of fertile fronds), and by the sori of the
upper pinnae being conspicuously decurved at the base. Material from the
type locality i is now under cytological ite in England, but information from
this source is not yet available
It is possible that a afore poles name already exists for this species,
if it occurs in ane et: s L. M. Underwood said long ago in a letter
to William Harris, ‘Ther a perfect “mg of Blechnum species in the
cecidentae-longifollun-unilaterale group”. These have never been properly
elucida i have named the Jamaican plant pata tab to emphasize that
the species being described is based strictly on West Indian material. Its
precise position must await a definitive monograph of the group.
YBLECHNUM PLUMIERI var. nigrocarina
polypodioides Swartz, Prodr. 127 (1788); ponent cere —
Kuhn, Fil. Afr. 92 (1868), non Raddi (1819); Struthiopteris polypodioides
(Swartz) Trev. in Atti Ist, Ven. III, 14: 571 (1869). TYPE from Jamaica,
wart.
a the study by Deaeeruawt: A ater a Bot. Club 39: 270-275, 1912)
oO Indian Struthinnteris S nlumie
216 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
as separate species. However, they are obviously very papain related, and an
examination of numerous collections shows that there is an intergradation in
Lesser Antilles area. This suggests that they are better treated as
variction v of a single polymorphic species (now placed in the ee meio
In considering possible epithets to use for the variety occurring in Jam
it is evident that the name polypodioides is available at the wareat level, tt
, based on ype of Swartz’
name would also oe available at the le oe for a same plant.
BLECHNUM POLYPODIOIDES Raddi in Opusc. Sci. Bol. 3: 294 (1819) Se
re Asis from Brazil, Raddi (not ee illustrated in yn Fi 1 We
1806
79, 3, f.1 (1810), ‘illeeit mate becaus perfluo publis
blechnoides (Lag. ex Swartz) C.Chr., Ind, Fil 151 F905? se "Keys neis73),
Under the present International Code of Bota nical Nomenclature, it
seems Bs wiped to take up Raddi’s name for this common and widespread
specie name cannot continue to ee used for the following
petite which requires a new epithet if recognized as a species.
/ CTENITIs harrisii Proctor, sp. no
Rhizoma decumbens, 7-10 eon agri ee apice paleis fuscis flavo-brunneis
plerumque uno tantum in quoque ultimo segmento; indusium firmum
pallide brunneum glandis destitutum.
Rhizome decumbent, 7-10 mm thick, densely clothed at apex with dark
yellow-brown, linear-attenuate scales to 1. 5 cm long, 0.4—1 mm broad, these
mot arene and with entire margins. Fronds several, fasciculate, up to 30
m long; stipes shorter than or equalling the blades, 6-15 cm long, deep
‘chdideteoon, essentially eglandular, bearing numerous spreading, dark
brown, filiform scales to 8 mm long, each fom a minutely swollen bas. the
cells eg Tee. = shethente;: Blades 3-pinnate, lanceolate or lance-
oblo cm broad below th , long-acuminate
t i plane -deliats, acuminate, at least the lower ones subopposite,
ote (up to 3 cm apart), 3—4.5 long, 2~3 cm broad at base, stalked,
su ilateral; sec divisions 7-10 a the lowest ones delta eo
divisions oblique, oblong-subfalcate, scutsh or unt,
appressed-ciliate with articulate hairs: costae and costu Dee ‘ben: oe bearing
small brown bullate scales with hairlike to: ultimate veins obscure, |-
PROCTOR: TAXONOMIC NOTES ON JAMAICAN FERNS 217
orked, bearing minute appressed articulate hairs on upper side; —_
pment and eglandular on both sides. Sori small, —. 1 to each ultima
segment; indusium firm, light brown, glabro us and without glands.
TYPE, Parish of St. Thomas: Old England, Blue ap Harris 12903
(JAM). The ent was collected at ‘3500 feet altitude, on She y slope near
the falls’’, Feb. 28, 1919. PARATYPES: a Andrew, Jo art,
JP 176, ake ete (BM, JAM). This material has slightly stele ie issue
but other agrees with C. haves: tatties than the following specie
Bositiey nie ated: Hart (P).
-CTENITIS paryula Proctor, sp. nov. Nephrodium pubescens Desv. in Mém
S Lite Paris 6: 361 (1827), nomen nudum; non Polypodium pubescens L.
(1801).
(1759), nec Aspidium pubescens Swartz
—8 mm crassum, apice paleis flavo-
brunneis lineari-filiformibus ad 7 mm longis 0.2-0.3 mm latis eglandulatis
integris dense vestitum. — plures fasciculatae 8-23 cm longae; —
ncaa breviores, 3-10 cm longi, fusco-brunnei minute glandulo-puberu
infra i 3-5 cm latae, He aA Legg pond pinnae
deltatae obtusae’ suboppositae, inferiores 2 srt (us
distantes), 1.5-2.5 cm longae, basi 1.5-2. ine petiolaine subaequi-
c
Paucis sparsis minutis brunneis bullatis — apice attenuatis praeditae;
] i i appressos et tos subtus
nacea minute
sparseque glandu
gla ndulis parvis porn on’ dense tectum.
= bent or mangers ng, 5-8 mm thick, _— clothed at
7 mm long, 0.2-0.3 mm
u
scales less than 2 mm long, the cells
2-pinnate-pinnatifid or barely 3-pinnate, lance-oblo
l 3-5 cm broad below the middle, a mente 4
obtuse, subopposite, the lower ones remote
long, 1.5-2.5 cm broad at base, s
3-6 pairs, the went ones chiefly oblong,
divisions all o tuse, the margins bearing a few, 1U
costae and costules beneath with a few scattered, a dus
scal m 4
be 5 eath. : supramedil;
i
TYP ena on w, “below” John Crow Peak, Harris 7335
(BM, isotypes JAM, K). This ps te was doubtfully cited by Christensen
(K.Dansk.Vid.Selsk.Skr.VIII, 6: 39, 1920) as heniets epeinions, but
appears to differ significantly from that rig wos Cuban species. PARATYPE:
Loca i jamai ux, apparently the basis of
Nephrodium pubescens Desv. bly represented
merely a misidentification by oat
218 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
A plant with somewhat similar oblong fronds from the Cockpit Country
of icleway (Proctor 4071) is probably only a juvenile igs of C. hirta, as
shown by the structure of the scales and other characters
EICHENIA PALMATA (Schaffn. ex Fourn.) C.Chr., Ind. Fil. 323 (1906).
(Mertens palmata Schaffner in Fée ém. Foug. 9: 32 (1857), nomen
Gleichenia palmata Moore, Ind. Fil. 380 (1862), nomen nudum
pe moana palmata Schaffn. ex Fourn. in Nyl. & Besch., Mex. Pl. 1: 137 ( 1872);
TYPE from Jalapa, orote le. ad 229 (isotype K). Dicranopteris palmata
Schaffner ex Underw n Bull. T Sane nay 34: 259 a TYPE from
Orizaba, Mexico, Pris 9 (NY tc.); Maxon in
es fern aN r. Fl. 16: 61 (1909); phite patloe (Underw.) * cout Gen. Fil.
This is a peculiar case of nomenclatorial confusion. Fournier’s effective
publication ‘of Schaffner’s name was overlooked by Underwood in his
treatment of ease plants, and the latter described this entity under Dicran-
opteris with the same specific name but based Jaca’ p different type. In ss
this, he created a taxonomic synonym which c t be treated as a nomen-
clatorial one, though the specific epit ar are achaleal" Chiisenien’ s
combination of Gleichenia palmata can be tre as new and valid because,
alth ngly cited ‘‘(Schaffn Moore” as authorities (they having
published mere nomina nuda with no standing), he also cited the Fournier
work in which a description of this species first appeared.
GRAMMITIS Swartz in Journ. Bot. Schrader 18002: 17 (1801).
_ When separating the so-called “grammitid”’ ferns from Polypodium _
which genus they were long included), there are two possible courses 0
action. One is to set up a series of essentially ‘“‘one-character’’ genera —
as proposed by Copeland (Genera Filicum, Ann. Crypt. Phytopath, es 1947)
and others; the other is to establish an inclusive genus for most of these
abviowsly ‘tated ferns, separating out only the most aberrant groups at the
generic level. I prefer the latter course, and some of the species occurring in
amaica have already been transferred (Proctor in Bull. Inst. erage Sci. Ser.
. 31-36, 1953; Rhodora 63: 35, 1961; Brit.Fern Gaz. 9: 8-9, 1962) to an
expande d g enus Grammitis. The remainder of the Jamaican sabe are
hereby canned as follows:
RAMMITIS aromatica (Maxon) Proctor, comb. nov., based on Polypo-
dium aromaticum Maxon in Proc. U.S. Nat. Mus. 27: 743 (1904); P. firmum
auctt. non Kaulf. TYPE St Jamaica, Underwood 1449 (NY).
brunneo-viridis (Baker ex Jenman) showed comb. es
based on Polypodium brunneo-viride Baker ex Jenman in Jou ot. 15: 2
(1877). Clenchrarts meridensis sensu Copel. in Phil. phar Sci. ‘84: ota (1950),
pro parte, non Polypodium meridense cloteach. TYPE from Jamaica,
Jenman 32, in 1877 (K).
/GRAMMITIS capillaris (Desv.) Proctor, comb. noy., based on Polypodium
capillare Desv. in Berlin Ges. Nat. Fr. Mag. 5: 316 (1811). P. graveolens
—_ ex Jenman in Journ. Bot. 16: 265 (1877); Ctenopteris graveolens (Baker
ex Jenman) Copel. in Phil. Journ. Sci. 84: 408 (1956). TYPE from Jamaica,
ex herb, qc CP).
Is chrysleri (Copel.) Proctor, comb. noy., based on Crenopteris
sear ‘Copel: in Phil. Journ. Sci. 84: 448 (1956). Polypodium che
: octor in Bull. Inst. Jam. Sci. Ser. 5: 48 (1953), nomen nudum. TYPE fro
maica, Maxon 1524 (
Y SGa Nears cretata (Maxon) Proctor, comb. nov., based on Polypodium
PROCTOR: TAXONOMIC NOTES ON JAMAICAN FERNS 219
cretatum eta in Amer. Fern Journ. 5: 51 (1915). Polypodium albopunc-
tatum Baker ex Jenman in Journ. Bot. 15: 265 (1877), non Raddi (1819).
TYPE from Jamaica, Jesihan = in 1877 (K
GRA M & Gal.) Proctor, comb. noy., based o
Polypodium delicatulum Mart. & Ga 1. in Mém. Aca d. Bruxelles 15: 547 f 1
(1842). TYPE from Oaxaca, Mexico, Galeotti 6378 (BR, fragm. US, isotype
“GRAMMITIs exornans (Maxon) Proctor, comb. noy., based on Polypodium
exornans Maxon in Amer. Fern Journ. 18: 47 (1928). P. asplenifolium sensu
Jenman in Bull. Bot. Dept. Jamaica II, 4: 123 (1897), non L. (1753), TYPE
n9
“GRAMMITIS aiden (Slosson) Proctor, comb. noy., based on Polypodium
insidiosum Slosson in Bull. Torr. Bot. Club 39: 287, t.23, f.4-8 (1912). TYPE
from Cuba, Shafer 8043 (NY). This species has not pre reviously been reported
rom Jamaica; it is perhaps not very — differentiated from G. trifurcata.
The Jamaican population occurs only in the John Crow Mountains at the
eastern end of the island. Specimens seen snatch Proctor 5566, 5641, 22710,
and K. Wilson and G. Webster 534.
RAMMITIS arya (Undiew: 4 Maxon) Proctor, comb. novy., based on
Polypodium jenmanii Underw. ex Maxon in Contr. U.S. Nat _ Herb. 16: 62
(1912). fo Soe a sensu Jenman in Bull. Bot. Dept. "Jamaica II, 4:
118 (1897), excl. ref. Guadeloupe, non Mett. ex Kuhn (1869). TYPE from
Jamaica, Jenman 25, in 1879 (
MITIS moniliformi s (Lagasca ex Swartz) Proctor, comb. nov.
on Polypodium moniliforme Lagasca ex Swartz, Syn. Fil. 33 (1806). "TYPE
from Peru, Lagasca (not seen
spiape
>|
is
a
&
ats
uJ
oO
re!
z
o
S50
a
25 |
a
: /
wi
(a)
afr4o
5
c o
35 i
30
+25 R
va
ws) N
> =
5
Ww 7
Zi
=
80
: N e
—
N
oO] °
. pee
\
+40 Be i
°o
+20
Sane! ae =
STRIP NO. eee Se. wr TUNE AEE 1 ae FS
BRAUNTON ASHYON COURT SHERE
FIGURES 1-3 Histograms showing % fertility of ned lopm
to shade; FIG. 2 sho “age foe prvem with gl : the pm
height in feet
diameter), FIG. 3 uaabasion with li
-- ity in num
arf: th I a intensity - ay of full hight (o—0).
226 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
green and show a marked reduction in thickness.
The Ashton Court graph shows a negative correlation between frond
density and fertility, but this trend is contradicted by the Shere data, so that
little can be concluded on the relationship between the two.
Thus there appears to be some degree of correlation between fertility
and the size and morphology of the fronds, but a more exact measure of
vegetative development is required before this can be assessed in any detail.
Conway (1957) suggests that restricted rhizome growth will retard both
vegetative growth and sporangial development, so that a direct measure of
rhizome size and vigour may have to be employed in future investigations.
Only a preliminary and rather unsatisfactory investigation of the
edaphic factors in the Ashton Court and Shere areas was carried out, in
which the soil reaction and the moisture content of the top-soil and sub-soil
were measured, The results indicated that the less fertile fronds were growing
in soils of high acidity and with high moisture and humus contents. In
view of previous work on bracken, it seems likely that moisture content is
the most important of these three factors, and it is possible that the slightly
higher moisture contents recorded for woodland soils caused a reduction of
rhizome vigour sufficient to account for the observed reduction of frond size
and fertility. This interpretation would imply a reduction in fertility at the
margins of waterlogged areas where rhizome growth should be similarly
restricted, and is supported by Conway’s observation that a sand-dune
population was sterile when exposed to a high brackish-water table (Conway
1957). A more detailed investigation of the degree to which edaphic factors
change at the edges of woods is also required.
The humidity of the air in deeply shaded situations will obviously be
meter were interesting in that they showed no such changes, but as they were
taken after a fairly long, dry spell, it is possible that more abrupt changes
might be recorded in more normal climatic conditions.
The intensity of the light is the most obvious of the possible factors
affecting the fertility of the fronds, but this does not necessarily mean that it
is the most important. If measurements of light intensity can be made in
different areas in such a way that the readings can be directly compared,
some idea of its importance in relation to other physical factors may be
obtained. The exact comparability of the readings is vital, and, as slight
the two areas. In Ashton Court light intensity was measured by means of
an Eel light meter, and at Shere a Weston Universal Exposure Meter was
used. The readings were taken at frond level at about noon on bright
DRING: SHADE AND BRACKEN FERTILITY 227
summer days, and the values plotted are the averages of fifty readings taken
in each strip. In order that the two sets of data should be more nearly com-
parable, the results are expressed as a percentage of the light intensity in
the o
However, it is doubtful whether a reading taken at any one time bears
much relation to the light actually received by the plant during the course of
the year. The use of hemispherical photographs to estimate both diffuse
light and direct sunlight, as described by Anderson (1964), is one very simple
and promising method of measuring the total effective light received by the
plant, and another is described by Dore (1958), but similar problems apply
also to the other constantly changing physical factors of a micro-habitat,
humidity and temperature, and until these are solved critical analysis of the
detailed relationships of these factors is impossible.
Bearing such reservations in mind, it is worthwhile to consider the
information which can be gleaned from the data obtained in this investi-
gation. Assuming that the two sets of data, when expressed as percentages
of light intensity in the open, are comparable, it is noticeable that the fronds
show different degrees of response to light intensity (Fic. 3). Thus the
Ashton Court fronds seem to show a greater range of fertility in response
to a smaller range of light intensities. This would suggest that light is not
the sole controlling factor, and that the differences in response are due to
some other factors, such as soil, climate or vegetative development, which
are contrasted in the two areas. If more detailed and reliable information
about two more directly comparable areas was obtained, this variation in
the degree of response to light could be confirmed, and the data could be
subjected to statistical analysis to determine the effects of pairs, or larger
combinations, of factors on the fertility of the bracken fronds.
REFERENCES
Se. M.c. 1964, Studies of the woodland ight climate I. The photographic
computation of light conditions, J. Eco : 27-41.
BOODLE, i A. 1904, The structure of the leaves of ibe ee a aquilina L.)
n relation ee environment, J Bot. 35: 6
BRAID, K. 1959, Bracken, a review of the literature, ose te Bureau
Pepa a Publications 3/1959
BURKE, ms Pp. T. 1953, Study of the influence of light and — properties on the
rowth of bracken, Mag. Blundell’s Sch. Sci. Soc 13-17.
ee E. 1957, Spore production in bracken, J. Ecol. 45: Fa oh
DORE, w. G. 1958, A simple chemical light meter, Ecology 39: 151-3.
GOEBEL, K. 1905, Organography of Plants (Part I), Oxford University Press.
SCHWABE, W. W. 1951, The mineral nutrition of bracken I. Prothallial culture and
the effects of phosphorus and — supply on leaf production in
the sporophyte, Amn. Bot. 15: 417
TANSLEY, A. G. 1939, The British Isles and their Vegetation, Cambridge University
Press.
WOODHEAD, 1906, Ecology of hg plants in the neighbourhood of
Huddersfield, J. Linn. Soc. Bot, 37: 333-406.
228 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
NOTES AND REVIEWS
SPORELING BRACKEN ON LITTLEWORTH COMMON
A NOTE ON ECOLOGY OF BRACKEN
er
able to make direct contact with a suitable substrate. Rarely, as at Little-
The summer of 1959 was very dry in England, only some 3.6 inches of
rain being recorded at Kew Observatory from May to mid-September;
normally, nearly 10 inches would have fallen in the same period. As a result,
dr
destroyed. This area of the common, about 50 feet above sea-level, consists
of fairly widely spaced birch trees with bracken and Molinia caerulea in the
herb layer. The soil is sandy and partially leached with a small dispersed
iron pan and about two inches of peat on the surface. The fire, which was
observed only in its dying stages on October 18th, progressed erratically
leaving “islands” of bracken in some places while burning away the peat to
expose the mineral soil in others. Some of the birch trees were blown down
by later autumnal gales, after having had their major roots (in the top 12
inches of soil) destroyed by the fire to leave only smaller roots penetrating
deeper. The immediate effect of the fire was to provide an open habitat ready
to be colonised anew and also to stimulate fresh growth on those plants
which had survived; Molinia caerulea and Potentilla sp. sprouted leaves,
seeds germinated and the remaining birches developed buds at the bases of
their trunks. (In the following spring and summer most of the bare ground
was colonised with seedlings, mainly by wind-dispersed seeds such as of
birch and rosebay-willowherb). On December 13th the first organism of an
unusual type seen was a fungus Anthracobia melaloma, a discomycete with
an orange ascocarp up to 5 mm in diameter, growing in small clumps on the
blackened soil; by the first week in February it was being overgrown by a
filamentous green alga Ulothrix sp., and a second species Anthracobia
macrocystis, lemon yellow in colour, was thriving.
¥:
weather proceeded to warm up, only 1°F of frost being recorded once at
Kew between then and the summer, further bracken prothalli were not seen
MELVILLE: ECOLOGY OF BRACKEN 229
until mid-April. The moss protonema appeared more hardy as they con-
tinued to gow. Conway found that bracken prothalli could survive several
degrees of frost and a covering of snow but that dry cold was quickly fatal.
It is possible that it was partly through desiccation that these prothalli died.
hey were usually found in small sheltered patches such as in a footprint or
under the collapsed dead fronds of the previous year’s growth where the
humidity was presumably higher. Bracken sporelings were up to two inches
high by the end of May and were growing in competition with a flush of
seedlings which included birch, rosebay-willowherb, groundsel, gorse and
poplar. The Anthracobia had disappeared and instead grew another orange
fungus of similar appearance Pyronema confluens and a small brown agaric
Flamula carbonaria. The first sporophyte fronds of bracken are pale green
and simple in shape; later ones are pinnate and there is a gradual increase in
size and complexity. After four months they were up to 18 inches high and
approaching mature fronds in shape though still more delicate and of a
lighter green. An interesting point is that each plant had two rhizomes set
rigidly at about 90°-100° to each other which grew downwards into the soil
and levelled off at their normal depth of six inches.
When area was revisited four-and-a-half years later it was pre-
dominantly covered by eight-foot saplings, mainly birch with a few willow
and aspen. Some of the original bracken was left, as well as Molinia and
birch and oak trees. Leaf-litter removed by the fire had reaccumulated and
the ground was partially moss-covered. There was no apparent way of
distinguishing mature sporelings from original bracken, except by locality.
Mature bracken fronds appeared between birch saplings on areas which had
previously been swept clear by the fire. It seems probable that this particular
area will develop into a thicket of trees much like some other areas on
Littleworth Common.
Ecology of the Bracken gametophyte
The sexual stage in the life-history of bracken is usually found only in
protected, sterilised and alkaline sites. Such sites, as noted by Conway, are
usually on burnt ground, in damp hollows (such as quarries and embank-
ments), and on old walls or rubble (such as bombsites). One reason given
by Braid for this scarcity is that flowering plants and mosses produce sub-
stances which somehow inhibit the growth of the bracken prothalli. Conway
found that bacteria, moss protonema and especially algae have mar ed
suppressing effects on bracken prothalli but she queried as to whether this
was due to competition for nutrients. Also, alkalinity is found to favour
their development (optimum range is pH 5-7) and potash is found in high
concentration in bracken tissues; it would be released into the soil when
the plant is burnt and this seemed to be corroborated by a higher pH on the
burnt area. In similar, but unburnt areas, the pH was 5.5-5.6 in the top
8 cms of soil The burnt land was pH 6 on the surface and pH 5.5 at 8 cms
depth. This measurement was taken in June 1960 and some leaching probably
230 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
occurred during the winter. However, the fact that bracken spores developed
on this burnt area and not elsewhere in the vicinity may be due simply to
the direct contact the spore was able to make with the soil. The spores are
small and are not equipped for prolonged growth without water and
nutrients. Normally, the soil is covered by other plants or loose leaf litter
which prevents the rhizoids of the prothallus from contacting soil or similar
moist substrate. Occasionally, as at Littleworth, this surface covering is
removed and the full life-cycle of the plant is completed.
REFERENCES
BRAID, K. Sug 1959, Bracken: a review of the literature, Commonwealth Agric.
au Mimeographed Publ. 3/1959: 69 pages
CONWAY, “% 1949, The autecology of Bracken (Pteridium aquilinum (L.) Kuhn):
e germination of the spore, and the development of the prothallus
and the young sporophyte, yey R.Soc.Edinb. (B) 63: 325-343.
121 Mortlake Road, Kew. J. D. MELVILLE
THE FERNS IN FLORA EUROPAEA
FLORA EUROPAEA Volume One. Lycopodiaceae to Plantanaceae; edite d
by T. G. Tutin, V. H. Heywood, N. A. Burges, D. H. Valentine, S. ss ——
and D aly eb with the assistance of P. W. Ball and A, O. Chat XXL
[+ii] +4 64 [+4] pages, 5 maps, 22 x 28 cm; Cambridge, Cadet Pris,
1964; 84 shillings net.
Review of the chapter on ferns
For ist
plants, je pie oe of the first of four volumes of Flora Europaea is 4
m e i
u
taxonomy which is both modern and practical in the sense that classical
methods based on morphology, a s far as possible, are suppleme ented by
results deduced from cytology and tibet imental taxonomy. For every species
keys, descriptions, geographical distributions ~ om if possible, chromosome
are given. Such data are also given recognised subspecies. Ihe
€ common hybrids are mentioned as well ~ taxa of doubtful status.
This should encourage further studies of the latter.
a Europaea begins with Pteridophyta ren ges 1-25), of which pages
8-25 are thn to ferns port py loner divide d into 21 families. To
following scientists (in alphabetical 0 rde ): a O. Chater, J. A. Crabbe,
- Fuchs, V. H. Heyw
ae , A. Lawalrée, J. D. Lovis, W-
Rothmaler(#, ea’ Tutin, °D. a Via T. G. Walker and D. A. Webb.
* F. A. Stafleu has reviewed the whole of Vol. 1 in Taxon 14: 105-107 (1965)
REICHSTEIN: THE FERNS IN ‘FLORA EUROPAEA’ 231
ope, when compared with other continents, is very poor in ferns.
Altogether, some 100 good species truly native to Europe are described, as
this a bigger number than expected. It is explained by the fact that about 20
are either very rare or very local or confined to remote places, so that hardly
work, giving most important information in condensed form. It will
indispensable for everybody interested in ferns, whether living in Europe or in
other continents. It is to be hoped that it may also help, at least to a certain
degree, to bring some stability in the nomenclature of ferns.
e following remarks from a non-professional but enthusiastic
amateur of ferns are offered for consideration for any addendum or
edition; provided interest in the Flora proves to be as great as is expected,
a new edition may be necessary quite soon.
Remarks
In some few cases the Flora gives extra-European distribution in
parenthesis. I regret that this very useful information is not extended to
more, if possible to all species. I would also welcome “‘apogamous”’, after
the chromosome number of obligate apomicts; it is given for Preris cretica
but not for all known cases; this property is often quite important to under-
stand particular distribution patterns and the forming of local clones with
distinct morphology. I wonder how long our well known Beech Fern
(Phegopteris) will really feel happy in Thelypteris. Under Dryopteris
abbreviata (DC.) Newman, the indication “sori usually . . . 1-2 (-4) on each
pinnule’’ corresponds to De Candolle’s original diagnosis but is correct only
for dwarf plants: well developed plants from Spain, France and Italy
invariably had 6-8 sori on each of the longer pinnules (see pictures, Reich-
stein 1962). Probably most Swiss botanists will not agree that the alpine
strains of Woodsia glabella R.Br. should be separated specifically, as W.
pulchella Bertol., from North-American and some Scandinavian elements, as
suggested by Poelt (1952, 1954); they tend to accept them as conspecific, in
agreement with Meyer (1959) and Brown (1964); experimental taxonomy
may well help to settle this question and find the reason for some morpho-
logical peculiarities in certain northern forms. Pteridologists in Switzerland
will also regret the revalidation of the name Polystichum aculeatum (L.) Roth
for the Hard Shieldfern; perhaps this could not have been avoided, in
accordance with the International Code of Botanical Nomenclature, and
may even correspond to British usage; it will, however, perpetuate existing
difficulties in evaluating the meaning in older continental European literature,
where the same name was not only used indiscriminately for both species
but, even when used specifically, was much more frequently attached to P.
setiferum and not to the species which we would have preferred to call
232 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
P. lobatum (Huds.) Chev. in agreement with Janchen (1963). In the distri-
bution of some species of Asplenium, a few minor errors may be mentioned.
For A. obovatum Viv., Portugal is given twice; according to Pinto de Silva
(1959) and Fernandes (1960) this species probably does not grow in
Portugal; if this is true, it would be confined to mediterranean coasts. For
A. billotti F. Schultz, Switzerland (where a few plants are still growing, see
literature and pictures in Manton & Reichstein 1962) was not given, and
Germany is given in error; the locus classicus (near Wasigenstein, Alsace
has been French territory since 1918; the fern is not known within today’s
frontiers of Germany (teste A. Schumacher in litt. 17.XII. 1964). Switzerland
is also not given for A. onopteris L. where it is not uncommon in the Tessin
and is also reported from the Misox (Becherer 1959) and the Puschlav
(Becherer 1950). It is true that in some cases a sure differentiation between
A. onopteris and A. adiantum-nigrum may require cytological investigation,
and the whole complex needs further, particularly experimental study;
therefore not all distribution records for these two species are reliable; in
well-developed plants, like those found in southern Switzerland, it is usually
quite easy to distinguish A. onopteris from A. adiantum-nigrum, but in small
or dwarf specimens this may become very difficult or even impossible;
macrosocopic morphology is not reliable in such cases; spore size is helpful,
but the chromosome number is most reliable.
There are some ferns which, for editorial reasons, are not included in
Flora Europaea. One such case is Dryopteris remota, and I take this oppor-
tunity to append the following remarks:
In spite of its doubtful status, D. remota (A.Br.) Druce could have been
mentioned, although this would have raised some unsolved taxonomic
questions (see Gatzi 1961). Not only have different taxa been described under
this name (see Manton 1938, 1950) but probably the same slightly variable
taxon is described under different names, including some of those given by
Rothmaler (1945, 1963). What I mean by D. remota is the triploid apogamous
viable spores (Fischer 1909, 1919; Waldmann 1928; Dépp 1932), thus
behaving like a true species. It most probably corresponds to the plants
collected by A. Braun in the Black Forest and also to var. subalpinum Borb.
(1876). It has nearly always been assumed to be a hybrid. This is most
probably correct, but so far nobody could prove that it is a recent hybrid
originating where it is found. Out of the many different pairs of species
REICHSTEIN: THE FERNS IN ‘FLORA EUROPAEA’ 233
species both of which are still growing in Europe. An apogamous hybrid
producing viable spores and therefore behaving to some extent similarly to
which tetraploid and pentaploid forms are known, as mentioned in the
Flora. Both have successfully been produced by experimental hybridisation
of D. filix-mas with diploid and triploid D. borreri respectively (Dépp 1941,
1955). This shows that the apogamous nature is inherited from D. borreri as
one parent in at least this particular case.
REFERENCES
BECHERER, A. 1950, Beitrige zur — des Puschlav, Jahresber. Naturforsch.
Ges rau ubiindens 82: 131-1
—— 1959 , Beitriige zur Flora des Misox, ibid. 88: 1-27.
BORBAS, Vv. DE 1876, Symbolae ad pteridographiam et Characeas Hungariae
raecipue AP Verh. Zool.—Bot. Ges. Wien 25:
BROWN, D. F. M. 19 A m Spy aie a of the fern genus Woodsia,
eihefte zur Nova Hedwieis
DOpP, w. 1932, Die Apogamie bei pf a m bes: AI.Br., Planta 17: 86-152.
cosmic? Ueber Dryopteris paleacea Christensen (D. Borreri Newm.), Ber.
eutsch. Bot. Ges. 59: 423-426.
—— 1955, Experimentell erzeugte Bastarde zwischen Dryopteris oe (i)
opteris paleacea (Sw.) C. Christensen. Planta 46: 70-91.
FERNANDEZ, R. 1960, Notas sobra a Flora de Portugal VIII, Bol. See. Brot. (2)
: 99-165.
FISCHER, - 1909, Ueber Aspidium we AI.Br.: Kreuzung oder Mutation ?-
neuer Fall von Apogamie, Ber. Deutsch. Bot. Ges. 27: 495-502.
—— 1919, coe mie bei Pace beaterdee, ibid. 37:
GAETZI, w. 1961, Ueber den pie ars Stand der Dryopterisforschung, Ber. St.
all, Natures, Ges. 17: 3-73.
JANCHEN, . 1963, Ge Anderte Namen von Gefasspflanzen Oesterreichs, Phyton
Bac 10: 1-1
MANTON, ar 1938, Hybrid Disa (Lastrea) in Britain, Brit. Fern Gaz. 7:
—— 1950, un aR Core and Evolution in the Pteridophyta, Cambridge
MANTON, GQ and seicwah ite T. 1962, Diploides Asplenium obovatum Viv.,
seat coal 79-91,
MEYER, D. 95 Die Chromosomenzahl der Woodsia glabella R.Br.
Mitte fileubenah, Willdenovia ~ 214-217.
PINTO DA SILVA, A. R. 1959, Plantas ete e novas areas para a flora de Portugal
, Agronomia Lusitana 20: —247.
POELT, J. 1952, Zur Kenntnis der cadens Woodsia in Europa. Mitt. Bot.
Staatssa amm. (Miinchen) 5: 167-174.
—— 1954, Woodsia pulchella Bert. — ein verkannter Farn der Alpen, Ber.
Bayer.
ene T. 1962, Dryopteris abbreviata (DC.) Newman in Apennin,
Bauhinia 2: 95-113. 4.
oe ~ 1945, Der Formenkreis von Dryopteris paleacea (Sw.) Hand.-
, Candollea 10: 91-101. rey
—-— 1963, “Eskursionsflora ae Deutschland IV. Kritischer Ergdadnzungsban
a Berl
ora han Pane zur ered ns. nde der Prothallien
einheimischer ‘sibpeadecoee Diss. Marburg
The University, Basle. T. REICHSTEIN
234 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
A FOURTH SUPPLEMENT TO CHRISTENSEN’S INDEX
INDEX FILICUM, SUPPLEMENTUM QUARTUM PRO ANNIS 1934-
1960 (Regnum Vegetabile vol. 37), prepared by Rodolfo E. G. Pichi-Sermolli
with the collaboration of F. Ballard, R. E. Holttum, H. Ito, F. M. Jarrett,
SAC Lk. i
Taxonomy; xiv+370 pages, x cm; Utrech; International Bureau for
Plant Taxonomy and Nomenclature, January 1965; price 40 Dutch florins.
Supplements to the Index Kewensis issued at approximately five-yearly
intervals keep phanerogamic botanists informed of new names and transfers
relating to the genera and species of flowering-plants. Pteridologists are less
ortunate. The third supplement to Christensen’s Index Filicum (1905 ’
index of the names of pteridophytes, followed by supplements in 1913, 1917
and 1934. The public-spirited Carlsberg Foundation of Copenhagen acted
‘INDEX FILICUM, SUPPL. 4’ (REVIEW) 235
regards citations, this supplement follows the New York Botanical
Garden guide to citation appended to the International Code of Botanical
Nomenclature. This means that it fails to distinguish between periodicals and
isolated independent works, a distinction of great importance to librarians.
Thus, we have as references: Ching, Sunyatsenia 5:261; Holttum, Rev.Fl.
Mal. 2:162; Praia, Stud.Bot.Cechosl. 1:94. A librarian will search in vain
Cechoslovaca or for a periodica sh title might be abbreviated to Rev.Fi.
Mal.; the insertion of ‘in’ before the name of the periodical in which the
author published obviates sinbionsi . Fortunately, — supplement ——
with a catalogue of literature for ae eriod 1934 t ea which not only
i also : number f icati
an index of abbreviations of names soheaemneng botanists concerned with
flowering-plants will also find these u
Compilation of the iciteeas was paren ted by April 1964.
accordance with the 1961 International of Botanical Nomenclature,
names of bigeneric hybrid groups originally published near a description
omina nuda a
otanical Cotigress at Edinburgh in roan 1964, the relevant article ao) of
s fourth supplement to nrbtouene. gettin ntal work reflects
great credit oa all ge er with its production. Its coverage seems to
have thorough and its compilation accurate and it is rong re
printed, ehiitg ales ita bedian tool both easy and pleasant to
British Museum (Natural History), London WILLIAM T. STEARN
236 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
THE SOUTHPORT FLOWER SHOW
26—28th AUGUST 1964: FERN CLASSES
Judges’ report (by R. Kaye, President of the British Pteridological Society):
Mr W. G. Mackenzie (Curator of Chelsea Physic Garden) and I were both
immediately impressed by the quantity, variety and improved appearance
of the fern exhibits which were better groomed and much cleaner than last
year, and in general were a credit to their owners; it is by no means an easy
task to lift, pack and travel with a collection of ferns without damaging them.
In several cases competition was keen and we had a difficult task to allocate
the placings.
Mr Brookfield had no competition for Class 8, but richly deserved
First Prize; his group included some magnificent Polypodium vulgare varieties,
several fine Athyrium filix-femina varieties such as ‘Clarissima’, ‘Victoriae’,
‘Plumosum Druery’ and ‘Plumosum Percristatum’; Polystichum varieties
included good examples of P. aculeatum ‘Pulcherrimum Bevis’ and ‘Gracilli-
mum Druery’; the exhibit attracted considerable interest and Mr Hayhurst,
who spent most of the three days in the Fern Tent, was kept very busy answer-
ing questions about it and the other fern exhibits.
e prizewinners and their entries in the other Classes are listed below.
There were some very fine plants among them, involving much careful
cultivation often for many years; among those which impressed us most
were: the British species of Mr Jackson (Classes 15 and 16); the hart’s-
tongues of Messrs Almond and Hayhurst (Class 10); the entries of Athyrium,
Dryopteris and Polypodium in Class 9 which was particularly popular;
Mr Hayhurst’s Adiantum ‘Kensington Gem’ (Class 19) which we recognised
from previous shows and which is a tribute to its owner; Mr Plowman’s
Adiantum ‘Fritz Luthi’ (Class 19); Mr Brookfield’s Polypodium vulgare
*‘Hadwinii’ (Class 11); and in Class 17 Mr Hayhurst’s Phyllitis scolopendrium
‘Crispum’ about three feet in diameter and in perfect condition. Class 18
attracted only two entries.
When entering a Class for more than one fern, make sure that each
specimen is as good as possible, because no specimen can gain more than its
proportion of the total points. Damaged fronds were evident on several
plants, particularly Polystichum in Class 12. A little grooming might have
saved the largest specimens in Class 18 from being placed Second because
of damaged and discoloured fronds. Some of the best plants in Class 13 were
spoilt by weathering of the fronds; although this is difficult to avoid in late
August, it downpoints in the judging.
The British Pteridological Society’s Information Bureau (report by R. Kaye
and N. Robinson): as in former years, the Society is indebted to the South-
port Show Committee for providing free space in the marquee devoted to
horticultural societies. Many thousands saw the Bureau and many hundreds
of enquiries about ferns and the Society were answered by Messrs Almond,
SOUTHPORT FLOWER SHOW 1964 237
Brookfield, Hayhurst, F. Jackson, J. Jackson and Robinson. For the first
time the Bureau was decorated with an L-shaped limestone rockery set with
dwarf varieties of British ferns. Great interest was aroused, many wanted
to buy Polypodium vulgare ‘Trichomanoides’, and it remains to be seen how
many new members were recruited. Despite a greatly increased interest in
ferns from a few years back, it seems that something additional is needed
to crystallize interest into membership; for this reason a discussion was held
during the Annual Excursion at Brantwood to find means of making the
Bureau more effectively attractive; as a result of the Brantwood discussion
hopes are high for the Bureau’s future achievements.
Prizewinners and their entries (reported by N. Robinson, Northern Meetings
Secretary of the British Pteridological Society):
CLASS: 8 ie group of hardy British ferns, with accent on diversity and variation,
rranged in a natural mann . ip i space 13 feet by 8 feet and not
higher than 6 feet above the
Ist Messrs J. Brookfield & Son hs pale to hold the Silver Perpetual
hoemat Trophy presented by the British Pteridological
CLASS 9 Six dissimilar hardy British ferns with accent on variation
Ist Messrs Brookfield: Athyrium filix-femina ‘Congestum Pa ul’, A.f-f
sum Druery’, Polypodium vulgare ‘Elegantissimum’, P.y.
*‘Hadwinii’, Poly tichum aculeatum ‘Gracillimum’, P.a. ‘Pul-
cherrimum
2nd_—sCF. Jackson: Athyrium ” fillx-femina ‘Acrocladon’, Dryopteris a
smun regal ‘Cristata’, Phyllitis scolopendrium ‘Grandiceps’,
Polypodium vulgare ‘Cambri icum’, Polystichum aculeatum ‘Pul-
cherrimum
3rd B. Hayhurst: Athyrium filix- spss ira ‘Frizelliae’, Dryopteris ‘Ramosis-
sima’, Phyllitis scolopen rummondiae’, Polypodium
yulgare iaicerrien um’, Polys Sti tichum setiferum ‘Divisilobum
nsum’, P.s. ‘Imbricatum Paes um’.
CLASS 10 Three distinct varieties of Phyllitis scolopendrium:
Ist B. Hayhurst: Beir. ‘ ‘Ramo-margina ata’, ‘Speciosa’.
2nd_=sONN.. H. Almond: ‘Crispa’, ‘Drummon ndiae’, ‘Ramo-marginata’.
3rd Messrs Brookfield: ‘Crispa’, ‘Cristata’, ‘Sagittato-cristata’.
CLASS 11 Three distinct varieties of Polypodium vulgar
Ist essrs Brookfield: eat nese ‘Cornubiense’ ee
2nd __—B. Hayhurst: ‘Grandiceps Cowan’, ‘Hadw winii’, ‘Pulcher
3rd N. Robinson: ‘Cambricum’ pea form), cote’; Cestatn’
(good form).
sare 12 hee distinct varieties of Polystichum:
Robinson: P. aculeatum ‘Pulcherrimum Bevis’, P. —
‘Divisilobum Grandiceps Jones’, P.s. “Div bos Ivery :
2nd Messrs Brookfield: P. aculeatum ‘Gracilimum’, P. sata Sali
bum 2th ‘Acutilobum
aa. C«@B. ig yhurs : P. aculeatum *Pulcherrimum Bevis’, P. setiferum
‘Divilobum Iveryanum’, P.s. *Hirondelle
238 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
CLASS 13 Three distinct varieties of Athyrium filix-femina:
Ist B. Hayhurst: ‘Frizelliae Capitatum’, ‘Frizelliae Cristatum’, ‘Vic-
toriae’. a
2nd =-N.. H.. Almond: ‘Cristatum’,‘Plumosum Superbum Druery’, ‘Victoriae
3rd Messrs Brookfield: ‘Acrocladon’, ‘Congestum Paul’, ‘Plumosum
Cristatum
CLASS 14 Three distinct varieties of PE ad (including Gymnocarpium and
Thelypteris) (all exhibits were Dryopteris):
Ist F. gpa tent “Crispatissima sleinane “‘Lineata Cristata’, ‘Ramo-
tata’
2G. 3. Hayhurst: ‘Angustata Cristata’, ‘Depauperata Cristata’, ‘Ramo-
He” ON, EL Alinatilt ‘Angustata’, ‘Angustata’ (dwarf form), ‘Lineata
ristata’.
CLASS 15 Three rac varieties of Asplenium (some distinct species were
submitted):
Ist ¥ Jackson: adiantum-nigrum, trichomanes ‘Cristata’, viride ‘Cristata’.
2nd =N. Robinson: ruta-muraria, septentrionale, viride.
3rd Messrs Brookfield: adiantum-nigrum, trichomanes, un-named variety.
CLASS 16 Three distinct normal species of hardy British ferns:
Ist F. Jackson: Asplenium ery: Dryopteris dilatata (‘‘sub-
species”), Woodsia ilve
2nd = Messrs Brookfield: E ecuiohan dryopteris, Polypodium vulgare,
Thelypteris palustris.
ara: NN; agate cate Blechnum spicant, Dryopteris carthusiana, Polystichum
set.
CLASS 17 One British fern, any kind or variety:
Ist B. Hayhurst: Phyllitis ca ae ium ‘Crisp
2nd = Messrs Brookfield: Athyrium filix-femina ‘Plimosum Cristatum’.
3rd N, H. Almond: Aihyriaa flix. -femina ‘Cristat
CLASS 18 Six dissimilar greenhouse ferns:
Ist Messrs Brookfield: Adiantum ‘Kensington Gem’, Davallia riper fre,
Nep oe behead tena Soh Ag Platycerium alcicorn
Polypodium aureum var. Ate
2nd__—sCiB..:_ Hayhurst: Adian selee ‘Cnt ium falcatum, Lomaria
—— Nephrolepis ‘Cristata’, Wesleeienes exaltata, Platycerium
CLASS 19 Three dissimilar greenhouse ferns:
Ist — se eam atte Gem’, Davallia cana-
2nd__—sCwBB._ Hayhurst: Adiantum “Kensington Gem’, Nephrolepis ‘Cristata’,
cum.
3rd Barnes: Adiantum cuneatum, Nephrolepis exaltata, Nephrolepis
mosum’,
CLASS 20 One greenhouse fern:
Ist B. Hayhurst: Adiantum ‘K,
2nd Plowsaa: Adiantum ‘Fri Ng eas a opeak
3rd Messrs Brookfiel Id: Adiantum ‘Kensington Gem’.
BRIT. FERN GAZ. 9 (6) 1965 239
BRITISH PTERIDOLOGICAL SOCIETY
MEETINGS IN 1965
IMPERIAL COLLEGE OF SCIENCE & TECHNOLOGY, LONDON sw7,
14 March. An account illustrated with colour-slides by Mr A. C. Jermy on
his plant-collecting visit to Trinidad in the summer of 1963, followed by a
conversazione and exhibits.
ROYAL BOTANIC GARDENS, KEW, surrey, 16 May. Mr H. J. Bruty
BEM conducted a tour of the ferneries, including the new filmy-fern house.
BROCKENHURST and LYNDHURST, HAmpsuirRE, 19-21 June. As in 1963,
noteworthy ferns among 18 species seen were Ophioglossum, Osmunda and
Pilularia, In addition we found Ceterach officinarum and Polystichum
Setiferum.
AUSTWICK area, MID-WEST YORKSHIRE, 4 & 5 July. Together with other
lime-loving species, several plants of Polystichum lonchitis were seen during
this Northern weekend meeting.
CHELSEA PHYSIC GARDEN, Lonpon sw3, 25 July. The Curator, Mr
W. G. Mackenzie, showed and explained the treasures of the Garden after
telling us its history from 1673. Among the collection of British fern species
in the conservatory, it was good to see our three filmy-ferns thriving in a
Wardian chamber among other ferns and liverworts, and also some fern
varieties remaining from the days of Thomas Moore’s curatorship 1848-1887.
In one of the glasshouses, part of the plant of Asplenium x murbeckii col-
lected by F. Jackson in Borrowdale (GAZETTE 9:49,110) was still flourish-
ing. Professor J. Ewan and his wife, of Tulane University, were with us; we
were charmed to have them and to strengthen the link between ourselves and
the American Fern Society of which he is a past-president.
TUNBRIDGE WELLS area, KENT, 23 August. After assembling at Eridge
Railway Station several localities were visited, including Eridge Green and
Saxonbury Hill, as we worked our w ay across country to Frant. Many
Species were recorded, including Hymenophyllum tunbrigense and Dryopteris
aemula, It was a great pleasure to have with us two members of the American
Fern oe Dr R. M. Tryon and his wife Dr A. F. Tryon, of the Gray
Herbar
THE eka EXCURSION, 12-19 September. Thirty-two members
(including Dr Patzak from Vienna) and three friends attended at Brantwood,
on the east side of Coniston Water in the Lake District. Formerly the home
of John Ruskin, Brantwood is now the conference and study centre of the
Council for Nature
We gathered on the Saturday in lovely weather, looking forward to a
happy week among the Lakeland ferns. It gave us great pleasure to see many
new faces and to welcome new members. The afternoon was occupied with
fern-hunting in the vicinity of the house, and in the evening we assembled
240 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
in the lecture-room (Ruskin’s former study) to hear an illustrated talk by
the Warden, Mr Jeeves, on ecology in the Lake District.
Our Meetings Secretary, Squadron-Leader Halligey, had prepared a
very full and active programme which kept us on the move all the week.
The only fault was that it covered so much ground that we had little time
to devote to our immediate neighbourhood, and this will have to be remedied
during our next visit. We shall be going there again, for the area is an excel-
lent one for ferns, and Brantwood makes an ideal centre—and a comfortable
one, especially in wet weather. We can vouch for this because, after a fine
weekend, the rain began on Monday and continued all the week. On only
one day, however, did it upset our programme, and on this day we made
good use of the time with talks in the lecture-room.
On the morning of Sunday we decided to follow the nature-trail laid
out in the Brantwood grounds which cover 250 acres. We started off con-
scientiously following the trail until we reached a ravine cutting deep into
the hillside, a veritable fern paradise, and—the fern hunt was on, the nature
trail forgotten! Polystichum aculeatum and P. setiferum flourished, Hymeno-
phyllum wilsonii was noted several times, Gymnocarpium dryopteris and
Dryopteris draped the steep banks and precipitous rocks. Altogether, within
the confines of the estate and within a small radius of the house we recorded
21 fern species, and in the lake below the house JIsoetes lacustris was found.
The attraction of Coniston Old Man across the Water could not be
ignored, and one of our first visits was planned for this area, as it offered us
a variety of habitats from the lakeside up to the high crags; bad weather
postponed it for two days. Starting off from Coniston, we recorded 16 species
on the way up from the village, including Osmunda regalis. On the fell itself
we added only Asplenium adiantum-nigrum, Lycopodium clavatum an
L. selago to the list.
A visit to Whitbarrow Scar made another memorable day—both for
finds and for rain! Leaving the cars at Witherslack Hall we followed a wood-
land track to the base of the Scar, noting on the farm walls Asplenium ruta-
filix-femina, Dryopteris borreri, D. filix-mas, D. dilatata, Polystichum aculea-
tum, Thelypteris limbosperma and, rather surprisingly, Dryopteris carthusiana
and Blechnum spicant. It was interesting to note the lime-hating B. spicant
growing here, indicating the presence of pockets from which the lime had
been leached.
The woodland area to the south of Coniston Water provided an inter-
BRITISH PTERIDOLOGICAL SOCIETY MEETINGS 241
lude on a day when the rain stopped for only a few hours. We listened to a
most ae lecture by Dr Holttum on the ferns of the world (see pages
the morning while the rain poured down, and returned about
four o sees when it recommenced to hear Mr Kaye give his presidential
talk on variations of the Ladyfern (see pages 197-204). We noted 16 species
during the day, including a fine colony of Ceterach officinarum.
A day was spent renewing our acquaintance with Borrowdale where
the Annual Excursion was centred in 1954. It was a hurried visit and we con-
centrated our attention on Castle Crag where we found Asplenium septen-
trionale. Dryo, opteris: abbreviata was growing in crevices on the top of the
dence, something we are often diffident about doing, because stunted plants
of D. filix-mas and D. borreri growing in rock and mountain habitats can
be very misleading. Gymnocarpium dryopteris and C. fragilis were seen on
the Crag, H. wilsonii in the woods at the bottom, and in the damper places
T. phegopteris was abundant. We also noted two polypodies, P. interjectum
and P. vulgare. In all, 18 species were recorded from this small area. Some
of us went on to see Mr Fred Jackson’s garden and fern collection at Stone-
thwaite, and it was a joy to see so many of our more interesting ferns so
well grown.
Humphrey Head is an interesting limestone spur jutting out from the
coast a few miles south of Grange-over-Sands, and supports a flora which
includes some rarities. Our visit was a short one so we were able to explore
only a small part of the Head, and not many species were seen. The varietal
find of the week was made here, a colony of Polypodium australe “Semila-
cerum’. On our way to Humphrey Head we visited the Nature Reserve at
Roudsea Wood where we were welcomed by the Warden who conducted
us round this large tract of woodland and heath. Many unusual plants were
seen, including a rare sedge Carex flava, which is one of the reasons for the
protection given to this locality. Ferns were prolific and we recorded 17
— including Ophioglossum vulgatum and Osmunda regalis. Our day in
e Grange-over-Sands district culminated in a visit to our member Miss
teas, who most hospitably provided tea in one of the local hotels in
Cartmel before we visited her fernery. Her most interesting collection of
British and foreign ferns is in a greenhouse and a conservatory and in the
garden borders; in her house she had laid out an exhibition of fern books
and herbarium specimens. We carried away with us gifts of plants as well as
memories of a very happy visit to this generous lady.
On the following day we set off on our homeward journeys but, before
our ways divided, we again received a gracious hospitality—this time from
our President and Mrs Kaye who invited us to their house and nurseries in
the morning before entertaining us to lunch at a Silverdale hotel. It is worthy
of note that in he te nurseries we were able to see one of the best varietal finds
made during the Society’s excursions in recent years, Asplenium trichomanes
242 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
‘Incisum Greenfield’, a wild find by P. Greenfield in the Quantock Hills in
Somerset in 1960, and without doubt the finest incised form of this species
yet found. The plant was successfully established in Mr Greenfield’s eae
in Surrey, but he felt it would be more likely to survive in Mr Kaye’s care
and so it was sent to Silverdale about two years ago. It was evident from
the fine development and vigorous condition of the lovely plant we were
able to admire that the decision to send it North was a wise one.
So ood fern varieties were found this year, the best being Polypodium
australe etre collected from Humphrey Head. Crested ladyferns
were found on Whitbarrow, and more were seen along the roadside near
Brantwood as well as crested forms of Dryopteris borreri. On a wall im
Keswick we were introduced to a colony of Asplenium trichomanes ‘Crista-
tum’, the numerous plants being a very fine form of this variety. Other
variations of a minor nature were noted, including a crenate form of Ceterach
officinarum.
Among the pteridophytes seen during the week, 13 were new records
and 4 were confirmations of pre-1930 records for 10-kilometre squares (we
visited seven) of the ATLAS OF THE BRITISH FLORA. A survey of
polypodies segregated 154 specimens into 116 Polypodium vulgare, 29
P. interjectum, 4 P. australe and 5 pentaploid hybrids. These records, together
with 34 segregates within the malefern complex (17 Dryopteris filix-mas, 15
D. borreri, 2 D. abbreviata), have been passed to the Nature Conservancy
for inclusion in the ATLAS and its forthcoming critical supplement.
In the following ~_ of pteridophytes seen during the excursion, figures
1-24 (bold =new record, * confirmation of pre-1930 record, for the ATLAS)
refer to localities:
\splenium na eae es
f 2,440, 24
A. ruta-muraria — 2, 4, 5, 7, 10, 72, 13 PESTS 17: 28
A. septentrionale —
f
f
anes — 2, 3, 4, 5, 7, 10, 12, 13, 14, 15, 17, 24
\. viri oe 4
Athyrium filix-femina — /, 2, 3, 4, 5, 7, 12, 13, 14, 16, 18, 19, 20, 22, 23, 2
Blechnum spicant — /, 2, 2 4, < 6, 9, 73, 14, 16, 20, 23, 24
Ceterach Fi Heme ah >a 14*, 15*
Cryptogramma crispa — 2, 3, 5.6, y 9,17, 20, 22,'23, 24
® 1 24°
"yeopodium cclavatum —7
ystopteris fragilis — 2, 14, 2
ryopteris abbreviata 24
orreri — /, 2 ee Ap A A ae 14, 16, 19, 20, 22, 23, 24
carthusiana — /, 13, 14*, 16* < 19
). dilatata — 2, 3, 4, CA 9, 12, 13, 14, 16, 18, 19, 20, 22, 23, 24
). filix-mas — /, 2, 3, 4, 5, 10, IZ. 13, 14, 15, 16, 17, 19, 20, 22, 23, 24
». villarii — /4
— a am 5, 13; 17
. telm —21
( iymnocarpium dryopteris — 2, 3, 24
G. robertia —I
] iymenophyllum wilsonii — 2, 23
Isoetes lacustris — 2,
]
]
BRITISH PTERIDOLOGICAL SOCIETY MEETINGS
Orme vulgatum — /3
Osmunda regalis — 8, /3
Phyllitis scolopendriuim — 2, 10543, Tei Se77
ypodium australe —
P. interjectum aap ne eee 17216, 20,22. 23 oe
PSV eon re — 1, 2, 3, 4, 5, 6, 7, 9. 12, 13, 14, 15, 17, 18,20, 22; 23, 24
P. interjectum X ‘vulgare — 3, iF,
A ate “aioe SOMERLOT 7 ds 10, 3. "4 16
P. setiferum — 2, 10
Pteridium aqu ilinu m — /, 2, 3, 4, 5, 6, ;, A 13,44, ).16.17, 7 19, 20, 22, 23, 24
Thelypteris limbosper ma — 2, 35342 55i 7, Oo. 75.4 14, 16, 20, 23.
T. phegopteris — 2, 3, 5, 9, 23, 16, me 23, 24
V.C. 69 Fs re yb gi (with North Lancaster) visited National
(O.S. 1” map ser. 7 sheets 88 and zt September Grid Ref.
1 pe "Sw co : Le ht 15 34/2991
2 BRANTWOOD 12-19 195, 3196
3. BURNEY 15 258
4 CONISTON Villa 12-17 309
5 CONISTON, near Kanwey? Stati tion. 16 2997
6 CONISTON, road above Railway S 16 2997 A
7 NISTON, moor track leading to ‘Church Beck 16 2798, 289
8 CONISTON, Churc 16-17 2998
9 CONISTON OLD MAN, cliffs near Low Water 16 2798
10 HUMPHREY 18 3973, 3974
11 LAKE WINDERMERE 14 3995
12 LOWICK BRIDGE aS a 3281, S 282,
13 ROUDSEA woop, Nature Reserve ae oe 3381, 3382
14. WHITBARROW SCAR 14 4485, ert
15 WITHERSLACK HALL farm walls .. 14 gr nag
16 ITHERSLACK HALL wo fa He ie 14 7189
17 WoopLaNpD Church vicinity 15 3489
18 WooDLaNpD Church to Row Ridding, roadside . 15 5489
19 WOODLAND vicinity, Row Ridding, wo 15 pod
20 WOODLAND vicinity, hillside above Raisthwaite 15 aEyS
21 WRAYSHOLME TowER, ditchbank by road 18
be Oe pa Cumberland
29 S. 1” map ser. 7 sheet 82) ‘i
BORROWDALE, between ee and Hollows ee 35/2417
23 pORROWDALE, between Hollows Farm and e 2416, 2 5 6
ah 2415, »
24 BORROWDALE, Castie Crag as ee ae 17 3513, 2516
ANNUAL GENERAL MEETING, 16 September. Held during the Annual
Excursion. Reported in NEWSLETTER . fence 1965.
BRITISH MUSEUM (NATURAL RY), LONDON sw7, 24 October.
Mr J. A. Crabbe conducted a visit to she Herbarium and Library of the
Department of Botany, followed by a conversazione and exhibits.
Amendment to account of 1963 Annual Excursion
lines 16
On page 191 of the 1964 GAZETTE the vice-county numbers on
and 18 should be transposed, i.e. locality 33 should be V.C.47 and locality
34 should be V.C.46.
244 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
BRITISH PTERIDOLOGICAL SOCIETY
MEMBERS
Honorary Members distinguished by : *%
ADAMS, Prof. WILLIAM, 4 Comin St, New Concord, Ohio, U.S
ALLEN, Mrs. B. E. G., Villa Coca, Los Barrios, Provincia de Cadiz, data
n.
ALLISON, BR 26 Ives Rd, Hewlett, eit Island, New York, U.S.A.
AMERICAN FERN "SOCIETY, © Brooklyn Botan c Garden, New Yo rk, U.S.A.
ASHBERRY, Miss ANNE, Chessins, Chipeal-Sensaley, Chelmsford, Essex.
BALLARD, F., BSc, FLS, c /o Royal Botanic Gdns, Kew, Richmond, Surrey.
ANGE a? 8 u W7
BEIJING WAIWEN SHUDIAN, P.O. Box 50, Peking,
ha Pte — BSs, 29 Duke of York's Royal Military. "SiicoL Guston,
Dov
BENEDICT. RALPH C., PhD, 185 Hall St, Brooklyn 5, New York, 2 =
BE OCAL ABSTRACT S (U.S.A), University of Pennsylvania, Philade
BISHOP. Ҥ N., 45 Hol ly Rd, Handsworth, Birmingham 2
BLOOM, JOHN, ‘Garden News’, 62 High St, Stamford, ey
BOLTON, T. H., Birdbrook, Halstead, Essex
BORUP, Mrs. E. M., 30 Earl’s Court ‘sooure London, mba
BOTANICAL SURVEY OF INDIA, 14 Madan St, Ca Icu 33
BOTANISCHER GARTEN & MUSEUM, Kénigin-Luise-Str. 6-8, Berlin
BRIGHTMAN, F. H., BSc, 2 Red Oak Close, Orpington, Ken
RIGHTMAN, JULIAN J., 2 Red Oak Close, Orpington, Kent
BRITISH MUSEUM, London, Wwcl
BRITISH MUSEUM (NATURAL ede es London, SW7
BROOKFIELD, Pe 102 Stamford Rd, Birkdale, Southport, sey
BR mmf 23 Da agmar Rd, Eo -on-Thames, Surr Id
BURNETT, LESLIE, 3 Beech Hill Rd, Wylde aan dens Coldfield,
BURNS, 1 Miss AILSA, 24 Basset St, Camborne, Cornwall
a peta Ne Mass onda cig in 4, California, U.S.A.
CAPPER, 148 eg era Ave, Crew hire
CARDEN, ony ILDA G., 33 Moor P ark rk DW OON Middx.
CA RTWRIGHT, 7 F. 143 Pittmansfield. tlow,
COR. _ BOUGHTON, Cobb-web, Under “Piet Rd, ‘Falls Village, Connecti-
US.
COCHRANE, The Hon. TC. A., Balgownie, 18 Cliff Terrace, Buckie, Banfishire
COLHOUN, Pro
EY, V., Pucketty Farm, Faringdon, Ber
COULTHARD, CHARLES E., 143 Muste = Rd, West Bridgford, Nottingham.
. URE, 41 Quee
on, sw7
ept. of Botany, British ek m (Nat. Hist ), London, 5
UCATION, Ay
NE, Mrs. FERN, W., 174 Summit Ave, , ai it, New Jersey, U.S.A.
CRAW, Mrs. G. ma A., LLB, Greenways, Sutton Place, Abinger Hammer,
Dorking, Surre
CROMPTON, Mrs. on Thriplow Farm, Thriplow, Royston, Herts.
DARTMOUTH COLLEGE, BAKER LIBRARY, Hanover, New Hampshire,
retin sion) J. MB ChB FRCP (Edinburgh), Linton Muir, West Linton,
Pee
DE J ONCHEERE, G. J., Koninginneweg 8, Wassenaar (ZH), Netherlands
BRITISH PTERIDOLOGICAL SOCIETY MEMBERS 245
DEKKER EN NORDEMANN’S nin C Hoe BOEKHANDEL,
U y
DYCE, JAMES W. Hilltop, 46 Sedley Rise, Loughton, Essex
DYER, Mrs. E. M., Rockfield, Alrewas, Burton- soa iad Staffs.
DYER. aif A., FLS, Rockfield, Alrewas, B esi sek ge Staffs.
ELLISON, P. J., 7 Tudor Walk, Wattord: Hi
EVANS, G. B. BA A PhD, Dept. of Botany: Lintietilty of Malaya, Kuala
EWAN Prof. JOSEPH, Dept. of Botany, Tulane University, New Orleans 18,
isian S.A,
FINCHER, F., Randan Wood, Woodcote, Bromsgrove, Worcs
FISHER, Cc. H,, MA, Drabbington, ‘Thornbury, Bromyard, Herefordshire
FOSTER, F. GORDON, 10 North Terrece, Maplewood, New Jersey, U.S.A.
FUCHS-ECKERT, H HANS Py DPhil, Clematislaan 61, Wassenaar (ZH),
herlands
GABE, HOWARD L., 17 Penylan Terrace, Penylan, Cardiff
GARDINER, J. C., FLS, FCA, 7 Stan hope Terrace, London, W2
GATZI, , Dufour-Str. 118, St Gallen, Switzerlan
GILBERT, "GEOFFREY, 116 Haverstock Hi ll, London, NW3
GI MOUR, ao Bs , FLS, Director, University Botanic Gdns., Cambridge
GLASNEVIN BOTANIC GARDE EN, Dublin, Eire
GLEGG, Miss N. CREINA, Chessins, Chignal-Smealey, Chelmsford, —
GOWIN NG-SCOPES, ERIC, "Ros ewood, Stonehouse Rd, Halstead, Ken
GRAY HER BARIUM, 22 Divinity Ave, Cambridge sa pnts USA.
GREENFIELD, H. F., Brittain’s Lane, Sevenoaks,
% GREENFIELD, P., h B u Rd, —— Surrey
HALL, NEILL, D,. 1225 NE95 St, Seattle 15, Washington,
’ n
HALL, Mrs. J. F. 6 John’s Close, Gorsewood Rd, eotes Dartfo rd, Kent
HALL, PETER C., 6 John’s Close, Gorsewood Rd, rtley, Da rtford, Kent
mgaag Poe i Ldr P., BSc AMIEE RAF (Ret), Territet, 5 Ravenscourt Rd,
, Hants.
HAMBLIN cm H. R., 16 Shaftesbury Rd, Romford, Esse
HART, D. J., BSc, Round Oak Lodge, Bishopsgate Rd. Englefield Green,
e
HAYHURST, BERNARD, 91 Windermere Rd, agg = pieiiaebiee
HEALEY, F. J., 38 Lockstile Way, Goring, Reading, Ber
HEALEY, Mrs. J. R., BSc, 38 Lockstile Way, Goring, so Berks.
HENDERSON, CAMPBELL, 2023 W48 Ave, Vancouver, B. C., Canada
HEWITT, A. H., 29 Balfour Crescent, Woiverhampton, taffs.
HODGES, Miss JANE, 12 Northview, ‘Londo on, SW19
HOLTT TTUM, Mrs. C. E. URSULA, 80 Mortlake Rd, Kew, Richmond, Surrey
HOLTTUM, R. E. MA ScD FLS, c/o Royal Botanic Gardens, Kew, Rich-
mo rre
HOME, Miss HM. LOGAN, Edrom Nurseries, Coldingham, Berwickshire
HOSHIZAKL Mrs. BARBARA JOE, 565 N. Westmoreland Ave, Los Angeles,
Cali
HULL UNIVERSITY, Cottingham Rd, Hull, Yorkshire
et JeKs University of eg fin porns —— Ness, ae ——
N
gowrie, Perthshire
IRONSIDE, E. TF, The Orchard, 27 Slade Rd, Newton, Mumbles, Swansea,
am.
JACKSON, FRED, Knotts View, Stonethwaite, Borrowdale, Keswick,
um :
JACKSON, JAMES F., Knotts View, Stonethwaite, Borrowdale, Keswick,
Cumberland
246 BRITISH FERN GAZEITE: VOLUME 9 PART 6 (1965)
JAMES, P. W. ie FLS, Dept. of Botany, British Museum (Nat. Hist),
ondon
JAPANESE’ FERN ge a rd o T. NAMEGATA (President), 481 Narita,
Narita-shi, Chiba-ken, Tok
JARRETT, Miss F. M., MA i, FLS, The Herbarium, Royal Botanic
Ga rdens, Kew, Ri chmon d, Sur
JENKINS, F. H., 88 Turkey St, Enfteld, Middx.
JERMY, A c.. BSc FLS, Dept. of Botany, British Museum (Nat. Hist.),
n, SW7
KAYE, Mrs. MARION, Waithman’s House, Silverdale, Carnforth, Lancs.
KAYE, REGINALD, Waithman Nurseries, hha ae Ea = Lanes
KELLEY, At 1 Tudo r Rd, Wheathamstead, St. A s,
KINGSTON, Miss MARGARET a Wesley paeag eaten Hay, near
Walsall, Sta
KIPPAX, Miss ANNE, 119 Rosehill Rd, Burnley, Lancs
KNOX, Mrs. WILLIAM C., 649 Isle of Palms, Fort y aedurdale beckery? U.S.A.
LAWALREE, r. ANDRE G. C., 3 Avenue Van i ia Bruxelles
LEICESTER UNIVERSITY. Universi Leic
ae. ml INGEBORG, otanioches Institut, Piirerutels 4, 3550 Marburg,
LINNEAN SOCIETY OF LONDON, Buriington House, London, WI
OYD, E. M., 47 Bond St Nort h, Hamilton, Ontario, Canada ‘
LLOYD, ROBERT M., Dept of ‘Botany, ‘tinwersity of California, "Berkeley
California, U.S.A
TU
LONDON NA HISTORY SOCIETY, c/o Botanical Sect. Secretary,
28 Te ge ae London, SW4
LOVIS, JOHN D., PhD, Dept. of Botany, The University, seni
MACDONALD, mdb ef A., Dept. of Botany, The University, St. Andrews, Fi 4
MACKENZIE, W. G., Curator, Chelsea Physic Garden, 66 Royal Hospita
WwW
oO n
MACLBAY, Prof. K. N. G., gem of ee Bechuanaland Protec-
torate and Swaziland, P. O. Roma, Mas Basut oland TE
MAGYAR dag carr ager AKADEMIA, ’GENETIKAL INTEZETE,
Herm o U.15, Budapest II, Hungary : >
% MANTON, Prof “IRENE, PhD FRS, Dept. of Botany, The University, Leeds
MARCHAN NIH, 3. Cardiff R.D.1., Stratford, Taranaki, New —
MASHITER, JOHN, Roseneath Harewood Rd, Skipton, Yorkshire
ir fag 72 Hampton Rd, Southport, Lancs
MAY s. MARGUERITE B., 58 Bouverie Ave, Salisbury, Wilts.
MIDWEST. INTER LIBRARY ‘CENTER (C.A.), 5721 Cottage Grove Ave.,
Chicago 37, Illinois, U.S.A. d,
apse “REDHEAD, E. "TD MA FLS,7 Ashley Gdns., Petersham, Richmon
MILSUM. J. N., OBE, FLS, Grays, Tilford, Farnham, Sur
MORGAN, Prof. DENYS, MSc FLS, Dept. of Botany, University College,
Roma, via Maseru, Basuto a
MORLEY. EUV: DiAscties, Stolford, near age a Somerset
MORLEY, R., 4 Monyash Close, Cha ddesden,
MORTON, C. 'V., Cu rator of Ferns, Dept. of Boviniy, U.S. National Museum,
ete teers Institution, Washingt on — U.S.A.
MOWAT, JOHN L., SHM, University Botanic Gdns., St. Andrews, Fife.
NATIONAL BOTANIC GARDENS, Linckcow, India
NATIONAL MUSEUM OF WALES, Cardiff.
'y inbur
NETHERCOTT, P. J. M., 6 Hazelwood, Hazelwood Rd, Bristol 9
NEW_ ZEALAND NATIONAL LIBRARY SERVICE, Botany Division,
pd oe rns Bag, Bo oe gies New Z G St,
rs. J. P., BSc, of Botany, lege, Gower
Loniok W. C1 pt. otany, aiversity College
BRITISH PTERIDOLOGICAL SOCIETY MEMBERS 247
NORMAN, Miss H. D. 57 Oakington Ave, Wembley Park, Middx.
NORRIE, Mrs. A. G; Stackadale Turriff, A Aberdeenshire
OLBY, R. C., School of Botan ny, South Parks Rd, Oxford
OWEN, OW. WEN, Orient Cottage, Llanrug, ara beau cigs
PARR, Mrs. PHYLLIS A., 11 :Belgrave Rd, Ches :
PATZAK, ALOIS, DPhil Sly Botanische Abies, Naturhistorisches
useum, Burgring yi nna |
PAYNE, R. M., 8 Hilltop. Locahtak: Ess
PENFIELD, PAUL Jr, 17 Bradford Rd, Weston, Mass,, 02193, U.S.A.
PERRY, B. J., BSc, 7 Church St, Exmouth, on
PERRY’S HARDY PLANT FARM, Theobalds Park Rd, Crews Hill.
iddx.
Enfield,
PICHI- SERMOLL: Prof. Dr. RB, E.G: {stitute : Orto Botanico ‘Hanbury’,
The University, Corso bogs ali, Genova, Ita
POLLARD, R. S. W., LAMTPI, 127 Park Rd, ys don, W4
PROCTOR, G. R., BA FLS, Institute of Jamaica, Kington. Jamai
%* RAMSBOTTOM, JOHN, OBE MA DrSc VMH FLS, c/o British gtaeee
(Natural History), London, W7
RAD ee R. F., Wiggin Hill, St. Ives, yeas
9 St. Johanns-Ring, asel,
RICHM ND, Miss T. M. Richmond Hill, Silverdale, Carnforth, Lancs.
RIJKSHERBARIUM, Nonnen nsteeg, Leiden, Netherlands
ROBINS 7 Yewlands Drive, Fu wood, Preston, Lancs.
urrey
ROYAL HORTICULTURAL SOCIETY, Vincent Square, London, SWI
RUCH, (B.K.W.Z.), ul. Wronia 23, Warsaw, Po
RUTHERFORD, Miss ALISON WwW. The Ciitein Cari Eskan Farm, near
Helensburgh, Dunbartonshire
SCHELPE, anlage pew MSc DPhil feoh one Herbarium, University of
ape Tow Rondebosch, South A
SCHULZE, Be GERHARD, Heer anietig “46, Ludwigshafen / Rhein, Germany
SCHUMACHER, Dr. ALBERT, 8 Eichbornweg, 522 Waldbrdl, German ny
SCOTT, WA ALTER, Jr., Easterhoull, Scalloway, Lerwick, Shetland
SCOTT. WILFRED, 18 Crakegarth, Dalston, Carlisle, Cumberland
SHAKESHAFT, I.E. M., BSc, 26 Moon St., Wolverton, Buc
SHAW, The Rev. C. E., Holy sy Vicarage, Waterhead, Oldham, Lancs.
‘alle .
ER
SIMS, Miss P. a 131 Ruskin Park House, London
SITWELL, ySACHEVERELL, Weston Hall, pater 9 Northants.
SERay tan, 100 7 Park Rd, Sheffie
sagt sm P. a B. OTWAY, VRD DPhil MP, Colebrook House, Win-
chester, Han
SMITHSONIAN 1 INSTITUTION, Redon pe 25, D.C., U.S.A
KA, R. R., 7824 Normal Ave, La Mesa, Calif. 92041, U.S.A
SOUTH LONDON BOTANICAL INSTITUTE. 323 Norwood Rd, London,
SOUTHPORT FLOWER SHOW, bee vies pony Lord St, Southport, Lanes.
SPENCER, Mrs. S. B., York Gate, Ade
ST. CLAIR- MORFORD, E., 5 Boulevard Edouard VII, Nice-Cimiez, A.M.,
ran
STRACHAN, P. W., _ Cantab.), 71 The Mall, London,
STUART, Mrs. M. “ tees Seid und Rd, Moulin, Pitlochry, Perthshire
SUMMERTON, JOHN, Ra ES, 81 Leeds Rd, Tadcaster, Yorkshire Aone
SWINSCOW, T. D. B BS MSc, FLS, Everley, London Rd, nee wo
Herts.
TAYLOR, H., 8 Cotterill Rd, Surbiton, Surrey
248 BRITISH FERN GAZETTE: VOLUME 9 PART 6 (1965)
TAYLOR, P. G., Royal Botanic Gardens, Kew, Richmond, Surrey
TEMPLE, MAX, 3 Woodlands Close, London, NW
TEMPLE, PETER LRIBA MInstRA, 42 Ho lly Park, London
TETLEY, Miss M. I., Valley Howe, Cartmel, Grange-over Sands, Lancs.
THOMAS, BA, BSc, fae of Botany, The University, Reading, Berks.
THOMAS, G. S., Briar Cottage, Fairfield Lane, West En d, Woking Surrey
THOMPSON, E., St. David’s, Lydwell Rd, Torquay, Dev
ee door GEORGE E., 593 Buchanan St, Hillside, N. iS, to, U.S.A.
TINNE, » AuM., Lo chwood Cottage, near Ga rtcosh, Glas
TREWREN. < 22 St. Barnabas Rd, Linthorpe, | Middiesiecaas. Yorkshire
TROTTER, R. D., Brin House, Flichity, Inver
baat Dr. ROLLA M., Gray Herbarium, oo Divinity Ave, Cambridge 38,
TULANE’ UNIVERSITY, St. Charles Ave., New Orleans 18, Louisiana, U.S.A.
TU yee ens DA, Dept. of Bi ology, Laurentian University, Sudbury,
ari net
UNIVERSITY COLLEGE OF NORTH WALES, Bangor, Caernarvonshire
UNIVERSITY COLLEGE OF WALES, Aberystwyth, Cardiganshire
VILLARET, PIERRE, DSc, Curator, Musée Botanique Catonal, Palais de
umine, Lausanne, Switzerland
eee H. STEWART, MRCS LRCP, 97 St. Stephen’s Rd, Canterbury,
ent
WAGNER, Prof. WARREN H., uF Dept. of Botany, University of Michigan,
Ann Arbor, Michigan 48104, U.S.A.
WALKER, STANLEY, BSc PhD, Dept. of Genetics, The University, Liverpoo
WALKER, TREVOR G., BSc PhD FLS, Dept. of Botany, The University,
N 8
%& WARBURG, E. F., MA PhD, FLS, South Hayes, Yarnell’s Hill, Oxford
WEBSTER, W. S., ik Station Rd, Steeton, Keighley, Yorkshire
WELCH, Mrs. 'B., Penyllan, Gadshill Rd, Charlton Kings, Cheltenham, Glos.
WILLIAMS, STUART L., 664 Rochester Way, London, SE9
G.
WILSON, Mrs. THEODORA, 91 Edith Ave, Piymouth, Devon
WwW F. W.
OOD, F. i ty Calstock Rd, Woodthorpe, Nottingham
WOOD, P. 9 Slievemoyne Park, Newcastle, Co. Dow
WOOD, Mise “VIVIEN L., Guiseley Lodge, High Ronis "Hospital, Menston,
Ilkley, Yorkshire
WOOLLEY, R., 22 South Broadway St, Burton-on-Trent, Staffs.
WRIGHT, W., 145 Monmouth Rd, London, NI
Please send any corrections to the above list tu the Secretary
We have a very comprehensive
collection of
BRITISH FERNS & THEIR VARIETIES
We also specialise in
Alpine and Herbaceous Plants
Dwarf Trees, Flowering Shrubs
House Plants, Water Lilies
and Greenhouse Plants
CATALOGUE ON. REQUESE
REGINALD KAYE LTD
SILVERDALE, LANCASHIRE
THE HARDY PLANT SOCIETY
provides a link between all wishing
to know and/or grow hardy
herbaceous perennials
Membership can be most rewarding:
subscription 12s 6d per annum
BROCHURE FROM THE HONORARY SECRETARY
MISS BARBARA WHITE
10 ST BARNABAS ROAD
EMMER GREEN
READING, BERKSHIRE
2a
BRITISH FERN GAZETTE
VOL. 9 PART 6 1965
Contents
Variation in Athyrium in the British Isles
R. KAYE 197
British ferns in a world perspective
R. E. HOLTTUM 205
Taxonomic notes on Jamaican ferns
G. R. PROCTOR 213
The influence of shaded conditions on the fertility of Bracken
M. J. DRING 222
Notes and reviews :
Sporeling Bracken on Littleworth Common
J. D. MELVILLE 228
The Southport Flower Show 1964
R. KAYE & N. ROBINSON 236
The ferns in Flora Europaea (review)
T. REICHSTEIN 230
Index Filicum, Supplementum Quartum (review)
W. T. STEARN 234
The Gardener’s Fern Book (review) R. KAYE 204
Coloured Illustrations of the Japanese Pteridophyta (review)
M. E
212
British Pteridological Society :
Reports of meetings... ee ae aoe =e 239
List of members eee 244
E. C. Freeman Ltd., Glennys Acre, London, S.W.11
THE
BRITISH FERN
GAZELLE.
VOLUME NINE PART SEVEN’
1966
Y
ty
Wissocr: BoTanrcw#e THE BRITISH
PTERIDOLOGICAL SOCIETY
AUG 2- 1966
GARDEN LIBRARY
BRITISH PTERIDOLOGICAL SOCTET.
Officers and Committee 1965-66
PRESIDENT
Reginald Kaye
VICE-PRESIDENTS
Thomas H. Bolton
J. Davidson MB ChB FRCP (Edinburgh)
Percy Greenfield
R. E. Holttum MA ScD FLS
T. D. V. Swinscow MB BS MSc FLS
SECRETARY AND TREASURER
J. W. Dyce
46 Sedley Rise
Loughton, Essex
EDITORS
A. C. Jermy BSc FLS and J. A. Crabbe
British Museum (Natural History)
London SW7
COMMITTEE
R. F. Cartwright P. Temple LRIBA MinstRA
143 Pittmansfield 42 Holly Park
Harlow, Essex London N 3
(Joint Meetings Secretaries)
T. A. Dyer FLS, B. Hayhurst
F. J. Healey, Jean R. Healey BSc, F. Jackson
Frances M. Jarrett MA PhD FLS, R. M. Payne
R. S. W. Pollard LAMTPI, N. Robinson
EDITORS
Peter Temple LRIBA MinstRA
The
: : FERNS and —
ALLIES upon an se Egco sin sauce lie TWENTY:
Full details will be sent upon application to the SECRETARY.
BRIT: FERN GAZ. 9 €7)° 1966 249
PSILOTUM NUDUM IN EUROPE
BETTY MOLESWORTH ALLEN
Villa Coca, Los Barrios, Prov. Cadiz, Spain
In January 1965 I found near Algeciras in southern Spain a colony of
Psilotum nudum (L.) Griseb. growing in a rock crevice. This is one of
the primitive fern-allies, belonging to the PSILOTOPSIDA and has
not hitherto been recorded for Europe. This class comprises two
genera: Tmesipteris confined to the southern hemisphere and Psilotum
which occurs in most tropical countries reaching Japan in the north
and southern Africa, Australia and New Zealand in the south; the
closest locality to Europe seeming to be the Cape Verde Islands.
he area where I found the Psilotum lies approximately 20 kilo-
metres from Algeciras at about 170m above sea-level where large out-
crops, probably sandstone, occur amongst the cork woods. The first
plants found were growing in an almost horizontal deep and very
narrow fissure, about one metre up the vertical face of a 30-metre
outcrop facing south. There was one clump, then a space of 17 cm,
then some small plants were spaced for a distance of 12 cm, and some
dead fronds extended for a further 11 cm. No other plants were mixed
with the Psilotum, but in more sheltered crevices on the almost bare
rock were a Ruscus (probably R. hypophyllum) and a small plant of
Asplenium adiantum-nigrum; about one metre above was a large
prostrate plant of Calluna vulgaris from whose humus-packed base
grew a Davallia canariensis. Above these on a small shelf were quite tall
ground beneath.
Later I found more Psilotum growing
vertical rock face, but about 2m above a she (
and quite separate patches, all round about the same level and all in
narrow fissures which with two exceptions were nearly horizontal.
The plants were in full sun, at least in summer, but usually a slight pro-
jection of the rock gave a little shelter from the west. The largest patch
t and thick and often
bunched together. About 20 cm away, but in the same fissure, was a
20m higher up the same
If. There were five distinct
lichens, a liverwort, and a small Se
no doubt other plants would also be there du
ES
THE BRITISH FERN GAZETTE Vol. 9 Part 6 was published 14 July 1965
aa0 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
shelf immediately below had sparse vegetation on a thin layer of soil:
one plant of Erica scoparia, one of E. australis, a large patch of the
stemless palm Chamaerops humilis, and a small cork oak which gave a
little shade to some of the Psilotum. The Sedum was common here.
The heat of this rock during the summer months is intense, for the
sun shines on it all day, but the deep narrow fissures in which the Psilo-
tum grows would probably remain rather cool. The stream apparently
dries up at this point in August/September.
Perhaps worth noting is that a few metres away to the west of the
face is a large and similar cliff, also with horizontal fissures; but it faces
due east and there was no Psilotum on it, and with one exception its
vegetation consisted of plants not noted on the former rock; these
included Cistus ladaniferus, Arbutus unedo, a Ulex and a Cytisus; the
only plant common to both places was Erica scoparia. :
Approximately 400m downstream of the Psilotum-bearing rock
and beyond a definite curve in the stream were three more patches of
Psilotum. These were near the base of a vertical face of a very large
rock outcrop, similar to the former except that the altitude was about
130m and the face received more permanent shade from tall Quercus
spp. which were closer to the rock face and in the winter would probably
be very damp. Two patches stretched for about 2m along a horizontal
crevice; the other was a smaller and denser tuft. oa tee
hilst it must not be overlooked that Psilotum escapes easily from
cultivation, this Spanish locality seems sufficiently remote and the
habitat relatively undisturbed to consider this species native. It may
therefore be found in similar areas here in the south, and it is still pos-
sible that further specimens from Spain may turn up in herbaria.
I am most grateful to the Director and Dr Frances Jarrett, Royal
Botanic Gardens, Kew, for assistance. Herbarium specimens have been
deposited at Kew and the British Museum (Natural History), London:
B. E. G. Molesworth Allen 5082 (K), 5088 (BM).
REVIEW
NATURE IN NEW ZEALAND series : NATIVE FERNS by Bruce
resi 64 pages, 13 x 19 cm; Wellington: Reed, 1963; price 8 shil-
ings.
The most striking feature of this little book is the 25 pages of line
drawings illustrating the 107 species of ferns described: the thumb-
nail sketches of the tree ferns are particularly fine and expressive.
Phe text gives the local name, Latin name (with translation) and Fi
i Ze
British Museum (Natural History), London SW7 ree Be JERMY
UaI[V “H O “A :o104g * } UJOYINOS ‘SBIIDIB]/Y IBVIU DOIADID YOOI V UL WNpnu wNnjOpISg
\o
Oo
an
(7)
9
ue is oh
Bee:
PERN: S23 AZ:
BRIT.
BRIT. FERN GAZ. 9 (7) 1966 251
FERNS IN A WARDIAN CASE
GEOFFREY GILBERT
116 Haverstock Hill, London NW3
About two years ago I decided to build a wardian case, chiefly for
the purpose of growing ferns. Such a case not only allows for the
growth of many specimens in a small area, but also permits a high degree
of control, so that exact observations can be made. The idea was not
new to me, for I could remember being inspired and delighted by a
wardian case owned by my grandmother, when I was a child. This case
was built outside the window, so that when you opened the sash you
looked into a moist, ferny world that contained a miniature pool and
an orchid that actually flowered, albeit somewhat erratically. The
pleasure this case gave to its owner in later years illustrated the value of
an idea developed by Dr Ward more than a century ago, but now largely
neglected.
The actual construction of my case, which stands in a sheltered
corner on the north side of my house, may be judged from FiGure 1.
It is six feet long and three deep, at present providing for 82 specimens
with space for more. The base is formed of corrugated asbestos sheet,
covered with broken brick and a layer of peat five inches thick. It
stands at table level and opens at both front and top to allow convenient
access. Some form of small lattice window, though not yet installed,
is urgently needed to allow access in cold weather without unreasonable
chilling.
HUMIDITY
The plan of the case was based on a series of observations, rather
than precedent. Important among these was the fact, familiar to every-
one, that if a patch of damp, shady ground is covered with a bell jar,
an abundance of ferns, liverworts and mosses, quickly springs up. I
associated this fertility with the moisture content of the alr, a hair
hygrometer showing that this was seldom far short of 100%. As will
be seen later, this was not the whole story, but fortunately my omis-
sion did not involve alteration of the structure.
HEATING
To obtain high humidity without heating is usually a simple matter
of stopping up cracks and providing a large evaporating surface, for
which the peat was likely to suffice. I wished however, to use higher
temperatures. As is well known, when a hot body is placed in an en-
Closed space, the humidity is reduced out of reasonable proportion to
the increase in temperature. In my case an electric light bulb will often
reduce humidity 20° without noticeable effect on the temperature.
B
252 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
In a conventional greenhouse this drawback is overcome by laborious
spraying whenever the pipes have to be particularly hot. Such a course
is out of the question in a case sited out of doors, where this would
involve deluging the plants with cold air as well as water.
To avoid this difficulty, I decided to use a warm pool as a source of
heat. Water heating on this scale can conveniently be provided by a
standard 200w aquarium heater with thermostat. I use a three-gallon
tank kept at 90° in winter and this gives a temperature of about 40° in
cold weather. A paraffin heater can be used under the peat bed, to
help out in very severe conditions. The tank steams freely into the air,
too freely without control, which is easily provided by means of metal
shutters to cover part of the surface. Only a small area is normally
exposed and the metal shutters provide a convenient site for raising
spores, which do best at a temperature of 75° or thereabouts. The tank
is surrounded by a miniature rock garden suitable for growing mosses,
lycopodiums, etc.
It might be supposed that such a high humidity level, usually 100 Yo»
would prevent evaporation and harm the plants, but in practice alr
moves continually, apparently picking up moisture in one place an
depositing it in another, so that evaporation never really ceases. Indeed
the 100% figure may be no more than a sort of microclimate in the
instrument itself, while considerable variations exist in the air currents.
One advantage of humidity relates to root aeration. Some ferns
such as the parsley fern, are very sensitive in this respect and only grow
satisfactorily when their roots are spread out among stones and moist
air circulates freely. Actually air in relation to root growth is mentioned
in every book, but in my experience, more often mentioned than ps
into practice, chiefly for the obvious reason that air dries out the soil
with irritating rapidity. Since this drawback does not appear In 4
humid fern case, composts can be very open, with a large amount of
broken brick, pot, etc. if desired. The pots should be raised, so that air
can flow freely through the hole in the bottom. The parsley fern and
other plants with the same requirement will then behave perfectly well.
THE CO, COMPLICATION
When the case was beginning to grow in a satisfactory way; I
repeated the bell-jar experiment, first clearing an area of peat, that I
meni
FIGURE 1 Geoffrey Gilbert’s wardian case
A) active compost to supply additional carbon dioxide, B) soil or peat for
ferns, C) cover sheet to control moisture, D) aquarium heater with thermostat,
E) perforated zinc sheet covered with gravel, F) corrugated asbestos sheet.
GILBERT»: FERNS: IN ‘Ae WARDIAN CASE
STORAGE SPACE
4
SO INERT RRNA) meramee
FIGURE |
254 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
judged well supplied with spores. Part of the surface was covered and
part kept as a control area. I confidently expected that both would now
yield the same result, but I was completely wrong. The area under the
jar was choked with young growth before the area outside even began
to grow.
After some experiments I obtained one of the indicator solutions
used when it is intended to increase the CO, content in commercial
greenhouses. Since soil is credited with producing as much as an ounce
of carbon dioxide per square yard per day, it was not surprising to find
a high concentration under the jar, probably sufficient to be toxic to
some plants. In deficit of other explanation it seems reasonable to
associate this concentration with the rapid development of spores of
mosses, liverworts and ferns and the attractive liveliness of young
growth. It may perhaps be pointed out that the concentration under a
jar soon diminishes as the plant increases in size and eventually be-
comes sub-optimal. In my experience, plants then weaken, lose colour
and are usually attacked by fungi, a point to watch in bottle gardens.
The CO, concentration in my case, though showing more diurnal
variation, was approximately equal to that outside and it seemed likely
that a greater concentration would be an improvement. An easy way
of producing clean CO, is by means of rotting vegetable matter So,
clearing a space under a shelf, I added a large pile of peat soaked in
suitable fertilizer. There was little room for doubt about the effect of
this measure, mosses and young ferns springing up in abundance and a
general liveliness to be noted in the vegetation. This improvement
however, still fell short of the effect produced by the bell-jar and it would
be interesting to experiment with an artificial source of carbon dioxide.
It would be useful to know if anyone has knowledge of the effect
of carbon dioxide on spores and prothalli, as it is these that appear to
be so noticeably affected. Traditionally they do best on soil that has
been lightly scalded, not sterilised, and covered with a glass. My first
sowing on elaborately sterilised soil, protected in every way, took seven
months to germinate. Can this have been connected with the lack of any
living things to produce CO,? The small amount of air under the glass
at the top of a pot is certainly heavily impregnated.
THE LIGHT FACTOR
Another point that must be thought about carefully before siting
the case is the light factor, as it is obviously annoying, especially from
a decorative point of view, to have to paint the glass and worse to get
inadequate growth. To make certain on this point, I established a
number of ferns under glass covers and placed them in various situa-
tions. Measurements of frond lengths were used as a rough indication
of growth and an ordinary exposure meter used to measure light values.
GILBERT: FERNS IN A WARDIAN CASE 25:5
This involved some trouble, perhaps out of proportion to the immediate
need, but of sufficient general interest to appear worthwhile.
Summarising the results briefly, I found that no fern seems to benefit
by an open situation, usually the reverse, and that they vary greatly in
relation to their minimum requirements. Some will grow well, develop-
ing large handsome fronds when the light is no stronger than 1 foot-
candle, as against 400 in sunlight. Others, particularly the large ferns
such as Dryopteris borreri, develop very slowly below 25 foot-candles.
By far the most adaptable fern I tried was Asplenium trichomanes.
Temperature and the CO, factor obviously confuse these results
but they may be taken as sufficient for the purpose. The conclusion
seems to be that a noon reading of twenty-five foot-candles is the best
compromise. This condition prevails in a well lit room near to the
window and I was able to arrange the same light value in a convenient
corner on the north side of the house.
APPEARANCE OF THE CASE
The charm of a wardian case lies in the richness of the vegetation
and the perfection of the green shades that respond so well to moist
conditions. Here ferns may reveal a beauty not seen in nature. Mosses
add greatly to this effect and can advantageously cover every stone or
area of soil.
On the practical side it should perhaps be mentioned that mosses, in
colonising stones, are usually handicapped by lack of nitrogen. For
instance a brick soaked in ammonium sulphate will cover with moss
in a matter of weeks, where years may be required without fertiliser.
The traditional and safer method fertilising is to paint with whey
from sour milk. Choise mosses such as Thuidium or Leucobryum should
be watered with rain water and of course must be introduced.
Flowering plants may be introduced for variety. Orchids and
epiphyllums are very suitable and many small woodland plants such as
Soldanella, Pyrola, Shortia, Trientalis are worth trying, remembering
that those coming from pine-woods may need an occasional dressing
of needles to maintain the acidity. Of course fern prothalli spring up
everywhere and one always hopes to find interesting hybrids. |
The introduction of mosses and other irrelevant specimens to
build up an effect may strike the more serious pteridologist as just time-
wasting beautification. I can point out however, that there is much of
scientific interest in a small area of mixed vegetation. In such a con-
venient place you are likely to look far more closely and observe details
that might escape you in a woodland area. You never quite know what
you will find when you open your case in the morning. It is interesting
even to find that something has died, though I myself, lacking true
scientific detachment, prefer the reverse to happen.
256 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
THE IDENTIFICATION OF BRITISH
HYMENOPHYLLUM SPECIES
GARETH B. EVANS
University of Malaya, Kuala Lumpur.
The confusion which has surrounded the identity of Hymenophyllum
tunbrigense (L.) Smith and H. wilsonii Hook. since the time of John Ray
has already been reported (EVANS & JERMY 1962). An account of the
biology of the two species will shortly be published in the Biological
Flora of the British Isles series in the Journal of Ecology. This present
paper describes those features of their morphology and anatomy which
can be used to distinguish them both in the field and herbarium.
The sporophytes of both species form moss-like carpets with
numerous overlapping and often pendulous fronds borne on the densely
interwoven thread-like rhizomes. The fronds are superficially alike,
as can be seen from the silhouettes in FiGures 1-13. The stipe 1s
thin and glabrous, about half the length of the rhachis, and the frond
lamina whilst very variable in size is nearly always longer than broad. It
bears numerous short pinnae around | cm long arranged alternately on
the rhachis. The pinnae are divided dichotomously into several seg-
ments, each segment 3-5 mm long and | mm wide. The sori are found
terminally on the segment nearest the rhachis, but occasionally two to
several may occur on one pinna.
In both species the translucent lamina is only one cell thick except
immediately around the nerve, yet the species differ in colour and tex-
ture, especially in older mature fronds. H. wilsonii is a darker olive
green with a reflecting surface whilst H. tunbrigense has a lighter blue
green hue with a matt surface. Moreover, the lamina in H. wilsonii
has a less membranous texture and also its stipe and rhachis are stiffer
than in H. tunbrigense. Stunted fronds in exposed colonies of H. wilsonil
are held almost vertically whereas those of H. tunbrigense usually have
a sharp bend in the stipe so that the fronds lie nearly horizontal to the
rhizome mat. They differ as well in the angle of attachment of the
pinnae to the rhachis; each pinna axis in H. tunbrigense grows approxl-
mately at right angles to the rhachis and the pinnae segments point
forward and are so held that the overall frond surface is flat or slightly
convex. In H. wilsonii, on the other hand, the axes of the pinnae depart
nearly at right angles from the rhachis but are deflexed and as the seg-
ments are not aligned along the plane of the lamina the frond appeats
in profile to be a one-sided series of tiers. It was for this reason that the
species was formerly called H. unilaterale.
In addition there are fewer segments per pinna in H. wilsonii (rarely
exceeding 3-5) and they are displayed almost palmately because the
FVANS: BRITISH HYMENOPHYLLUM SPECIES fn
points of dichotomy lie close together. In H. tunbrigense there are more
segments per pinna, usually between 5-11, and they lie predominantly
on the acroscopic side, i.e. that side of the pinna nearest the apex. The
pinna segments have nearly parallel sides in H. tunbrigense and the
lamina is continuous from one segment to the next. In H. wilsonii the
segments are ovate-lanceolate and taper to their base, remaining scarcely
connected to each other by any lamina. Successive pinnae in H. wilsonii
are almost the same size and length, but in H. tunbrigense they rapidly
shorten so that the frond shape is broadly ovate-lanceolate contrasting
with narrowly lanceolate in H. wilsonii.
The combination of these features of shape, orientation, colour
and texture enables one with a little experience to distinguish by eye
most colonies at distances up to twenty feet! In well pressed herbarium
Specimens most of these features are lost, but the relative number and
Shape of the segments can still be used to separate them on most
occasions, although other criteria must be used for confirmation.
When sori are present the species can be very easily distinguished.
In both, the sorus consists of two green indusial flaps (or valves) which
freely articulate at the base enclosing a cluster of sporangia arranged
compactly on a central column. 4H. wilsonii generally bears more,
around 10-15, sori per frond than H. tunbrigense which has around
5-10. They are pear-shaped and swollen in H. wilsonii resembling a
duck’s bill and invariably project perpendicularly to the plane of the
lamina. In H. tunbrigense they are sub-orbicular and only slightly
swollen, lying almost in the plane of the lamina. The indusial valves
gape widely at maturity in H. wilsonii but this is less conspicuous in H.
tunbrigense.
However, the most reliable and convenient criterion of all for
identifying fertile material concerns the indusial valves. In H. wilsonii
the edges are quite entire, whereas in H. tunbrigense the distal edges
are raggedly toothed. This character was first noticed by William
Wilson and taken up by w. J. HOOKER (1830) in the first validly pub-
lished description. During my examination of hundreds of specimens
I found no reason to doubt the reliability of this character.
The anatomy of the fronds provides additional distinctions. One
that is occasionally quoted, e.g. HYDE and WADE 1962, refers to the
distal point of the pinna segment; the vein is said to end before the tip
in H. wilsonii, i.e. c.4 cells behind, but continue to the tip, 1.¢. c.2 cells
behind, in H. tunbrigense. Unfortunately, I have observed that this
distance is dependent on the growth phase of the segment, the vein
appearing nearer the tip in young vigorously expanding segments of
H. wilsonii and in my experience, whilst it holds true in many cases, it Is
not a reliable character.
te
58 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
FIGURES 1-7 H ymenophyllum Ln emeenre, W =wilsonii) from average con-
1) desiccation- overtop Spey ee of : teas m exposed rain 2) T bem crevice
within boulder scree; 3) Ta iss
EVANS: BRITISH HYMENOPHYLLUM SPECIES 259
@ 2
FIGURES 8-13 Hymenophyllum (T= tunbrigense, W=wilsonii) from very lush
conditio
8) laboratory culture of T with low light and above 90% humidity; 9) frond of T from
Killarney; 10) three-year-old frond of T cultured in high light, humidity and tem-
perature, cf. Fig. 13; 11) W as 8; 12) W as 9; 13) exceptional specimen of W from
Fermanagh (coll. Praeger 1904, hb.Nat.Mus.Dublin). ALL NATURAL SIZE
EXCEPT FIG. 13 HALF SIZE.
c
260 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
tunbrigense
5Ou.
FIGURE 14 Frond cells of Hymenophyllum
Differences in the surface appearance of the lamina are also ap-
parent; in the words of W. J. Hooker (Brit.FI.: 450. 1830) “[wilsont
is] . . . more loosely cellular and coarsely reticulate than the last [tun-
brigense] .. .”
In fact, the cells in H. wilsonii are far larger, rectangular and con-
spicuously arranged in rows running obliquely to the segmental vein.
In H. tunbrigense, they are smaller, more square, and appear to be
arranged haphazardly. See FiGure 14. To be more precise, the mature
cells of H. wilsonii are 0-094-+-0-002* mm long, and those of H. tunbrig-
ense 0-060-+0-01 mm. The width of cells in surface view in both species
is the same at 0-039+-0-001 mm. GULLIVER (1863a, b) made an observa-
tion on the sizes, but not the arrangement, of these leaf cells when he
also remarked on the inadequacy of J. C. Sowerby’s figures in English
Botany (HOOKER 1831). The gametophyte cells are likewise very dif-
aes in size, having the same dimensions as the lamina cells described
above.
* These dimensions are quoted along with the standard error of the mean.
EVANS: BRITISH HYMENOPHYLLUM SPECIES 261
It is interesting to note that the spores of each species also differ
in size, and JESSEN & ANDERSEN (1959, p. 31) quote a range of 62-74
diameter in H. wilsonii and 40-48p diameter in H. tunbrigense.
One further convenient distinction applicable to both sporophytes
and gametophytes lies in the number and size of the chloroplasts. H.
wilsonii has very numerous small chloroplasts so that about 60-80 of
them line the periphery of the cells as seen in surface view and this is
easily separated at a glance from H. tunbrigense where only 30-40 can
be counted. This criterion has proved to be especially valuable in
identifying sterile fragments of old herbarium material in which, afte
soaking in water, chloroplasts become visible again.
The gametophytes of each species are very similar in form, being
strap-shaped with many short rhizoids at least in culture, for they have
never yet been detected free-living in nature apart from a few contained
within the indusial valves of old sori. Shortly after germination they
both show marked tri-radial symmetry, and this, according to STOKEY
(1940) is a constant feature of the genus Hymenophyllum and commonly
found in Trichomanes as well.
As both species are highly plastic in their sporophyte morphology,
depending on the environment, a brief summary will be given here of the
ranges encountered. (For a fuller account, see EVANS 1964).
The fronds of H. wilsonii are usually from 3 to 6 cm long and about
2 cm wide, but exceptionally this range can extend from 0-5 cm to 30
cm and to 15 cm in H. tunbrigense (see Fics. 1-13). The smallest
fronds are found in colonies on rocky surfaces liable to periodic droughts
in very exposed positions in the mountains. In rather dark sites, where
very high humidities are maintained constantly even during prolonged
droughts, apical growth of the fronds persists from season to season in
both species. In this way long etiolated fronds are produced often with
rather distant pinnae. Not infrequently under these conditions the
fronds are found with sori in spaced groups, each group corresponding
to a season’s crop; but note that there may be two growth seasons in
exceptionally favourable years. On a few occasions I have found five
such groups on a single frond. H. wilsonii produces longer fronds in
lighter sites under constantly high humidity.
However, whilst the fronds may be potentially of unlimited growth,
the great majority in practice must last for not more than two years,
because periodic desiccation afflicts most colonies and kills the growing
apex if not the frond itself. For instance, after four years not one of the
many fronds that I had marked in North Wales in 1960 had survived.
In certain circumstances, such as in very humid places where diffuse
light is abundant but where direct sunlight never falls on the fronds,
H. tunbrigense can produce fronds of such unusual morphology that
they cannot be separated easily from those of H. wilsonii without
262 BRITISH PERN GAZETTE: VOLUME O°-PART 7-619060)
recourse to anatomical criteria (see Fics. 10 & 13). This problem arises
not infrequently with herbarium material as collectors often prize
unusual specimens. A very rare form of tunbrigense that I have only
once succeeded in developing experimentally (in low light and above
90% humidity) possessed exceedingly numerous squarrose pinna seg-
ments, and each pinna seemed of indefinite growth so that the frond
resembled a small bush. (see Fic. 8); similar fronds occur rarely in the
field.
In conclusion I would add that there are important physiological
differences between the two species, particularly in their reactions to
low light intensities and to desiccation. H. wilsonii is sensitive to low
light but very resistant to desiccation whereas tunbrigense is the oppo-
site, being very tolerant to low light and very sensitive to desiccation.
As a consequence the two species occupy relatively distinct micro-
habitats. Only very rarely does one come across a patch containing
intermingled fronds of both species. When the two are grown together
in low light then H. tunbrigense wins the battle aided by its smothering
growth form, but in conditions subject to periodic and severe drought
tunbrigense succumbs and wilsonii is the victor.
I am indebted to the Director of the National Museum of Ireland
for permission to reproduce Fig. 13; to the Editors for arranging the
figures; and to Prof. P. W. Richards “he supervised my research. Also,
I wish to acknowledge assistance from Abdul Samat b. Abdullah in
typing and from Ja’affar b. Haji Abdullah in preparation of silhouettes.
REFERENCES
EVANS, G. = 1964, Autecology of British species of Hymenophyllum. Ph.D. Disser-
tation Univ. Co il. N. Wales, Bangor (unpublished).
EVANS, G. B. & JERMY, A. C. 1962, Notes on the history of British fern species 1:
onfusion in the filmy ferns. Brit. Fern Gaz. 9: 81-
GULLIVER, = 1 oe , On the Teaficelis of ee British species of Hymenophyllum.
Hist. ser. Il 12:
—— 1863b, ‘Comparison c of the leaf-cells in the British Hymenophylleae. J. Bot.,
Lond.
HOOKER, W. J. 1830, The British Flora. London
1831 , Supplement to the English Botany of t the late Sir J. E. Smith and Mr
Sow erb (Tab. 2686 H. wilsonii—text by W. Wilson).
HYDE, H. A. & WADE, A. E. 1940, Welsh iets Cardiff.
JESSEN, = & ANDERSEN, S. T. (with A. FARRINGTON) 1959, The interglacial om
ear Gort, = Gabae: Ireland. Proc. R. Irish Acad. sect.
STOKEY, ra G. 1940, Spore germination and vegetative stages “Of the ei neophyte
of Hiyrsenppie tans and spread ty Bot. Gaz. 101: 759-790.
BRIT. FERN GAZ. 9 (7) 1966 263
DRYOPTERIS VILLARII IN BRITAIN
OLIVER L. GILBERT
Botany Department, The University, Newcastle-upon-Tyne
Dryopteris villarii (Bell.) Woynar has rather precise habitat require-
ments which restrict it to a small part of north west England with an
outlying station at Llangollen in North Wales. It is abundant in only
three localities: the hills of Ingleborough, Whitbarrow Scar and the
Hutton Roof—Farleton Knott massive which all lie within 25 miles
(40 km) of each other in the area where Yorkshire, Lancashire and
Westmorland meet. Here the populations can be measured in hundreds
or in the case of the Lancashire one in thousands of individuals. Many
other hills in this general area carry smaller populations of the fern,
for example it can be found on Scout Scar, Orton Scar, Great Asby
Scar, around Malham Tarn on Wild Boar Fell, etc. It has recently
been seen in twenty-one 10-km grid squares (see map on page 265).
This part of the country is the main area of outcrop of the Great
Scar Limestone, a bed of very pure Carboniferous limestone over four
hundred feet (130m) thick which produces typical karst scenery with
much outcrop. The distribution and abundance of the fern can be
correlated with its exposures to a remarkable degree. The mid and
south Craven fault scarps which mark the junction of the Great Scar
Limestone and the impure basin limestones of the Craven Lowlands
support the most southerly colonies in England, while the most easterly
ones are on the limestones overlooking Upper Wharfedale before they
vanish under the Yoredale and millstone grit cover of the Pennine dip
slopes. To the north, as limestones gradually become subordinate in
the succession, the fern decreases and the last major appearance of the
massive beds at Great Asby and around Brough correspond with its
northern limit.
Great Scar Limestone country provides three types of habitat
Suitable for D. villarii: cliffs, screes and limestone pavement. If all
localities known to the author containing twenty or more plants are
scored, the following picture of habitat preference emerges: pavements
(11 sites) 64%; scree (4) 24°; cliffs (2) 12%. As all these habitats are
Widespread these figures probably give a true picture. A comparison
of them, together with other limestone habitats available to but un-
Occupied by the fern should give some indication of the reasons for its
limited distribution.
Soils from these habitats are reasonably uniform being rendzinas
grading into mull rendzinas where there is drift contamination. — In
appearance they are very dark with a well developed fine to medium
264 BRITISH FERN GAZETIE: VOLUME 9 PART 7 (1966)
crumb structure. Mineral grains are intimately mixed with the black
well decomposed humus and there is excellent incorporation by the
abundant soil microfauna. At four to five inches (10-13 cm) this
typically grades into a very stony, highly calcareous, humus silt loam
which is usually moist. Analysis of several samples from around the
roots of D. villarii have shown that it is a strongly alkaline soil with a
high proportion of calcium carbonate and organic matter. The base
exchange capacity is high and the carbon/nitrogen ratio of around 10
shows that it is very fertile (TABLE 1).
TABLE 1—Soil analysis from three localities of D. villarii on Ingle-
orough, Yorkshire.
Sample Organic Calcium Carbon Nitrogen Carbon/
No.
matter carbonate nitrogen
(per cent) (per cent) (per cent) (per cent) ratio
1 15-2 26:8 7-6 0-70 10-8
2 22°6 29-4 lid 1-03 11-0
3 29-5 55:4 14-8 1-25 10-9
Sample pH of | Exchangeable Exchangeable Base
No. fresh calcium* potassium exchange
soil capacity
(in mille-equivalents per 100g of soil)
1 7-9 20-3 0-25 21-9
2 7:8 — 0-26 38-3
3 78 0-49 38-1
* only one observation made.
Sample No.1 Mull rendzina. South facing scree slope.
No. 2 As for No. 1 but north facing block scree.
No. 3 As for No. 1 but limestone pavement.
In Britain this precise soil type is almost confined to mountain
limestone areas, but similar soils can be found on other pure lime-
stones such as the Chalk, Magnesium and Jurassic, indeed Thelypteris
robertiana a fern closely associated with D. villarii grows on them all.
o for this and other reasons such as its behaviour on garden soils
and its occurrence near Brough on a mixed limestone and sandstone
scree, it seems unlikely that soil factors do more than restrict it to well-
drained base-rich soils.
The altitudinal range of the fern varies from sea level in Silverdale
where the vegetation contains a strongly southern element (including
Adiantum capillus-veneris) to 1600 feet (490m) on top of the Pennines
where the highest D, limestones occur. Here snow is lying on an
6 .
IVBER TT: DRYOPTERIS. VILLARIP IN SRBRATAIN 265
— forty-five mornings in the year and ground frosts are
: a every month (MANLEY 1957). When other limestone areas
aaa : oo compared for rain days, Meyer’s precipitation/saturation
quotient, mean temperatures and sunshine, the sites which fail
match a part of the Silverdale—Craven Highlands belt are those
istribution in the British Isles of —, —_ Each dot represents at
: ; nal Grid. The distribution of
oo limestone is shown by toi ai gone much o
ured by glacial drift or intercalated with non-calcareous strata
266 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
lying south of the Exe-Tees line. This leaves Derbyshire as a large
limestone area where one might reasonably expect to find the fern.
It does not follow that climatic factors are operating to exclude the
plant from calcareous habitats south of the line, it just means they could
be. An examination of its distribution outside Britain is of no help in
determining its climatic limits as there is evidence that our plant 1s an
endemic cytotype.
Topography seems to be the most important factor shaping the
distribution of D. villarii. The plants are sensitive to wind exposure and
grazing, so only sites offering sufficient protection are available to it
and these are widespread only on the harder Palaeozoic limestones.
A fewer simple experiments have verified this. In 1960 a colony on
block scree near Malham Tarn was investigated. It had been noticed
that by the end of summer most plants had been grazed, so stones were
removed from around several clumps and replaced in such a way that
the fronds were exposed to increased grazing but not to increased wind.
Sheep graze sporadically on this barren scree slope and in 1961 they
damaged fronds in all the experimental areas. Damage varied from
being eaten to ground level to fronds with just the lowest pair of pinnae
left and others still intact. The experiment was recorded yearly till
1965. It showed that increased grazing gradually weakened the plants,
leaves annually getting fewer and smaller till after five years of existing
for most of the time below the compensation point, reserve food in
rhizome and leaf-bases was used up and many were dead while most of
the rest were producing only a few stunted leaves.
Exposure to wind was also investigated after observing that on
Whitbarrow Scar where fencing against deer has recently eliminated
grazing, the tips of fronds were shrivelled and brown by mid-July.
The experiment, also carried out at Malham Tarn, involved moving
well grown ferns from the scree slope to a nearby ungrazed limestone
pavement where they were planted out into grykes (fissures) of different
depths. Results indicated that at least six to eight inches (10-15 cm) of
sheltered air above the rhizome is needed if plants are to survive. Those
in situations where this requirement was met were still growing healthily
and producing spores five years later, while those in more exposed sites
were either dead or growing with very reduced vigour. These produced
typical crops of leaves for the first year or two but by August they were
always brown and withered. A second crown of small (2-3 inch, ae
cm) rather leathery leaves was sometimes produced and lasted until
autumn. After two or three years all those in exposed sites were pro-
ducing these as the first crop. The plants must then be photosynthesis-
ing at a net loss throughout the year and be incapable of maintaining
the species on these sites.
GILBERT: DRYOPTERIS VILLARII IN BRITAIN 267
The absence of D. villarii from the fragments of ash (Fraxmus
excelsior) and other deciduous woodland which have survived on these
northern limestones serves to illustrate its intolerance of all but the
lightest shade. Where it does occur on partly wooded pavement it is
always in glades which receive a lot of sunlight.
So D. villarii has as its basic requirements a well drained calcareous
soil in an open or only lightly shaded site which offers protection from
grazing and strong winds. This is an unusual combination for a fern
with fronds which are typically about eighteen inches (46 cms) long,
and must be largely responsible for its restricted distribution. The only
habitats where they are fulfilled are (i) on block limestone scree with a
good proportion of the blocks over two feet (60 cms) in diameter—
strikingly demonstrated on the Eglwysen Mountain cliff complex at
Llangollen where the colonies are located on the only streaks of coarse
scree; (ii) limestone pavement with fairly wide, broken sided grykes
which allow reasonable light penetration and good opportunities for
rooting; (iii) —and most restricted— low sheltered cliffs with a north
to south-east aspect. :
These habitats support the same very uniform plant community.
It is extremely fern-rich and associates vary from shelter-demanding
woodland plants, to grassland and tall herb species. Bryophytes are
well represented and many are calcicolous species. The following
species were over 80°% constant in ten widely separated colonies which
covered at least two examples of each habitat type: FERNS: Asplenium
ruta-muraria, A. trichomanes, A. viride, Cystopteris fragilis, Dryopteris
villarii, Thelypteris robertiana (70%), Phyllitis scolopendrium; ANGIO-
SPERMS: Festuca ovina, F. rubra, Geranium robertianum, Mycelis
muralis, Oxalis acetosella, Sesleria varia, Taraxacum officinale (agg.),
Viola riviniana; BRYOPHYTES: Camptothecium sericeum, Cono-
cephalum conicum, Ctenidium molluscum, Encalypta streptocarpa,
Fissidens sp., Grimmia apocarpa, Mnium undulatum, Neckera crispa,
Tortella tortuosa. :
No worthwhile discussion of the history of the species as a mem-
ber of the British flora can be undertaken without considering reports
of it from three mountainous areas: Snowdonia, Arran, and the Lake
District. The Snowdonia records made by J. E. Griffith from Cwm
Idwal (1884) and Cwm Glas (71887) are quite reliable, the former being
backed up by correctly named herbarium specimens. These two cwms
contain Ordovician rocks which at certain levels are strongly basic
(WILLIAMS 1927) and it was doubtless on these strata that the fern was
found. It appears that Griffith is the only person to have seen D. villarii
in this area, but local botanists are confident it will be rediscovered.
The only information available from Arran is a note in the Proc. Nat.
Hist. Soc. Glasgow, Vol. 4, pt. 2. “Mr Stewart exhibited Lastrea rigida
from Arran, its first discovery in Scotland, 1879.” The presence of
o
268 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
thick beds of very pure carboniferous limestone on the island mee
the record must be taken seriously. The Lake District records were a
made this century from Honister Crag. Small plants have been a
on three or four occasions on quarry debris and the suggestion is —
spores must come from a larger colony on some inaccessible part : an
crag. The rock is not unsuitable, being Borrowdale Volcanic ve
with local heavy calcite veining and many of the usual associates 0 : .
villarii occur on the cliff. These records are important because they
give the fern a much wider distribution than could be imagined from
(PROCTOR 1958). These occur in reasonably close proximity to pepe
of the present-day localities. It is possible that some of the an
limestone pavements never had a complete tree cover, and the prese es
of Polystichum lonchitis growing with D. villarii on three such gfe
ments in Craven and an old record from Farleton Knott would se
to bear this out. cele
The history of the fern in Britain is of some interest as we app on
ly have an endemic tetraploid race (n=82; Farleton Knott, Whit ee
Scar, Ingleborough, Brough, Leck Fell), while the few counts W fe
have been made on the Continent show it to be diploid (n=41; France,
Switzerland). ae
PANIGRAHI (1965) who has looked at the cytogenetics of speci a
from Ingleborough considers the British plant to be an allopolyp :
to which the continental diploid is ancestral. If this is so, our plant,
sexual fern of hybrid origin, probably deserves specific rank, myo
Should continental material prove to be exclusively diploid ther
would be a case for considering part of the D. villarii complex having
survived at least the last glaciation of Britain. Further specu
is unwise until more is known about the continental populations an
about the Ice Age in Britain (DUNHAM & JOHNSON 1963).
REFERENCES
DUNHAM, K. C. & JOHNSON, G. A. L. 1963, The geology of Moor House, a Naver
Nature Reserve in northeast Westmorland. Monographs of the Na
onservancy 2, H.M.S.O. London. : 5-56:
MANLEY, G. 1957, The climate at Malham Tarn. Rep. Field Studies Council 195 ,
PANIGRAHI, G. 1965, Preliminary studies in the cytotaxonomy of the Dryopteris
villarii (Bell.) Woynar complex in Europe. Amer. Fern J. 55: 1-8. he
PROCTOR, M. C. F. 1958, Ecological and historical factors in the distribution of t
British Helianthemum species. J. Ecol. 46: 371. 1
WILLIAMS, H. 1927, The geology of Snowdon. Quart. J. Geol. Soc. Lond. 83: 346-431.
BRIT. FERN GAZ. 9 (7) 1966 269
ORIGIN OF THE FERN FLORA OF
TRISTAN DA CUNHA*
ALICE F. TRYON
Gray Herbarium, Harvard University, Cambridge, Mass.
The islands of the Tristan group are remarkable for their isolation in
the south Atlantic about 1800 miles west of South Africa and 2000
miles east of South America. They are also remarkable in that nearly
half of the indigenous vascular plants are pteridophytes. I will present
here the distribution of these and of some of the flowering plants and
animals and from this evidence draw conclusions on a source of origin
and a manner of dispersal of the Tristan biota.
GENERAL DESCRIPTION AND HISTORY
There are three main islands in the Tristan group—Tristan,
Inaccessible and Nightingale and I also include Gough Island, some
200 miles south-east, which has a similar flora and fauna that has
recently been surveyed by British scientific expeditions.
The climate is oceanic. The average temperature of Tristan is 54°
F. with a range of 38-75° F. Rainfall of 66 inches is evenly distributed
through the year and the prevailing winds are from the west. There are
frequently violent storms particularly affecting the western portions of
the island. The islands are believed to be of Tertiary age and the main
masses are basaltic lava. There are evidences of older, local eruptions
similar to that which occurred in the fall of 1961, when the settlement
was evacuated.
The vegetation of Tristan has been studied by WACE and HOLDGATE
(1958) and that of Gough by wACE (1960, 1961). I am much indebted
to Dr N. M. Wace for the photographs used here as well as for data
from his vegetational studies. There is a marked similarity in the flora
associations within the larger vegetational divisions appear to be com-
mon to all of the islands studied. Differences are ones explained mainly
in the diverse topography. Gough is smaller, rectangular, with topo-
graphy irregular and the highest point, Edinburgh Peak, is 2730 feet.
Tristan, the largest island, has a circular form and the vegetation is
strongly defined by topography and altitude into four main zones. The
lowland plains—a limited area along the coast on which the settlement
is located that has been farmed and grazed. There are narrow beaches,
grassland and mires, and many introduced species occur here. The
cliffs form an abrupt barrier around most of the island, are nearly
vertical to 45°, and extend up to 1000 feet. Heath, scrub and grass-
land associations occur on the flanks or ridges and level terraces. On
steeper slopes there are frequent eroded areas caused by peatslips. The
* This study is supported by a National Science Foundation Grant GB 1693.
270 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
base forms a plateau about a mile wide between 1000 and 3000 feet
completely surrounding the island. This is the richest area for ferns and
about half of the species on the islands are found here in the vegetational
unit—the pteridophyte association. The peak from 3000-6760 feet,
the highest point, has a crater containing a lake and the vegetation
consists of grassland, heath and alpine desert. The varied topography
within each of these zones provides an abundance of habitats suitable
for pteridophytes. Thirty-three species are reported from Tristan and
6 from Gough.
Spores of Ophioglossum, Hypolepis, Vittaria, Grammitis, Hymeno-
phyllum, Eriosorus and others not readily identifiable occur in peat
samples taken in studies of the Pleistocene vegetation by HAFSTEN (1960).
The lowest level of peat has been radio-carbon-dated at nearly 5000
years. Two pollens of special interest, Nothofagus and Ephedra, also
occur in these samples although neither genus occurs on the islands.
The Nothofagus pollen has been identified as N. pumilio which occurs
in Patagonia and has occurred there throughout the Quaternary. This
and the Ephedra pollen are considered records of long distance dispersal
from South America.
The earliest collections from Tristan were made by Aubert du
Petit-Thouars in 1793. His Esquisse de la Flore de Tristan d’Acugna,
including descriptions of 16 pteridophyta, was published in the Mélanges
de Botanique et de Voyages (1811). It has recently been noted by Mrs
M. J. Van STEENIS-KRUSEMAN (1964) that a printed copy of the Esquisse
was presented to the French Academy as early as 24 October 1808 and
thus the descriptions of 13 species of pteridophytes based on this work
should have the earlier date. Collections of Thouars are in the Muséum
National d’Histoire Naturelle at Paris, and those I have examined are
ample and well prepared specimens.
The revision of the Tristan pteridophytes by CHRISTENSEN (1940),
one of the results of the Norwegian expeditions to Tristan, is the most
complete account of the vascular plants. The following list is largely
a compilation from this work and also that of wAce (1961) wit
emendations from recent studies and comments upon the species OF
specimens that I have examined. It is not possible to make final
taxonomic decisions in many cases as this awaits comprehensive studies
of species throughout their ranges. It is, however, possible to draw
reasonably accurate conclusions on the geographic relationships of the
The pteridophytes of the Tristan islands can be grouped ge0-
graphically into three main classes: South American—those with the
same or closely related species in South America and not in Africa,
these may also be more widely distributed especially in Australia and
New Zealand; Amphiatlantic—those which occur or have close rela-
tives in both America and Africa; African—those in Africa and else-
TRYON: FERN FLORA OF TRISTAN DA CUNHA 274
where but not South America.
SOUTH AMERICAN PATTERN
Species occurring in the Tristan group and the same or closely
related species in South America and usually elsewhere, but not in
Africa
Ophioglossum opacum R. Br. is a rare species occurring on the
higher slopes of the mountains, on coarse, rocky soils of the moorlands.
On the basis of a collection from St. Helena, CLAUSEN (1938) maintained
the species but noted that it has a bulbous stem similar to O. crotalo-
Phoroides Walt. and that it was treated by Prantl under this species
which ranges from the south-eastern United States south into Chile.
Hymenophyllum aeruginosum (Poiret) Carm.—on shaded rocks or
fallen tree fern trunks. This is one of the Sphaerocionium group having
branched trichomes on the leaves and is very close to if not the same as
the Chilean H. ferrugineum Colla.
Eriosorus cheilanthoides (Sw.) A. F. Tryon*—among rocks in fern
scrub or forest edge. With the exception of this species in the Tristan
group the genus is wholly American, occurring from southern Mexico
to Bolivia and Brazil. The unique spores are found throughout the
samples from the lowest levels to the recent and record the occurrence
of this species throughout the Pleistocene. Specimens from Tristan
are not distinguishable from some occurring in the Andes, formerly
called E. elongatus (Grev. & Hook.) Copel. or E. flabellatus (Grev. &
Hook.) Copel.
Vittaria vittarioides (Thouars) C. Chr.—a forest species occurring
on rock ledges or crevices; Christensen regarded it as an endemic
related to American species. It is distinct from those of the Andes in
having deeply sunken soral grooves some distance from the margin,
ranched paraphyses and large, yellow, tetrahedral spores.
Blechnum penna-marina (Poiret) Kuhn—a herbaceous member of
the Lomaria group, it has the widest ecological tolerance of any of the
Tristan ferns for it occurs on the strand as a colonizer on up to the peak
where it may be a dominant (Plate IIB). The species has been reported
to be diploid n=34 in New Zealand.
Asplenium alvarezense Rudmose Brown—an ephiphyte, frequent
on Blechnum trunks. Christensen considered it endemic but closely
related to 4. magellanicum Kaulf. (A. dareoides Desv.) which grows on
arborescent ferns in Chile. :
Dryopteris aquilina (Thouars) C. Chr.—forms dense layers with
leaves up to a metre tall, under Phylica trees. It is one of the Ctenitis
*Eriosorus cheilanthoides Sw.) A. F. Tryon comb. noy. oe
Grammitis cheilanthoides Sw., Syn. Fil.: 23, 219, 419 (1806), ‘Ins. Mauritii’ in error.
ZT? BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
group very close to, if not the same as, the Chilean D. spectabilis (Kaulf.)
C. Chr. Christensen regarded the plants of Tristan as distinct on the
basis of their coriaceous leaves with strongly enrolled margins and a
mixture of hair and scales, but the specimens that I have examined from
Chile cannot be differentiated on these characters.
Grammitis magellanicum Desv.—among mosses, epiphytic on rocks
or dead tree trunks. Christensen noted the relationship of these plants
to G. billardieri of Australia and New Zealand but regarded them as
distinct because of their nearly sessile and glabrous leaves. A broader
concept of the species is taken by sora (1960) in a treatment of the
genus in Argentina. The South American material is included as a
variety of G. billardieri Willd. and the Tristan plants appear to belong
here as var. magellanica (Desv.) Sota f. nana (Brack.) Sota on the
basis of the small size of the leaves. ; ;
Lycopodium diaphanum Sw.—in scrub and forest, usually in drier
sites. This is one of the clavatum group closely related to L. contiguum
Klotz. of South America as shown by the sessile strobili with long,
attenuated yellow sporophylls.
Other species having distribution patterns similar to those of
species noted above are: Trichomanes angustatum Carm., Asplenium
obtusatum Forst. f., Dryopteris parallelogramma (Kunze) Alston,
Polystichum mohrioides (Bory) Presl, Azolla filiculoides Lam., Lyco-
podium magellanicum Sw. Fifteen species appear to be represented in
South America by the same or closely related species.
AMPHIATLANTIC PATTERN
Species occurring in the Tristan group and the same or closely
related species in South America and Africa are usually elsewhere.
Histiopteris incisa (Thunb.) J. Sm.—one of the predominant
species in moist, open places of the fern bush community occupying
up to 90% of the field layer. A beetle, Coninomus, occurs a most
exclusively on this fern and is abundant on Gough where the Histiopteris
is most frequent.
Adiantum poiretii Wikstr—on wet rocks, walls and cliffs. The
species was described from a collection of Thouars from Tristan but 1s
widely distributed in the American tropics and Africa. Christensen
indicated that the species was likely an endemic but also noted that
some of the collections he examined agreed with forms from the conti-
nents. PICHI-SERMOLLI (1957) considers this distinct from the African
material which he calls A. thalictroides Willd. & Schlecht. Chromosome
reports of 2n=114 and n=60 add further complexities, all of which
may be resolved by studies of the plants throughout their range.
PLATE I
A, above) Phylica bush with Blechnum palmiforme in foreground, about 4 feet high,
Sandy Point, Tristan. B) Elaphoglossum succisifolium and Blechnum penna-mari
at 1000 feet in the Glenn, Gough. Both photos by N. M. Wace.
PRit FERN GAZ 9.107) 19.66
iM
a
ie?
a
TIN
a *Z > tbs
TRYON: FERN FLORA OF TRISTAN DA CUNHA 27:3
_ _Blechnum palmiforme (Thouars) C. Chr.—often forms tree fern
thickets (PLATE IIA) dominating extensive tracts sometimes with the
other arborescent plant, Phylica arborea Thouars, a member of the
Rhamnaceae. The trunks are up to 3 feet (1 m) and the crown about 3
feet in diameter. It belongs to the Lomaria group along with B. tabulare
(Thunb.) Kuhn of South Africa and B. magellanicum (Desv.) Mett. of
Chile. SCHELPE (1952) in his revision of the South African blechnums
indicates that the plants of this complex in Tristan and South America
are probably distinct from the African.
Blechnum australe L.—a herbaceous species of the Blechnum group
tolerant of moist or dry sites and salt spray belongs to a closely related
group occurring in both America and Africa. The American plants
have been segregated as B. auriculatum Cav., on the basis of the adnate
condition of the pinnae, but this is hardly a reliable character. A revi-
sion of the South American species is in progress by Maria Teresa
Murillo, Curator of Ferns at Bogota.
Asplenium monanthes L.—among rocks in exposed places or in tree
fern scrub. It is surprising that this species has a relatively limited
distribution in the islands for it is commonly occurring and wide ranging
in Africa and America. Plants from Madeira have been reported to
be apogamous triploids; thus, studies of the species in other parts of its
range should be of much interest.
_ Rumohra adiantiformis (Forst. f.) Ching—on broken terrain on
drier places in the fern bush and on talus receiving salt spray. The size
and division of the leaves are highly variable and several geographical
varieties have been recognized. The circular form of the sori an
numerous, broad petiole scales denote its relationship to Polystichum.
_ _ Elaphoglossum hybridum (Bory) Moore—on moss-covered rocks
in the forest. Plants from these islands are identical with specimens from
Réunion and from southern Brazil. This and the following more fre-
- Other pteridophytes having amphiatlantic distribution patterns
similar to those discussed above are Hymenophyllum peltatum (Poiret)
Desy., Hypolepis rugosula (Labill.) J. Sm., Asplenium erectum Bory,
Athyrium medium (Carm.) Moore, Thelypteris tomentosa (Thouars)
Ching, Elaphoglossum obtusatum (Carm.) C. Chr., E. insulare C. Chr.
and Lycopodium insulare Carm. Seventeen species have the same or
closest relations with species in both America and Africa.
AFRICAN PATTERN ;
Asplenium platybasis Kunze.—known from only a single collection
from the Phylica forest on Inaccessible Island. Christensen proposed a
274 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
variety, subnudum to distinguish this from the typical form on St. Helena
having denser indument. He indicated the relationships of these to A.
friesiorum C. Chr. in Africa and the wider ranging A. falcatum Lam.
This is a notable record for the islands as it is the only species among the
pteridophytes having its closest relatives in Africa without any American
relatives. These relationships however should perhaps be regarded as
tentative until the species from both of these areas are more closely
studied.
OTHER BIOTA ‘ego
It is of interest briefly to review pertinent data on the distributional
ranges of certain flowering plants, invertebrate animals and birds
where these give indications as to the origins of the flora and fauna.
Th fei 8 lant s* of Tnstan
have not been adequately surveyed but these are estimated at between
30 to 40 species about half of which are considered to be endemic.
Eight are the same or have closely related species in South America
and four occur only in the southern part of the continent; five occur in
Australia and New Zealand and only one occurs in Africa. The follow-
ing few illustrate these ranges. :
Lagenophora nudicaulis (Comm.) P. Dus.—a minute composite of
the tribe Asterae, usually less than 3 inches tall, occurs in Tristan and
Chile. BEAMAN & DE JONG (1965) in a recent paper on the American
species of Lagenophora, which have a disjunct distribution in the Andes,
note that the fruits or sticky glandular achenes are well adapted for
long distance dispersal.
mpetrum rubrum Vahl—a heath-forming plant of the Empe-
traceae. It occurs frequently with Blechnum penna-marina, is ubiquitous
in all vegetational zones up to the peaks and has reddish-black drupes.
It grows only on these islands and southern South America and adjacent
islands.
Phylica arborea Thouars (Rhamnaceae)—the dominant arborescent
plant in the islands which often occurs with Blechnum palmiforme. This
is the only species of flowering plant which has largely African relations.
The species also occurs in the New Amsterdam and Mascarene islands
but the genus centers in Africa. :
Acaena stangii Christophersen (Rosaceae)—resembling Potentilla
in its habit, has fruits with retrorse barbs. The genus is mainly of the
southern hemisphere but with disjunct species in California and Hawall.
Hydrocotyle capitata Thouars (Umbelliferae) is a small, creeping
plant considered to be endemic to the islands, but belongs to an austral
complex having members in South America and New Zealand.
The f a u n a of the Tristan group, like the flora is rela-
ray - ind-
ng about 600 plants (inclu
an account listing P paca
WACE ;
1965. The biological report of the Royal Society Expedition to Tristan da Cat
1962, Part II The terrestrial botany of the Tristan da Cunha islands. Phil. Trans:
R. Soc. London Ser.B. 249 :273-360.
TRYON: FERN FLORA OF TRISTAN DA CUNHA PA Be
tively sparse. There are no mammals, amphibians, reptiles or fresh
water fishes, but mostly invertebrates. HOLDGATE (1960) reports the
composition of the fauna as 107 marine forms and 87 terrestrial ones,
only eight of which are considered to be good faunistic indicators and
all of these show strong affinity to American species. Three different
groups Turbellaria (Procerodes ohlini), Diptera (Ornithomyia remota)
and Coleoptera (Lancetes varius) show closest relations with forms in
southern South America. Of 37 terrestrial and freshwater species having
clear affinities outside of Tristan, 31 are closest to South American
forms and only six are close to Old World or African species. There is
a high correlation of the invertebrates on Tristan with forms captured in
high altitude trapping including such flightless ones as springtails,
spiders and mites.
The birds are perhaps the best known group and there is record
of 33 species—21 breeding there and seven endemics, including a flight-
less rail on Inaccessible. There are five finches, four closely related, and
on a smaller scale these show the effect of adaptative radiation much as
the Darwin finches in the Galapagos islands. Non-resident birds in-
clude seven American species, among them the Barn Swallow and the
Purple Gallinule, both having their nearest stations in Brazil some 2000
miles and about 40 flying hours away. There are also seven American
Lepidoptera on Tristan.
CONCLUSIONS
It is well established that the islands are of volcanic origin and
probably of late Tertiary age and thus they have been available for
colonisation only since that time. It is generally accepted that if South
America and Africa were at one time united they reached their present
position by the Jurassic—too long before the Tristan islands were
formed to have had a role in the origins of their present flora. The
position of the islands on the mid-Atlantic Ridge suggests the possibility
that there may have been past isthmian links over which migrations
from the continents could have occurred. However, the paucity and
composition of the pteridophyte flora, as compared to those of South
America and Africa, do not support this possibility. There 1s a notable
absence of such families as Cyatheaceae, Gleicheniaceae and Schizaea-
ceae and of such genera as Pteridium, Pteris and Polypodium, and
particularly of the heterosporous genus Se/aginella. All of these families
and genera are prominent members of the fern floras of South America
and Africa. Migrations across land connections or island archipelagos
would be expected to result in a richer flora than the islands have and
one that contains a better representation of the predominant elements
of the adjacent continents. :
The poverty and composition of the pteridophyte flora is consistent
with migration by wind dispersal over long distances where the element
276 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
of chance becomes an operational factor. The kinds of propagules of
the plants, as the light fern spores produced in copious quantities, the
sticky achenes of Lagenophora and barbed seeds of Acaena, are also
consistent with long distance dispersal by air. It is evident that the biota
of the islands is composed of forms that are at one stage capable of
transport by air, being light, adapted for flying or clinging
The prevailing westerly winds which bring violent storms across the
south Atlantic are also consistent with this means of dispersal and the
biotic relations. ;
he source of the flora and fauna may be determined by relation-
ships with those of other areas. It is evident that these relationships are
predominately with South America and this is especially clear in the
pteridophyta. Among the 33 species, all but one have relatives or occur
themselves in South America, and 15 of these do not occur in Africa;
eight are only American having species in Tristan and America or are
endemic on Tristan and other oceanic islands with close relatives ex-
clusively in America. Seventeen of the 33 species have African relations
as well as American ones and a single species has wholly African re-
lations. The relationships of the pteridophyta, a large number of the
flowering plants and the known faunistic indicators are predominately
with South American species. Thus it is reasonable to account for the
largest part of the Tristan biota by means of air-borne immigration
from South America.
REFERENCES
BEAMAN, J. H. & DE JONG, D. C. D. 1965, A new species of Lagenophora (Compositae)
from Guatemala. ‘Rhodore 67:
on = 1940, The pteridophytes of Tristan da Cunha. Res. Norw. 1937-8
ped.
CLAUSEN, . me wt A ‘monograph of the Ophioglossaceae. Mem. Torrey Bot. Club
19: 1-177.
HAFSTEN, U. 1960, Pleistocene development 2 Ph rage = climate in Tristan da
Cun pe and Gou gh Island. Univ. Bergen Arb. Natury. 20: 1-45. a
HOLDGATE, M. auna of the eet Atlante islands. er se Roy. Soc. Lonao
Ser. B. 152: 550-567.
wall L. M. A. DU 1808, Esquisse de la Flore de Vile de Tristan d’Acugna,
—— 1811, Melee: de ae et de Voyages, Par
PICHI-SERMOLLI, R. E. G. 1957, Adumbratio flora Aethiopieae 5: Parkeriaceae, Adian-
e, Vittariaceae. Webbia
SCHELPE, E. A. S L. a bys e ‘ean of the Abicon species of Blechnum. J. Linn.
Soc. Bot.
SOTA, E. R. DE LA 1960, Polypodiaceae y Grammitidaceae Argentinas. Opera Lillo-
STEENIS-KRUSEMAN, M. N 1964, Dates of publication and bibliographical notes.
Fl. Males. Bull. ‘19: 1142,
WACE, N. aos or of the southern oceanic islands. Proc. Roy. Soc. London
er
—— 1961, The vegetation of Gough I sland. Ecol. Aone. 31: 337-367.
—& eres M. W. 1958, The vegetation of Tristan da Cunha. J. Ecol. 46:
BRIT. FERN GAZ. 9 (7) 1966 277
SOME RECENT WORK IN WALES ON THE
POLYPODIUM VULGARE AGGREGATE
P. M. BENOIT
Pencarreg, Barmouth, Merioneth.
INTRODUCTION
The existence of three cytotypes within Polypodium vulgare was de-
scribed by MANTON (1950), and sHIVAS (1961, 1962) published a study of
these cytotypes in which she treated them as species—P. australe Fée
(diploid), P. vulgare L. sens. str. (tetraploid) and P. interjectum Shivas
(hexaploid). The present writer came to study the group while working
on the Merioneth Flora and the result is some information that is
thought to be new and worth putting on record, concerning annulus
characters, size of stomatal guard-cells, habitat, and geographic distri-
bution. All the material studied was identified from morphology and
spore fertility.
HYBRIDS
Early in the study it became clear that the pentaploid hybrid P.
interjectum X< vulgare is common in those parts of Wales where both the
parent species grow; plants which from their morphology and sterility
were obviously hybrids were continually found, and they are now known
from sixteen stations in ten 10-km National Grid squares in Merioneth
intermediacy between the two closely similar wu
recording of these species unwise without observation of spore fertility
or a chromosome count. :
Fortunately Polypodium spores are produced in great quantity and
are relatively large; even with a strong hand-lens one can make out
the difference between those of the species (more or less uniform in
size and bright-yellow colour) and those of the hybrids (a mixture
varying from normal in size and colour to small shrivelled and white,
with some abnormally large). :
No plants suspected to be either of the other two possible hybrids
within the group (the triploid P. australe x vulgare or the tetraploid
P. australe x interjectum) were met with.
278 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
ANNULUS CHARACTERS
MANTON (1950) and sHivAs (1961, 1962) refer to differences between
the three species in the number of indurated (hinge)
¢ ells in the annulus of the sporangium. This is one of the most
useful distinguishing characters, though not an absolute one; Shivas
shows that there are overlaps between the species and suggests that a
determination should be based on a minimum of 25 annulus counts,
and ROBERTS (1966) has found that the overlaps between P. australe and
the other two species are even greater than was thought.
However, in examining a large amount of material for this charac-
ter the writer came to recognise that the colour he
annulus is probably the most reliable non-microscopic charac-
ter for distinguishing between P. interjectum and P. vulgare, the two
members of the aggregate most commonly met with. In P. vulgare the
dehisced sporangia are pale-buff with red-brown annuli and they gape
wide open, exposing the annuli and making them yet more conspicuous;
even with a hand-lens the annulus can clearly be seen as a dark line on
the sporangium. In P. interjectum on the other hand, the dehisced
sporangia are pale golden-brown with the annuli hardly differing in
colour, and even in dry conditions the two halves of the sporangium
remain more concave and half-closed, partially concealing the incon-
spicuous annulus. Annulus colour is of little use for distinguishing P.
australe, which falls more or less between the other two species. The
annuli of P. interjectum x vulgare are usually similar in colour to those
of P. vulgare.
The three species and the pentaploid hybrid all have different ranges
of annulus width (see Fics. 1-5). Here again the dif-
ferences, though a useful guide, are not absolute, except possibly
between P. vulgare and P. interjectum which can usually be separated
by a minimum of five annulus measurements per plant. The following
figures are the ranges of means of ten plants of each species and the
hybrid, obtained by measuring the width of five annuli (pressed flat)
in each plant: P. vulgare 67-75 u, P. interjectum x vulgare 74-80 HL,
P. interjectum 76-86 1, P. australe 81-100 . The individual measure-
ments are shown in the histograms Fics. 4 & 5. In P. australe the 1n-
durated cells are shorter than in the other taxa; that, combined with
their smaller number, gives the annulus a distinctive closely-packed
appearance (see FiG. 3). ROBERTS (1966) has studied this characters
and produced statistical evidence.
STOMATAL GUARD-CELLS
The three species and the pentaploid hybrid also have different
ranges of stomatal guard-cell length. Figures were obtained by measur-
ing a single stoma, visually selected as typical, in each of 250 plants—
80 each of P. vulgare and P. interjectum, 50 of P. australe, and 40 of P.
279
THE POLYPODIUM VULGARE AGGREGATE
BENOIT:
‘wnipodajog ut sisjovreyo |[90-pren8 pue snjnuue jo sweidosiy L-» SAUNOW
‘ajpajsnp “gq (¢ pue wnjoafsajul ‘gq (7 ‘aavdjna wumipoddjog ({ :1nuUeK dysLID\9RIeYO oY} Sulmoys wIBuBIodg ¢-]|
(tl) TISO-cdavno fO HLONAT (1) ‘TTHO-aquvnp JO HLONGT
r
a
—
Or
7
3. £2 OL L9 ¥9 19 8S SS tS 6h M EF OF OL EL OL 19 ¥9 19 BS SS TS br % E
; 1 : 3 Sano ame : omome
2 :
winaralsaquy s+ aueBjna > WNaDa[4aqUL ——— ajesgsne —-—.— auezina
(1) SMINNNY dO HLGTM (11) SMIQNNY JO HICIM
89 +9.
wo
o
a
SWIdNVS dO YaaNNN
280 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
interjectum x vulgare. A piece of epidermis from the underside of the
basal pinna in each frond was placed in a drop of water on a microscope
slide. The epidermis specimens were taken from fresh fronds, except in
the case of the rarer P. australe where observations were made from
dried material but, to counteract the shrinking effect of drying, the
epidermis was soaked in cold water for some hours before being
examined. Care was taken to sample poor and robust plants as well as
typical ones. The histograms Fics. 6 & 7 were constructed from the
results.
Figures obtained for stomatal guard-cell length are: P. vulgare
43-58 1, P. interjectum x vulgare 53-65 yw, P. australe 48-65 1, P. inter-
jectum 58-71 yp. The ranges of P. vulgare and P. interjectum, the two
common (and indeed in many parts of Britain the only) species are
almost discrete; in 80 samples of each of these two, the only overlap
is of two samples of P. vulgare and four of P. interjectum, all with 58 pt
guard-cells. P. australe and P. interjectum x vulgare are both inter-
mediate in this character though their ranges differ.
Thus, leaving aside P. australe, which is rare or absent in most
districts and has a distinctive frond shape that is fairly easily recog-
nised, stomatal guard-cell length would be an almost infallible guide to
barren Polypodium material but for the existence of the pentaploid
hybrid. Unfortunately, that is too abundant to be ignored. Neverthe-
less the length of the stomatal guard-cells remains a useful supple-
mentary character.
The almost exact intermediacy of P. interjectum x vulgare in this
character is noteworthy. In the histogram Fic. 7 the modal class of
the hybrid is the same as the uppermost class of P. vulgare and the
lowermost of P. interjectum; the extreme classes of the hybrid are the
same as the modal classes of P. vulgare and P. interjectum respectively ;
and 27 samples of the hybrid come within the range of P. vulgare and
26 within the range of P. interjectum. The distribution of annulus
width in the hybrid (Fic. 5) shows similar features. This is excellent
evidence of hybridity.
Although P. interjectum vulgare is so common and produces
some normal-looking (fertile?) spores, the histogram for guard-cell
length (see FiG. 7) shows an approximation to a normal distribution
with a range similar to those in the parents and supports Shivas’s view
that the majority of individuals are first-generation hybrids. The distri-
bution of annulus width in the hybrid shows similar features. Neverthe-
less, the spore fertility of the hybrid varies. One does find individuals
with a higher proportion of apparently normal spores, and it is probably
significant that they more closely approach P. vulgare in their mor-
phology; such plants may well be back-crosses or segregants, and they
deserve cytological study.
BENOIT: THE POLYPODIUM VULGARE AGGREGATE 281
The order of increasing length of the guard-cells, vu/gare—australe
—interjectum, shows that the length is partly a basic character of, or
inherited from, the various diploid species of Polypodium, as well as
partly a mere expression of ploidy, just as spore length (which has the
same order) was shown to be by Shivas. An increase, or at least not a
decrease, in the size of stomata is commonly associated with higher
ploidy, yet the allotetraploid P. vulgare has distinctly smaller stomata
than the diploid P. australe. This suggests that one or both of the
diploid progenitors of P. vulgare must have distinctly smaller stomata
than P. australe, despite the same level of ploidy—i.e. stomata at least
as small as those of P. vulgare itself—a point that may be of some taxo-
nomic value in the elucidation of the diploid species. P. interjectum
has the largest stomata of all, as one would expect in the species with
the highest chromosome number. Stoma measurements for the various
American diploid relatives of the group and of hybrids of them with
P. interjectum would be most interesting.
HABITAT
P. australe is well known to be a calcicole on the Continent, and it
obviously is so in Wales where almost all the stations are on highly
calcareous wall mortar or limestone rock. Observations in Wales show
that there is an ecological difference between P. interjectum and P.
vulgare; P. interjectum has clearly inherited the calcicoly of the australe
typically associated with Asplenium ruta-muraria, A. trichomanes ssp.
quadrivalens, Ceterach officinarum, and the mosses Encalypta strepto-
carpa, Barbula unguiculata and B. recurvirostra.
writer seen P. interjectum off wall mortar here, and then in most cases
as isolated plants that were probably strays. But in the Carboni-
ferous limestone country of Anglesey it is abundant along roadside
banks—a habitat where it apparently never grows In acidic country—
and it is to be expected that P. interjectum is frequent in natural habitats
in calcareous districts.
P. vulgare on the other hand is abundant and normally the only
species in acidic habitats—dominant Quercus petraea woodland on base-
deficient soils of the hill-country, overgrown bushy drystone walls and
banks, mountain rocks, and tree trunks. It is uncommon on wall
mortar, where it is pale-coloured, sickly-looking and poorly grown.
DISTRIBUTION
According to sHIvas (1961), P. australe is a southern European
species reaching its northern limit in the south-west of the British re :
P. vulgare is a northern species which extends to the Arctic; and f.
282 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966
interjectum is intermediate, extending as far north as the souther:,
altic.
A corresponding distribution pattern is to be seen in Wales. Here
P. australe is a local plant of lowland, mainly coastal districts, at all
common only in Pembrokeshire, where DAVIS (1964-5) has had con-
siderable success in searching for it; significantly this is the mildest part
of the Principality, coming within or near the 4° C mean February
minimum isotherm. P. vulgare is common everywhere, even in the
mountainous interior; indeed it is often the only species inland. P.
interjectum is intermediate—common on roadside walls in the mild
coastal strip but rare more than about ten miles from the sea. Lack Oi
mortared walls probably partly accounts for its absence from the acidiv
hill country; yet it is seldom found even on apparently suitable walls in
the valleys inland.
ACKNOWLEDGMENTS
The writer would like to thank Mr R. H. Roberts for freely making
available the results of his related studies, for criticising the draft 0
this paper, and for his encouragement; Mr T. A W. Davis for his
useful information about Polypodium in Pembrokeshire, and for speci-
ally collecting the material on which much of the data for P. australe
are b Mrs T. G. Walker (Dr M. G. Shivas) for her helpful remarks;
and the Keeper of Botany at the National Museum of Wales for the
loan of specimens from the Welsh National Herbarium.
REFERENCES
DAVIS, T. A. W. 1964-5, Field notes: Plants: Pembroke. Nature in Wales 9: 28, 70, hg
MANTON, I, 1950, Problems of Cytology and Evolution in the Pteridophyta, Cambri “at
ROBERTS, R. a “i ome observations on Polypodium australe. Brit. Fern Gaz. 9:
283-28
1961 , Contributions to the cytology and taxonomy of species of Poly-
SHIVAS, M. G. edge”
podium in Europe and America, I: Cytology, If: Taxonomy. J. Linn. 90¢-
Bot. 58: 13-25, 27-38.
—— 1962, The Polypodium vulgare complex. Brit. Fern Gaz. 9: 65-70.
acer
EQUISETUM: REQUEST FOR MATERIAL
Living material of Equisetum variegatum, E. ramosissimum, E.moorel,
E. x trachyodon, and E. x litorale is required for research purposes
involving experimental cultivation techniques.
Anyone who could supply small amounts of any of these species
from wild habitats is invited to communicate with C. N. Page, Depart-
ment of Botany, The University, Newcastle-upon-Tyne 1. Material of
any foreign species of Eguisetum would also be most welcome.
BRIT. FERN GAZ. 9 (7). 1966 283
SOME OBSERVATIONS ON POLYPODIUM
USTRALE
R. H. ROBERTS
51 Belmont Road, Bangor, Caerns.
INTRODUCTION
Three of the more distinct varieties of Polypodium have been recorded
from the extreme north-east corner of Caernarvonshire, v.c. 49,
These are var. cambricum (L.) Lightfoot (PENNANT 1784; Lowe 1865;
HYDE & WADE 1940), var. semilacerum Link (HYDE & WADE 1940) and
var. serratum (Willd.) Milde (HYDE & WADE 1940; GRIFFITH 1895). It
is worth noting that all three have been found at different times at
Conway, while two of them—var. cambricum and var. serratum—are
also recorded from the area of Carboniferous limestone near Llandudno.
AS MANTON (1950) has shown, both var. serratum and var. semila-
cerum usually correspond to the diploid cytotype, of which the var.
cambricum is thought to be a sterile variety. This being so, the occur-
rence of these three varieties at the same locality was not unexpected.
It also implied that Conway and the limestone outcrops around
Llandudno were likely localities for the diploid.
Following the appearance of the papers by sHIVAS (1961), a search
was made at both places. At Conway plants were found which answered
in many respects to the description of P. australe Fée. They occurred
In considerable quantity on the walls of the castle and on parts of the
town walls, though a few plants grew in crevices of the highly calcareous
rock on which the castle stands. A brief search around Llandudno was
Unsuccessful, but the presence of P. australe on the adjacent limestone
has been confirmed recently by Mr Eirian Griffiths of Deganwy.
Examination of the specimens gathered from Conway at once
presented some difficulties. While some of them seemed to answer to
the description of P. australe fairly closely, others were not so easy to
identify with certainty. These had rather longer and narrower fronds,
and their pinnae were not so strongly serrated. What proved more
perplexing was that in some plants the number of indurated cells in
the annulus of the sporangium was frequently greater than the range
of 3 to 8 found by Shivas. As the specimens had been collected in
March, it was difficult to determine whether they possessed paraphyses
or not. At the time it was suspected that some of these plants might be
hybrids of P. australe with either P. vulgare L. or P. interjectum Shivas.
Pieces of rhizomes of some of these doubtful plants were brought
into cultivation and more fronds were gathered from Conway later in
the year. A closer study of this material yielded some interesting
284 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
results. Paraphyses were found among the young sporangia in all of
them. They were also seen to be fully fertile, producing a full comple-
ment of good spores. The possibility that they were hybrids was there-
fore regarded as highly unlikely. Shivas has shown that the presence
of paraphyses is diagnostic for P. australe in Europe; they do not occur
in its Fl hybrids which, moreover, are apparently sterile and produce
mostly empty mis-shapen spores. Gradually it became clear that these
problematic plants all belonged to P. australe, which in Conway seemed
to be much more variable than the descriptions of this species indicated.
A more detailed study of the variation of some of the characters which
have been used in its diagnosis was consequently made.
THE NUMBER OF INDURATED CELLS IN
THE ANNULUS OF THE SPORANGIUM
Counts of the number of indurated cells in the annulus were made
in random samples of sporangia obtained by scraping a few sori from
each frond. The range of values differed from one plant to another.
While in some plants it was only 4 to 8 and agreed closely with the
results obtained by Shivas, in others the total range was from 5 to as
much as 14, with a mean value around 8. This was the case, for ex-
ample, in a plant whose fronds in all other respects answered to the
description of var. serratum. But other specimens had various inter-
mediate ranges connecting these extremes, and it was remarkable that
many of them had sporangia with 9, 10 or 11 cells in the annulus. The
aggregate data from the Conway specimens are shown in FIG. bs
In view of these anomalous results further counts of the number of
indurated cells in the annulus were made on specimens from various
other localities. Some of them, kindly lent by the National Museum of
Wales, were from widely separated places: from Glamorgan and Mon-
mouthshire to Somerset and Wicklow. Others were obtained from
about a dozen localities in south-west Wales, where P. australe has
been detected in many places recently by the enthusiastic field-work of
Mr T. A. W. Davis.
The data obtained from them show quite clearly that the Conway
plants are in no way exceptional. Many of those from south-west
Wales are evidently just as variable. As the histograms (Fics. | & 2)
show, the number of cells in the annulus has the same range in both the
Conway and Cardigan samples. The counts from a single frond from 4
Somerset locality (Fic. 3) also correspond closely with them. The moda
class in these three is identical, but it must not be assumed that this 1s
the case in all specimens. While it is true that, whenever a representa-
tive sample of plants from any single locality was examined, the mode
was usually around 6 or 7, several individual plants were encountered
in which the mean number of cells in the annulus was between 8 and 9.
In one of the plants from Conway the average, based on 95 sporangia,
POLYPODIUM AUSTRALE
RGBER TS
‘unipodéjog JO saisads 9a1y} dy} Ul snjnuue Jad s{j99 poyenpu jo WSus] ULE; (¢ ‘sojdures
[[@ JO ayeBa133e oy (p ‘orew-1odns- -U0}SO AA, WOI} Puory o[SuIs v (¢ ‘URZIpred wos} URId sUO (7
‘ABMUOD WO1] ool Udd}XIS aft: :UO pase ‘ajp4jsny ‘g UI snjnuue eon S]]99 payeinpul jo Joquinyy
(11) STIZO CHEVENGNI JO HLONTT STISO CHLVENGNI @o usanaN
SEELZI Ol 6 G2 9 6 6 ez PLEL TI 1101 68.2 9.5 Fe
@ 0 B86 9€ ve TE “OF 807 SF 40 222 0e 8 Soh aoe aoe eoneme ] amore: eeagmeemmmme ri
Les s
beg &
'
! ol
' L Ol
' S|
i
| . .
: y ois 2) .
had 4
as b 0% &
1 mo
wine Ly | rlEIZIL NOL 68 L9S HET weit Olé 8 LIS HEL
| | ayezysne = CEC oe 9%
j of ; S
. q PO!
I . SI
oie3[NA L OF
0
: | : :
$ “DJ i ¢ ‘Old eee F
286 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
was as high as 9-6. The overlap with P. interjectum is thus seen to be
much greater than previous studies have indicated; the means from a
few individual plants even approach the lower mean values found in
P. vulgare. The pooled results from a total of 520 sporangia are shown
in Fic. 4.
THE LENGTH OF THE INDURATED CELLS
During the course of this study the usefulness of another micro-
scopic character became apparent. This is the size of the indurated cells
of the annulus, to which attention has already been drawn by Manton.
When the annulus of a dehisced sporangium is flattened by moderate
pressure between two glass slides its length can be measured quite
readily. The mean length of the cells in the annulus is then found by
dividing this value by the number of cells it contains. Measurements
of 20 annuli were made from each specimen of P. australe, P. interjectum
and P. vulgare from a number of localities. The results obtained so
far (Fic. 5) show that there is a considerable difference between P.
australe and P. interjectum in this detail. Means based on ten annuli
fall into two sharply separated groups: 21-26 p for P. australe, 28-35 »
for P. interjectum. In practice this has proved to be one of the best
means of separating these species. As BENOIT (1966) has shown, there
is little difference between them in the width of the annulus. As a result
the flattened annulus of P. australe has a more contracted look than
that of P. interjectum. In most cases this difference is readily perceptible
under the microscope, and has been found especially helpful in identi-
fying specimens of P. australe with a mean number of indurated cells
greater than 7. On a few occasions it has also proved useful in the
determination of specimens of P. interjectum in which most sporangia
have as few as 6 indurated cells, with a total range of only 4 to 8, thus
by this character proving inseparable from P. australe, even when the
minimum of 25 annulus counts suggested by Shivas was made.
However, the difference in this character between P. australe and
P. vulgare is too small to be of use in this way, as can be seen from the
histograms. But there is never the same difficulty in separating these
two species, as the mean number of cells in the annulus of P. australe
is rarely as great as in P. vulgare, and even then the wide difference 1n
frond shape is sufficient to distinguish between them.
FROND SHAPE
Another character used in the diagnosis of P. australe is the shape
of the frond. This is described as broadly ovate to triangular, and the
longest pinnae are stated to be the second or third pair from the base
(SHIVAS 1962). It was found in the present studies that many of the fronds
did not fit this description and some were even markedly oblong-
lanceolate; the longest was most often the third pair, but there were
several where it was fourth, fifth or even sixth. Even so, as long as this
ROBERTS: POLYPODIUM AUSTRALE 287
greater variation is borne in mind, frond shape remains a good diag-
nostic character of this species.
Thus, while these results suggest that the variation in British
material of P. australe is considerably greater than was realised hither-
to, its specific status is in no way affected. Increasing familiarity with
it has merely served to confirm that in all respects it is a very distinct
taxon.
ACKNOWLEDGMENTS
I wish to thank Mr P. M. Benoit for much useful discussion of various
points, for criticism of the grat a and for the loan of herbarium
specimens; also Mr T. A. W. Davis, whose diligent field work made a
wide range of material available Fein m many new localities; and the
eeper e ee at the National Museum of Wales for the loan of
specimen
REFERENCES
BENOIT, P. wa 1966, Some recent work in Wales on the Polypodium vulgare aggre-
Brit. Fern Gaz. 9: 277-282.
eid : 1895, The Flora of Anglesey & Carnarvonshire, Bangor.
YDE, . & WADE, A. E. 1940, hades h Reo re cs
LOWE, " ; 1865, Our native Fer.
MANTON, 1. 1950, Probes of Citolaey and Eoakeion in the Pteridophyta, Cambridge.
PENNANT, T. . A Tour in Wales vol. 2, 40, London
SHIVAS, M. G. 1961, Contributions to the cytology and taxonom my of species of fee
odium in Europe and America, I: Cytology, II: Taxonomy. J.
Pe
Soc. Bot. 58: 13-25, 27-38.
—— 1962, The Polypodium vulgare complex. Brit. Fern Gaz. 9: 65-70.
REVIEW
THE FERNS OF PERU: POLYPODIACEAE (DENNST AED-
TIEAE TO OLEANDREAE) [Contrib. Gray Herb. Harvard Univ.
CXCIV] by Rolla Tryon; 253 pages, 16 x 24 cm; Cambridge, Mass,
1964.
There are those who on seeing : _ like the above may dismiss it
yY Saying “I shall not be collecting ferns in Peru’. Such persons,
however, if they have a interest in growing ferns forget they may
and
and those to Adiantum (41 taxa), Pityrogramma (9) and Cheilanthes
(15) are especially helpful. Dr Tryon reflects his _— field serra
pre poe (Natural History), London SW7 A. C. JERMY
DYCE: -PHE- BP:S2 AUSTRIAN “EXCURSION 289
noted on walls several Asplenia — A. adiantum-nigrum, A. ruta-
muraria, A. septentrionale, A. trichomanes, along with Cystopterts
fragilis, Phyllitis scolopendrium, Gymnocarpium dryopteris and G.
robertianum. In the ravine larger ferns were plentiful, including
Athyrium filix-femina, several of the Dryopteris species including the
hybrid D. x tavelii, Pteridium aquilinum and, most interesting of all,
Polystichum aculeatum, P. braunii, and P. setiferum along with hybrid
forms. Oak and beech ferns were abundant, as indeed they were wher-
ever we went during the Excursion, while Blechnum spicant and Poly-
podium vulgare were also common.
In contrast to the sunshine on Monday, the following day dawned
dull and wet, and we delayed our start till about 10.30 when the rain
showed signs of abating. The weather decided some of our company
to spend the day sight-seeing in Graz, and our guide led a reduced
party up among the vineyards where the retaining drystone walls of
the terraces were full of ferns, chiefly the smaller species, and we spent
a busy time studying and photographing Asplenium hybrids. The spec-
ies A. adiantum-nigrum, A.ruta-muraria, A.septentrionale, A.tricho-
manes, A, viride were all present, and growing happily with their par-
ents were the hybrids A. x alternifolium (septentrionale trichomanes)
and a possible A. adiantum-nigrum trichomanes. Cystopterts fragilis
was also common in the walls.
Above the vineyards we traversed the hillside along one of “
rough hill roads, and dropped down again by a sunken Jane full o
large ferns. A fine vigorous D. filix-mas with crispate pinnules was
collected. The lane was the home of many salamanders, and some time
was spent watching and photographing the slow-moving beautifully
Returning to the valley we followed the river back towards
Deutschlandsberg, passing a large colony of Matteuccia strato
growing in boggy ground at the water’s edge. Some of the party electe
to return to the town while a few of us with Herr Otto set off up the
opposite hillside where in the woodland Thelypteris limbosperma was
common and in a boggy hollow D. carthusiana was at home in the
midst of an extensive colony of Eguisetum. These “‘fern allies were
well represented in the area, and we found E. arvense, E. hyemale,
E. palustre, E.sylvaticum, and E. telmateia. Lycopodium clavatum
was also seen and Herr Otto was able to show us some plants of
L. complanatum.
Later in the afternoon we said goodbye to our guide and [0
Deutschlandsberg and set off for our next stop, Pernegg, about 20
miles north of Graz. We joined our sightseers at Graz station, and a
meal there was made memorable by delay which led to a hectic rus
to catch our connecting train. Our hotel in Kirchdorf, an adjacent
288 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
THE B.P.S. AUSTRIAN EXCURSION*
J. W. DYCE
46 Sedley Rise, Loughton, Essex
For the first time in the history of our Society, and in keeping with
our wider field of interest, we ventured abroad in 1965 — to Austria
— for our Annual Excursion. On Saturday 21st August, 17 members
and friends assembled at Victoria Station in London to catch the boat
train, looking forward to a fortnight among the Austrian mountains
and the pleasure of finding new ferns. Our party included Miss S.
Archer, J. W. Dyce, G. Gilbert, Mr & Mrs A. C. Jermy, Miss
M. Norman, Mrs. P. Simmonds, P. Temple and W. Wright from
London, H. J. Bruty, Dr F. M. Jarrett and Professor R. E. Holttum
from Kew, Mrs E. M. Brightman and J. J. Brightman from Orpington,
Mrs G. L. A. Craw from Dorking, P. Halligey from Lymington, and
K. Trewren from Middlesbrough. In Austria we were joined by Mr
& Mrs Allen from Los Barrios in South Spain, Dr. Alois Patzak from
the Vienna Naturhistorisches Museum, and Professor Helmut Melzer
from Judenburg in Austria.
Travelling on the Tauern Express, we reached Graz in Steiermark
on Sunday evening, and from there journeyed south to Deutschlands-
berg, a small pleasant town in the valley of the Lassnitz River. Here
we were out of the wilder mountain area and surrounded by wooded
hills, some with vineyards clothing their lower slopes, and the town
was gay with colourful flower-beds and window-boxes.
On the outskirts of Deutschlandsberg the Lassnitz emerges from
a deep wooded ravine full of ferns, and this was the obvious choice
for our first day. By strange good fortune, as we started our hunting
we met a local botanist, Heinz Otto, who is working on the ecology
of the valley and has a particular interest in ferns. He joined forces
with us during our two day’s stay, and we would put on record our
appreciative thanks for the enthusiastic help he gave us. He has a wide
knowledge of the ferns of the area and their distribution, and this was
put freely at our disposal so that we were able to see most, if not all,
of the ferns of the district, including the rarer ones. Without his assist-
ance we could not have hoped to achieve so much in two days.
Following the path along the river from Deutschlandsberg we
* This describes the ANNUAL EXCURSION. :
The Newsletter will report other MEETINGS IN 1965 viz 13 February
(London SW7), 30 May (Canterbury), 3 July (Kew), 23-25 July (Ashdown
Forest), 18 September (A.G. ondon 9-10 O
Humphrey Head, Kendal), 23 October (London NW3
Amendment to heading of last year’s reports
The heading “Meetings in 1965” in Gazette 9:239 should have read
1964
“Meetings in oF
290 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
village to Pernegg, was reached at bedtime, and next morning we were
up betimes to make an early start in spite of heavy rain which fortu-
nately did not last long.
€ were now in the valley of the Mur with mountains of pre-
cipitous rock rising on the eastern bank but those on the west, although
high and steep, were tree-covered and less rugged. A long pleasant
walk took us into the wooded ravine valleys of those western moun-
tains, where we split into several parties probing in different directions.
The conditions were ideal for ferns, and the large-growing woodland
species were abundant. Along the tracks and on the banks of their
accompanying streams grew Dryopteris filix-mas, D. borrert, D.
dilatata, D.carthusiana, with the hybrids D. x doeppii (borrert *
carthusiana), D. x tavelii (borreri = filix-mas) and others which ap-
peared to have even more involved parentages. Athyrium filix-femina,
Matteuccia struthiopteris, Thelypteris limbosperma, Pteridium aqul-
linum were noted, also Polypodium vulgare, Cystopteris fragilis, and
the oak and beech ferns. We also recorded Equisetum arvenseé, —
palustre, E. pratense and E. sylvaticum, Selaginella helvetica creeping
happily over wet rocks, and the clubmosses Lycopodium annotinum
and L.clavatum. Gentiana asclepioides in full bloom was common
throughout the area and was greatly admired, as were colourful patches
of Cyclamen europwum growing by the wayside. : :
Stiff climbing took us above the valleys with magnificent views
opening out over the wooded hills. The wet morning had cleared up
into a bright sunny day and the alps with their scattered chalets, perched
high above in the clearings, were bathed in warm sunlight. It was a JOY
to rest after the labours of climbing and drink in the beauty of the
peaceful scene. On the high ground many other ferns of interest were
seen, including Asplenium septentrionale, A. trichomanes, a0
Gymnocarpium robertianum. :
The next day was spent on the east side of the valley, and starting
from the village of Mixnitz we followed the road to the Barenschutz
Waterfall. As we climbed, the road deteriorated to a track leading UP
the narrowing valley between high cliffs, on which we found new a
Asplenium lepidum, A. trichomanes subsp. inexpectans, and the hybri
A. javorkae (lepidum x ruta-muraria). A large stream poured down
the valley which continued to narrow till we reached an almost vertical
cleft in the rock-face, down which the water cascaded in numerous
waterfalls and rapids. Up this cleft our path led, and for over 600 feet
we climbed by a series of step-ladders and bridges, a most amazing
construction made from logs, the bracing supports driven into crevices
in the rock or jammed across the narrower parts of the chasm. In places
the ladders clung, seemingly precariously, to the rock-face, sometimes
bathed in the spray of the waters swirling past us. It was an impressive
DYCE: ‘THE B.P.S.: AUSTRIAN EXCURSION 294
climb, made more interesting by the many ferns luxuriating in the
humid conditions. Athyrium and Dryopteris species were abundant,
with other ferns, but most exciting was Cystopteris montana, that very
rare fern of the Scottish mountains, growing in large vigorous colonies
on the wet rock-face. It was an exciting moment when we were able
to admire this fern at close quarters. Above the waterfall the valley
opened out into alpine meadows, with many alpine flowers and ferns,
among them Polystichum lonchitis.
Mixnitz would make a good centre for a much longer stay, and so
it was from here that on our third day in the district some of us set
off up the eastern mountains once again, to the DrachenhGhle, while
others had a restful day in the valley, and some explored the opposite
hills where, on patches of serpentine rock, Asplenium adulterinum was
found. The Drachenhéhle, the Dragon’s Hole, an immense cavern
running deep into the mountain, was reached after a steep climb
through woodland and then up a rock chimney. Coming out on top
of the cliffs high up the mountain-side, we had magnificent views of
the valley below with the Mur River, and strung along its banks the
villages Mixnitz, Pernegg and Kirchdorf. We had been joined on the
previous evening by Professor Helmut Melzer from Judenburg who is
working on the fern hybrids in the area, and he was able to lead us to
many interesting Asplenium hybrids growing on the cliffs. In addition
to A.lepidum, A. ruta-muraria, A. trichomanes subsp. quadrivalens
and A. viride, we saw A. trichomanes subsp. inexpectans x viride (A.
* adulteriniforme). Cystopteris fragilis and C. regia, Gymnocarpium
dryopteris and G. robertianum were also seen.
In the morning our luggage had been left at Mixnitz Station, and
from here we travelled during the evening to our next stop, Kraubath,
further up the Mur valley. The main attraction here was the serpen-
tine rock with its Asplenium hybrids, and on the following morning
(Saturday) we were off early, led by Professor Melzer. A narrow valley
leads into the hills from the nearby hamlet of Chromwerk, and in a
short time we were in the serpentine area. To us, accustomed to
regarding Asplenium hybrids as rare treasures, it was an exhilarating
experience to see them growing abundantly on the rocks, and in the
Screes and grass. The only species seen were A. adiantum-nigrum (often
referred to as A. forsteri, a species of problematic status, by local
botanists), A.ruta-muraria, A.trichomanes and A.viride, but they were
outnumbered by their hybrid progeny. A. forsteri (v.s.), a fern of the
serpentine, was common and we found a few plants of A. x woynaria-
num, a hybrid between A. adiantum-nigrum and A. viride. A. adul-
‘erinum, another fern restricted to the serpentine, was also common,
but more interesting were the crosses of adulterinum with the
Subspecies of trichomanes and with viride.
292 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
After lunch we returned to the main valley and explored around
a large quarry at Schaffer Hube. We found Asplenium hybrids here
again, and in addition many flowering alpines, as well as another new
fern, Cheilanthes marantae, growing in loose rock accumulated at the
base of the cliffs. In the wooded slopes below, A. viride grew vigorously
and a large colony of huge plants was found with bifid and trifid
fronds.
Sunday was a day of rest -— spent sitting in trains as we travelled
westwards and in the evening we reached Krimml, not far from the
Tirol border. We had left Steiermark behind, a quiet peaceful country
unspoilt by tourism and were now in a popular area where everything
was much more sophisticated — and expensive. The attraction of
Krimml is the magnificent Waterfall, the finest in Austria — the waters
come from the glaciers above and precipitate themselves down the
mountainside in three successive falls totalling about 1400 feet, a most
impressive spectacle.
Needless to say, it was to the Waterfall that all our footsteps
led on the following morning — a wet one, which is not to the liking
of tourists so we had the place more or less to ourselves. Later when
the sun came out so did all the sightseers, their attire and footwear
more fitted for city streets than the rough track which zigzagged
up the mountain through the woods alongside the Falls, with many
viewpoints from which to admire and photograph the tumbling waters.
Below lay the valley and the village of Krimml, caught occasionally in
fitful sunlight forcing its way through breaks in the creeping mists.
We duly admired, photographed, and were impressed by the magnifi-
cence of the scene, but there was much more of interest to us in the
abundant plantlife, with ferns and their allies predominant. The con-
ditions were ideal for these, and they flourished accordingly, giving
us a very rewarding day.
It was obviously not Asplenium country, and only A. viride was
noted. The Lady-fern and the common Dryopteris species were abun-
dant, including the hybrid D. x tavelii, and the oak and beech ferns
were everywhere. Polypodium vulgare was seen and we looked in vain
for P.interjectum; Matteuccia and Thelypteris limbosperma were
looking superficially very similar. Four Polystichum were there
— P. aculeatum, P. braunii, P. lonchitis and magnificent plants of P.x
luerssenii (aculeatum x braunii). C ystopteris fragilis was common
among the huge rocks littering the mountainside and some specimens
with fine attenuated pinnules were collected. Only one Equisetum, E.
sylvaticum, was seen, also Selaginella selaginoides and Lycopodium
selago and L. annotinum. Extensive fruiting colonies of the lattet
provided some good colour slides.
DYCE: THE B.P.S. AUSTRIAN EXCURSION 293
After our brief visit to the Krimml Waterfall we were again on
our way, and left Krimml at 5 o’clock by bus, to travel by the new
High Alpine Road over Gerlosplatte to Mayrhofen. This road zigzags
up the mountains above Krimml in a series of hairpin bends, and in
one place winds completely round passing over itself. Impressive views
of the Waterfall across the valley opened out as we climbed. Darkness
had fallen before we descended again steeply to Mayrhofen, so we
were unable to admire the beauty of this small town in its superb
mountain setting. Another bus run took us up the Zemm valley to
Ginzling tucked away among the hills. From here only a rough track
led steeply up the Zemmtal for about 6 miles to Breitlahner, and up
this we were transported by minibus — quite a hair-raising experience
— to the Breitlahner Gasthof, where we were to remain for the rest
of our stay in Austria
Breitlahner, a hamlet of only a few houses scattered over a small
alp at the junction of two valleys, the Zamsertal and the Zemmtal, at
an altitude of over 4000 feet, is the end of the road for wheeled traffic.
From here one walks, and pack-horses are used for transport to the
habitations further on. These narrow valleys between precipitous
mountains run southwards into the high Tirol hills, the Zemmtal
climbing for about 7 miles to the Berliner Hiitte at about 6700 feet,
and the Zamsertal much longer, but giving an easier climb of about
4 miles, to the Dominikus Hiitte. Mountain torrents race down to meet
at the junction of the valleys, and at this spot stands the Gasthof in a
fine setting.
We were looking forward to four days scrambling, climbing and
plant-hunting, and we started off eagerly on the Tuesday morning in
fine weather, each intent on spending the day in his or her own way.
In the afternoon the rain began, and continued incessantly all week
till Friday morning when it suddenly stopped, the sky cleared, and
we saw the sun again. This was the storm which caused so much chaos
and havoc throughout the Tirol and the north of Italy, but it did not
disturb us unduly apart from cramping our activities somewhat. Cas-
cades poured over the mountain precipices and surged down the
valleys. Everywhere was sodden underfoot, and there were muddy
landslides over the tracks, bridges were swept away, and the occasional
reverberating roar of falling cliff loosened by the rains echoed round
the hills. Intermittent glimpses of the high peaks through the clouds
Showed them becoming whiter and whiter with fresh snows. And on
the Friday afternoon, when they glistened in the sunshine against a
blue sky, there were magnificent scenes to use up the colour film in
our cameras.
There was much coming and going during our stay, the tougher
members venturing further afield, some to climb to the Berliner Hiitte
2.94 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
where they stayed for one night, some to the Dominikus Hiitte, and
others to scramble on the wet mountain slopes bringing back ferns and
alpine plants. Our less energetic members, more averse to water and
mud, sallied forth during the short intervals when the storm decreased
in intensity. The drying-room was working overtime, full of dripping
clothing and soaked boots, but our spirits remained cheerful, undamped
by the weather.
Ferns were plentiful in the valleys and on the slopes, as well as
alpine plants many of them still in bloom and creating a counter-
attraction to the ferns. One delightful find was Moneses uniflora, the
Single-flowered Wintergreen, which is a very rare plant of the north
of Scotland. This was in flower on the rocky wooded slopes of the
Zamsertal. Sheltered valley bottoms were the habitats of the larger
ferns and we noted Athyrium filix-femina, Dryopteris assimilis, D.
borreri, D.carthusiana, D.dilatata, D. filix-mas, D. X tavelii and
several other problematic hybrids in this genus, Matteuccia struthiop-
teris, Polystichum aculeatum, P. braunii, P. x luerssenii, Thelypterts
limbosperma and Pteridium aquilinum. With them were Blechnum
spicant, Polypodium vulgare, Thelypteris phegopteris and Gymnocar r
pium dryopteris. “Fern allies” were well represented with Equisetum
arvense, E. palustre, E. pratense, E. sylvaticum, E. variegatum, Lyco-
podium annotinum, L. clavatum, L. selago and Selaginella selaginoides.
Higher up grew L. alpinum, and another selection of ferns, the
more interesting being Asplenium x alternifolium, Botrychium lunarta,
P. lonchitis, P. illyricum (aculea‘um = lonchitis), Cystopteris mon-
tana and Woodsia alpina. It was most interesting to see Our ap
P. lonchitis growing abundantly on the hillsides. Only a few plants .
the very handsome strong-growing hybrid, P. x illyricum, were =
Other spleenworts noted were A. ruta-muraria, A. septentrinnts:) :
trichomanes and A. viride, along with Cystopteris fragilis and Athy
rium distentifolium. ft this
n spite of all the rain it was with great regret that we le t a
lovely spot early on Saturday morning, needless to say in brillian
sunshine, setting off down the track by minibus to Ginzling and Mayt-
hofen, where we caught our train en route for Innsbruck. Shopping
and sight-seeing occupied the time there before boarding our train for
home and in the late Sunday afternoon we were again at neelaae
Station and departed on our separate ways with happy memories ©
a most successful Excursion. : ?
e thing remains — to give tribute to those whose special contri-
butions made the Excursion successful and to whom we are all greatly
indebted. Particular mention must be made of the preliminary pla
by Miss Anne Sleep on which our whole itinerary was based, and ne
the tremendous amount of work put in by Mr. Peter Halligey bo
BECE: “THE BPS) AUSTRIAN EXCURSION aa35
before and during the Excursion. To him is due the credit for the
Organisation which ensured that everything went off without a single
hitch from start to finish and we are deeply grateful to him. Dr. Patzak
gave us invaluable assistance and guidance which has been greatly
appreciated, as was the help given by Professor Melzer whose know-
ledge of the ferns and in particular the hybrid forms, made our visit
to the Mur valley so very rewarding.
FERN CLASSES AT SOUTHPORT FLOWER SHOW
25-27 AUGUST 1965
JUDGES’ REPORT
It does not seem a year ago since W. G. Mackenzie and I were walking
round the fern exhibits at Southport and as always it is a great pleasure
to note the ever increasing standard of perfection the exhibitors have
achieved in growing and presenting their favourite specimens. This
year, through ‘‘circumstances beyond their control’’, one or two of the
major exhibitors had not entered their usual contributions and their
a great surprise in view of the conditions : a strong gale was blowing,
One or two marquees actually were blown down, the side canvas was
billowing uncontrollably, the roof canvas was whipping and cracking
as though it would split asunder any second. The noise was nerve
racking, yet the following morning all was well, the ferns stood serene
and beautiful as though nothing had happened.
_ Noel Brookfield showed his usual ground group full of first class
varieties. All the classes were entered, one or two very strongly, giving
the judges some close pointing and much deliberation before making
the awards. In the greenhouse ferns I noted a fine pan of Phymatodes
diversifolia — this grows very freely in the open up the trunks of palm
trees at Logan — and there were good examples of Davallia and
Platycerium. The superb specimen of Adiantum ‘Kensington Gem
shown by Bernard Hayhurst was better than ever. One gets to recognise
some of the exhibits as old friends. Such fine specimens should do much
to stimulate public interest in our hobby and encourage new exhibitors
to try their luck.!
The British Pteridological Society is indebted to the South-
port Show Committee for providing such a comprehensive set
of classes for British ferns and the judges would like to see even more
members coming forward as exhibitors at future shows.
R. KAYE
Waithman Nurseries, Silverdale, Lancs.
296 BRITISH FERN GAZETTE: VOLUME 9 PART 7 (1966)
PRIZEWINNERS AND THEIR ENTRIES
CLASS 8 group of hardy British ferns, with accent on diversity and
ie garg arranged in a natural manner in . space 13 feet by 3 feet
Pt fee ot higher bers 6 feet above the grou
Ist Mediate J. Brookfield & Son who co pias to hold the Silver
Perpetual Challenge Trophy presented by the British Pteridolo-
gica
say 2 9 Six ong v8 hardy British ding with accent on variation :
M okfie Athyrium filix-femina ‘Capitatum’, A/f.
‘Pritchardit. ‘Ad. ‘Shannan Superbum Druery’, Phyllitis
sco lopendrium iy Ae Fongodle um vulgare ‘Hadwinii’, Poly-
stichum aculeatum ‘Gra
2nd =J. Pye: Athyrium De veniss ‘Capitatum’, A.f. ‘Cruciatum’,
Dryopteris filix-mas, ‘Cristata’, Polystichum setiferum ‘Con-
spicuilobum’, P.s. ‘Cristatum’, P.s. ‘Divisilobum’.
3rd_sO#€B.. Hayhurst: Athyrium filix-feminia ‘Cruciatum’, Dryopteris yet
ccisa’, Osmunda regalis ‘Cristata’ Phyllitis scolopendr
‘Ramo-marginata’, Poly podium vulgare ‘Pulcherrimum’, Poly sti-
chum setiferum ‘Divisilo bum’
cages 10 Three distinct varieties of doctusl cia
essrs Broo ee “Marginata’, ‘Crispa’, ‘Cristata’ :
dnd B. Hayhurst: ‘Albo-variegata’, “ Bist. LOWE 303
that many botanists were at that time = hae pote: regarding
specific distinctness of Polystichum aculeatum and P. setiferum.*
clearly = (v. 13), thatan interspecific hybrid sould be Hose
to be sterile, e did evidently take extreme pains to demonstrate
vans ake the a = of whom experienced the same phot ly
negative result.t Lowe further informs us that this hybridisation
experiment was tee: iene successfully ors only by himself (p. 80),
but also by E. F. Fox and Colonel A. M. Jon es (p. 31).
mm see no seep to doubt the genuine success of Lowe’s experi-
ment. To the best of my knowledge, this is then the first known
sioesful artificial synthesis of an inter- san ve hybrid, and as
such ji remarkable anticipation, by ixty years, of ~
Polysti ara aculeatum x setiferum Suheide coi ed by MAN
( 1950), although the modern hybrids were produced for a totally
different reason, as will be shown in a later paper.
In order to understand the extent of Lowe’s achievement, and
equally to comprehend the reasons for his mistakes, it is necessary to
consider his work in its historical context. The date of his first experi-
ments on the mixed sowing of spores from pairs of fern varieties is
variously stated by him to be 1851 s re and sits (p. 25). oe
date is correct, in all conscience the date of commencement must be
sastied as very early, in view of the vi that the presence of fern ale
sex organs on fern prothalli was not sree until 1848 by Lesczyc-
Suminsky. Previously, in 1844, Nageli had seen spermatozoids
released from a fern prothallus. In 1849 Hofmeister provided an
are nen account of the function of the gametophyte in eg ag
bas his own observations of sexual biology in Pilularia, Salvinia
me Selaginella. His conclusions were confirmed for more typical fern
gametophytes by ayy cre in 1851. (The source of this historical data
Is SACHS (1890) pp. 436-444.) Furthermore the entire span of Lowe’s
work lies fi aa to the rediscovery of Mendelian heredity (1900),
the discovery of the reduction division (1895), an nd the tg effective
h m
s the firm establishment of the chromosome theory of
co Which finally established beyond any doubt that the act of
*Polystichum aculeatum= Aspidium aculeatum: P. setiferum= =A. angular
Is there in this a partial a for the rejection of Lowe’s pda he by the
ry uae Societ ? The genuineness of the distinction between Pobiatican aculeatum
and P. setiferum was cruci is experimen e specimens he sent (p. 63),
specimens. The — ers distinguts hing these two species ‘are never so cleat in
Pressed specimens aioe in the living plants. What was clear enough in
ne s pails at Shirenewton Hall may have been Jess than Sivicin in the rooms
o Linn
tHowever, a M. Jones (Gard. Chron. ser. II. 3:458. ea geagend states that
~~ example given to him by Lowe was not barren, “. . has been proved by
yself and others to be easy of reproduction from spores
304 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
fertilisation consists of the fusion of one ovum with a single sperm.
us we know that cells in the sporophytes of Asplenium viride, for
example, all have 72 chromosomes, but these sporophytes produce,
by means of a reduction division, spores with 36 chromosomes. These
spores germinate to give rise to gametophytes (the cells of which contain
36 chromosomes), which produce spermatozoids and ova which
wo
three, or more spermatozoids participated in the fertilisation of a single
egg. This alternation of stable chromosome numbers, 72— 2,
is consistently maintained through an indefinite number of life-cycles,
thus demonstrating that multiple fertilisation has no place in the normal
pattern of life. Although this type of proof was not available in
Lowe’s time, nevertheless the general principle of one egg, one sperm
was already accepted. However, Lowe (p. 70), writes ““ May we
think the established law that only one sperm acts on the same germ does
not hold good with ferns, . . .”’ (his italics). Why should Lowe have
suggested that the ferns were an exception to this rule? The answer
lies in his own experimental work. Lowe had in many of his hybrid-
isation experiments sown together not just two parent forms, but three,
four, or more, and even eight on one occasion (p. 84), all in the same
pan. He had become convinced, on examination of the progenies of
certain of these experiments, that the morphology of some 0 the
hybrids he produced could only be explained if more than two parents
had participated in their production. In other words, these plants
were of multiple, rather than dual, parentage.
this point on, the argument becomes more complex, because
the outcome of some of his experiments led Lowe to suggest that
multiple parentage might not necessarily originate by the more obvious
means of the multiple fertilisation of a single ovum by two or more
spermatozoids, but instead through several independent fertilisations
of one gametophyte either contributing to the production of a single
sporophyte, or resulting through reciprocal influence in the produc-
tion of two or more sporophytes of identical morphology. This fusing
or mingling effect Lowe termed “ assimilation ’’. Lowe proposed this
hypothesis because he could conceive of no other way of explaining
the outcome of certain hybridisation experiments wherein he obtaine
several examples of the same unique and novel variety from one
prothallus (pp. 37, 64, 84, 102, 115-116). There is no doubt that
Lowe was entirely mistaken in his interpretation, for although one
rothallus can certainly sometimes sustain more than one simultaneous
fertilisation and successfully produce two or more sporophytes (this
as occurred many times in my own hybridisation experiments on
Asplenium), the heredity of these sporophytes is independent.
then can th b
.
ese particular results obtained by Lowe be explained?
°
=
single prothallus can be interpreted as the consequence of the behaviour
of a single genetically unbalanced hybrid zygote in having produced
EGViDS:- FERN HY BRIDISTS Lb) BR LLOWE 305
more than one shoot apex before being able to achieve sufficient co-
ordination for - sib ose sitgoh Se fronds. Behaviour of this kind has
been observed on occasion eeds, as ny result of lowe gene
between seletntae ae compatibie parent In this respect it
significant that concerning var. Tea ; -~ of the forms a
question), Lowe comments (p. 102), that = “. . has taken longer
(1.e. five ser to develop than in most case
The cept of multiple fertilisation: as the normal process in
ferns is totally incompatible with modern prsirerie ct but the possibility
that on rare occasions, more than one spermatozoid can take part in
the fertilisation of a single fern ovum does ist deserve to be eed
with ridicule, or without proper consideration, however much on
instinctively does not believe in it.
Ithough one might expect that the volume of chromosome
studies syne ae ane would enable us to exclude, at least as a shea
consideration of any significance, the possibility of the rare occurrence
of sdcaeera multiple ee curiously this is tte in fact na
f course, in any hybridisation between parents of known chro
number, the chromosome number Sk the progeny is credictabls; eat
hybrid obtained by Dr Janet Emmot oa Mrs So uter), bet =
aealga but rather for the interpretation of the results. The ee
series of experiments described (pp. 30, 87, 92-96, 117-119), started in
1888, concerns four varieties of Phyllitis scolopendrium, namely
Undulatum ’ with a wavy frond, ‘ Spirale’, a dwarf form with a spiral
*Teste Dr A. Braithwaite (Asplenium aethiopicum complex).
306 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
tions of pairs, (series B-G), and all combinations of three varieties
wie = an
e reports that the ones of three of the four pans in me A
were “ enanaie like”’ the par The ‘ Keratoides ° pan * aried
slightly ’? between ‘ Kerato ides” ped its parent, * Victoriae °. in the
case of series B-G, at least a thousand plants were raised, “ all combina-
tions of those sown in the same pan, and no single one was exactly like
either of the parents’’. If Lowe’s iaainabres of the nature of the
progenies from series B-G is accepted at its face value, it is a little
difficult to understand, since in progenies totalling a thousand plants
obtained in this way, one would expect some prothalli to be fertilised
by spermatozoids of the same variety. However, in epee
experiments conducted simply by dense sowing of m
the progeny — will not be a random sample of on. fertilisation
our may well have distorted the composition of the Shad
actually raised by Lowe, so that they did not in fact represent the
portions of — lisations occurring in the population of esd! 32 Fi
ins in the
The oe ac pan is that one which contained prothalli of all four
varieties. Lowe claims that out ofa promeny of between two and three
hundred es four plants “ unmistakably show the character of all
the four par x saw from these four oes and the four parent
varieties pong al illus rated. How should we interpret these experi-
ments? It is easier to epitige at a confident conclusion if we concentrate
our attention on aces similar but simpler experiment (pp. 99-102),
involving the mixed sowing of three varieties of Phyllitis, namely
poi ive’, ‘ Rosabelle’, and ‘Corale’. Conc ncerning the % sii
Lowe writes of one plant, which he named ‘ Desirée’, that it “ con "
clusively shows the projections of ‘ Khedive ’, the crispate character 0
orale’, and the fimbriated margin of ‘ Rosabelle ’.”” (p. 100). He
ontin nues, ““ None of the experiments show multiple parentage more
plainly than this”’. In view of this last statement, it is reasonable to
sirée’ as critical for evaluation of Lowe’s claim to have
obtained oo. of = parentage. Further pare, this is the
only experiment where we are provided with photographic illustrations
of both the hybrid and its postulated parents. It is evident to me that
owe was clearly mistaken in his interpretation of the origin of
*Desirée’, which might readily be cane from the cross ‘ Khedive ’
x *Rosabelle’, and indeed arom o be a more or less perfect
pun tigen between these va
andid assessment of ‘the limited evidence available to us is
that in his anxiety to obtain his desired evidence, Lowe unwittingly
permitted his enthusiasm to dull his faculty for objective judgment,
for it is clear in the case of ‘ Desirée ’, that the fronds illustrated do not
justify the claim made. Therefore, and regretfully, Lowe’s assessment
of his own progenies must be regarded as suspect. Undoubtedly, he
LOVIS : FERN -HYBRIDISTS 45.3: LOWE 307
raised some PaO hybrid forms. Without our modern know-
ledge of the immense power of genetic recombination, it is not really
surprising that sat came to believe that some of the remarkable plants he
raised can only have originated by the participation of more than two
= rents. Indeed, we still cannot be sure that certain of his hybrids
not in fact arise in such a way. Although arguments for multiple
fertilisation as a normal event now have significance only for the
history of biology, the case for the occurrence of iced ie
shape ig as a rare accident remains equally unrefuted and u
The entry of more than one male nucleus ah the egg cell sui clearly
been observed as a rare phenomenon in several different organisms.
What is less certain is how often this act esate in the development of
a polyploid but otherwise normal embryo. (See: AUSTIN, 1965, pp.
109-117). If it were to prove demonstrable, it could only be done by
genetic experimentation in a manner not perhaps in essence very differ-
ent from Lowe’s own experiments, but the marker characters used must
be more different from one another than those used by Lowe, affecting
distinct aspects of the plant, and the manner of their inheritance needs
to be known. Perhaps the most satisfactory demonstration would be
the production of an interspecific hybrid simultaneously involving
on a scale sufficient to allow a nethey will e chance of success (assuming
olog
extreme optimist fori attempting 1
The criticisms in the above appreciation are not intended to carry
any reflection on what was a remarkable achievement, or to detract
from the credit due to an extraordinary pioneer. — "The sheer volume of
experimental work on fern hybridiendon” carried out by Lowe is
pba © and has never yet been exceeded by one person. Thus we
ead (p. 73), that in one period of five years, from 1887-1892, Lowe
fostigetea nearly 500 separate hybridisation experiments. Although
the labour rte ne in instigating a hybridisation attempt by mixed
sowing of spores is much less than that concerned when using more
direct modern methods, the task of cultivation and classification of the
progeny is very much more difficult. Some of the ay — ane
from these sowings were extremely large, amounting o occasio
(p. 100), to two ee eS It would st ie re caaahie
for one man to giv any experiments, including some of a hig
degree of complexity, ‘he attention and critical analysis they would
roperly re
: No ; ono has yet been made of the fact that Lowe’s hybridisa-
tion experiments had a secondary pur which was the production
of new and more magnificent varieties for horticultural purposes and
for exhibition. It is very clear from the account given (see pp. 42-62)
that in this field he was pre-eminently and inordinately successful.
Finally, perhaps the most curious feature of Lowe’s work on fern
hybridisation i is that in spite of its great volume, no records or results
of his work appear to have survived, apart from his ee
Does anyone know the whereabouts of Lowe’s herbarium?
308 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
REFERENCES
AUSTIN, C. R. 1965. Fertilisation, New Jersey: Prentice Hall.
OTT, JANET I. 1964. A cytogenetic investigation in a Phyllitis-Asplenium
complex. New Phytologist 63:
LOWE, E. J. 1868. Thea abnormal forms of ferns. Rep. 37th Meeting Brit. Assoc. Adv.
ci.,
—— 1895. Fern Hite, ifty years’ experience in crossing and cultivation|with a
list o most important varieties and a history of the discovery of multiple
sabehianes te London: Nimmo.
—— 1890. Hybrid ferns and crossed varieties. J. Roy. Hort. Soc. 12: 505- ae
—— 1892a. Facts Digg anes Sheree rod the propagation of ferns. Rep. 61s
Meeting Brit. Ass 7-688.
—— 18925. On fern ind: their multiple Salena Loc. cit.: 688-690.
LOWE, E. J. & JONES, A. M. 1889a. Abnormal ferns, hybrids, ee their parents. Rep.
per Meeting Brit. Assoc, Adv. Sci.: 7
9 i
ga 3: 27-32.
Sere 1. 1950. Problems of cytology and evolution in the Pteridophyta, esi
, F. G. J. VON 1890. History of Botany, 1530-1860 [translated by H. E. F.
Garnsey; revised by I. B. Balfour], Oxford.
REVIEW
AN ANNOTATED BIBLIOGRAPHY OF MEXICAN FERNS by
George Neville Jones, pp. 286, University of Illinois Press, 1966: price
$5
In this bibliography of Mexican Ferns which includes Psilotopsida, el nba i
and spi ges over 1200 references are cited in the Author Index (pp. |-
237). maining pages (238-286) consist of four major ‘t Finding
Indexes” whieh are classified as follows: I GENERAL 1 ibliceraphy. ;
2 Collections, Expeditions etc.; 3 Economic and ash ihwte Botany;
4 Geographical distribution, Floristics; 5 Herbaria; 6 Annotated lists,
F
. : Keys; 8 nc
Illustrations; 9 Vegetation, ecology; 10 Mo rphology Il GEOGRAPHICAL
(29 division of Mexico). III BIOGRAPHICAL (collectors and works on
Mexican seri IV SYSTEMATI AG:
the nanenieions and this book is the result of obvious oor in this field.
are included if Mr Jones has evidence of them being found in Mexico sub-
sequently. Where any new species is described aaa of type specimen pe
where it now resides is given in the annotation
One _ annot review a book like this. ei can say however that it will
duced tyepwritten MS) one would have expected and hoped for a shorter
period between one Acar of the manuscript (1962) and date of publication,
A. C, JERMY
BRIT. FERN GAZ, 9 (8) 1967 309
BOLIVIAN PTERIDOPHYTES COLLECTED BY
WINIFRED BROOKE
J. A. CRABBE
British Museum (Natural History), London, S.W.7
ABSTRACT
A collection of 194 pteridophytes made by Miss W. M. A. Brooke in
Bolivia in 1948/50 is listed, together with an introduction to collectors
in Bolivia and a bibliography which aims to cite all major works dealing
with Bolivian pteridophytes. Twenty-one new nomenclatural combina-
tions (in Adiantum, Eriosorus, Microgramma, Thelypteris, Xiphopteris)
are made, and one Selaginella is in course of publication elsewhere as
a new species.
INTRODUCTION
This account is based on a manuscript left by the late A. H. G. ALSTON
in 1958. I have made several additions and corrections, but the
botany is much as Alston left it. My sole motive in publishing under
my name alone is to simplify the future citation of the new nomen-
clatural combinations. Most of the credit for the naming of the
specimens and for recognizing the sparseness of literature on Bolivian
pteridophytes should be accorded to Alston. In view of differing
Opinions on family and generic affinities, the genera and species are
listed alphabetically. ce
Of previous collections of pteridophytes from Bolivia, those of
Herzog and Stiibel have been completely identified and published,
complete but somewhat inaccurate lists of those made by Bang and
Rusby have appeared, and those made by Buchtien are incompletely
listed. The collections of Bridges, Cuming, Kelly and Pentland lacked
localities. The fine collections of Mandon and D’Orbigny were never
published as enumerations. Other collectors in Bolivia have been
Pearce, Mexia, Fries, Fiebrig, Tate, Pflanz, Gunther, Williams, Weddell
and Cardenas. Specimens from some of the Bolivian collections have
been quoted in the works of Fée, Hooker, Christ, Christensen and
Maxon.
Miss Brooke was encouraged to make her trip by Dr John Rams-
bottom, then Keeper of Botany at the British Museum (Natural History)
and she was in Bolivia from December 1948 to November 1950. The
places she visited are given on the map (page 310) drawn by Joyce
Parker, and a descriptive account is in BROOKE (1952); field numbers
3002-6949 were collected in Bolivia, after which she went to Chile
(numbers 6950-7086); field notes, determination lists and seed lists
are at the British Museum (Natural History). In addition to collecting
seeds for seed shareholders, Miss Brooke prepared herbarium specimens
which are now deposited in the following institutions (in order of size
of set): London (BM) [just over 2000 numbers], Chicago (F), New
York (NY), Utrecht (U), Geneva (G), Stockholm (S), Washington
(US). The specimens listed below are those in BM;; attention should
BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
BOLIVIA cd
and the collecting localities of
Winifred Brooke
ae”
o
gah ttstees, .'
oe
#008" heneeay
. +
f ogeet test
.
oO
ucre 14, Tacacoma 1, Tako Tako 13, Tana
, paca 7, Taquina 11, Tres Cruces Pass 7,
Tunari 8, Ulmapalca 3, Unduavi 5, Villa Eufronio aeee 13, Viloco 7, Zudanez 15.
CRABBE: W. BROOKE’S BOLIVIAN FERNS a £3
be drawn to the occurrence of mixed gatherings.
NUMERATION OF SPECIES
In order to reduce the printed matter, geographical details are listed
here at the beginning; under the species, localities are listed in alpha-
betical order, together with information from Miss Brooke’s plant
labels.
LOCALITY (Department, Province) latitude; longitude
AGUAS CALIENTES (Cochabamba, Arque) 17.488; 66.37W
AGUAS DE CASTILLAS (Oruro, Cercado), erg ‘of Oruro
CARACOLES (La Paz, Inquisivi) 16.598; 67.24W
yo (La Paz, Nor Yungas) 5-10 miles down the valley from
CAXATA, (La Paz, Inquisivi) 17.078; 67.20W
CHALLANA (La Paz, Larecaja) 15. 57S: 68.12W
CHORETI (Santa Cruz, Cordillera) 20. O1S; 63.32W
CHORO (La Paz, Nor Yungas) 16.178; 67. 40W
CHULUMANI (La Paz, Sud Yungas) 16. 24S ; 67.31W
GRAN tee (La Paz, Larecaja), ’gold- mine penne Sorata and
Taca
HEREDIA ‘chamsenes Azero), 19.488; 64.08W
HUATARATA (La Paz, Omasuyos) 16. 10S; 68. ese)
INCACHACA (Cochabamba, Chaparé) 17. 148; 65.4
LA FABULOSA (La Paz, Larecaja), tin-mine at ead oF valley NE of
Challana
LANGUNILLAS (Santa Cruz, Cordillera) 19.38S; 63.43W
LOCOTAL (Cochabamba, Chapa ré) 17.118; 65. 48W
LURUNI (Cochabamba, A sort ang hot springs near Tunari
MIZQUE (Cochabamba, Mizque) 17.56S; 65.19W
ORURO (Oruro, Cercado) 17.598; 67.09W
PAMPAS Le po ‘pena Cruz, Cercado), an estate about 12 miles
from Santa Cru
PONGO (La Paz, Inquisivi) 17.038; 67.17W
QUIME (La Paz, Inquisivi) 17.02 ; 67. 1SW
viteco (La Paz, Loayza) 16.578; 67.33W
£2 BRITISH FERN GAZETTE: VOLUME: 9 \PART. 8 (GE967)
Adiantum baenitzii Rosenst. CHORO, 2700m, on river bank, 23 Jan
50, 6058, UNDUAVI, 1800m, on damp slope above road in valley
below town, 15 Oct 50, "6882,
A. ne ee cs Presl. pn 2700m, on shady bank above river,
Feb 50, 6 QUIME, 2490m, on wet bank under trees, 4 Apr 49,
5429 Feith rey A. poiretii]. SORATA, 2550m, in shady lane in
damp thicket, 31 May 50, 6416.
A. orbignyanum Mett. ex Kuhn. QUIME, 2400m, in sheltered gorge,
13 Apr 49, 5487; 2550m, in rock in gorge, 13 May 49, 5490. VILA
VILA, 2100m, in crevices of rocks above river, 17 Dec 49, 5859.
A. pilosum Fée. CHORO, 2850m, on steep grassy slope in valley,
21 Jan 50, 6025. LURUNI, 2790m, in plantation, 10 Sep 50, 6791.
QUIME, 2400m, in wet plantation sabe river, 16 Mar 49, 5301.
A. poiretii Wikstr. var. poiretii. QUIME, 2490m, on wet bank under
trees, 4 Apr 49, 5429A [with 5429 A. Disviohall:
A. poiretii var. glaucescens (Mett. ex Kuhn) Crabbe comb. nov.
crenatum Poir. var. ieee Mett. ex smh in Abh. Naturf. Ges.
Halle 11:29.1869). UNDUAVI, 1800m, on damp roadside bank in
valley below town, 6883A [ an 6883 A. williamsii},
A. tinctum T. Moore. ROSAL, 2010m, on rocks in glen, 30 Sep 49,
5725.
A. williamsii T. Moore. CHORO, 2700m, under trees, 16 Mar 50,
6272. UNDUAVI, 1800m, on damp roadside bank in valley below
town, 15 Oct 50, 6883 [with 6883A A. poiretti var. glaucescens].
Asplenium auritum Swartz. INCACHACA, 3000m, 22 Aug 50, 6784.
A. bangii Hieron. ROSAL, 1950m, in damp shady glen, 3 Oct 49,
5740.
A. castaneum Schlecht. & Cham. QUIME, 2850, beside path, 4 Apr
49, 5432A. VILOCO, 4440m, in rock crevices on wet slope, 18 Mar
49, 5318.
A. davallioides Hook. CARMEN, 2400m, on trunks of trees in warm
forest, 11 Feb 50, 6129.
A, foeniculaceum Kunth. CARMEN, 2400m, on trunk of tree in
warm forest, 11 Feb 50, 6132A. GRAN PODER, 3000m, in dense
forest, Jun 50, 6295A; 3000m, on shady bank, Jun 50, 6438.
A. gilliesii Hook. AGUAS DE CASTILLAS, 3690m, in damp crevices
of rock in dry district, 1 Mar 49, 5251. LAGUNILLAS, 3000m, in
wet forest, 7 Mar 50, 6276.
A. oe Kunze. GRAN PODER, 3000m, on shady bank, 3 Jun
nay Mead INCACHACA, 2400m, on tree in forest, 13 Aug. 50,
A. lorentzii Hieron. SORATA, 2250m, on river bank, 1 Jun 50, 6415.
QUIME, 2700m, on shady bank, 4 Apr 49, 5430.
A. monanthes L. SORATA, 2400m, on bank of stream, 30 May 50,
CRABBE: W. BROOKE’S BOLIVIAN FERNS 313
6407. QUIME, 2400m, on wet rocks in sheltered gorge, 13 Apr 49,
5488; 2400m, between rocks by river in gorge, 13 Apr 49, 5491.
A, etait Swartz. COROICO, 1500m, 5 Jun 50, 6646. QUIME,
3 Apr 49, 5409; 2400m, between boulders by river in sheltered
aries, 13 Apr 49, 5493.
A. rhizophorum L. CHULUMANI, 2100m, in dense wet forest, 29
Jun 50, 6576. INCACHACA, 2400m, in forest, 17 Aug 50, 6715.
A. squamosum L. CARMEN, 2400m, in warm forest, 11 Feb 50, 6125.
A. woodwardioideum Gardn. GRAN PODER, 3300m, 9 Jun 50, 6483.
Azolla filiculoides Lam. CHORETI, 858m, on still water in backwater
of river, 10 Sep 49, 5607. HUATARATA, 3840m, on shallow
water at edge of Lake Titicaca, 19 May 50, 6376. MIZQUE,
1800m, on pool in river, 21 Dec 49, 5902.
Blechnum sie ie (Baker) C. Chr. INCACHACA, 3300m, on wet
rock, 15 Aug 50, 6721.
B. estar Rosenst. GRAN PODER, 3000m, on wet grassy slope,
3 Jun 50, 6445. LAGUNILLAS, 3000m, on top of cliff, 7 Mar 50,
6153.
B. ieee. os gs INCACHACA, 3300m, on roadside bank,
15 Aug 50, 6
B. fink : iF lee 2250m, on river bank, | Jun 50, 6412.
Botrychium schaffneri Underw. CHORO, 3150m, on path through
pasture, 19 Feb 50, 6135A.
Campyloneurum aglaolepis (Alston) Sota. CHORO, 3090m, on shady
bank, 18 Jan 50, 6005. QUIME, 2400m, in crevices of rocks on far
side of river, 5 Apr :
C. lorentzii (Hieron.) Ching. ROSAL, 1950m, at edge of glen, 30
ep 49, 5716.
Cheilanthes marginata Kunth. QUIME, 2550m, common on shady
wet rock wall, 31 Mar 49, 5402.
C. myriophylla Desv. Seagal 2550m, common on shady wet rock
wall, 31 Mar 49,
C. pilosa Goldm. LA eeuiniues 4500m, in rock crevices, 26 Apr
50, 6323. ayn PONGO and QUIME, 2490m, below rock beside
road, 7 Apr
C. pruinata abe AGUAS DE CASTILLAS, 3900m, in rock crevices
on dry sunny hilltop, 8 Mar 49, 5260.
6 Ag sale fragilis (L.) Bernh. CHORO, 2850m, on dripping rock
by stream, 23 Jan 50, 6056. QUIME, 2550m, on dripping rock in
pce 3 Apr 5489
Dennstaedtia glauca (Cav.) C. Chr. QUIME, 2400m, on wet ground
beside river in a side valley, 5 Apr 49, 5426.
Elaphoglossum aff. accedens (Mett.) Christ, probably undescribed.
314 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
CARMEN, 2400m, hanging from tree, 11 Feb 50, 6227. Kew have
a specimen from Colombia (André 2362) which is similar but has
shorter scales on the under surface. The type-specimen of Acrosti-
eee —" Mett. Mandon 8-bis, (BD; photo BM) is from
Bolivi
E. angustisimum ee - Chr. INCACHACA, 2400m, on rock in
open, 13 Aug 50
E. crassipes (Herds: : si QUIME, 2400m, among rocks in sheltered
orge, 9 Apr 49, 5463 [with 5463A E. leprosum); 3300m, on rock,
9 Apr 49, 5465.
E. ec, sae Christ. GRAN PODER, 3000m, on steep grassy
slo Jun 50, 6447. UNDUAVI, 3120m, among shady rocks,
13 Oct 0, 6842 (with 6842 E. lindenii\.
E. te (Mett.) Christ. QUIME, 2400m, among rocks in sheltered
gorge, 9 Apr 49, 5463A [with 5463 E. crassipes].
E. paid (Bory) T. Moore. INCACHACA, 2400m, on rock, 13
Aug 50, 6674. UNDUAVI, 3150m, among rocks in ‘cloud district,
3 Jul 50, 6621; 3120m, among shady rocks, 13 Oct 50, 6842 [with
68424 E. engelii].
E. vag cng Christ. LAGUNILLAS, 3000m, in wet forest,
7 Mar
E. rested ea T. Moore. CHORO, 3150m, on rocks near stream
on open slope, 18 Jan 50, 5992A with "5992 E. unduavense].
ULMAPALCA, 4050m, by river, 27 Apr 50, 6337.
E. mettenii (Kuhn) Christ. UNDUAVI, 3150m, on rocks in cloud
district, 3 July 50, 6620. A specimen from Ecuador (Jameson 315)
at Kew is rather larger and stouter with paler scales and frond a
little more cuneate at base.
E. unduavense Rosenst. CHORO, 3150m, on rocks near stream on
open slope, 18 Jan 50, 5992 ee 5992A E. mathewsii|.
—— gg Kunth. QUIME, 2400m, on wet ground near
9, 5440. SORATA, 2400m, on wet shady bank of
sr 30 Say 50, 6402
Eriosorus elongatus (Hook. & Grev.) Copel. LAGUNILLAS, 2850m,
on tree, 6 Mar 50, 6196A [with 6/96 Jamesonia scalaris and 6196B
Ri peruvian a|\.*
E. mathewsii (Hook.) Crabbe comb. nov. (Gymnogramma mathewsii
Ho p. Fil. 5:128.1864) INCACHACA, 3300m, on wet bank of
road, 15 y Hel 50, 6725.
E. orbignyanus (Mett.) A. F. Tryon. INCACHACA, 2400m, in damp
tangles on slope in ravine, 19 Aug 50, 6768.
*6196 is a mixture at BM. Alston had split it into 6196 Jamesonia scalaris and
6196A Eriosorus elongatus. 1 have se separated 6/96B as Jamesonia peruviana.
A. E. Tryon (Contrib. Gray Herb. Hary. Univ. 191: 168, 178. 1962) cites
6196 at U as J. per uviana, and at US under both J. peruviana and J. scalaris.
CRABBE: W. BROOKE’S BOLIVIAN FERNS 315
E. retrofractus (Kunze ex a Crabbe comb. nov. (Gymnogramma
retrofracta Kunze ex Mett., Fil. Lechl. 1:10.1856). UNDUAVI,
3000m, on shady roadside peak 1 Jul 50, 6598.
E. hi ey (Fée) A. F. Tryon. LA FABULOSA, 4500m, hanging
mp shady rocks, 30 Apr 50, 6343. This seems to be a well
Getclenad example of this fore which was described from smaller
plants from Peru: Lechler 2255.
Gleichenia bifida (Willd.) Spreng. CHULUMANIT, 2100m, on banks
above road at edge of forest, 18 Jun 50, 6508. COROICO, 1440m,
on open sunny slope, 5 Jul 50, 6632.
Hymenophyllum abietinum Hook. & Grev. LAGUNILLAS, 3000m,
on trees in forest, 7 Mar 50, 6165. UNDUAVI, 3150m, on wet bank
in deep shade in cloud district, 3 Jul 50, 6617.
H. elegantulum Bosch. GRAN PODER, 2850m, on wet mossy bank,
n 50, 9. nerseatin eget 3150m, on wet bank in deep shade,
Jul 50, 6618; 3060m, wet rock in pugs district, 15 Oct 50,
pian (with 6894 H. apeiliaaiclt and 6894A. H. myriocarpum).
H. ferax Bosch. UNDUAVI, 3060m, under wet rock, 15 Oct 50,
6893 [with 6893A H. speciosum).
H. a venides Swartz. GRAN PODER, 3600m, by spring in grotto,
n 50, 6473. INCACHACA, 3300m, on wet rock in glen, 15
ce "30, 6727.
H. myriocarpum Hook. CARMEN, 2400m, in moss on trunks of
trees in warm forest, 11 Feb 50, 6130. GRAN PODER, 3600m, on
wet dark bank of stream, 10 Jun 50, 6471. INCACHACA, 3300m,
on wet rock in glen, 15 Aug 50, 6726. LAGUNILLAS, 3000m, on
trees in wet forest, 7 Mar 50, 6190B. UNDUAVI, 3060m, on wet
rock in cloud district, 6894A [with 6894 H. speciosum and 6824B
H. elegantulum|.
H. pyramidatum Desv. INCACHACA, 2400m, on bank in dense
forest, 14 Aug 50, 6689; 2700m, on tree, 15 Aug 50, 6746
H. secundum Hook. & Grev. UNDUAVI, 3150m, on ius rocks in
misty district, 3 Jul 50, 6619; 3060m, on wet shady rocks, 13 Oct 50,
6831.
H. speciosum Bosch. CARMEN, 1800m, hanging from branch of
tee) in warm forest, 11 Feb 50, 6131. UND eet, 3060m, unger
wet rock, 15 Oct 50, 6893A (with 6893 H. ferax); m, on wet
rock in cloud district, 15 Oct 50, 6894 [with 6894A i dijribontouh
and 6893B H. een
LAGUNILLAS, 2850m, on trees,
Jamesonia peruviana A.
6 Mar 50, 6196B [with 6 196 7. oaierts and6196A Eriosorus elongatus).
J. scalaris Kunze GUNILLAS, 2850m, on trees, 6 Mar 50,
6196 {with 6/ 96A Eriosorus elongatus and 6196B Jamesonia peruy-
iana\.
Lycopodium cernuum L, EL PARADISO, 2100m, by path through
316 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
swamp in dense cloud aye “ ee 50, 6569. HEREDIA, 1200m,
on rocks in gorge, 22 Sep
L. clavatum L. CHORO, nein near stream in glen, 18 Jan 50, 6015.
GRAN PODER, 3000m, on wet shady bank, 4 May 50, 6424.
INCACHACA, 2700m, on = by road in forest, 15 Aug 50,
6720. UNDUAVI, 3000m, on dam mp slopes in partial shade in
cloud district, 1 Jul 50, 65974 with 6597 L. jussiaei and 6597B L.
reflexum|.
L. complanatum L. CHULUMANI, 2100m, hanging down bank in
wet forest, 18 Jun 50, 6511. INCACHACA, 3000m, on damp bank
in forest, 24 Aug 50, 6781.
L. cruentum Spring. CHORO, 4050m, in marsh around a spring, 24
Feb 50, 6260.
L. jussiaei Desv. meliscayetet geht -igties under bushes on slope, 25
Jun 50, 6562. UNDUAVI, 3 n dam mp slopes in partial shade
in cloud nom 1 Jul 50, 6597 are 65974 L. clavatum and 6597B
L. reflexum); : 2850m, on wet banks beside road, 3 Jul 50, 6611.
L. lechleri Hieron. GRAN PODER, 3150m, on bushy slope beside
river, 10 Jun 50, 6492.
: magellanicum Swartz. CHORO, 3750m, 6 Feb 50, 6118.
mettenii Hieron. CARMEN, 2400m, hanging from tree in warm
ger sien 11 Feb 50, 6132.
L. reflexum Lam. Between COROICO and UNDUAVI, 3150m,
on open slope by road, 16 Oct 50, 6907. UNDUAVI, 3000m, on
damp slopes i in partial shade in cloud district, 1 Jul 50, 6597B [with
6597 L. jussiaei and 6597A L. clavatum).
L. saururus Lam. CHORO, 3900m, on wet open rocky slope, 3 Feb 50,
6095. LA FABULOSA, 4650m, on open slope, | May 50, 6347.
Microgramma chrysolepis (Hook.) Crabbe comb. nov. (Polypodium
chrysolepis Hook., Ic.P1.8 :t.721.1844). CHORO, 3000m, on tree and
rock, 27 Jan 50, 6081.
Bs —. (Desy.) Crabbe comb. nov. (Polypodium ee Desv.
. Linn. Paris 6:226.1827). INCACHACA, 2400m,
tibiae up bush at edge of forest, 12 Aug 50, $66 61. Between
PONGO and QUIME, 3150m, encrusted on wet shady rock by
road, 7 Apr 49, 5456.
M. galatheae (C. Chr.) Crabbe comb. nov. (Po/ypodium galatheae C.
Car. in Bot. Tidsskr. 25:79.1902). PAMPAS VERDES, 4500m, on
tree, 11 Nov 49, 5786.
M. piloselloides ) Copel. UNDUAVI, 3150m, on wet shady bank,
3 Jul 50, 663
M. squamulosa iia: ) Sota. QUIME, 1500m, hanging down over
rocks in subtropical gorge across bridge far below town, 21 Apr 49,
ei OSAL, 2010m, among roots of tree in shade, 28 Sep 49,
CRABBE; W. BROOOKE’S BOLIVIAN FERNS si7
saree pendula (Raddi) J. Sm. CHULUMANI, 2100m, in dense
, 18 Jun 50, 6597; 1800m, on damp shady bank, 24 Jun 50,
Vtahane aurea (Poir.) Desv. QUIME, 2850m, beside path, 4 Apr
49, 5432. ROSAL, 2010m, on shady bank by road, 30 Sep 49,
5727.
N. reenter Rosenst. QUIME, 2400m, on bank near river, 5 Apr 49,
a nivea (Poir.) eg EP AGUAS DE CASTILLAS, 3600m, in rock
crevice, | Mar 49,
Ophioglossum Adbssial Walt. CONA CONA, 4200m, in short
grass in marsh, 8 Feb 49, 5219. TRES CRUCES, 4800m, in short
wet grass, 11 Apr 49, 5272.
Pellaea ovata (Desv.) Weatherby. QUIME, 1800m, on rocks and banks
in hot wet gorge across bridge far below town, 21 Apr 49, 5509.
P. tenera (Gill.) Prantl. AGUAS CALIENTES, 3420m, in wet rock
crevices, 30 Jan 49, 5183.
Fe, Le One (Cav.) Link. AGUAS DE CASTILLAS, in rock crevices
p of hill, 27 Feb 49, 5229.
=. crassifolia (L.) Underw. & Maxon. QUIME, 2400m,
under boulders near river, 5 Apr 49, 5436; 2400m, between rocks in
Sheltered gorge, 5 Apr 49, 5500. ROSAL, 1890m, on rock by
stream in dark glen, 3 ‘Oct 49, 5736.
Pityrogramma austro-americana Domin. CHULUMANI, 1800m, on
shady roadside bank, 23 Jun 50, 6545 | with 6545A P.calomelanos}.
P. calomelanos (L.) Link. CHULUMANI, 1800m, on shady roadside
bank, 23 Jun 50, 6545A [with 6545 P. austro-americana).
Polypodium bryopodum Maxon. QUIME, 2400m, on rock near river,
5 Apr 49, 5442.
P. crystalloneuron Rosenst. CARMEN, 2400m, on trees in forest,
11 Feb 50, 6275.
P. filicula Kaulf. ROSAL, 1890m, on damp rocks in shady glen, 3 Oct
49, 5739.
P. lomariiforme Kunze. On short cut to PONGO from QUIME,
50m, between rocks, 7 Apr 49, .
P. loriceum L. CHULUMANI, 2100m, in dense damp forest, 29 Jun
50, 6568.
P. semipinnatifidum (Fée) Mett. INCACHACA, 2400m, in partial
shade at edge of forest above river, 13 Aug 50, 6687.
P. sessilifolium Desy. CHORO, c.3300m, on trees, 21 Jan 50, 6032.
Polystichum polyphyllum — CARACOLES, 3900m, among
boulders by river, 23 Apr 49, 5517. LA FABULOSA, 4500m, in
rock crevice, 26 Apr 50, 6366 6307; 4500m, in damp rock cave,
30 Apr 50 6342.
318 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
P. yungense Rosenst. COLOMI, 3600m, on marshy grass slopes, 7
Jan 49, 5095 [with 5905A Ti helypteris pavoniana|.
Selaginella cruegeri Jonman. LAGUNILLAS, 3000m, in wet wood, 7
Mar 50, 6256.
S. macrophylla A. Braun. About 5 miles along road from INCA-
CHACA to LOCOTAL, 1800m, beside forest road, 20 Aug 50, 6765.
S. marginata (Humb. & Bonpl.) Spring. SANTA CRUZ, 4500m,
covering ground near river in forest, 25 Nov 49, 5
S. microphylla (Kunth) Spring. SORATA, 2400m, Jun 50, 6649.
S. novae-hollandiae ee: ee INCACHACA, 2400m, under
rock in ravine, 13 Aug 50, 6673.
S. trisulcata Asplund. es helian tt 2100m, in wet glen in cloud
district, 22 Jun 50, 6531. INCACHACA, 2400m, beside path in wet
gorge, 12 Aug 50, 6659.
S. sp.nov.aff. S. popayanensis Hieron. INCACHACA, 3000m, 22 Aug
50, 6783.
Thelypteris pavoniana (Klotzsch) Crabbe comb. nov. (Po/ypodium
pavonianum Klotzsch in Linnaea 20:386.1847). COLOMI, 3600m,
on marshy grass slopes, 7 Jan 49, 5095A [with 5095 Polystichum
yungense |.
Trichomanes tenerum Spreng. ROSAL, 1920m, close to stream in glen,
30 Sep 49, 5729.
Trismeria trifoliata (L.) Diels var. aurea Fée. Twelve miles below VILA
, 2040m, in wet sunny ground, 25 Mar 50, 6223.
Woodsia crenata (Kunze) Hieron. CHORO, 3300m, hanging down
wet rock in shady glen, 30 Jan 50, 6091. LA FABULOSA, 4500m,
- Seed rock crevices, 28 Apr 50, 6326. QUIME, 3000m, on rocks
n mountain slope, | Apr 49, 5409A. VILOCO, 4500m, in rock
crevinss. 20 Mar 49, 5344: 4350m, in rock crevices, 27 Mar 49,
Xiphopteris allosuroides (Rosenst.) Crabbe comb. nov. (Polypodium
allosuroides Rosenst. in Meded. Rijks-Herb. 19:16.1913). LA
FABULOSA, 4500m, under rocks in shade, 2 May 50, 6371.
X. aromatica (Maxon) Crabbe comb. nov. (Polypodium aromaticum
axon in Proc. U.S. Nat. Mus. 27:743.1904). CHORO, 3000m,
on tree, 30 Jan 50, 6088; 6089A [with 6089 X. subscabra|.
be a (Desv.) Crabbe comb. nov. (Polypodium capillare Desv. in
Berl. Mag. 5:316.1811). INCACHACA, 3300m, on damp roadside
bank, 15 Aug 50, 6731.
X. choquetangensis (Rosenst.) Crabbe comb. noy. (Polypodium choque-
tangense Rosenst. in Meded. Rijks-Herb. 19:18.1913). CHORO,
3300m, hanging down wet rock in wooded glen, 2 Feb 50, 6090.
X. heteromorpha* var. dependens (Baker) Crabbe comb. nov. (Poly-
*Xiphopteris heteromorpha (Hook. & Grev.) Crabbe comb. nov. Polypodium hetero-
morphum Hook. & Grev., Ic. Fil. 1: t. 108. : 1829). er
CRABBE: W. BROOKE’S BOLIVIAN FERNS 319
poet dependens Baker in Hook. & Baker, Syn. Fil.: 335.1867).
GRAN PODER, 3600m, in wet dark grotto, 10 Jun 50, 6472.
UNDUAVI. 3150m, in deep shade on wet bank, 3 Jul 50, 6615.
X. lanigera (Desv.) Crabbe comb. nov. (Polypodium lanigerum Desv.
in Berl. Mag. 5:316.1811). About 5 miles along road from INCA-
CHACA to LOCOTAL, 1800m, on tree in forest, 20 Aug 50, 6762.
X. moniliformis (Lag. ex Swartz) Crabbe comb. noy. (Polypodium
moniliforme Lag. ex Swartz, Syn. Fil.: 33.1806). CHORO, 3000m
in forest, 2 Feb 50, 6278. GRAN PODER, 3000m, on wet bank,
10 Jun 89,
X. sareieukeiia (Mett.) Crabbe comb. noy. (Polypodium ran Pec
INCAC
coe Fil. Lechl. 1:6.1856). CHACA, 3600m
wet rock on damp roadside bank in forest, 15 Nog 50, Pon.
gf cre (Desv.) Crabbe comb. nov. (Polypodium peruvianum
esv. in Mém. Soc. Linn. Paris 6:231.1827). QUIME, 2700m, on
wet bank 5 Apr 49, 5428. VILOCO, 4440m, in rock crevices on
wet slope, 18 Mar 49, 5316.
X. pilosissima (Martens & Gal.) Crabbe comb. cna (Polypodium
gaa Martens & Galeotti in Mém. Acad. Brux. 15:39.t.9. f.2
1842). About 5 miles along road from INCACHACA toLOCOTAL,
1800m, on tree in forest, 20 Aug 50, 6761.
X. pozuzoensis (Baker) Crabbe comb. noy. (Polypodium pozuzoense
Baker in Hook., Ic. Pl. 17:t.1672.1886). CARMEN, 2400m, hanging
from tree in warm forest, 11 Feb 50, 61
X. — (Christ & Rosenst.) Crabbe Sil, noy. (Polypodium sodiroi
Christ & ee in Repert. Nov. Spec. Regni Veg. 5:14.1908)
ba? FAB OSA, 4500m, in shady rock crevices, 28 Apr 50, 6325.
X. eels ‘ Klotzsch) Crabbe comb. nov. tar, ngosie
Klotzsch in Linnaea 20:377.1847). CHORO, 3000m, on tree, 30
Jan 50, 6089 [with 6089A X. aromatica).
X yee (Rosenst.) Crabbe comb. noy. (Polypodium yungense
osenst. in Repert. Nov. Spec. Regni Veg. 5:236.1908). LAGUN-
ILLAS, 3000m. on trees in wet forest, 7 Mar 50, 6166.
IBLIOGRAPHY OF BOLIVIAN FERNS
ciedins general works like those of Fée and Hoo
ker)
BAKER, J. G., 1901. Decades Kewenses 349 (Polypodium bangii). Kew Bull. 1901:
BENEDICT, R. C. A revision of the genus Vittaria J. E. Smith. Bull. Torrey
Bot.
914.
Club 41: 391-410.
BERGDOLT, E. 1933. Pteridophytenstudien, I: Die heterophyllen Trichomanes-Arten.
Flo fora, Jena 127: 251- Fl
BRAUSE, Hag ycopodiaceae, Polypodiaceae in J. PERKINS, Beitrage zur Flora
49:17
n Bolivia. aps ai 9-180.
BRITTON, Py G. 1888. An enumeration of the plants collected by Dr H. H. Rusby in
2S uth America 18851886, IL: Prssdophyta. Bull. Torrey Bot. Club 15:
oe 1895, ‘Filices, Lycopodiaceae in H. H. RUSBY, On the collections of Mr
Miguel Bang in nape Il. Mem. Torrey Bor. Club 4: 271-273.
320 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
—— 1896. rire’ in Idem, I Op. cit. 6: 128-129
BROOKE, W. M. A. 1952. Some Boligas plants. Proc. "§. Lond. Ent. Nat. Hist. Soc.
1951 3: 137-143.
BUCHTIEN, O. 1910. ous peiggye a la flora de Bolivia |Filices pp. 5-59}, La Paz
CHRISTENSEN, C. & ASPLUND, E. 1926. Pteridophyta in ie We Contributions to
e flora of the 1 Bolivian Andes. Ark. Bot an F
ELAND, E. B. 1941. Tropical American ferns. Univ. C ali if P Pal Bot. 19: 287-340.
FOSTER, R. C. 1958. ‘A catalonte of the ferns and flowering plants of Bolivia. Contrib.
Gray Herb. ld Univ. 184: 1-223.
GILBERT, B. D. 1897. © new Bolivian ferns of the Miguel Bang collection. Bull.
Torrey Bot. Club 24: 258-259.
—— 1907. Filices in H. H. RUSBy, An enumeration of the plants collected in
Bolivia by Miguel Bang, 4. Ben New York Bot. a ae 4: 465-467.
HERZOG, T. 1915. Lycopodiaceae i ie von Dr zog auf seiner zweiten
Proee durch Bolivian in ‘den Jahren 1910-1911 perio ass Pflanzen, II.
Meded. Rijks-Herb. 27: 1-2.
cache. G. 1896. Beitrage zur Kenntnis der Pteridophyten-Flora der Argentina
einiger angrenzender Teile von Uruguay, Paraguay und Bolivien.
Bot. Jb. 22: 359-420.
—— 1906; 1907; 1908; 1909. Plantae Stiibelianae, Pteridophyta, von Dr Alphons
Stiibel auf seinen Reisen nach Siid-amerika, besonders in Columbien,
Ecuador, Peru und Bolivien gesammelte Pteridophyten (Gefasskrypto-
gamen), 1; Il; UI; IV. Hedwigia 45: 215-238; 46: 322-364; 47: 204-249;
48: 215-303.
—— 1913. In G. BRAusE, Polypodiaceae, in J. PERKINS, Beitrage zur Flora von
Bolivia. Bot. Jb. 49: 179- ev
—— 1915. Selaginellaceae in Die von Dr Th. Herzog auf seiner zweiten Reise
durch Bolivien in pte satieo. 1910-1911 gesammelten Pflanzen, Il. Meded.
Rijks-He ek isi. ;
MAXON, W. R. ORTON, C. Vv. 1939. New ferns from Bolivia and Peru. Bull.
Torrey hor: Club 66: 39-45.
PERKINS, J. 1913. Equisetaceae in PERKINS, Beitrige zur Flora von Bolivia. Bor. Jb.
ROSENSTOCK. E. 1908; 1909; 1911; 1912: 1913; 1928. Filices novae a Dre O. Buchtien
n Bolivia collectae, ae = Ill; IV; V; VI. Repert. Nov. Spec. Regni Veg.
8- : 56-64.
5: 228-239; 6: 308— : 342- 344; ll: 53-60; 12: 468-477; 25: 5
—— 1908. Filices novae iv ye rt. bee ‘Spec. Regni Veg. 6: 175-176
—— 1913. Filicales, Isoétaceae in Die n Dr Th. Herzog auf seiner zweiten Reise
~ Bolivien in den Jahren 1910 u. 1911 pesistehelien Pflanzen. Meded.
Rijks-Herb. 19: 3-26.
RUSBY, H. H. 1895; ee Marsileaceae, Selaginellaceae in RUSBY, On the collections
f guel Bang in Bolivia, Il; Il. Mem. Torrey Bot. Club 4: 271;
eet AOE, Equ taceae, Lycopodiaceae in rusBy, An enumeration of a
plants collected i in Bolivia by Miguel Bang, 4. Bull. New York Bot. Gar
WAGNER, Ww. H. 1954. A Bolivian Elaphoglossum [E. cardenasii] of unique leaf
Pace ay Bull. Torrey Bot. Club. 81: 61-67. pees ee
BOOK NOTE
THE bg Sheba ENCYCLOPEDIA OF PLANTS AND FLOWERS,
London: Paul Hamlyn, New York: Crown Publishers, 1966. This book is
deployed around. F. A. Novak’s marvellous collection of 1120 photographs.
c
cS
Ae,
a.)
a
= |
&
ory they
through the plant kindom, from bacteria to Ses o
world of its own. The ferns and fern-allies are iftuatrated by 27 species, 1
of which are well-known foreigners, the remaining 16 occu rring wild in the
British Isles. At 25 dilaes this is wondechia value, as also is the companion
volume on the kingdom
J. A. CRABBE
BRIT. FERN ‘GAZ,.'9'(8) 1967 321
A CONTRIBUTION TO THE CYTOTAXONOMY
OF ASPLENIUM MAJORICUM
ANNE SLEEP
Botany Department, University of Leeds
Asplenium majoricum Litard. is a rare plant, probably endemic to the
island of Mallorca*, where it pie only in a few localities in the
mountains bordering the north-west coast. It is a small fern of rock
crevices, and was first described by Cores (191 1) from walls near
the town of Séller. It is still found in this neighbourhood, at eh
from 100-350 + although h elsewhere it is known to occur u
groves; it prefers light shade and northern - e. It is usually
accompanied by much A. iy asa! L. subsp. quadrivalens D. E.
New growth s
towards the en e # September, and ss kaos actively cating the
winter and earl
general similarity in appearance between A. majoricum and
some forms of A. fontanum (L.) Bernh. has been pointed out by several
authors, Ey LITARDIERE (1911), while others have stressed a
These oy taxa are all included in
t A. majoricum, at that time
*Regarding the possible occurrence of A. majoricum outside seer ig two possibly
relevant reports from the mainland of Spain are given by artinez (in
MARTINEZ, M. 1934. Mem. Soc. Esp. Hist. Nat. 14: 421 and 425-6) from the a) ond
: pe be
KNOCHE in 1921 (flora Balearica 1: 095) tice the » codibsination A. lanceolatum Huds
nd
Thi or does however state that the Mallorcan plant is not hairy and from his
description he is undoubtedly referring to the plant ibed by de Litardiére as
A yim be: ; g erent plants are apparently involved, the rata ue
on rian peninsula is doubtful, and until fu i i
A,
available it is s perhaps best to regard A. majoricum as probably endemic ' to the island
Mallor
322 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
F
‘P
ah ait
a>, 1"
wi i
a Vey
ES, fs
Cc
Me eden 1 Silhouettes (natural size) of fertile fronds from plants in cultivation
t Leeds: — majoricum (4x) from Sdller, Mallorca. B) Synthesize
triploid hybrid A. fontanum3 x A. majoricun? in its second year mel growth. C) A.
pr seal (2x) from Isére, France
the cytology of = amajoria ies and showed it to be a Pore
e
n= 72. Discussion of its possible origin was however wit
i igati It can now be
mow h ence presented below, that A. majoricum is yet
anorher sllotetranioes gf sia and that A. fontanum is one of t
SLEEP: ASPLENIUM MAJORICUM 323
two ancestral types.
THE EXPERIMENTAL INVESTIGATION
The observations recorded here were obtained during the course ot a
extensive study of synthesized Asp/enium hybrids (SLEEP, 1966) o
few of which are relevant to the problem of A. majoricum. Five ads
specific hybrids combining A. majoricum with other diploid or tetra-
ploid species have however given significant information and the
nurebnal details of inseminations versus hybrids raised are given in
Taste I. A single strain of A. majoricum from Soller, Mallorca,
was used Spal ages although sometimes as male and sometimes as
female. r the other species there was usually a choice of strains,
and eras possible ny Orga attempts were made with material
only the collective figures for any one type of hybrid are paperean the
table. Further information concerning the sources of material sore
fully omnes into hybrids is given as an appendix i 32
noteworthy that very large numbers of prothalli must in meet cases yo
inseminated i in order to obtain even a modest yield of hy
Illustrations of the morphology of the various plants involved i :
this investigation will be found in Fics 1-3. A. majoricum (4x) a
A. fontanum (2x) are shown in Fic. 1 A me C, and other species ee
fully crossed with A. majoricum in FIG. 2. The corresponding hybrids
are illustrated in Fic. 1 B and Fic.
TABLE I
Numerical wore of successful hybridisations involving
oricum (4x) with the species listed.*
&
a
— r= a» Percentage
2 ee = success
Se oo Se
GR ee Bee ug -
a ki HYBRID %
A. fo ee ey oe ee 0 Bee
2 A. MEE (2x) ‘ a { ; ay 2.86%
3. A. viride (2x) GO ise ese cites ic
sean HYBRID: :
4. iacedenaalan aa. 16 78 132 7%
of se Jortsenté (4x) 3 9 2 0 0 2 anit
rying degrees of persistence, to synthesize hybrids
*Attempts have been made, with 4
between a majoricum and certain other meets of Bsa nah Sm er iy fevecaat sid
tia andiezii
(2x), A. trichomanes subsp. tric ork neater over 400 prothalli were
(4x) but without success,
insemina
324 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
The chromosome pairing encountered at meiosis in the a
listed in TABLE | is summarised in TABLE 2. In all cases a minimu
6 cells was analysed and hess alternative fee pecs were available
more than one hybrid plant was investigated.
xamples are illustrated in PLATE I and in Fics 4 and 5.
TABLE II
Chromosome pairing in hybrids involving A. majoricum (4x)
with the species listed.
(Note—In Asplenium the monoploid chromosome number is 36)
No. plants
analysed Chromosome pairing
TRIPLOID HYBRID Bivalents | Univalents
ee” porn (2x) 4 36 3
5 aero onopteris (2x) 2 0 108
3. A. viride (2x) 1 0 108
TETRAPLOID HYBRIDS
4. A. macedonicum (4x) 5 c. 36 72
5. A. forisiense (4x) 1 36 evF2
INTERPRETATION
resale hsm yay of chromosome pairing among the triploid hybrids.
Had it A. majoricum x A. fontanum) stood alone there would
hav
represent A. fontanum chromosomes which have found partners
among half the aeieat complement of A. majoricum, or (b) that the
pairs are autosyndetic, i.e. that they involve only A. ab rgb pager
somes. On the first interpretation A. majoricum would b me
polyploid, with A. fontanum representing one ancestral wpe
second interpretation, on the other hand, shag indicate an citiaily
autopolyploid origin for the i ae speci
The evidence provided by hybrids 2 sat 3 in TABLE 2 eliminates
the second alternative. Antciyndete pairing is not displayed when
A. majoricum is crossed with genuinely unrelated species since in two
cases we find 108 univalents.* These facts thus strongly favour the
view that A. majoricum is an atlopolyploid and that A. fontanum or a
form with similar chromosomes is one of its parents.
The nature of the second parent is less easily determined, although
certain species can definitely be excluded. Thus A. onopteris L. an
A. viride Huds. can both be ruled out by the failure of chromosome
pairing displayed by hybrids 2 and 3 discussed above. A. obovatum
Viv. can almost certainly be excluded by the evidence from the tetra-
ploid | hybrids (items 4 and 5 in TABLE 2), although the reasoning here is
indirect. There is strong reason to believe (SLEEP 1966) that both
*Identical results have been obtained _ both these hybrids under varying weather
conditions and at different times of year
SLEEP: ASPLENIUM MAJORICUM 325
A. macedonicum Kiimm. (4x) and A. forisiense Le Grand (4x)
contain genomes corresponding to the same two iploid species,
A. fontanum and A. obovatum, and if this interpretation is
correct the 36 bivalents observed in hybrids 4 and 5 would in each case
Tepresent A. fontanum chromosomes, those of A. obovatum being
without partners.
Some additional evidence similarly excluding A. trichomanes L.
subsp. trichomanes can be quoted from a hybrid recently synthesised
by Lovis (1966). This is a tetraploid cross between A. majoricum and
A. adulterinum Milde (4x). It shows complete failure of pairing,
forming 144 univalents at meiosis. This fact still further confirms the
allopolyploid nature of A. majoricum, but since the genomic composi-
tion of A. adulterinum is known it provides further negative evidence
regarding the second parent of A. majoricum. A. adulterinum is also
allopolyploid with A. viride and A. trichomanes subsp. trichomanes
as the two parental types (Lovis, 1955, 1966). The new hybrid thus
excludes A. trichomanes subsp. trichomanes as well as confirming the
lack of affinity with A. viride which is already known. oe
an earlier stage in the enquiry attempts at hybridizing A.
majoricum directly with A. trichomanes subsp. trichomanes (2x) had
been made without success. This new indirect evidence is therefore
important, more especially in view of the black petiole in A. majoricum
and the fact that CHRISTENSEN (1913) interpreted hybrids described in
the literature as A. trichomanes x A. fontanum as equivalent to A.
majoricum. Morphology alone is thus an insufficient guide to affinity
in this case. }
There is thus at present no evidence regarding the nature of the
second species involved with A. fontanum in the parentage of A.
majoricum beyond the exclusion of A. viride, A. onopteris, A. obovatum
and A. trichomanes. However, Professor T. Reichstein (personal
ll age
communication) and myself and shown to be tetraploid. A diploid
nevertheless exists, recently described by MEYER (1964) under the name
of A. glandulosum subsp. bivalens D. E. Meyer (A. glandulosum Lois.
is a synonym for A. petrarchae(Guérin) DC.). The affinities of this new
form are therefore clearly pertinent to this problem and are currently
under investigation by other workers in Leeds.
ACKNOWLEDGEMENTS ;
Thanks are due to Dr J. D. Lovis for supplying the material of
A. majoricum used in this investigation and for helpful discussion, to
the gardening and technical staff of Leeds University Botany Depart-
ment for their constant attention in the cultivation and pote 8
326 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
wife, both for their hospitality and for showing me Asplenium majoricum
in its native habitat. ae y, I wish to ex pl ress my thanks and
appreciation to Professor I. Manton for her advice and encourage-
ment, and for her halisdn in preparing the manuscript for publication.
SUMMARY
Five types of interspecific hybrids involving the tetraploid A. majoricum
A Cc
FIGURE 2 Silhouettes (natural size) of fertile fronds of other species of Asplenium
incorporated into hybrids with A. majoricum. All poe are es in cultivation a
eds. A) A. onlinseesin tn from Aritzo, Sardinia. B) A. viride (2x) fr og m Hutton
Roof, Westmorland. C) A. macedonicum (4x) from Prilep, Matedots
SLEEP: ASPLENIUM MAJORICUM 327
Litard. with other diploid and tetraploid European species have been
synthesized and their chromosome pairing analysed. The evidence
indicates that A. majoricum is an allotetraploid species with A. ata
as one of its parents. The other parent is still unknown although A.
viride, A. onopteris, A. obovatum and A. trichomanes subsp. trichomanes
(2x) can be excluded on positive evidence.
A B C
FIGURE 3 Silhouettes (natural size) of fertile fronds from 1 mamas Sa bee
involving A. majoricum. All — id h yorid Asya 1 s io righ major ‘cums
A iori $x te hth triploi rid, ; 4
) A. majoricums hybri ei eT : ) As Sitadinicun’ 3x majoricum?
0. 4.
x viride?, triploid hybrid,
tet toid hybrid, TABLE 1
BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
=) 08
©
a, ‘5% Ve aro kip
Ta)
a) ° eg SUS a
oO
Pre ot (ens
[5 & ae’
O )
MoT §
9 ° *&o Boe Og SS a
4)
0 “he e” c#5 o® Sis Fao,
q &° 09G Slay
A ae B
Ss 9
goo ® is
Ad 9
re) as
° S
Aes 0% "owe
aes
Hr ghee * ,
. RW 2° 9 =
Oo BS 0 0
oO sg o 0 .@)
.S) e) 5
. i x .
onopteris? (cytology not otherwise illustrated) showing 108 univalent chromosomes
C) Tetraploid hybrid A. i majori
PLate ID showing 35 vivalents and 74 univalents.
BRET, PERN: GAZ. 9: €8) 1967 PLATE A
PLATE |
Photographs from permanent aceto-carmine squash preparations of spore mother
cells in meiosis, all reproduced at a magnification 0 of x 1000.
A) heating = Bolger hybrid between A. majoricum from Séller and A. fontanum
5 de showing 36 pairs and 36 univalents. For explanatory diagram
B) Synthesized triploid hybrid ant A. Ahad from Westmorland and A. majoricum,
108 univalent chrom
C) aa in Asdlonia ans ma Prilep, Macedonia. n=72.
hybrid een A. majoricum from Soller and A. mace-
D) ae kage. ey betw
from Prilep, showing approxim
For nouns diagram see Fic
ate 7 n paired and 2n cnnaied chromo-
to face page 328
SLEEP: ASPLENIUM MAJORICUM
APPENDIX
Details of the sources of material successfully incorporated into hybrids
Locality of origin
Source of spores
Collector
Species
eee
TT nw
A, majoricum
A, fontanum
A, fontanum
A, onopteris
C’as Patro Lau,
Barranc, Sdller,
oche, Vaud,
Switzerland
Villard de Lans,
sere, France
Aritzo, Sardinia,
Italy
Stock plants at
Leeds
Stock plants at
AS/24,
Stock plant at
Leeds. Spores
from Herb. Mus.
Brit.
Plant cultivated in
Basel. TR 114
J. Orell
A. Sleep
A. H. G. Alston
T. Reichstein
A, viride Hutton Roof, Wild collection A. Sleep
Westmorland,
Great Britain
A, macedonicum Markovgrad, N. of Stock plants at A. H. G. Alston
Prilep, Macedonia, Leeds. Alston
S. Jugo 17776
A. forisiense Brissago, sh Plant cultivated in T. Reichstein
Switzerland Basel. TR 34
REFERENCES
E, J. A., JERM 1964. Flora ph ig 14-17, Cambridge.
c. & LOVIS
CHRISTENSEN, C. 1913. bees Filicum, Susann ent I. Ke
JERMY, A. c. & LOVIS, J. D. 1964. Asp lenium majoricum Litardiére. Brit. Fern. Gaz.
3-16
LITARDIERE, M. R. DE 1911. Contribution a l’étude de la flore ptéridologique de la
péninsule ibérique. Bu r. Bot. 21:
LOVIS, J. D. 1955. tae at siesieann and its probable parents. Proc. Bot. Soc.
Brit. Isles | Be
peer rtifieial reconstruction of Asplenium adulterinum. Nature. (In
the ae
MEYER, D. E. c pres. Seg neue und seltene Asplenien Europas III. Ber, Deutsch. Bot.
Ges. 77:
ORELL, J. & scabies Asplenium majoricum R. Litardi¢re; Su area de
expansién en la ‘Sierra Norte de Mallorca. Collectanea Bot., Barcinone.
dn the pre
SLEEP, A. 1966. Rees cytotaxonomic problems in the a genera Asplenium and
Polystichum. Ph.D. thesis, University of
BOOK NOTE
compiled by Brummitt, has been published by the International Bureau
or Plant Taxonomy and Ni firing (106 Lange Nieuwstraat, san
25/- mongst some 2250 references to vascular plant taxonomy,
A. C. JERMY
330 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
JOHN SMITH OF KEW
R. E. HOLTTUM
Royal Botanic Gardens, Kew, Surrey
John Smith was one of the great pteridologists of the 19th century,
and he was the only one of them who based his ideas mainly on
observation of living plants. It was his misfortune that W. J. Hooker,
under whom he served at Kew, did not take kindly to Smith’s new
ideas. He had also many other misfortunes. Today he is little known
except to those who are concerned with the taxonomy and nomen-
clature of ferns, and his work for Kew has been too little recognized.
This situation has been recently brought to the attention of the Kew
community by an article by R. G. C. Desmond in the Kew Guild Journal
of 1965 (pp. 576-587) entitled ‘“‘ John Smith, Kew’s first Curator ”’.
The biographical data which follow are summarized from Mr Desmond;
the main object of the present article is to call attention to John Smith’s
work on ferns.
He was born at Aberdour in Fifeshire on 5 October 1798, son of a
gardener. After serving an apprenticeship he obtained a post at the
Edinburgh Botanic Garden in 1818, and there took the opportunity to
study botany. He came to London in 1820 and worked at the Royal
forcing garden at Kensington. He was appointed to the Kew staff
in 1822. In 1823 he became foreman of the hot-houses and propagat-
ing department, and was responsible for receiving new importations
of plants from various places in the tropics. It was then that his
interest in ferns began, but it was not his only interest. After the
death of Sir Joseph Banks in 1820 Kew began a decline which led
to the appointment of a Royal Commission in 1838 to consider the
future of the Garden, and ultimately to a new lease of life for Kew as
a national botanical institution from 1841 with Sir William Hooker
as Director. During this difficult period John Smith was largely
responsible for the maintenance of the whole collection of plants, and
he evidently had a very wide knowledge of them. The Commission
commended him, and he was appointed Curator, under the new regime,
from 1841, a post he retained until in 1864 he was obliged to retire
owing to failure of his sight. In 1880 he published Records of the Royal
Botanic Gardens, Kew, in which he gave lists of species of plants with
which he was concerned during his whole period of service, with many
historical notes of varied kinds. He relates, for example, the history
of early attempts to cultivate epiphytic orchids under glass, and also
gives notes on the early cultivation of palms and succulents, with lists
of the species grown in his time and the history of individual plants.
Before proceeding to deal with his work on ferns, I must record
something of his private life. His tombstone, near the west entrance
BRIT: FERN: G:A‘Zi-0 €e7o1967 PLATE 1]
PLATE II, John Smith (1798-1888)
(Copyright: Royal Botanic Gardens, Kew)
a
to face page 330
HOLTTUM: JOHN SMITH 331
to Kew parish church, provides a summary of domestic tragedy. Of
six children, only two survived childhood, and their mother died at
the age of 35. The only son, a promising boy who was appointed to
take charge of the museums at Kew, had to cease work at the age of
25 owing to ill-health, and died at 33; the last surviving daughter
died at the age of 42. John Smith outlived them all, and though blind
occupied his last years in writing a series of books which showed the
wide range of his knowledge and interests. He also left an extensive
manuscript history of Kew, in which some of his private grievances
are more fully expressed. He died at Kew on 12 February 1888.
John Smith records that when he went to Kew there were about 40
species of ferns in cultivation. Fifteen years later there must have been
a considerable collection of tropical species, as shown by some of the
drawings made by Francis Bauer (artist at Kew from 1790 to 1840)
which were the basis of Hooker’s Genera Filicum (1838-1842), to
which I will revert later. After Hooker’s appointment as Director
at Kew he began his great work Species Filicum, covering all known
species of ferns, the publication of which occupied twenty years (1844-
1864); during this period the number of ferns in cultivation greatly
increased. John Smith published a list of almost 400 species living in
December 1845. His next published list was in 1857, and included all
those known to be in cultivation, the number being 600. In his book
Ferns, British and Foreign (1866) he enumerated 1084 species which
he had seen alive.
A study of Smith’s successive lists indicates the development of
his thought on the inter-relationship of ferns and thus on classification.
His first important publication was in 1841-43, in Hooker’s Journal
of Botany, vol. 2, ‘‘ An arrangement and the definition of the genera of
ferns, with observations on the affinities of each genus’. He records
that he had long thought about the subject, and had come to the
conclusion that venation and other vegetative characters should be
used in the definition of fern-genera (mentioning the influence of Robert
Brown’s stray comments on this subject), and that when PRESL’s Tenta-
men Pteridographiae (1836) came into his hands he found much agree-
ment between his ideas and Presl’s. Prior to this, the shape and
position of the sorus and of its indusium were the sole characters
admitted in definition of genera. John Smith saw that this brought
together species which, as judged by the whole structure of the plant,
were not at all nearly related (for example, Polypodium and Phegopteris).
His successive schemes show an increasing insight, and mark an
important contribution to thought on the subject, thought that received
insufficient recognition owing to Hooker’s reactionary attitude. In
Species Filicum, and in the later summary entitled Synopsis Filicum,
Hooker reverted to the crudely artificial concept of genera, and for half
332 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
a century these great works were the only books covering all ferns in
the world.
In the introductory part of Smith’s last book on ferns (Historia
Filicum, 1875) he stated: ‘‘ the living collection, requiring daily care
and attention, led me to become acquainted with the special habits and
modes of growth of the different species; and this has enabled me to
draw up the following natural arrangement’. He went on to disclaim
finality ; but it is undoubtedly true that his scheme was in advance of any
produced until more than fifty years later. We may also note here
W. J. Hooker’s generous tribute to John Smith, in the introduction to
Synopsis Filicum (1865), written shortly before Hooker’s death. In
referring to the living collection of ferns at Kew, he wrote that it was
the finest in cultivation, and ‘‘ the formation of this fine collection is
mainly due to the exertions and ability of Mr John Smith, who for 43
years held an important position among the officers of the Royal
Gardens. His knowledge of ferns, and his writings on them, justly
entitle him to rank among the most distinguished pteridologists of the
present day ”’.
A number of the illustrations in Hooker’s Genera Filicum (1838-
42) are accompanied by names and latin generic diagnoses provided
by John Smith, who had studied the plants in question and had evid-
ently provided material of them for the artist. Some were drawn from
living plants, some from herbarium specimens; among the latter are
representatives of Hugh Cuming’s collection from the Philippines,
which were enumerated by Smith and included a number of genera
recognized by him as new. As noted above, Smith’s ideas on classifica-
tion at this time were set forth in his publication of 1841-43. He
divided the major family Polypodiaceae into seven tribes, based mainly
on soral structure. Within each tribe he had separate sections for
genera with free veins and anastomosing veins, except in Pterideae
(where the Preris and Blechnum sub-groups are distinguished by soral
characters) and in Dicksonieae (which included Lindsaea, Davallia and
the filmy ferns). In the tribe Polypodieae Smith still retained Phegopteris
(including Gymnocarpium of current works) with the free-veined
Polypodium species.
This scheme was improved in the catalogue of 1857, where he
incorporated the idea of a distinction between Eremobrya and Des-
mobrya (two elegant new terms coined by him from the Greek and
used by no-one else—he must have had a good grounding in both
Latin and Greek). These two terms denoted the distinction between a
creeping rhizome the tip of which advances beyond the existing fronds
and to which all fronds are jointed (e.g. Polypodium) and a rhizome or
caudex of which the apex does not advance beyond existing fronds,
HOLTTUM: JOHN SMITH 333
the fronds not being jointed to it. In 1857 only Polypodium and its
immediate allies were included in Eremobrya.
The scheme of 1857 was developed further in Ferns, British and
Foreign (1866), with explanatory notes and illustrations. Here Ere-
mobrya is made to include not only the whole Polypodium group of
ferns (Polypodiaceae in the strict sense of modern works but omitting
Platycerium) with both free and anastomosing veins, but also the
Davallia group and Oleandra. In Desmobrya there is a tribe Phego-
pterideae which includes all Thelypteroid and Dryopteroid ferns,
whether indusiate or not (but it includes also Nephrolepis). Hymen-
ophyllaceae is recognized as a distinct family, of equal rank to
Polypodiaceae, though Cyatheae and Dicksonieae are still ranked as
tribes within Polypodiaceae.
The final scheme of Historia Filicum (1875) gives the same content
to Eremobrya and Desmobrya, but in the former there is some further
subdivision of the Polypodium group and Platycerium is included.
Desmobrya is divided into a larger number of tribes; for example
Blechneae and Adianteae are separated from Pterideae. Gleichenieae,
Hymenophylleae, Schizaeae and Osmundeae are all ranked as tribes
of Polypodiaceae, which thus comprises all Leptosporangiate ferns, a
quite logical arrangement. The introduction to this book gives a
careful survey of the history of classification, and is well worth study
by modern pteridologists.
The contemporary author whose ideas most nearly resembled
John Smith’s was Mettenius (1823-1866) of Leipzig. Mettenius made
very careful anatomical and morphological studies of many kinds, and
was more attentive to the details of scales and hairs than any other
author for the next fifty years. Had Mettenius lived (he died of cholera
at the early age of 43) it is certain that he would have altered the history
of pteridological thought. His only completed scheme of classification,
dated 1856 (in Filices Horti Botanici Lipsiensis) is much on the same
lines as Smith’s, but the later arrangements of Smith are certainly an
improvement. Neither author made a good subdivision of the great
mass of terrestrial ferns included by Smith in Phegopterideae; this
had to wait for Christensen (1912-1919) and subsequent authors,
and indeed the problem is not yet solved.
John Smith was a man of intense industry. He not unnaturally
had a sense of grievance when his work was undone by the plans of
those in authority over him, and this is shown in his manuscript
record; it is clear that in many cases his special knowledge was dis-
regarded. To his subordinates he was probably a severe master, but
evidently at least two of them (whom he mentions by name) had received
sufficient inspiration from him to help him in making records after
334 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
his sight failed. Mr Desmond reports a statement by Botting Hemsley
that Smith ‘“‘ was not of a sympathetic nature... he r requ uired no
relaxation himself . . . and gauged others’ needs by his own”. This
statement should be read in conjunction with the testimony of W. J.
Hooker of the excellence of John Smith’s services to Kew. Looking
back, we can see him as a man with both originality of mind and also
a strong sense of duty; Kew may well hold him in honour and remember
his many services with gratitude.
REVIEW
THE OXFORD BOOK OF FLOWERLESS PLANTS. Text by F.
Brightman, illustrations by B. Y. Nicholson, pp. viii + 208, aa
University Press, 1966 price £2.5.0. 25 x 18cm
This book illustrates in vivid pacaptied almost 690 species of pipeidonatet,
bryophytes, lichens, fungi and marine algae with its main aim “‘to help t
beginner with identification’’. Short descriptive stad sh given opposite the
drawings with the same aim in view and in many instances mention is made
of closely related ase not illustrated. To caliente the use of the book the
stematic order but in habitats under the following
main headin ngs: seashor ore, grasslands, uplands, wet places and woodlands;
each of these is subdivided so as to be more precise a guide. The plants are
om teeth of Dryopteris dilatata are not well snows in ine outlined
seup. The robustness of The/lypteris limbosperma (p, 189) is well executed
but the detailed drawing does not show the submarginal sori and gives an
recognisable, and the bent stipe gives a false impression of a creeping, slender
rhizome. On the question of colour dinticting I think p. 185 is too blue;
one useful field re is that Dryopteris aemula is more yellow green
than D. dilatata but on pp. 185 and 186 D. pice. wl far too yellow, and with
its gingery scales it rese Basha D. assimilus. The of Asplenium cart
(p. 1) is too reddish and the shiny black = of Rabati on the same page
th and there are some “excellent
—e
=
gz
eelel
5
eo
o
a.
are not true but saa 7 os isolated and exceptional case. The scale, often
only mentioned in ext, is sometimes misleading, e.g. Athyrium filix-
femina and Sana Poms are given as having fronds 24 pie 18 inches
ica SO vely.
A, C, JERMY
BRIT. FERN GAZ. 9 (8) 1967 RR)
SPORELINGS OF EQUISETUM ARVENSE
IN THE WILD
Cc. N. PAGE
Department of Botany, University of Newcastle-upon-Tyne
Little is known about the establishment in the wild of the short-lived
spores of Equisetum, although much is tacitly assumed. Recorded
occurrences of wild Equisetum prothalli are very few, and observations
on the subsequent development of the sporeling plants in nature are
virtually lacking. Thus it was with some interest that a large number
of small plants of Equisetum were recently located which, it was felt,
could be demonstrated to represent sporeling plants at the end of their
first season.
The plants were growing at Whittle Dene Reservoirs some ten
miles west of Newcastle-upon-Tyne, to where my attention was first
drawn by Mr O. L. Gilbert in September 1966. The reservoirs had
been drained in early April of that year for the construction of new
sluice gates, and by September, the damp mud surface which covered
most of the reservoir basin was found to be carpeted with small plants
of Equisetum arvense L.
The reservoir previously had not been drained for thirty years and
a fine deep silt of pH 6.2 was now exposed. The silt merged into a
muddy gravel of pH 8.5 towards the periphery of the reservoir which
was bounded by sloping stone retaining walls extending to some depth
above and below the normal water level. Three species of Juncus,
together with a variety of small annual weeds, were already beginning
to establish themselves near the gravel edges. The muddy bottom had
shrunk as it consolidated and had developed a network of deep cracks,
although the surface had remained damp. The presence of large
shells of the freshwater mussel Anodonta in the muddy bottom right
up to the flanking walls confirmed that the whole of the reservoir had
been continuously covered for many years by a good depth of water.
The Equisetum plants were already well established, and carpeted
the whole of the bottom of the empty reservoir, often a dozen plants
to the square foot. Each formed a loose rosette of three to eight simple
or sparsely branched decumbent shoots, 4 to 14cm long and | mm
diameter, no particular shoot dominating over the others. Only
occasional plants had developed their first leader shoot, which grew
more or less erect, up to 20 cm tall, and 5-sided. These bore several
regular whorls of five short 4-sided branches. Each plant examined
had produced between two and five rhizomes varying from 2.5 to 35 cm
in length, and 1.2 to 1.5 mm in diameter. These penetrated the mud
at a steep angle, only occasionally producing a lateral branch rhizome.
336 BRITISH FERN GAZETTE:*VOLUME 9 PART 8 (1967)
"
UT)
Ss e of sporeling E. arvense gathered in early September. Large e broken-line
See rt position of soil and fissure etn A) tubers, B) leader-shoot developing,
C) shoot developed from a rhizome which penetrated a fissure. mall lateral
shoot also emerges from the first sode of the uppermost rhizome. Natu ral size.
PAGE: EQUISETUM ARVENSE 337
Where, however, the rhizome struck the surface of a deep fissure a single
4-sided aerial shoot was often produced. The rhizome internodes,
which were white or pinkish in colour, varied in length from 1.2 cm
close to the aerial rosette, and increased to 2.5cm at some distance
from it. Unlike those of the aerial shoot, each rhizome internode
became fully extended immediately behind the apex. Regular whorls
of four or five long slender unbranched roots originated from almost
every underground node. Larger roots occasionally could be found
which bore simple laterals at frequent and irregular intervals. The
young rhizomes usually grew downward to a depth of at least 20 cm
before starting to level off, and their roots penetrated to 25 or 30cm
below the soil surface.
By early November the plants were still green and had grown to
form bushy tufts about 30 cm across, with up to two dozen shoots.
Some of these sparsely branched semi-decumbent aerial parts reached
a length of 25cm and had up to fifteen nodes. Many plants had
developed, at the end of only their first season, three to five small
solitary oval tubers from some of the older rhizome nodes just beneath
the mud surface.
MATZKE (1941) found gametophytes of Equisetum arvense growing
under very similar conditions to those pertaining here, in the bottom
of an old iron-ore mine which had been flooded and redrained. By
extremely large numbers of young plants observed. Furthermore,
these pieces would have had to have been available for uniform dis-
persal over the reservoir by the water before it was completely drained
off in early April. At that time of the year, aerial shoots of E. arvense
are not in evidence in Northumberland and therefore could not have
been available to provide a source of fragments in this manner.
Similarly, the stone retaining walls of the reservoir would have minim-
338 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
ised, if not totally prevented, fragmentation of any rhizome material
into the water, and no other source from which such abundant pieces
of rhizome as would have been required, could be found.* On the
other hand, very shortly after the reservoir had been drained, many
colonies of Equisetum arvense in the surrounding area are known to
have been shedding their spores.
From the available evidence it is concluded, therefore, that these
hundreds of horsetails clothing the bottom of the reservoir had arisen
as the result of the successful establishment of spores earlier in the
year. The wet mud surface exposed by the retreating water formed a
virgin habitat free from the competition of established plants, and
hence immediately available for the germination and development
of the short-lived green Equisetum spores being shed in great abundance
nearby. It is therefore believed that under these apparently ideal
conditions the prothalli would have developed vigorously, soon
producing these young sporelings. Matzke and Feigley both record
that commonly more than one sporophyte was produced on each
prothallus, but judging by the distance of several inches between most
of the present plants, one sporophyte only must have ultimately arisen
from each prothallus, and unless there were initially a great many
more prothalli than even the present number of sporophytes would
lead us to believe, most must have arisen as the result of self-fertilisation.
By the end of their first season these plants dominated the habitat
with nearly 100°% cover over the surface, and an impressive develop-
ment of roots and rhizomes beneath it. Had the reservoir not been
re-flooded, there is every reason to believe that they would have been
permanently established, and that the Equisetum arvense had very
effectively pioneered this new bare habitat. :
I would like to thank Dr T. G. Walker for his help in the preparation
of this paper. REFERENCES
FEIGLEY, MARGARET, 1949. An occurrence of greene A of Equisetum in
Cheboygan inane Michiga n. Am. Fern Jour 39:
MATZKE, E. B. 1941. he Torreya 41: 181-187.
*The water is screened of all such debris before entering the reservoir.
BOOK NOTE
INDEX SELAGINELLAR UM by Clyde F. Kid (Mem. Soc. Brot. 18. 1966).
Dedi A ton and based on his manuscript index, it was
inevitable that the major part of Reed’s icktex is Alston’s. Comparison
eared serious delay. As it is, we have a long-wanted index and nomenclator,
even though finding the answers may sometimes require oe and
inspired rummaging among its 287 pages.
BRIT. FERN GAZ. 9 (8) 1967 339
GRAMMITIS IN SAMOA
W. A. SLEDGE
Botany Department, University of Leeds
Species of Grammitis are all small ferns with simple, entire fronds and
naked sori. They commonly grow epiphytically in rain forest or mist
forest usually at considerable altitudes and are world wide in distribu-
tion within the tropics or subtropical regions to which they are almost
wholly confined. They are rarely if ever seen in fern houses and are
reputed to be almost impossible to grow in cultivation.
Seven species of Grammitis are included in CHRISTENSEN’S “* Re-
vision of the Pteridophyta of Samoa” Bull. Bishop Mus. 177:1-138,
1943). In this monographic account of the genus (in Philip. Journ.
Sci. 80: 93-271, 1952) COPELAND attributed eight species to Samoa.
The Society Islands are credited with five species, Hawaii with four,
Fiji with three and New Caledonia with three. Copeland considered
the number of species present in Samoa to be “‘ most remarkable ”
and he adds that the paucity of collections from there is also remarkable.
This is certainly born out by the fact that the most frequently collected
species of Grammitis in Samoa is not known to have been collected
more than half a dozen times; other species are known from one, two
or three collections only. Part of the reason for the small number of
collections of these Samoan ferns is connected with their being in-
conspicuous plants which are largely confined to the mountain forests
in the most inaccessible regions of the interior. But part of the reason
is also due to the fact that all the species in Samoa appear to be remark-
ably localised. One may see several specimens of a Grammitis growing
epiphytically on a tree trunk and search neighbouring trunks in vain
for further specimens; indeed the rest of the day may be spent in
traversing mossy forest rich in ferns and apparently ideally suited for
species of Grammitis without another specimen being seen.
During 3 months’ fern hunting in Samoa in 1965 I was fortunate
to find all the species of Grammitis though only two species were
collected on more than one occasion. As would be expected in a
group which favours high level rain forests, the island of Savaii which
has the highest mountains is richest in species. Four species of
Grammitis on present evidence would appear to be confined to Savaii.
The lowest altitude at which I encountered any species was 2000 ft.
Since Christensen and Copeland were dependent for their accounts
on herbarium material, much of it both old and somewhat scanty, it is
scarcely surprising that the improved knowledge of these little known
Samoan ferns which my collections have afforded has necessitated some
revision of the previous accounts. One of Copeland’s species is here
340 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
reduced to a synonym but as one new species is described the total
number of Samoan species remains unchanged.
The missionaries Powell and Whitmee who collected ferns assidu-
ously in Samoa nearly a century ago unfortunately rarely cited localities
whence their specimens had been gathered. Information about
distribution within Samoa is therefore scanty and I have not thought it
necessary to repeat the information which is already given by Christ-
ensen in his fern flora of the islands. Specimens of all my gatherings
have been deposited at Kew.
The following key, modified from those given by Christensen and
Copeland, will serve to distinguish the species.
Fronds with black margins a. es +
Fronds lacking black margins:
1. G. vaupelii
Stipe glabrous... ae se ot .. 2. G. graminella
Stipe hairy:
Sori near the midrib:
Sori immersed in deep craters ¥ .. 3. G. insularis
Sori superficial or slightly immersed:
Marginal hairs over 1 mm. long:
Fronds glabrous above save on costa 4. G. monticola
Fronds hairy above .. es .. 5. G. hookeri
Marginal hairs very short 6. G. conformis
Sori medial to supramedial:
Sori in a regular submarginal row page sot & whitmeei
Sori irregularly biseriate or scattered .. 8. G. samoensis
1 GRAMMITIS VAUPELII (Brause) Copel., in Philip. J. Sci. 80:
265 (1952)
Polypodium Vaupelii Brause, Notizbl. Bot. Gart. Berlin 8: 140
(1922
P. marginellum Baker in J. Bot. Lond. 14: 12 (1876) non Swartz
1788).
My single gathering of this—three specimens and only one frond
fertile—is only the fourth known collection, all of them from the
island of Savaii. The fronds are very narrow, up to 17 cm long and
2-3 mm wide
I have seen no specimen of P. marginelloides J. W. Moore from
Tahiti, which Copeland thought probably the same species. It has
wider fronds (7 mm) and is said to have forked glandular hairs on the
frond margins which is not the case in my Samoan plants.
SAVAIL: epiphytic in mossy forest, Maugaloa, alt. 1050-1200 m, 6 March
1965. Sledge 1738.
2 GRAMMITIS GRAMINELLA (C. Chr.) Sledge comb. nov.
Polypodium graminellum C. Chr., Index Fil. 530 (1906). Bull.
SLEDGE: GRAMMITIS OF SAMOA 341
Bishop Mus. 177: 108 (1943).
P. simplex Baker in J. Bot. Lond. 14: 12 (1876) non Sw. (1801).
P. ligulatum sensu Hieronymus in Hedwigia 44: 78 (1905) non
Bak. (1867).
Grammitis ligulata sensu Ching in Bull. Fan Mem. Inst. Biol. Bot.
10: 240 (1941); sensu Copel. in Philip. J. Sci. 80 (2): 150 (1952)
non Polypodium ligulatum Bak. (1867).
This is the fern which Hieronymus and Copeland considered the
same as Baker’s Polypodium ligulatum, a supposedly Fijian species
based on a gathering by Brackenridge. Both authors used Baker’s
epithet for Samoan plants. But the identity of Baker’s species had
never been settled by reference to the type and when Copeland wrote
to Alston asking him to look into the matter the reply he received
stated (Joc. cit. 151) that ‘he found at Kew only an empty cover
bearing the name Polypodium ligulatum’’. Yet if Alston had looked
a little further he would have found the specimen, and just where one
would expect to find it, ic. amongst the specimens which Brackenridge
sent to Hooker as his P. conforme.
No doubt later authors, in trying to place Baker’s species, have
been troubled by the fact that no known Fijian Grammitis would agree
with Baker’s description. But Copeland has pointed out (Joc. cit. 150-
151) that localities were ‘‘ of very minor importance to Brackenridge
who in publication and distribution gave one number to what he
regarded as a species no matter how far apart the specimens were
collected”. This judgment is certainly born out by Grammitis ligulata
(Bak.) Ching which Baker founded (Syn. Fil.: 320) on a specimen of
Brackenridge’s sent as P. conforme. This is one of three specimens
mounted on one sheet, two of which have setose stipes and short
marginal hairs on the fronds, and are G. conformis. The third specimen
is glabrous and has narrower, longer and thinner fronds. Two speci-
mens adjacent to one another on the sheet had already been annotated
by Hooker “ more like P. subspathulatum Brack. Tab. | f. 1” and this
note is followed by ‘‘ = P. ligulatum Syn. Fil ’’ in Baker’s writing.
It is evident therefore that Baker founded his new species on two
specimens one of which was typical G. conformis. His description
(Bak.) Ching. In describing the texture of the fronds as ‘
Grammitis, but with the species which Mettenius described, from
specimens collected on Tahiti, as P. trachycarpum, two years after
342 BRITISH FERN GAZETTE: VOLUME: 9% PART. 8 (1967)
Baker had—inadequately and in one respect misleadingly—described
the same species from a single specimen which Brackenridge had almost
certainly collected in Tahiti but had failed to distinguish from his
P. conforme.
Baker’s Polypodium simplex was founded on Whitmee 123 ex parte
and the Kew sheets of this gathering and Powell 192, which is manifestly
identical, carry thirteen good specimens. My own gathering agrees
exactly with Whitmee’s and Powell’s specimens.
Christensen states that the sporangia of G. graminella are glabrous
but {Copeland describes them as “‘ bearing minute, dark-chestnut setu-
lae and’s early ber he iscorrect. Onthe other hand Copeland’s
reference 19 cit. 151) to the paleae on one gathering as “ bearing 2
or 3 unicellular hairs at t the tip”’’ is inexplicable. Hairs are never
present a the scales of G. graminella. 1 have examined under the
binocular microscope all the Kew specimens of this species and every
one of my own and the paleae are uniformly naked. Indeed of all
Samoan species of Grammitis only in G. py Slionaty are the paleae
provided with long apical hairs, sometimes solitary but often two or
oe Bem eo at their tips. Copeland’s observation was therefore, if
, made on a specimen which was ae not G. graminella
neety ioe well fave been an aberrant G. samo
It was evidently on account of this 1 etal that Copeland
placed G. graminella next to G. samoensis in s monograph. In other
respects he considered it “not very near’’ to that species. I see no
relationship ‘hat tever between them. The affinity is clearly with the
plant from the Society Islands which has for so long been called G.
trachycarpa but for which the name G. /igu/ata must now be substituted.
In his key to the Pacific Island species (/oc. cit. 140-141) Copeland
uses the herbaceous versus the subcoriaceous texture of the fronds as
the character whereby to separate G. trachycarpa from G. graminella.
Doubtless if he had not been misled by Baker’s false statement he
would long ago have referred P. ligulatum to the former species and not
to the latter. The rather stiff texture and opaqueness of the fronds of
Samoan plants is just as constant as is the thin texture of Tahitian
plants in which the fronds also have a greater tendency to be repand
ut in other respects I see no significant differences.
SAVAII: epiphytic in mossy forest, Maugaloa, 990 m, 4 March 1965.
Sledge 1722.
3. GRAMMITIS INSULARIS Copel., in a} Calif. Publ. Bot. 18:
223 (1942); Philip. J. Sci. 80: 149 (195
Polypodium subspathulatum Brack., var. “f sensu C. Chr., in Bull.
Bishop Mus. 177: 108 (1943).
This was based on Vaupel 451 collected from “ Hinterland
Maugaloa ”’ on 27 March 1906 and distributed by Vaupelas P. conforme.
Neither Whitmee nor Powell appear to have found this species and
no other gathering was known until my two collections both made in
Savaii and one of them in ats near Maugaloa. Christensen cites the
SLEDGE: GRAMMITIS OF SAMOA 343
Berlin label of Vaupel 451 as from “‘ Mataana 1600 m, epiphyte ’’.
(Mataana and Maugaloa are both volcanic cones situated about three
miles apart on the eastern Side of central Savaii). Copeland states
that the Vaupel collection “is uncited by Ch ee” when in fact
it is cited and under the species where one would expect “ find it cited.
op in fact pointed out the differences between the Samoan
plant and P. subspathulatum, which was described pan the Society
eae but oe from naming it since he had seen “ only one
rather old pla
This is tre separated from all other Samoan species of io alan
by jee wei ocr sori. Copeland describes the fronds as up to
dow an by Groner, cred branch vein which terminates
about half way betw costa and frond margin. The basiscopic
branch forks at about the level of the ending of the acroscopic branch,
and their endings are situated just within the margins of the frond.
On the upper surface of the frond all the vein endings appear as con-
a A punctate markings and hence within each margin there is a
of dots and another row of dots but only half as numerous occurs
satel midway between costa and margin.
Copeland states that G. insularis differs from G. subspathulata
Imm
FIGURES 1 & 2 Rhizome scales of Grammitis: 1) G. hookeri, 2) G. monticola
344 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
99
in “the inframedial sori, more naked fronds and 8
prow is also distinctive as a comparison of mae fig. winches insular)
and fig. 26 (G. subspathulata) shows. The i differ n only
olitior in the two species but those in G. insular are meni in ans sesel
craters. As regards the “ more naked fronds”’ of G. i nsularis there
are certainly very few marginal hairs on the type specimens but
marginal and surface hairs are fragile and easily abraded in old fronds.
y own specimens sho cepa moderate numbers of hairs are distributed
along the margins much as in G. subspathulata. 1 see no difference
in the oy of the two aa Although Copeland claims that these
afford one of the specific custincens he does not state what the
= Pi are nor are these indicated in his descriptions under the
two species where the ee are referred to in virtually identical
terms.
SAVAII: epiphytic in mossy forest, Maugaloa, 1200 m, 6 March 1965.
Sledge 1739. _ Epiphytic in mossy forest, Tuisivi Range above
Kono: 1350 m, 17 March 1965. Sledge 1797.
4 GRAMMITIS MONTICOLA Sledge sp. noy. PLATE III
Rhizoma breve, crassum, paleis rufo-brunneis, lineari-lanceolatis,
acutis, 4-5 mm longis munitum (Fic. 2). Frondes approximatae,
rith ad t al sepia ORS ROAR EEANS: eae ad 2 i longa, pilis
castaneis, squa , 2-3mm longis Lamina usque ad
20 cm longa, c 132 m lata, ide herbacea “last, integra, ali-
ue ma c
circa extremum rami acroscopici furcatus, aliquando una e
divisionum denuo furcata. Sori uniseriati superficiales prope coasta
setis paucis longis circ cumdatis; pe Ser PaneNe sed orbiculascentia,
sporangia setulis minutis brunnei
Differt a G. hookeri statura maiore, a. pessoa oe
acutioribus et pilarum superficialium in frondibus defici
Rhizome short, thick, covered in red-brown, —reapiornen acute
scales, 4-5 mm long (see Fic. 2). Fronds clustered close together ; stipes
variable in length up to 2m, with chestnut-brown, squarrose hairs
2-3mm long. Lamina up to 20cm long, I-1.5cm broad, stiffly
ng,
stipe, the surfaces elsewhere glabrous; veins obvious, twice, rarely
three-times, — simple in the upper part, the fertile branch acro-
or less extended beyond the sorus, the sterile branc
BRIT, FERN GAZ. 9° (8) 1967 PLATE Ahi
se?
TYPE SPECIMEN
PLATE Il. Grammitis monticola Sledge
to face page 344
SLEDGE: GRAMMITIS OF SAMOA 345
HOLOTYPUS, vupotu: epiphytic, summit ridge of Mt. Fito, alt.
1080 m, 2 April 1965. Sledge 1835 (K).
Very different from any other Samoan Grammitis. It is one of
the hookeri-conformis group of species and most closely related to the
former species from which it differs in its larger size, longer and darker
scales (see Fics. 1 & 2) and in the absence of hairs (other than para-
physes) on the faces of the fronds. I have seen two Samoan gatherings
referred to by Copeland (in Philip. J. Sci. 80: 143 (1952)) under G.
wattsii. They are not the same as this species and I agree with Copeland
in referring them to G. hookeri.
5 GRAMMITIS HOOKERI (Brack.) Copel., in Philip. J. Sci. 80:
157 (1952).
Polypodium hookeri Brack., U.S. Expl. Exped. 16: 4 (1854).
P. setigerum Hook. & Arnott, Bot. Beech. Voy.: 103 pl. 21A (1832)
non Blume (1828).
his is the most widely distributed of the Samoan species of
Grammitis, occurring also in Hawaii, whence the type was described,
and Fiji. Records from elsewhere are not accepted by Copeland.
Within Samoa it is evidently very rare as stated by Christensen. [
collected it once only. The two specimens of this collection match
well the type gathering of P. setigerum from Hawaii.
There is one Samoan collection by Powell in the Herbarium of the
British Museum not referred to by Christensen. The only Samoan
gathering at Kew is a single specimen from Tutuila without the col-
lector’s name.
SAVAII: epiphytic in mossy forest, Tuisivi Range above Aopo, alt.
1350-1500 m, 17 March 1965. Sledge 1806.
ON
GRAMMITIS CONFORMIS (Brack.) J. Sm., Hist. Fil. 181 (1875).
Copel. in Philip. J. Sci. 80: 147 (1952).
Polypodium conforme Brack., U.S. Expl. Exped. 16: 4, pl. 1 fig. 2
(1854); C. Chr. in Bull. Bishop Mus. 177: 109 (1943).
Grammitis christophersenii Copel., in Occ. Pap. Bishop Mus. 15:
89, fig. 8 (1939). Philip. J. Sci. 80: 147 (1952).
The type sheet (U.S. Nat. Herb. 60714) carries two somewhat
dissimilar and damaged specimens. These have been described by
Copeland (/oc. cit.) but he was incorrect in his guess that they repres-
ented ‘* probably the entire collection’’, for three specimens bearing
the same label as the type sheet were sent to Hooker and are in the
Kew collection. The three Kew specimens are also not uniform; two
of them are G. conformis, the third is a glabrous plant which was
subsequently described by Baker as Polypodium ligulatum.
Copeland (/oc. cit. 148) expressed doubt as to whether Bracken-
ridge’s collection of G. conformis came from Fiji as stated on the label
and he considered a Samoan origin more probable, at least for the
smaller of the two specimens on the type sheet. Through the kindness
346 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
of Mr C. V. Morton I have been able to examine the type sheet of
G. conformis and to compare the specimens with those sent to Hooker.
The larger plant on the type sheet exceeds in size any of my specimens
and—quite Apert: from the evidence afforded by the specimen of
ligulatum Bak, on the Kew sheet, which in all probability came from
Tahiti—I think it ‘improbable that the four other specimens of Bracken-
ridge’s G. conformis represent a single gathering from one locality; for
in my experience different populations of G. conformis in Samoa sho
small differences yet i nobly, uniform within each local population,
Copeland’s surmise as to the origin oR Brackenridge’s specimens of
this species ey Guecetors well be correct though G. conformis
certainly does occur in Fiji; there ie in gatherings from there at
Kew — referred to this speci
ave examined the fragmentary type of G. christophersenii in
Conslanie herbarium (No. 16411). This differs in no respect from
G. conformis and Copeland himself evidently had doubts as to its
distinctness since, in his later work, he admits that “it is not unlikely
that, when better known, they will be found to blend’’. The venation
characters used to separate the two in his key (Joc. cit. 147) are not
significant since once and twice forked veins may both be present in
the same frond of G. conformis. Thou gh the fronds of G. christopher-
senii are said to be remarkable for the minuteness and denseness of their
marginal setulae, Copeland’s illustrations (/oc. cit., fig. 19, fig. 20) show
no difference whatever in the denseness or length of the marginal hairs
in this and G. conformis. In his key on p. 141 the two are separated
by the colour ‘of their paleae but no reference to this supposed distinc-
tion is to be found in the discussion under G. idee Spartans I see
no ae Hate in scale characters in any of my specim
Ithough the fronds of G. conformis are jessie as glabrous
save on the midribs and margins, young fronds are hairy above and
a few surface hairs are usually present on the upper side in the lower
part of mature fronds and occasional hairs are often to be found
elsewhere, at least close to the margins. In one of my gatherings
(Sledge 1696) hairs are frequent on the upper surfaces of the fronds
though in no other respects do the specimens differ from other collec-
tions of the same species. The maximum frond dimensions of my
specimens are 12cm long and 6mm wide. Though the fronds are
normally ateeniies to shortly stipitate, the stipes may sometimes
reach 2 cm in length.
n Samoa this appears to be the most frequent species of Grammitis.
I collected it in three localities in Upolu and in two in Savaii, whereas
of the seven other species I only found G. insularis on more than one
occasion.
UPOLU: coiniaty in forest on Mt. Fao, alt. 600m, 17 Feb. 1965.
Sledge
Ent in rain forest, Afiamalu, alt. 720m, 22 Feb. 1965.
SI 167:
Summit of Sigaele, epiphytic, alt., 780 m, 25 Feb. 1965. Sledge
SLEDGE: GRAMMITIS OF SAMOA 347
SAVAII: epiphytic, Matavanu, alt. 600-690 m, 3 March 1965. Sledge 1710.
Epiphytic in mossy forest, Maugaloa, alt. 1050-1200 m, 6 March
1965. Sledge 1740.
7 GRAMMITIS WHITMEEI (Baker) Copel., in Philip. J. Sci. 80 (2):
150 (1952).
Polypodium whitmeei Baker in J. Bot., Lond. 14: 12 (1876). C. Chr.,
Bull. Bishop Mus. 177: 109 (1943).
Easily recognised by its large size and submarginal sori. The
fronds are normally about 20 cm long and often longer and I-1.5 cm
broad. It is apparently confined to the island of Savaii.
SAVAII: epiphytic in mossy forest, Tuisivi Range above Aopo, alt.
1350-1500 m, 17 March 1965. Sledge 1799.
GRAMMITIS SAMOENSIS (Baker) Ching in Bull. Fan Mem.
Inst. Biol. Bot. 10: 241 (1941).
Polypodium samoense Baker, Syn. Fil. 321 (1867); C. Chr., Bull.
Bishop Mus. 177: 110 (1943).
P. savaiense Powell in Baker, J. Bot., Lond., 14: 344 (1876).
This is also endemic to the island of Savaii and not related to any
other Samoan species. The veins are mostly forked twice with a sorus
at the end of the shorter acroscopic branch and of both forks of the
basiscopic branch. The sori then appear irregularly biseriate, one
row more or less medial in position and the other intramarginal. But
frequently one of the basicopic divisions forks again and the sori are
then scattered rather than seriate. The linear-lanceolate scales differ
from those of other Samoan species in their ciliate margins and in
bearing hairs, up to 2 mm or even 3 mm long, at their tips. These are
either solitary or, more commonly, two or three are borne together.
Powell’s original gathering of G. samoensis was made in August
1863. Twelve years later he sent to Baker another gathering which he
and Whitmee had made in October 1875 on trees at 4600 ft in Savaii.
These specimens are somewhat larger than those of his original gather-
ing and the irregular arrangement of the sor! Is correspondingly more
evident on the broader fronds. Believing it to be a new species he
suggested the name P. savaiensis to Baker in a note which is attached
t
co
described it as new. There is however no doubt that the two gatherings,
each represented by a single sheet at Kew, represent the same species.
The distinctive scales are identical in the specimens on both sheets
and such differences as exist in the disposition of the sori are simply
an expression of the minor differences in frond width in the specimens.
SAVAII: epiphytic in mossy forest, Maugaloa, alt. 1050-1200 m, 6
March 1965. Sledge 1737.
I am grateful to Mr C. V. Morton who sent me three type speci-
mens of Grammitis from the United States National Herbarium, also
to Prof. Warren H. Wagner, Jnr., Ann Arbor, Michigan, for sending
me Copeland’s type specimen of G. christophersenii. 1 wish also to
acknowledge with thanks the grants from the Royal Society and the
John Lewis Trust which made possible my visit to Samoa.
348 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
THE IDENTITY OF TWO MADAGASCAN
SPECIES OF BLECHNUM
E..A. C..L. E. SCHELPE
Bolus Herbarium, University of Cape Town
Blechnum bakeri C. Chr. is based on a mixture of sterile fronds of this
species, sterile and fertile fronds of B. ivohibense C. Chr. and of mature
and immature fertile fronds of B. microbasis (Bak.) C. Chr. A lecto-
type is chosen for B. bakeri and the nOnY RY and known iikabanen
of B. bakeri and B. ivohibense are given
Although Sager ria (1932) recognised Blechnum bakeri C. Chr.
and B. ivohibense C. Chr. as distinct species, BONAPARTE (1925) and
TARDIEU-BLOT (1960) sien the latter to be a variety of the former.
B. bakeri is founded on Lomaria pubescens Bak., the type of which
was collected at Antananarivo, Madagascar, by Po ol. On examination
at Kew, this type collection proved to be a mixture of three different
taxa on two s
Most of Baker’s description of his L. pubescens agrees with sterile
fronds in this collection with pilose rachises and with pilose veins
below; consequently, these fronds have been taken as representing
B. bakeri. There is also a sterile frond and a fertile frond of B. ivohib-
ense as well as a mature fertile and an immature fertile frond of B.
microbasis (Bak.) C. Chr. in the type collection.
The two sheets forming the type ~scely of L. pubescens Bak.
at Kew have been annotated as follow
Sheet I M4 microbasis (Bak. C. heen frond (A)
B. ivohibense C. Chr.—sterile frond (B)
B. bakeri C. Chr.—sterile fronds (C, D)
Sheet II B. biker C. Chr.—sterile fronds (E, F)
B. microbasis (Bak.) C. Chr.—immature fertile frond (G)
B. ivohibense C. Chr.—fertile frond (H)
In view of the fact that most of Baker’ s description applies to the
pilose sterile fronds of this collection, it is proposed that the specimens
annotated C, D in the type collection should constitute the lectotype
of B. bakeri C. Chr. (L. pubescens Bak. non Kunze).
Consequent upon this choice of a lectotype, some discrepancies
in Baker’s description require explanation. The stipes in the proposed
oe (though obviously incomplete) vary from 1} to 14 inches,
to 3 inches as stated by Baker; this can be partly explained by
ek longer stipes on the specimens of B. ivohibense (B, H). Baker’s
description of the fertile frond appears to have been Bieter er from
the glabrous fertile fronds of B. ivohibense and B. microbasis in the type
ee considering his description of ‘erecto-patent pinnae, 1-3
inches
The type of B. bakeri var. glabrum is D’Alleizette 82 from
SCHELPE: MADAGASCAN BLECHNUM 349
Mandraka, Madagascar, presumably at Paris, has not been seen but
Tardieu’s description leaves little doubt as to its identity
he question now arises as to whether B. bakeri oad B. ivohibense
should be regarded as two separate species. The only differences
between them appear to be that the rachis and veins are ‘ioe e in B.
bakeri and glabrous in B. ivohibense, but no intermediates have been
seen as yet. However, both taxa seem to have been collected in close
proximity at two localities in Madagascar, at Antananarivo by Pool
and on the Pic d’ Ivohibe by Humbert. On the other hand, only B.
ivohibense of this “‘pair’’ of species has been collected in continental
Africa i _ widely separated localities in Rhodesia, norther Mozambique
and Tanzania. Consequently it seems advisable on the available
sap to treat these two taxa as separate species.
The synonymy and know distribution of these two species is
given below.
Blechnum bakeri C. Chr., Ind. Fil.: 151 (1905); Tardieu-Blot in
Hu gee Fl. Madag., Polypod. 2: 7, t.2, f.4 (1960).
aria pubescens Bak. in J. Linn. Soc. Lond. 15: tay
non cra (1843) nec Blechnum pubescens Desv. (1827) n
Hooker (1837). Blechnum ivohibense C. Chr. var. nth C: Chr.
[in Perrier, Cat. Pl. Madag., Pterid.: 44 (1932) nomen nudum] in
Dansk Bot. Arkiv 7: 106 (1 1932 ).
MADAGASCAR: Antananarivo, Pool s.n. (K lectotype; sterile fronds
of L. Aarne a Forét de Analamaitso, Perrier de la Bathie 7625
(BM syntype of B. ivohibense var. hirsutum); Pic d’Iv ohibe (Bara),
Hames 3299 BM wiiype of B. ivohibense var. hirsutum).
Blechnum ivohibense C. Chr. in Arch. de Bot., Caen 2, Bull Mens.:
211 (1928). Blechnum bakeri C. Chr. var. glabrum R. "Bonap. [ Not.
Ptérid. 16: 72 a 925) nomen nudum] ex Tardieu in Humbert,
loc. saat 8, t.2, ff.1-3 (1960). Blechnum umbrosum A. Peter in
Note Rep. Sp. Nov., Beih. 40: 82, t.3, ff.5—8 (1929).
ESIA: INYANGA distr., Pungwe Gorge, Chase 6659 (BM, K,
srcib, ‘Sekibesaide 2411 (BM).
BIQUE: Namuli, Makua Country, Last s.n. ‘oak Serra do
Giri, Marreau elo, R. Namitué, Mendonca 2162 (BM
ANZ : W. Usambara, Peter 16489 (BM, K, ates of B
faa anelings: iin, Mtns., Lupanga-Berg, Schlieben 2959 (BM)
MADAGASCAR: Antananarivo, Pool s.n. (K) in part; Pic d’Ivohibe,
Humbert 3300 (BM type of B. ivohibense)
REFERENCES
BONAPARTE, R. 1925. Notes Ptérid. Fougéres de Madagascar, 16:11 -221.
CHRISTENSEN, C. ee The Pteridophyta of Madagascar. Dansk Bot. Arkiv, 7:1-253.
TARDIEU-BLOT, M In H. Humbert, Flore de Madagascar et des Comores,
Po toluadlandon 2: 1-133, Paris,
350 BRITISH FERN GAZETTE: VOLUME 9 PART 8 ((1967)
VARIATION IN ADIANTUM IN THE
BRITISH ISLES
S. L. WILLIAMS
664 Rochester Way, London, S.E.9
Perhaps this article is the saddest in this series reviewing variations in
British ferns. In the latter half of the nineteenth century, many
variants of A. capillus-veneris were in cultivation, but alas, so far as the
writer is aware none has survived to the present day.
It seems that during the nineteenth century, the British Maidenhair,
our only indigenous species of a very large genus which centres on
South America grew in many localities in western England, Wales,
the Isle of Man, Eire and the Channel Isles. The distribution was
maritime, growing in rock fissures and caverns often at sites actually
facing the sea, which would suggest a surprising degree of tolerance to
saline spray having regard to the thin texture of the pinnae. A fern
not reliably hardy, it is of interest that it survived at these localities
which experienced the mildest winter climate when the remainder of the
country was enduring the very cold winters which occurred during the
second half of the 1800s.
Fern hawkers and collectors have been blamed for the decline or
elimination of the species from former relative abundance and, judging
from the quantity of 4. cuneatum types which are commercially disposed
of today, A. capillus-veneris perhaps because of its pleasing colour and
delicate form, is still much in demand.
In the BSBI Atlas of the British Flora (PERRING & WALTERS, 1962)
twenty three 10-km grid squares are marked as possessing the species.
There are others, however. In the last few years it has been found at
Cheddar Gorge (Somerset), in two localities in Kent, one in Surrey (a
score or more plants on a brick wall supporting a railway), and at
Whitewebbs Park, Middlesex.
It can frequently be found growing in old mortar walls in green-
houses—especially the supporting walls and footings. The writer
has observed it thus growing in Perry’s old nursery at Enfield (Middle-
sex), Stuart Low’s nursery at Bush Hill Park, Enfield, both of which
now no longer exist, and the Chelsea Botanic Garden (London) where
by virtue of the consideration shown towards it by the Curator, it still
flourishes; likewise at Hockley (Essex) and ina derilict greenhouse at
Swanley (Kent) it has been found growing, but always in the brick-
ork
__ Given the right conditions, and these appear to contain calcium
either as natural limestone or mortar, there appears to be no shortage
of spores in the air from plants already in cultivation to prevent this
WILLIAMS: VARIATION IN ADIANTUM Ke
fern from establishing itself. In common with other plants, when
growing under adverse conditions the growth is stunted, but fertile
fronds are nevertheless produced, even though perhaps only an inch
or less long overall and comprising one to three pinnules. Under these
conditions the number of pinnules and length of the stipe is affected
to a much greater extent than the actual surface area of the lamina
compared with a plant growing under better conditions. The rhizome
firmly attaches itself to the supporting medium and creeps into any
available crevice in the brickwork or rock, thus affording it a degree
of protection from inclement weather.
In some of the Royal Horticultural Society Shows at Vincent
Square, Westminster, A. capillus-veneris has been used in display work
staged by municipal authorities and some nurserymen, though it is
feared sometimes without knowing what the plant was. The Alpine
Garden Society shows occasionally produce a specimen, but the
writer has only seen very poorly grown plants with fewer than six
fronds.
Although the author has examined carefully every new plant
brought to his notice he has never observed anything to suggest
deviation from the typical form. As one would expect from different
environments changes do occur both in size of frond and pinnules.
In cultivation the application of fertilisers does not appear to influence
the resulting growth to the same extent as temperature and humidity,
and these factors must be borne in mind when assessing a new “find”’,
As one might expect E. J. Lowe developing his “multiple parent-
age” theory, did more work endeavouring to raise new forms of this
species than any other person and was most eminently successful, as
demonstrated by the number (in excess of fifteen) of his cultivars which
received First Class Certificate awards from the Royal Horticultural
Society or Royal Botanical Society between 1868 and 1872.
>
Hayle. In British Ferns and where found (E. J. LOWE, 1891, p. 26) it is
352 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
described as a large plumose variety, pinnules one inch or more broad
and an inch long, deeply divided into ten or twelve narrow fringed
divisions, indusium and sori—absent (yet it was fertile!).
Early wild finds also included:
‘Incisum Footii’—from Co. Clare, an incised form which looked
slightly depauperate
“Magnificum’—Co. Clare, a foliose form
‘Padleyi’-—from Devon, pinnules concave
‘Ramulosum’—another pyc Mpc Eire which divided several times
at the top forming a t
‘Rotundatum’—from the Isle ‘of Man, having rounded basal pinnules
Forms raised in cultivation, usually from sowings of mixed spores
were:
“Admirable’, F.C.C. 1871 (Fic. 8*)—large pinnules
‘Angustatum’, F.C.C. 1872—dwarf fronds “literally covered with
young plants’”’
‘Autumnale’, R.H.S. Cert. of Merit, 1890—erect, golden tint
‘Clorinda’ (Fic. Siem eed cut pinnules
‘Daphnites’, F.C.C. 1871—proliferous crowded pinnules
‘Digitatum’ "fronds unsymmetrical
‘Fimbriatum’—pinnules distinctly fimbriate
‘Fissum’—pinnules cut into shreds
‘Grande’—ultimate pinnules large
Rasiniardt as F.C.C. 1872, Certificated R.H.S. 1886—dwarf and
imbrica
“‘Incisum’ icghiiei large and much incised
‘Kalon’, F.C.C. 1868 (Fic. 9)—very large pinnules, overlapping
“Lowae’ "slender pinnules
“Magnificum’, Certificated R.H.S. 1869—large and bold
‘Multiceps-dentatum’ F.C.C. 1872— a nching
“Nobile’—golden pee eae pinn
‘Optandum’, F.C.C. 2 (FI a cut pinnules widely separated
‘Sibyl’ (Fic. i) ineised Laas pinn nules
‘Spectabile’—very large and incised pinnules
‘Undulatum’, Certificated R.H.S. 1869—fronds undulate.
Although the writer has raised several hundreds of plants from
spores of the typical form, as previously mentioned, he has not had the
good fortune to secure a variant. It seems very strange that none of
these plants which were outstanding even in their day, found their
way into University and Botanic Gardens throughout the world.
*The Figures refer to Fern Growing by E. 3. Lowe, 1898
BRIT. FERN GAZ. 9 (8) 1967 353
THE BPS 1966 EXCURSION TO BOVEY TRACEY
33 Wi DYCE
“ Devonshire lanes ... The prevalence of the motor in all shapes and forms,
from the great chars- a-banc and omnibuses which sweep the sides, to the impetuous
cycles which tear up the centres, has revolutionised the roads. No longer do the
ferns rise in their Byrne attaining the stature and decked with the grace which the
.
climate of Devo n give; everywhere the billhook has been usy— Make the
way clear for de iisoacgs is ‘the cry, and beauty has to bow before the ‘ Juggernaut ”
raffic.
Canon H. Kingsmill Moore, 1923, B.F.G. Vol. 5, No. 2.
After a lapse of several years the Annual Excursion was again held in
the West County from 10-17 September 1966, centred at Bovey Tracey
in South Devon, but gone is the peace and quiet of the country lanes
which we were able, only a few years ago, to hunt leisurely without
disturbance. Now the holiday-makers are tearing through them in
their cars and coaches, and in places it has become quite dangerous to
dally over the hedgerow ferns. I wonder what our worthy Canon
would say if he could revisit the Devon lanes he walked in 1923!
maller number of members* than usual assembled on Saturday,
10th September at the Edgemoor Hotel, situated on high ground about
a mile from Bovey Tracey on the way to the moors. It is worthy of
record that this hotel is the finest the Society has stayed at during the
many years the Secretary has been organising the Excursions since the
Second World War. It is beautifully situated in well-wooded country
with roads leading from it in several directions, it is very comfortable,
well organised, and the food is plentiful and good. The proprietors,
Mr. and Mrs. G. S. Kendon, are courteous and helpful, the staff is
Cheerful and efficient, and we were given every facility to pursue what
must have appeared to the other guests eccentric activities. For anyone
wishing to visit this lovely part of Devon, we can recommend whole-
heartedly the Edgemoor Hotel, Bovey Tracey. For hunting ferns it
makes an excellent centre, for they are very plentiful in the vicinity,
and even in the Hotel grounds where we found twelve of the more
common species.
With recollections of a pre-war visit when magnificent Royal Ferns
literally choked the channel of the Dart in the ravine south of Dart-
meet, and young plants flourished on every mossy boulder, the Secretary
led the party on our first day to this area, but alas! thirty years have
*We were nine in all—Reginald Kaye and Mrs ioe Dr Frances M. Jarrett, Mrs
G. Neilson, H. J. Bruty, Max and Peter Temple, W. Wright and the Secretary.
In addition Mrs E. M. Borup was accommodated at the Prestbury Country How
Hotel, and Dr James Davidson and Mrs Davidson were staying with their daughter
who resides in the nei ur On the Wednesday G. Gilbert arrived for the
A.G.M. and stayed for the rest of the week.
354 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
wrought big changes. Now the river bed is clear of the ferns, although
they still continue to grow abundantly along the banks. We enjoyed
the scramble through the ravine although the going was very rough
at times. Ferns grew in abundance and most of the common woodland
ones were noted as well as some spleenworts on the bridges. Fourteen
species were recorded.
Through the courtesy of the Nature Conservancy we were given
permission to visit the Nature Reserves in the area, and supplied with
maps and information. We availed ourselves of this opportunity to
visit three Reserves, Wistman’s Wood near Two Bridges, Yarner
Wood about a mile from our Hotel, and Bovey Valley Woodlands near
Manaton. The Conservancy welcomes visits by specialists and
responsible bodies to their Nature Reserves to study the many facets
of their natural history, and invites reports and records which can
make useful contributions towards the experimental work being carried
on in these special areas. Our visits were hurried ones, but we have
passed on to the Conservancy our records and observations regarding
the ferns.
Wistman’s Wood was visited on Monday morning. It lies along
the eastern slope of the West Dart valley at a height of 1500/1600 feet
about a mile north of Two Bridges from which it is reached by a
footpath. The Wood, already regarded as ancient and unusual, early
in the 17th century, was established as a Forest Nature Reserve in
1961. At one time the trees were very stunted, heavily branched and
festooned with epiphytes, but growth is now more vigorous. Although
the trees are not high, really stunted specimens are not so numerous,
and the epiphytic growths are reduced. The principal tree is the
pedunculate oak. Moss-covered boulders make movement treacherous,
but although the rock crevices provide shelter for many ferns, only
six species were seen here and we noted Athyrium _filix-femina,
Blechnum spicant, Dryopteris borreri, D. dilatata and Polypodium
vulgare. Most interesting was the luxuriant growth of P. vulgare on
the trees, accompanied by D. dilatata very much at home in this unusual
habitat. Pteridium aquilinum was abundant on the open slopes above
the Wood.
After lunch we travelled further west to Merrivale, and climbed
Cox Tor and Staple Tor. Some years ago in the Gazette (Vol. 8 No. 4)
Dr T. D. V. Swinscow described a visit to these Tors and reported
finding Hymenophyllum wilsonii and Thelypteris phegopteris, the Beech
Fern. The latter together with the Oak Fern, Gymnocarpium dryopteris,
is very uncommon in the West Country, and during our many visits
since the War to this part of the country we have never been able to
include either of these ferns in our records. The opportunity to see
T. phegopteris therefore, could not be missed, and we were able to
DYCE: BPS EXCURSION 1966 355
find it on both of the Tors, as well as H. wilsonii growing in a most
unusual habitat, just as Dr Swinscow described it, in the peaty soil
close to the foot of the rocks. Many other ferns were seen in the
sheltered corners among the rocks and nine species were recorded.
On the way home we made a short stop in the small hamlet of Pons-
worthy tucked away in a hollow of the hills. The village walls were full
of ferns large and small, but nothing unusual was noted.
On Tuesday we spent most of the day at Yarner Wood Nature
Reserve, a very different place from Wistman’s Wood, much larger
with two streams running through it. The altitude ranges from 275 to
1025 feet with a corresponding range of plant habitats. We devoted
most of our time to the lower and wetter parts where ferns flourished
abundantly and we recorded twelve species. In spite of hard going
through bramble and bog, a scramble along the Yarner Stream was
made enjoyable by the magnificent growth of the ferns, and the
discovery of Osmunda regalis which was flourishing happily in ideal
conditions. Athyrium filix-femina, Dryopteris borreri, D. filix-mas
and D. dilatata were also there, including the hybrid D. filix-mas x
borreri. Throughout the Reserve the dominant fern seemed to be
Blechnum spicant growing to a size seldom seen, and the variety hunters
in the party quickly noted a tendency for the pinnules to be serrate and
divided. Another plant was very close to the homophyllous variety
anomalum Moore in which all the fronds are similar and mostly
fertile. Other ferns seen were Asplenium adiantum-nigrum, A.
trichomanes, Phyllitis scolopendrium and Polypodium vulgare. In
places Pteridium aquilinum was growing vigorously, and on the higher
ground we found Thelypteris limbosperma. A fern looked for through-
out the week but not seen, was D. carthusiana. We were sure we would
find it in Yarner Wood for the conditions were right, and we did see
some plants which showed possible crossing between D. dilatata and
D. carthusiana. It is very possible that further search will reveal the
latter to be present in the Reserve
A quick look at Becka Falls, a few miles further north, was made
before we finished hunting for the day, and this proved most disappoint-
ing. Apart from the fact that there are very few ferns there, this
‘* beauty-spot ”’ is very much over-rated.
Our third Nature Reserve—Bovey Valley Woodlands—was
visited on the Friday, and this again provided hunting in lush wet
conditions which suited the fern population. We expected to see the
Royal Fern here, but did not find it, nor did we find D. carthusiana, in
spite of seemingly ideal requirements for its successful establishment.
Only nine species were found, but some of these were in abundance,
particularly along the river banks, where Athyrium filix-femina,
Dryopteris borreri, D. filix-mas, D. dilatata and Blechnum spicant
flourished along with Pteridium aquilinum and also the hybrid D.
356 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
filix-mas x borreri. Other species noted were Asplenium adiantum-
nigrum, Polypodium vulgare and Thelypteris limbosperma. Theapproach
to the Reserve from Water is a difficult one down steep wet and over-
grown slopes, and this seemed typical of the whole terrain, so that it
was impossible in the time at our disposal to cover much of the ground.
Our records therefore may not give a true picture of the ferns to be
found there.
We emerged from the valley near Manaton, and hunting on the
higher ground between Water and Manaton discovered in addition to
the species already mentioned, Asplenium trichomanes, Phyllitis
scolopendrium and Polystichum setiferum. A quick visit to Manaton
Rocks produced very little apart from acres of bracken through which
we forced our way with difficulty. Our return journey was through
Lustleigh on the opposite side of the river, which entailed quite a
hazardous drive across the valley by a very narrow lane—at the very
moment when a concentration of Devon’s holidaymakers seemed intent
on making the same journey in the opposite direction! A brief stop
was made by the bridge crossing the Bovey. The map indicates a
clapper bridge here, but we did not see it, nor did we find a large
variety of ferns. One of the most enjoyable moments of the day was
the discovery on a farm wall near Lustleigh of a magnificent colony of
Ceterach officinarum.
The River Webburn joins the Dart at Holne Chase, and for its
last three miles flows through a narrow wooded valley. Two very
brief visits in the past by the Secretary, when Hymenophyllum tun-
brigense had been found in quantity, gave the promise that further
hunting would be fruitful, and on the Wednesday we found, with
great difficulty, parking space for the cars near the junction of the
rivers, and set off up the Webburn valley by a rough path. The low
ground was boulder-strewn, with rocky wooded slopes above, and
ferns abounded. Asplenium trichomanes was noted along with Phyllitis
scolopendrium and Polypodium vulgare. The large-growing ferns were
common, Athyrium filix-femina large plants in great abundance,
Dryopteris borreri, D. filix-mas, D. dilatata and Pteridium aquilinum,
also Blechnum spicant, Thelypteris limbosperma and some Polystichum
setiferum. On the river banks we admired some large plants of
Osmunda regalis, and further up where the valley narrowed to a shaded
ravine, we found H. tunbrigense clothing a high perpendicular rock
overhanging the water. As there are old records for H. wilsonii in the
district, we persisted in our search for this seemed to be a suitable
habitat, and not far from its relative we found it, although not in such
quantity. Again D. carthusiana eluded our search despite the presence
of plants showing evidence of crossing between this fern and D.
dilatata.
Sea cliffs are always an attraction, and so, on the Thursday we
DYCE: BPS EXCURSION 1966 357
travelled south to the Kingsbridge peninsula, with the area between
Bolt Head and Bolt Tail as our destination. Leaving the main Salcombe
road we paused at Malborough and had a quick hunt while our party
assembled. Several wall ferns, Asplenium adiantum-nigrum, A. ruta-
muraria, A. trichomanes, Phyllitis scolopendrium and Polypodium vulgare
were seen, but not in great numbers. Narrow lanes with Phyllitis
scolopendrium on their banks, led us to Bolberry, and down to the coast
between Bolberry Down and West Cliff. Pteridium aquilinum covered
large areas on top of the cliffs, but very few other ferns were found.
Some Athyrium filix-femina, Dryopteris filix-mas and D. dilatata were
seen, with colonies of Polypodiu um vulgare among the loose stones and
in the grass, but our main attraction to the place, Asplenium billotii,
was abundant in rock crevices, while lower down the cliffs we found
A. marinum growing vigorously.
On the Saturday morning after an active week, blessed with good
weather, a sine qua non for our activities, the Excursion ended and we
wended our separate ways home with our finds. Several ferns were
collected during the week, but very little of great interest apart from the
homophyllous Blechnum spicant, some crested Asplenium trichomanes,
a small Polystichum setiferum which promises to develop into a cristu-
late specimen, and P. setiferum ‘ Percristatum’ found on the journey
down near Tiverton.
It will be noted how seldom we found Polystichum setiferum, and
this in the West Country, the ‘‘ home ”’ of this species. Its epicentre
lies further east on the Devon/Dorset/Somerset borders and north to
the Brendon and Quantock Hills, and there it is the dominant fern,
although in places it can disappear altogether. This disappearance
quickly becomes widespread as we move westwards, so that in the
Dartmoor area it is quite infrequent and only occasionally did we find
it in numbers of more than a few plants. The vicinity of our Hotel
was the one place where it was common, and strangely enough its pres-
ence here is not recorded in the Atlas of the British Flora.
In the following list of ferns seen during the Excursion, figures
1—25 (bold=new record, *=confirmation of pre-1930 record, for the
Atlas of the British Flora) refer to localities:
Asplenium | adiantum- -nigrum — 3, 4, 5, 8, 10, 11, 13, 15, 16, 17, 18, 25
A. illo sd 20
A. mari
Pm rta-murari 75.6.8, 11, 12, 13, 15, 16, 17, 18, 22, 23, 25
Athyrium. divie-tocee edi J, 4, 6 ref 8, 9, 10, iL i314, 16, 17, 18, 20; 21, 22, 23,
Blechnum seine 3, 4, 7, 8, 9, 10, 11, 13, 14, 17, 18, 21, 22, 23, 24, 25
Ceterach officinarum — 5, /2, i
Dryopteris hace oid, my 4, 6,7 8, 9, 10, 11, 13, 14, 16, 17, 18, 22, 23, 24, 25
D. dilata ta— I, 3,4, 6,7, 8, 9 10, 11,13, 14, 16,17, 18, 20, 21, 22,23, 24, 25
ry Stearns 3 46 7.8. 16. 17. 13, 14, 16, "17, 18, 20, 22, 23, 25
Himensphylioay tunbrigense — 23*
358 BRITISH FERN GAZETTE: VOLUME 9 PART'8 (1967)
H. wilso as
Osmu a pB Mah bd 2d. 25.
Phyllitis scolopendrium on i, Oy 10, ble 10s 751s ees ed, a.
olypodium vulgare sensu stricto — is S, 1471022,
, 24, 25
P. Vulgare aggregate — 4, 5, 6, 7, 8, 9, 10, 13, i5, i6, 17, 18, 19, 20, 21
Polystichum setiferum — 8, 10, 73, 7. ey
Pteridium aquilinum — /, 3, per 6, 7,8, 9, 10; 11,13, 14, 16; 10; 2, 44, 23; Ot 2
Thelypteris limbosperma — 3*, 8, 9, 23, 25
T. phegopteris — 7, 2/
V.C. 3 South Devo Date :
(O.S. 1 in. shes ser. 7 sheets 175, 176 and 187) visited National
September Grid. Ref.
1 Becka a a ee st i = 3 20/ 7580
2 Bolberr 5 6939
3 Bovey Valiey National Nature Reserve 6 7680, 7681
4 Cheston, coe be § 6858
5 Chud dleigh Kn : 4 8477
6 Clapper Bride, River Bovey, n near Manaton 6 7582
T Cox ie ni 2 12 5276, 5376
8 Dart 11
9 rp mvalley between Dartmeet/Holne New 11 6772, 6773,
Bridge Gis se s ea a 6872 6970,
6971, 6972,
7170
10 Edgemoor ee Bovey Trace te re 10-16
11 flan New B 5! a on tie 1 7070, 7170
12 Holne Paik, ace road near 5 7370
13 orsham . es 6 7581
14 ix est Cleave : 7581, 7681
15 Malborou ie
16 Manat “a 6 7481, 7581
17 ey 5 6759
18 Ponsworth 2 7073
es =a ng te rm, near = sine : ae
i oint, near Bolberry ..
21 Stap ie Tor se :: 2 5375, 5475
22 Wate 6
23 Webburn V alle 4 7172, 7173
24 Wistman’s Wood Nature Reserve , 177
25 Yarner Wood Nature Reserve 3 7778, 7779,
7878, 7879
BOOK NOTE
tion or apres in 7 wu => book we tod 250 pesetas (32/-) from the author,
Cc 9, , Las Palmas de Gran Canaria, or from the
publishers (Cabiido Ty a ‘de Gran eaasiny.
J, A. CRABBE
BRIT. FERN GAZ. 9 (8) 1967 359
NOTES, NEWS AND ABSTRACTS
A SHELTER FOR SMALL FERNS IN POTS
I have achieved great success with two shelters like the one described,
and at present have the following Seip all healthy and flourishing in
them: Adiantum pedatum var. aleutic Shae adiantum-nigrum,
A.a.‘Variegata’, 4 * alternifolium, y “font num, A.f. var. halleri, A.
macedonicum, A. “ murbeckii, A. r site inanapin, A, septentrionale,
A. trichomanes, A ‘Cristata’, A, t. risa Moulei’, A. viride, A.y.
Imbricatum’ dN Multifidum’, Blechnum penna-marina, B. p. ‘Cristata’,
Ceterach officinarum, be ieee 4 eris alpina, C. dickieana, C. fragilis,
C.f. ‘Cristata’, C. montana, Woodsia ilvensis
The first thing you pes is a container. A strong wooden box
would do, provided it is no less than eight inches deep. One of mine
is a disused teak washing-up sink which came from the local hotel, the
other I constructed myself out of three-quarter-inch boards from an
engine packing-case obtained from a local garage. Having got your
container, bore a few drainage holes which are covered by pieces of
perforated zinc. The next things needed are four upright pieces of
wood, one at each corner of the container, the front ones standing
fifteen inches and the back ones twelve inches above the edge of the
container, thus giving fall for the roof. ee are fixed at the
front, back, and ends. The ends and back can be filled in with per-
forated zinc. The roof is composed of wood sints with a half-inch gap
between each one, thus Jetting rain through and also providing shade.
The container, the size of which is ruled by how many pots you wish
it to hold, is now ready for setting up. Put a layer of sand on the
bottom. The pots are now placed on the sand so that their rims are
about level with the edge of the container. It is essential that clay
iy plastic) pots are used. Pack the pots round and in between with
ome water-retentive material such as sphagnum moss or peat or
vertanliee. Thus, the packing keeps the walls of the pots damp and
the sand provides drainage. The potting compost I use is one part each
of the following: fibrous loam, leaf-mould, peat, fine scree or sand;
plus a handful or so of ned meal for every cubic foot. For Asplenium
trichomanes, A. ruta-muraria and Ceterach a handful or two of crushed
mortar is helpful but not essential.
FRED JACKSON
Knotts View, Stonethwaite, Cumberland
FERN CLASSES AT SOUTHPORT FLOWER SHOW
24-26 AUGUST 1966
In 1966 three new classes (13 and 16 for foreign hardy ferns, and 11 for
a smaller ground group) were introduced to encourage more exhibitors
and to widen the opportunities, but there were only one or two entries
for them. In conversation with members manning the Society’s advice
360 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
bureau, it was mentioned that eae e difficulties were an important
factor for rae! potential ex
M eld’s ground peg (Class eo was a memorable a
expert aerunamhct of many beautiful specimens and rare varieties i
top condition. Mr Hayhurst’s smaller Saat group (Class ee a
also a fine exhibit. In view of the particularly handsome prize money
e Bae. several old friends pout the plants exhibited,
but would like to see much more competition. Even thoug it may
difficult to compete with wonderful specimens of many years’ growth,
it often happens that perfectly grown smaller specimens can score over
larger plants not quite at their best.
R. KAYE & W. G. MACKENZIE, Judges
PRIZEWINNERS AND THEIR ENTRIES
CLASS 10 group of hardy British ain with accent on diversity of species and
Ist Messrs J. Brookfield & Son who continue to hold the Silver ic gy
Challenge Trophy presented by the British Pteridological
CLASS 11 A display of not more than 20 diverse hardy British ferns, in a yc
not at 8 feet by 5 feet:
Ist B. Hayhur:
2nd iuure Brookfield
CLASS 12 Six dissimilar hardy British ferns with accent on variatio
Ist Messrs Brookfield: Athyrium filix-femi na ‘Congestum Paul’, Dryopteris
filix-mas ‘Askewii’, Phyllitis st an a ‘Crispa’, ‘Polypodium
Mppcety “Hadwinii’, Poly. sti ichum aculeatum ‘Pulcherri mum Bevis’,
P. setiferum ‘Divisilo bum
2nd __-B. Hayhurst: Athyrium inthe ‘Percristatum’, lis sapigd
‘ Angustata Crispa Cristata . dilatata ‘Succisa’, Phyllitis
Seclbunesalh teks * Ramo-marginata ’, Polypodium vu nlgar re
*Pulcherrimum’, Polystichum siferum ‘Iveryanum
CLASS 13 Six foreign Site hardy in Great Brit
Ist B. Hayhurst: Adiantum pedatum var ee um, A. venustum, Blechnu.
‘abulare, Onoclea phere iy p dlaadin diversifolia, Acienichee
acrosticho ides
CLASS 14 Six dissimilar greenhouse fern
Ist Messrs Brookfield: Vaiaieiee cdsaluinan ‘Kensington Gem’, Cyrtomium
falcatum, Microlepia oe k Platycerium alcicorne, Polypodium aureum,
Pteris c gd “Wimset
equal B. Hay Adiantum great a Gem’, A. eatum
Ist Davallia go eg Nephrolepis ‘Rosea’, N. exaltata, Plasycerium
CLASS 15 Three delice normal species of hardy British ferns
Ist Messrs Brookfield: —— capillus-veneris, Lewin trichomanes,
chenpd tae Phegopte /
2nd oN. Robin mn: Coneberts fragilis, Gymnocarpium dryopteris, Poly-
are
CLASS 16 Three es ferns sic in Great Britain: .
Ist B. Hayhurst: Asplenium adiantum-nigrum ‘Himalaya’, Dryopteris
wallichiana, huceueie. struthiopteris : :
3rd rookfield: Adi antum venustum, Matteuccia struthiopteris,
Taockis sensibilis
FERNS AT SOUTHPORT FLOWER SHOW 1966 361
ua 17 Three distinct varieties of Phyllitis sha dee
ol eld: rispa’ ,; Ramo-cristata’ ule syed
ae “‘Drummondii’, ‘Ramo-cr slats “Ramo- sabithade’
3rd N. mabineeas! “‘Crispa’, ‘Marginata’, oe Cristata’
CLASS 18 Three distinct varieties of Polypodium vulgare
Ist Messrs setae eae ornubiense’, adwinii’, “Pulcherrimum’
2nd : arandiaiee Co moat § ‘Hadwinii’, ‘Pulcherrimum’
3rd N. Rebitsed: ‘Longicaudatum’, ‘Prestonii’, ‘Pulcherr rrimum’
CLASS 19 Three distinct varieties of Polystichum
Ist Messrs Broo : P. aculeatum “Gracilimum’, P.a. *Pulcherrimum
Bevis’, P. setiferum ‘Wakeleyan
2nd N. Robi son: P. setifer fn ‘Cristatum’ P.s. ‘Divisilobum Densum’,
P.s. ‘Divisilobum Iv
3rd B. Hayhurst: P. aculeatum “Pulcherrimum Bevis’, P. setiferum ‘Crist-
atum’, P.s. ‘Imbricatum Cris
CLASS 20 Three distinct varieties of age filix-femina
Ist essrs aon okfiel ‘Clarissima’, “Cigeaeutiat Paul’, ‘Plumosum
Cristatu
2nd -B. Hayhurst: ‘Clarissima Jones’, ‘Cruciatum Cristatum’, ‘Frizelliae
Capitatum
CLASS 21 Three merci) edged bal Dryopteris:
Ist B. Hayhur. i Em a0 Cristata’, D. dilatata “Succisa’,
ix: as ‘Depauperata yee
2nd Messrs Brookfie Id: D. filix- ee *hakonet , D.f. ‘Fluctuosa Cristata’,
D.f. ‘Linearis Cristata’
3rd N. Robinson: D. borreri ‘Ramosissima’, D. filix-mas ‘Depauperata
Cristata’, D.f. ‘Fluctuosa Cristata
CLASS 22 Three ll eel or Thelypteris:
(one entry, n
CLASS 23 Three distinct varieties or species of Asplenium:
Ist Messrs Brookfield: A. adiantum-nigrum, A. porae, A. ruta-muraria
CLASS 24 One British fern of any kind or variety:
Ist essrs Brookfield: Adiantum capillus-veneris
2nd_—sé iB. Hc haat: peat filix-femina ‘Victoriae’
CLASS 25
Ist
2nd
3rd
One a fern
hur: + ees cuneatum ‘Kensington Gem
Messrs Brookfield: Adiantum cuneatum agi gerator Gem’
. M. Howarth: Adiantum cuneatum
N. ROBINSON
RESYNTHESIZING ASPLENIUM ADULTERINUM
Dr. J. D. Lovis writes that he has at last succeeded in resynthesizing
i ‘
go
himself in a su iihebeient issue (1957, Vol. 8: 182).
Lovis states that he has finally put the record straight by in fact achiev-
ing this synthesis. He believes this to be the first time that a natural
fern species has been reconstructed in this way. Further details are
promised for inclusion in a future part of the Gazette
362 BRITISH FERN GAZETTE: VOLUME 9 PART 8 (1967)
ASPLENOPHYLLITIS MICRODON REFOUND
r P. J. Girard has written from Guernsey, sending us photographs of
this species (cf. Brit. Fern Gaz., 9:35. 1961), which he has recently
rediscovered in the an We hope to publish a paper on this discovery
in the next issue of the Gazette
ABSTRACTS
Further work on gereoy ty aspects in Asplenium is continuing at Leeds
University. LOvIS, MELZER & REICHSTEIN report (Bauhinia 3:87-101. 1966)
sage aden lepidum x A. trichomanes subsp. yar which
e A. x aprutianum; the holotype, collected by H. Kun ue
Reichstein (TR700) in Abruzzo, Italy, is in cultivatio n and a herbarium
specimen is in Previous to this (Bauhinia 2 231.237 1965) these
hree authors had described A. x adulteriniforme trichomanes subsp.
trichomanes * A. viride) collected by H. Malzer near Gutenstein, Nieder6ster-
reich; again the holotype is in cultivation and a speci
E. Ber. Schweiz. : Tore ats ee poherey and described
a further hybrid A. x valgannense (A. seelosii x trichomanes subsp
quadrivalens) from Vareae. Italy: the eneiaag is Sopa at Ziirich (ZT).
Contributions “i the study of Dryopteris assimilis have come from w.
GATZI (Ber. Schwe Bot. Pa 76:146-156. 1966) reporting on Swiss
populations, and fro A. & D. LOVE (Flora, J. Icelandic Bot. 4:5-9. 1966)
confirming the diploid i in Iceland. J. A. NANNFELDT (Bot. Not. we tet
aterial. Ny es Univ. Sci. Budapest Rolando
Eétvés (Biol) 8:275- 284. | 966: in pant with Russian summary) give
an a D. assimilis in Europe with a dot map; the record for Eire
1S doubtful (vida s nd is based on the length of the lower
basiscopic pinn C.- IDE . sorsSA (Hereditas 56:377-3
1966) report Sredicniter’ chromat tographic investigation into four members
of the Dryopteris “‘spinulosa’’ complex, in cluding D. assimilis. R. M. TRYON
M. BRITTON (Rho Sorn 68 :59-92. 1966) give a thorough appraisal of
the variation of the cytotypes of D. tpt diritti showing by
silhouette the range of frond shape and dissec D. M. BR
SOPER (Canad. J. Bot. 44:63-78. 1966) give the distribution of Dryopt
species in Ontario, aontait out that all the D. ‘“‘dilatata” is diploid, Lee
conspecific wit ith D. assimilis.
G. BENL is ert his studies on the fern flora of Macaronesia: Nova
Hedwigia 12:137-148. 1966 ees rpm Mitt. Bot. Staatssamm. Miinchen
e 33-45. 1966 (Cheilanthes and es hols puree de Botanica 1: 25- ie
967 Sen ee tunbrigense). ad KEL (Cuadernos de Bota
i: :29-34. 1967) give: ag fehl abioscuphy “or recent works on the poteay
of the Canary Islan
The generic cae Gymnocarpium was decided on a vote by the General
oo of I.A.P.T. to be illegitimate (rR. Ross in Taxon 15:287. 1966).
A. & D. L6vE (Univ. Peivcade eg (Biol.) 24:8. 1966) proposed a Rigo
novum, Carpogymnia, to Biscay
. KA R (Nucleus 9: eR 1966) give a chromosome
count ‘for Ophioglossum eon (O. vulgatum subsp. ambiguum) from Ice-
60 and suggest it is an alloploi derived from an occasional
hybrid etwas O. lusitanicum and O. vulgatu
A. C. JERMY
BRIT. FERN GAZ. 9 (8) 1967 363
REVIEWS
THE ILL soit TED ght OF ILLINOIS: FERNS by R. H. Mohlenbrock,
xv+191 p I5% arbondale and Dewardsville: Southern Illinois
Miata y a hace wee ppbeensg Feffer & Simons Inc, 1967. $8.00
The ferns and “allies” have always made a convenient unit for monographing
ona nce basis and the 81 species and 6 lesser taxa treated in this attractive
volume are no exception. It is interesting to note that during the research
se the oe for the volume 13 additional species were added to the
inois lis
is a most useful key to the 28 genera described (including all
British aan except H. ymenophyllum) based solely on vegetation characters
with thumbnail sketches in the margins to help the descriptive text. Each
species is illustrated with a neatly executed, accurate full-page line drawing.
ommon names, synonymy, geographic range and habitat given are valuable,
and the author’s field or other notes either elucidate or summarize the
present taxonomy of the species under discussion. € gives chromosome
—— for some species, but unfortunately does not always give the source
of his information. He describes a new variety, Lycopodium y teow age var.
tryonii, on p. 26.
This is an up-to-date account of a good cross-section of N. American
ferns and gai worth buying if one is at all interested in temperate ferns.
FERNS AND ct oped OF CALIFORNIA by Steve J. Grillos, illustrated
Rita Whitmore, 104 pp. 11x 18.4 cm: University of California Press,
Berkeley, 1966. $1.50.
This is another of the more recent State Floras to be pr roduced covering an
n in California. The commoner fifty-five species are illustrated, often as
thumbnail sketches which are very effective although not so s successful with
the larger species e.g. it hg and Polystichum. The keys are good and
easy to use. The descriptions, unfortunately, are not standardised and
important characters Ag as whet her the rhizome is erect or creepin
often omitted; others are included twice both in the generic description and
in that of the species. In ik telmateia var. braunii sterile stems are
given as “white or pale green”; in fact this variety is distinguished by the
stomata on the greenish sterile a and I have never seen in merican
plants the ivory white stems so characteristic of our oe E. Saprearrgira
I suspect Mr Grillos is compromising between his own experience and the
sees gro % earlier literature. For just over 10s. this little book i is ; cortiaty
worth buyi
A. C. JERMY
OBITUARIES
EDMUND FREDERIC WARBURG
Dr Warburg, an Honorary Member, died at the age of 58 on 9 June 1966;
he was born on 22 March 1908. He was a product of Cambridge and his
first cocaine away from there was at Bedford College, London, but
the war found him interpreting aerial photographs in the R.A.F. In 1945
364 BRITISH FERN GAZETTE: VOLUME 9 PART 8'(1967)
he was invited to join the staff of the Department of Botany at We sti: as
Curator of the Druce Herbarium, later as lecturer and in 1964, a Reader
Heff, as he was known to his friends and the botanical fraternity at
large, is ene: known for his part in Clapham, Tutin and Warburg, Flora of the
British a pe in particular, cog og’ other families for which he will =
bere
long remem e.g. Rosaceae, he was responsible for writing up t
Pterido ean His approach was aagularly one of practicability ; he disliked
era out of one, e.g. in recognising Gymnocarpium a
Thelypteris, when on the face of it they are so si ikewise with lati
n
names he was adamant in refusing to accept Alston’s revival of /anceolato-
cristata for such a well known plant as Dyopteris spinulosa on the grounds
that it was cumbersome and a change for the worse. He had seen so many
=
r
wisdom of this refusal is easy to understand. Wiens was the President
of the Botanical Society of the British Isles at the time of his death an
believed evantind in the part that such Societies as ours can do to foster
interest in botany
GARETH BEVAN EVANS
On | October 1966 The Times reported a Dr G. B. Evans had been found
dead in his room in the University of Malaya, Kuala Lumpur. He had
a lecturer in botany at that University wid "Januar ry 1965.
Gareth was born in Cadoxton, Glam. 3 September 1935, the only
son of Mr and Mrs D. G. Evans. After attending Neath G.S., he served his
National Service in the R.A.S.C. and went up to Trinity — Cambridge
in October 1956. On praduadhe in 1960 he had been accepted by the
University College of N. Wales, Bangor, to study under Profes ssor P. W.
Richards. Al i
in 1964, discussions at Bangor stimulated an interest in tropical botany.
In 1961 he went on a student expedition to Morabali Creek, British matopert
and again in 1963 he led another party to do statistical ecology in the
area. ooh the intervening year he had joined an Oxford expedition to yi
a s where he collected further data for a statistical analysis of the
tropical gone rest. His interest in filmy ferns was always to the fore and he
made epecudlat collections in Africa, B. G. and in Malaya where he was
working on their physiology. va meer eo his life as a teacher had only
just begun, but he had gained a t from his students and inspired
perhaps more than a few to (0k ‘ioe 5 eeeniy fi into a plant’s reaction to its
own environment.
A. C. JERMY
We have a very comprehensive collection of
BRITISH FERNS & THEIR VARIETIES
We also specialize in alpine and herbaceous plants, dwarf trees, flowering
shrubs, house plants, water lilies, and greenhouse plants
REGINALD KAYE LTD
SILVERDALE, LANCASHIRE
CATALOGUE ON REQUEST
THE AFRICAN SUCCULENT
PLANT SOCIETY
In just over one year, this Society, enjoying the patronage of 22 of the
most distinguished people in the world of succulent plant study, under
the Presidency of Gordon D. Rowley, has attracted members in 32
countries. Volume 2 of the BULLETIN commenced in April 1967, and
will be sent on receipt of the Annual Subscription of £1 ($3.50).
Junior members 15s. ($2.50). Seedlists, with free seed distributions in
Spring, and when interesting seed is available.
Full particulars from the Hon. Secretary:
C. A. E. PARR, 30 Wray Crescent, London, N4 (ref. BPS)
THE HARDY PLANT SOCIETY
provides a link between all wishing to know and/or grow
hardy herbaceous perennials
Membership can be most rewarding: subscription 12s 6d. per annum
Brochure from the Honorary Secretary:
Miss BARBARA WHITE
10 St. Barnabas Road, Emmer Green, Reading, Berkshire
THE
BRITISH FERN GAZETTE
VOL. 9 PART 8
Contents
LOVIS, J. D. Fern hybridists and fern hybridising. I. The
work of Edward Joseph Lowe (1825-1900) ;
CRABBE, J. A. Bolivian Pteridophytes collected by Winifred
Brooke
SLEEP, ANNE. A contribution to the cytotaxonomy of
Asplenium majoricum
HOLTTUM, R. E. John Smith of Kew
PAGE, C. N. Sporelings of Equisetum arvense in the wild. .
SLEDGE, W. A. Grammitis in Samoa
SCHELPE, E. A.C. L. E. The identity of two a
species of Blechnum
WILLIAMS, S. L. Variation in Adiantum in the British Isles
DYCE, J. W. The BPS 1966 excursion to Bovey Tracey
NOTES, NEWS AND ABSTRACTS:
JACKSON, F. A shelter for small ferns in pots
KAYE, R. et al. Southport Flower Show
Abstracts and correspondence to Editors
1967
301
REVIEWS AND BOOK NOTES 308, 320, 329, 334, 338, 358, 362
OBITUARIES
363
_ Printed by the Devonshire Press, Torquay, England
THE BRITISH FERN GAZETTE
Volume 9
Index
New taxa or combinations in bold; authors in SMALL CAPITALS
Adiantum, in British Isles 350
ALLEN, B. M. 249
Alston, A. H 7
Anglesey (records) 9
raclinlotdes see (Swartz) Proctor 213
rigidissima (Hook.) Proctor 213
Asplenium y wind 305
font
y essen 324
lovisii 158
ruta-muraria 141
subsp. eset Lovis & Reichstein 143, tab.
iadophilum Proctor 213
trichomanes
subsp. inexpectans Lovis 155
subsp. quadrivalens 152
subsp. trichomanes 15
trichomaniforme 158
viride 304,
x Asplenophyliitis 305
microdon 362
Athyrium, cultivars 197
Austria (excursion) 288
Austwick (meeting) 239
egg: re 57
Barnes
asus | a
ss (meeting) 186
BENO! 77
Birdbrook ‘enon 86
Blechnu antillanum Proctor 214
rit
png 349
ieri 0 —e Proctor 215
Bolivia, fern collect n, 309
W. Brooke’s fern fre a 309
Bolton, T. H. (meeting
Borrowdale (meeting) 57
Breconshire (records) 190
Brentwood (meeting) 57
British ferns (distribution card) 184
axonomy) 205
British Pteridological Society
Annual Excursions:
1959 Ambleside, Westmorland 28
1960 Williton, Somerset 54
1961 Prenteg, rte’ 87
1962 Aberfeldy, Perths. 129
1963 Builth Wells. ois. 186
Lancs. 239
196 Bovey Tracey, Devon 353
Constitutio
eeting nats: 1960 57, 1961 86, 1962 128, 1963 186, 1964 239*
Membership lists 32, 62, 96, 135,
Secretary and Treasurer’s — 1958 31, 1960 60, 1961 92*
Brockenhurst (meeting) 186, 2.
Brooke, Winifr [oe
BRUTY, H. J. 23,
Buchanan, Ww. 2 Tobit) 194
Caernarvonshire ( sre rds) 91
Cardiganshire (reco:
Census list (British peridophytes) 4
Merit 183
Channel Islands 34
ntbes catanensis (Cos.) H. P. Fuchs 45
fragrans 44
hispanica 47
prsehrsaa 42
Chelsea Physic Garden (meeting) 239
Cochran, J. (obit.) 95
162
CRABBE, J. A. 26, 51, 123, 178, 309
CRANE, F. W. 37
tenitis harrisii Proctor 216
parvula Proctor 217
cultivated ferns 178
cultivars, of Adina 353
of At,
of ee sagen” 97
cultivation, = -
"399
Cumberland econ) 29. 243
Cyathea hook
sinuata
Cystopteris alpina 85
dickieana 85
*Subsequently reported in B.P.S. NEWSLETTER.
Devon, S (excursion. records) 353
Diplazium prrseerny (Cav.) Jermy 161
aistribution on (rts ferns) 184
Dixon, J. D. (obit.) 59
mare M. J. 222
Dryopteris aia 137
mota 232
villarit 263
DYCE, J. W. 50, 97, 183, 184, 288, 353
East Grinstead (meeting) 86
Eillot, E. A. (obit.) 58
Equisetum arvense 5
fluviati eg bad palustre 117—
reques
Eridge Rocks, wae
Eriosorus chelates (Sw.) A. a ag ty 271
thewsii (Hook.) Cra
trofra Kunze ex ee ) Crabbe 315
EVANS, G. ask 81, 256, 364 (obit.
Mates Poe see British Pteridological Society
Fern hybridists 301
Ferndene, Kendal 297
Filmy ferns
Flora Europaea, addition to 163
new 249
review) 230
Forfarshire (records) 133
Fox, E. F. 303
FUCHS, H. P. 38
GILBERT, 4 251
LBERT, O. L. 263
Gleichenta 194
Gough Island 269
G itis aromatica (Maxon) Proctor 218
cance ye octor 76
brunneo-viridis (Baker ex Jenm.) Proctor 218
capillaris (Desv.) Proctor
eggersii (Baker) Proctor 77
ex (Maxon) Proctor 219
graminella (C Sledge 339
monticola Sledge 3 ok
sr ier (Maxon) eee 219
Maxon) Proctor 219
rigens tdakon) Proctor 219
suspensa (L.) Proctor 77
GREENFIELD, P. 25, 85
Harris (Outer Hebrides) 117
herbarium (of cultivars) 26
E. 2, 205, 330
Hungershall Rocks, Ken t 84
hybrids, a ion mes spores 37
phils
Tidhcsepentin! ber ety 84
igense 81, 256
won 81, 256
JACKSON, F. F. 49,
Jamai can ferns ae my) 213
coset A. C. 4, 81, 119, 137, 160, 163
5. de 114
INCHEERE, G
iene: Col. rs M. 303
KAYE, R. 120, 197, 295, 359
Kew Gardens (meeting) 86, 129, 186, 239
Lake District (meeting) 28, 191, 239
Lancashire, N. (records) 29, 243
LANG, F. A. 1
Lastrea dilatata var. alpina 80
chest wey
LLOYD,
Long ‘tate (esting 128
Lowe, E. J. 30
LOVIS, J. D. 16, a 141, 147, 163, 301, 361
Lyndhurst (meeting) 186, 239
Mapplebeck, J. E. 302
CCLINTOCK, D.
semen pega "aula 91
MELVILLE, J.
Miietend fie ware . rhe 316
cordata (Desv.) C
theae (C. Chr.) conte 316
Mill Green Woods, meetings 129
Moly, J. 302
MORGAN, D. 120
Montgomeryshire (records) 191
New Zealand (flora) 16
Obituaries:
Buchanan, W. C. 194
Cochran, J. 95
Dixon, J. D. 59
Elliot, E. A. 58
Evans, G. B. 364
ton, F. G. 195
Warburg, E. F. 363
Whiteside, R. 133
ee pee 194
PAGE, C. N.
Perthshire santas 133
Phyliitis tee goatee *‘Corale’, ‘Desirée’, ‘Keratoides’, ‘Khedive’, ‘Muricatum,’
, ‘Rosabelle’, ‘Spirale’, ‘Undulatum’, ‘Victoriae’ 305, 306
trea! i
Pilularia globulifera rt an 123
Pityrogramma ebenea (L.) Proctor 219
williamsii Proctor 220
Plant varieties and seeds Bill 179
Polypodium australe 65, 283
illare 161
californicum 168
vulgare, N. America 168
ulgare, Wales 65, 277
Polystichum pir
culeatum ‘Densum’ 302
aculeatum ‘Sul dieihinae? 50
ae ee 303
iferum en 302
riko F, G. “obit .) 195
PROCTOR,
poreling 228
ae (records) 190
Registration of cultivars 181
jurtetenk T. 141
Reviews: Ashberry, A. Bottle Gardens and Fern Cases 93
Bri an, F. H. The Oxford Book of Flowerless Plants 334
Leatherman & Behrends. w 300
Raidebock: R. H. Iilustrated Flora of illinois: > Fecka 363
Parsons, F. T. How to singe on Ferns 122
Pichi-Sermolli et al. Index Filicum, Suppl. 4 234
Tagawa, M. Coloured sation of the Japanese Pteridophyta 212
and Mosses
Taylor, P. G. Britis 5
Taylor, T. M.C. Ferns gine’ Fern-allies of British Columbia 54
Tutin et al. Flora Europaea 230
Tryon, R. The Ferns of Peru: Polypodiacae (Dennstaedtieae to Olean-
Wherry, E. T. The Fern Guide 122
ROBERTS, R. H. 283
Samoa 339
Sardinia, ferns
Savill Gardens Sauna 128
A. C. L. E. 348
Sn owdonia (excursion) 87
Somerset (meeting, records) 54, 128
Southport Flower Show 1960 ay 1962 124, 1963 191, 1964 236, 1965 295, 1966 359
Spain (Psilotum in) 249
Stigmatopteris jamaicensis (Desv.) Proctor 221
St Kitts 71
Tasmania (flora) 18
Taylor’s Nursery (meeting) 12
Thelypteris pavoniana oye Crabbe 318
Thursley (meeting) 57
tree-ferns 194
Tristan da Cunha 269
TRYON, A. F. 269
Tunbridge Wells (meeting) 57, 239
varieties, names of 180
vegetative characters of ferns 2
Victoria (flora) 18
bene oe 87, 186
,»M ~
atkine S
Warburg, = Fr, eam ) 363
Wardian case
babowatiomet fea 29, 243
w (meeting) 128
Whiteside, R. (obit.) 133
liso tle — i pie 335
Wisley (n (eting) 186
Wo
[pina
Xiphopteris atlosrides (Rosenst.) Crabbe 318
tica (Maxon) Crabbe 318
choquetangensis (Rosenst.) Crabbe 318
heteromorpha aig & Grev.) Crabbe 318
ar. dependens (Baker) Crabbe 318
a (Desy.) Hanes 319
liformis Swartz) Crabbe 319
myriophylla (Mett. ) Crabbe 3 319
peruviana (Desv.
yungensis (Rosenst.) Crabbe 319